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Geological Society, London, Special Publications

doi: 10.1144/GSL.SP.2004.228.01.19p419-453.

2004, v.228;Geological Society, London, Special Publications Jorge F. Genise ants and termitestrace fossils in palaeosols attributed to coleopterans, Ichnotaxonomy and ichnostratigraphy of chambered

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Ichnotaxonomy and ichnostratigraphy of chambered trace fossils in palaeosols attributed to coleopterans, ants and termites

J O R G E F. G E N I S E

CONICET, Museo Paleontolrgico Egidio Feruglio, Av. Fontana 140, 9100 (Trelew),

Chubut, Argentina (e-mail: jgenise@mef org.ar)

Abstract: Most recorded trace fossils in palaeosols are burrows and chambers attributed to bees, ants, termites and coleopterans. Ichnogenera attributed to bees are grouped in the ichnofamily Celliformidae, whereas those attributed to ants, termites and coleopterans are included herein in the new ichnofamilies Pallichnidae, Krausichnidae and Coprinisphaeridae respectively. Shape, type of wall, fillings and associated burrows of chambers are the main morphological ichnotaxobases used for this classification; they are weighed with regard to the behaviour and architecture of the supposed trace-makers. Coprinisphaeridae are spherical, pear-shaped or ovoid structures, having active or passive fillings and constructed walls. The ichnogenera included are: Fontanai, Coprinisphaera, Eatonichnus, Monesichnus, Teisseirei and Rebuffoichnus, attributed to coleopterans. The similar Pallichnidae show lined or structureless walls, and include Pallichnus, Fictovichnus and Scaphichnium, also attributed to coleopterans. The Krausichnidae constitutes trace fossils composed of chambers of different shapes interconnected by burrow systems of inconsistent diameter or isolated chambers associated with burrow systems of different diameters. The Krausichnidae include Attaichnus, Parowanichnus, Krausichnus, Archeoentomichnus, Tacuruichnus, Vondrichnus, Fleaglellius, Termitichnus and Syntermesichnus, attributed to ants and termites. The stratigraphic ranges of insect ichnotaxa in palaeosols are reviewed and compared with the body fossil record of potential trace- makers, revealing that in most cases insect trace and body fossils show similar ranges. As stated by earlier authors, the Cretaceous was a critical period during which the oldest body fossils of dung-beetles, bees, termites and ants are recorded, whereas the trace fossils of these groups are recorded from this period or shortly after, during Cenozoic times.

The grouping ofichnogenera in ichnofamilies is an uncommon but commendable trend in some branches of ichnology, where highly significant ichnotaxobases can be used to make coherent groupings of ichnotaxa (e.g. Genise 2000; Bertling et al. 2003; Rindsberg & Martin 2003). Roselli (1939), a pioneer in insect palaeoichnology, first suggested a higher taxonomy for the hymenopteran and coleopteran trace fossils that he described. In his contribution he grouped the supposed hymenopteran nests in the family 'Nidus Himenopterogenosidae' and those of coleopterans in 'Nidus coleopterogenosidae' (Roselli 1939). Recently, Genise (2000) created the first ichnofamily for insect trace fossils in palaeosols, Celliformidae, to include Celliforma and allied ichnogenera. This contribution represents an attempt to classify many of the remaining chambered trace fossils in palaeosols, attributed to Coleoptera, Hymenoptera and Isoptera (Genise 1999), and to provide a general picture of the stratigraphic ranges of insect ichnotaxa in palaeosols. It also represents an attempt to identify the major behavioural features of these groups as reflected in the morphology of their trace fossils.

Few trace fossils constructed by organisms other than insects can be compared morphologically with insect traces. The more complex the

architecture, the more difficult it is to find analogous structures outside the insect realm. Probably the most similar trace fossils are those produced by another group of arthropods: crustaceans. Tunnels associated with chambers are known from crayfishes, whose trace fossils are included in the ichnogenus Camborygma Hasiotis & Mitchell 1993. However, Cambor- ygma shows a distinctive bioglyph composed of scrape and scratch marks, knobby and hum- mocky surfaces, pleopod and body impressions, all of which distinguish it from similar ichnogenera attributed to social insect nests. In addition, interconnected tunnels of very different diameters are absent, as in other known modern crustacean constructions (Bromley 1990). The ichnogenera Ophiomorpha and Thalassinoides, also attributed to crustaceans, commonly show burrow systems devoid of chambers (Bromley 1990), but Verde & Martinez (2004) described chambers having tiny tunnels radiating vertically from the upper part of the chambers, in connec- tion with both ichnogenera. Spongeliomorpha shows burrow systems in association with chambers in one ichnospecies: S. sicula D'Alessandro & Bromley 1995. The presence of large vertical shafts, small chambers below the maze of tunnels and the typical criss-cross pattern of grooves in

From: MCILROY, D. (ed.) 2004. The Application of lchnology to Palaeoenvironmental and Stratigraphic Analysis. Geological Society, London, Special Publications, 228, 419-453.0305-8719/04/$15.00 �9 The Geological Society of London.



420 J .F. GENISE

Spongeliomorpha (D'Alessandro & Bromley 1995) and the upper radiating tunnels in the chambers associated with Ophiomorpha and Tha- lassinoides separate these trace fossils from those of social insects.

The morphology of insect ichnogenera devoid of tunnel systems has few similarities with other known ichnological structures. The specimen illustrated by H/intzschel (1975) of Amphorichnus papillatus Myannil 1966 superficially resembles Fictovichnus (Johnston et al. 1996); however, the former is a filling of an amphora-like hollow ending in a distinct apical protuberance (Pemberton et al. 1988; Edwards et al. 1998). Another case is that of the cocoons of earth- worms and leeches described by Manum et al. (1991) attributed to the genera Burejospermum, Dictyothylakos and Pilothylakos. The former two were previously considered as a seed and a palynomorph respectively. These cocoons are more likely secretions of organisms (Manum et al. 1991) than structures resulting from their activity, and as such they are ruled out as trace fossils (Bertling et al. 2003). Also, the structure of these cocoons differs from that of insects (Manum et al. 1991), whose cocoons are true trace fossils. The ichnofossil Lithoplaision ocalae Diblin et al. 1991 may superficially resemble an insect chamber, but its conical shape and marine invertebrate remains in the wall are important differences from insect chambers.

Continental ichnology, and particularly insect palaeoichnology, is an exciting topic that is developing quickly in a changing scene, in which new discoveries occur daily. Thus this contribution is written with the conviction that the classification and stratigraphy of trace fossils proposed herein will probably need to be updated in the near future. However, a first impression is presented herein, with the understanding that it will help to order the somewhat chaotic ichnotaxonomy of insect trace fossils, providing a new standpoint from which to observe and analyse insect behaviour as reflected by trace fossils.

Theoretical background

Even the most complex insect trace f6ssils in palaeosols can be morphologically divided into two components: burrows (tunnels, shafts and galleries) and chambers. The latter term has no specific definition in the ichnological literature and glossaries (e.g. Ekdale et al. 1984; Bromley 1990). However, it is commonly used to name distinct enlargements of burrows in the entomological literature (e.g. Stephen et al.

1969; Halffter & Edmonds 1982; Grass6 1984; H611dobler & Wilson 1990). These excavated chambers are used without further modifications for nesting or pupation, and in other cases they are used to house more complex constructions for nesting or pupation. The knowledge of insect nest architecture was developed mostly through entomological studies, with each group of insects having its own nomenclature for excavated chambers and constructed structures of different functions. Some terms used in the entomological literature are pupation cell (e.g. Scholtz 1988; Skelley 1991), brood cell (Stephen et al. 1969; Batra 1984), brood ball chamber (Halffter & Matthews 1966; Halffter & Edmonds 1982), fungus garden chamber (H611dobler & Wilson 1990) and royal cell (Grass+ 1984), among others. The functions of these chambers are diverse, but usually they are related to nesting activities or pupation - in sum, to the successful development of larvae. Fossil nests and pupation cells of Coleoptera, Hymenoptera and Isoptera respectively are the most common insect trace fossils in palaeosols (Genise et al. 2000), a fact that was related to the high potential of preservation of these constructed or lined structures (Genise & Bown 1994a).

Females of most solitary bees and wasps nesting in soils, as well as dung-beetles, prepare chambers or structures constructed inside them, in which they provision food (pollen, nectar, prey, carrion or vertebrate excrement), lay an egg, and close the entrance immediately after. A single larva feeds on these provisions and com- pletes its development without the assistance of adults in most cases (Evans 1963; Halffter & Matthews 1966; Batra 1984). In other groups of solitary insects having recorded trace fossils in palaeosols, such as weevils (e.g. Lea 1925; John- ston et al. 1996; Genise et al. 2002b), the larvae are not restricted to chambers prepared by adults. Their development takes place in the soil, in which they move freely, feeding on vege- table matter until their pupation, whereupon they prepare a cell that protects them during this critical period before emergence as adults (e.g. Loi~icono & Marvaldi 1994). In turn, social insects such as ants, termites and some bees provision and inhabit the underground nests in which they lay eggs, and rear larvae that are confined to the interior of chambers (Michener 1974; Grass6 1984; H611dobler & Wilson 1990).

These main behavioural differences, and a large number of more specific ones, gave rise to the great morphological diversity of insect nesting and pupation structures in soils and palaeosols. Thus the available ichnotaxobases by



ICHNOSTRATIGRAPHY OF TRACE FOSSILS IN SOILS 421

which to classify ichnogenera in ichnofamilies depend on the features of chambers and associated structures, especially shape, wall, fillings and burrows. Accordingly, the former ichnofamily proposed earlier for insect trace fossils in palaeosols, Celliformidae Genise 2000, was based on various characters of cells, the usual name that hymenopterists give to the brood chambers made by wasps and bees (Evans 1963; Stephen et al. 1969; Michener 1974).

lchnotaxobases

Four ichnotaxobases, the most common characters used as the basis of ichnotaxa, were listed and analysed by Bromley (1990): general form, type of wall structure, type of branching, and nature of the fill. Of these ichnotaxobases, branching is a less important character for classi- fying insect fossil nests, than the cross morphology of the entire burrow system is considered herein as an effective ichnotaxobase for insect traces.

Shape

Excavated and constructed chambers show a morphological continuum that ranges from flat and tabular shapes to spherical ones. Fortu- nately, discontinuities exist within this spectrum and also some complementary characters that favour the separation of the range as a whole into discrete units such as ichnogenera. The shape of the chambers is an important character for termite and ant nests, but it is even more critical for separating trace fossils attributed to bees and beetles, because in the latter cases the associated burrows are rare. Celliformidae (attributed to bees) can be recognized by the presence of cells having rounded backs and flat or conical tops showing a spiral design that was constructed from the outside by the adult bee (Fig. 1). In contrast, closed pupation cells attributed to beetles have both extremes rounded and smoothed because the larvae themselves construct them from the inside. Among trace fossils attributed to beetles, there is a clear morphological discontinuity between spherical or pear-shaped traces, namely Coprinisphaera Sauer 1955, Fontanai Roselli 1939 and Pallichnus Retallack 1984, and ovoid ones: Monesichnus Roselli 1987, Fictovich- nus Johnston et al. 1996, Teisseirei Roselli 1939, Rebuffoichnus Roselli 1987 and Eatonichnus Bown et al. 1997. Among the ovoid forms, it is impossible to distinguish different shapes, with the single exception of Teisseirei, which shows a

depressed outline in cross-section (Fig. 3c). Sca- phichnium Bown & Kraus 1983 is unique because it has a peculiar hamate or lunate shape.

Fossil termite and ant nests show a more or less continuous spectrum, from flat chambers in Krausichnus Genise & Bown 1994b, Fleagletlius Genise & Bown 1994b and Archeoentomichnus Hasiotis & Dubiel 1995a, to spherical chambers in Attctichnus Laza 1982, Termitichnus Bown 1982 and Vondriehnus Genise & Bown 1994b. In most cases, other complementary characters are needed to separate these ichnotaxa. However, some morphological discontinuities can be recognized in the spectrum of shapes. Krausich- nus and the unnamed termite nests from the Plio- cene and Pleistocene of Africa (Coaton 1981; Schuster et al. 2000) display low, flat, tabular chambers, whose floor and roof are parallel. Vertical pillars commonly accompany these chambers, probably to reinforce the whole structure (Fig. 5b). These tabular chambers are clearly distinguishable from other flattened, but more oval, high chambers (e.g. Fleaglellius) in which the roof is more arched with respect to the floor (Fig. 6b). Tabular chambers are apparently also present in Archeoentomichnus (Hasiotis & Dubiel 1995a) and in Termitichnus namibiensis Miller & Mason 2000. However, in the latter the tabular chambers seem to be more likely the result of a tiered arrangement of meniscate burrows, probably made by another organism.

The complex architecture of social insects may result in secondary chamber systems within a primary chamber system. This is true, for instance, in Krausichnus trompitus, where the tabular chambers are grouped in spindle-shaped structures that are, in turn, connected with other spindles (Genise & Bown 1994b). Similarly, in Tacuruichnus, a single chambered trace fossil supports a boxwork with chambers in the thick peripheral wall (Genise 1997) (Fig. 5c).

Types o f wall

This ichnotaxobase is the most difficult to analyse because of the different common usages of the term 'wall' (e.g. Retallack 2001b) and the complexity that walls can reach in insect traces. Commonly speaking, the term 'wall' is applied indistinctly to two different structures: two- dimensional surfaces (e.g. 'burrow boundary' of Bromley 1990, 1996) and discrete three-dimensional constructions. This fact introduces some confusion, and is discussed below. In an excavated chamber the wall is the boundary between the cavity and the soil - for instance the brood ball chamber wall in dung-beetle



422 J. F. GENISE

Fig. 1. Schematic drawings to demonstrate common features in nesting structures, showing different types of wall built by some dung-beetles, bees and termites. In all cases the insects first excavate a chamber in soil and then construct within it one or more brood balls (dung-beetles), a cell (bees) or a complete nest (termites), respectively. Constructed walls may show an inner lining (bees) or may bear a system of galleries and chambers with lined walls (termites). In dung-beetles and termites, constructed structures are separated from the excavated chamber by a space, whereas in bees the constructed wall is built against the chamber. Drawings are based on data from Coprini (Scarabaeidae) (Halffter & Edmonds 1982), Emphorini (Apidae) (Hazeldine 1997; Genise & Poir6 2000) and Nasutitermitinae (Termitidae) (Grass+ 1958; Grass6 1984).

nests (e.g. Halffter & E d m o n d s 1982) (Fig. 1). In these nests one or more b rood masses, each one having its own const ructed wall, may be located inside this excavated b rood ball chamber (Halffter & E d m o n d s 1982) (Fig. 1). In a

const ructed bee cell, the wall is c o m m o n l y a three-dimensional s tructure removable f rom the soil. This const ructed wall is in turn conta ined within an excavated chamber with its own two- dimensional wall (e.g. Hazeldine 1997) (Fig. 1).




Furthermore, in some termite nests the external wall of the central part of the nest is a construction that contains a boxwork that has lined walls separated by a space (paraecie) from an excavated cavity (e.g. Grass6 1958) (Fig. 1). In sum, each constructed wall in a soil produces at least three surfaces: the inner and outer surfaces of the constructed structure and the surface of the bearing cavity, each surface of which may in turn be considered to be a wall. Different types of wall may be present and related in various ways within a single nest and potentially the same trace fossil. Confusingly, in nests of different insect taxa the wall may bear different names, particularly in termite nests, in which the specific nomenclature is more developed (e.g. Grass6 1984).

The complexity of chamber walls in insect traces is related to the necessity of maintaining very specific environmental conditions inside nests to protect larvae and provisions. To achieve these conditions insects commonly line or construct walls by adding different types of organic matter to the soil material (Michener 1979; Grass6 1984; Cane 1991; Hanski & Cambefort 1991; Genise 1999). Genise & Bown (1994a) proposed that insect fossil nests are the most common trace fossils in palaeosols because of the original incorporation of organic matter in the wall structure, which enhances the probabil- ity of being consolidated by diagenetic processes.

Bromley (1990, 1996), when describing the most common taxobases for invertebrate ichnotaxonomy, recognized seven categories of wall structure: no structure (no lining), dust film, constructional walling (constructional lining), zoned fill, wall compaction, diagenetic haloes, and wall ornament. At least five of these are represented in insect architecture and have been recorded in different insect trace fossils in palaeosols. In two ichnogenera, both attributed to ants - namely Attaichnus Laza 1982 and Parowanichnus Bown et al. 1997 - the chamber wall shows no particular structure, a fact that probably reflects the absence of linings in the walls of ant nests (Bown et al. 1997).

However, most recorded insect trace fossils in palaeosols have lined or constructed walls. To line a wall is to cover a surface (e.g. excavated chamber wall) with a 'plaster' or coating (e.g. secretions, clay, faecal material), whereas to construct a wall is to add 'bricks and mortar' (e.g. soil pellets or coarse grains and fine material) to produce a discrete, three-dimensional structure (Fig. 1). A further complication in insect architecture is that insects may also actively line the inner surface of constructed walls with fine material (Noirot 1970; Hazeldine 1997). Bromley

(1990, 1996) considered those lined walls covered by 'dust films' to be collected passively in mucus- lined burrows. Insects actively produce similar linings with fine material by two main methods: the addition of transported material, or fluidization. Among those that transport material from elsewhere, many species of termite usually line the internal surfaces of constructed or excavated chambers with faecal and/or regurgitated material (e.g. Noirot 1970; Grass~ 1984). This lining is well preserved in the ichnofossil Krau- sichnus tromp#us Genise & Bown 1994b (Fig. 5b). Some halictine bees are known to line cells with clay material transported from the tunnel (Sakagami & Michener 1962). The clay lining of Rosellichnus arabicus is probably derived in this manner (Genise & Bown 1996). Bees commonly line cells with water-repellent lipids (Cane 1991) after smoothing the chamber wall (Batra 1984). This smoothing behaviour triggers a fluidization process in the soil adjacent to the chamber that results in a distinct lining of fine material sourced from the same surrounding soil (Genise & Poir6 2000). The bee's movements against the water-saturated soil pellets of the wall during its construction increase the pore pres- sure, which in turn produces the escape of water, drawing the fine material towards the inner surface of the wall (Genise & Poir6 2000). Fossil bee cells included in Celliforma Brown 1934, Uruguay Roselli 1939, Ellipsoideichnus Roselli 1987; Palmiraichnus Roselli 1987 and Cellicalichnus Genise 2000 show this type of lining. Similar layers of clay material are found in the walls of the coleopteran pupation chambers Pallichnus Retallack 1984 and Fictovichnus Johnston et al. 1996, who proposed that they were built by coleopteran larvae. Lined walls are easily recognizable because they show a clear discontinuity and a smooth internal surface, whereas externally they intergrade gradually with the host sediment (Retallack 1984; Johnston et al. 1996). Accordingly, trace fossils having lined walls are usually preserved as detached casts of smooth surfaces, or they are preserved in situ in rocks, from where they can be distinguished because of the different texture and colour of the lining.

Of all the categories of wall distinguished by Bromley (1990, 1996), the constructed ones are the most common in insect fossil nests. Usually, their producers first excavate a chamber and then build a wall through successive addition of soil pellets or sand grains (e.g. Halffter & Mat- thews 1966; Stuart 1969; Noirot 1970; Lee & Wood 1971; Hazeldine 1997). There are several important differences between lined and constructed walls that arise from the particular



424 J .F. GENISE

behaviours involved. Usually, linings are thin in comparison with constructed walls; however, intermediate thickness may produce some confusion that would preclude a clear distinction. Lin- ings are usually made from fine soil or plant material, secretions, or excretions that are applied as a coating on an excavated or constructed surface (e.g. Sakagami & Michener 1962; Stephen et al. 1969; Stuart 1969; Noirot 1970; Lee & Wood 1971; Cane 1991). In contrast, constructed walls are made from unsorted soil material, within which coarse grains or dry faecal pellets are added one at a time and bound by fine soil or faecal material (e.g. Halffter & Matthews 1966; Stuart 1969; Noirot 1970; Hazeldine 1997; Cosarinsky 2001). In lined walls the insect interacts with only one (inner) surface of the chambers, whereas in constructed walls both inner and outer surfaces result from the behaviour of the insect. Thus the outer as well as the inner surface may show a bioglyph, as in Eatonichnus (Fig. 2d). Constructed walls are discrete and resistant structures that can be removed entirely from soils and, when preserved, from the host rock. Accordingly, many trace fossils bearing them are preserved not only as casts or in situ in palaeosols, but also, and more frequently, as complete structures, removed from the rock matrix, showing internal and external bioglyphs. In addition, insect trace fossils having constructed walls can be transported and re-deposited (e.g. Andreis 1972, 1981) in contrast to most trace fossils, which are generally in situ (e.g. Ekdale et al. 1984). These walls are characteristic of Palmiraichnus Roselli 1987, Rosellichnus Genise & Bown 1996, Uruguay Roselli 1939, Coprinisphaera Sauer 1955, Fonta- nai Roselli 1939, Monesichnus Roselli 1987, Teis- seirei Roselli 1939, Rebuffoichnus Roselli 1987, Eatonichnus Bown et al. 1997, Termitichnus Bown 1982, Fleaglellius Genise & Bown 1994b, Vondrichnus Genise & Bown 1994b, Tacuruich- nus Genise 1997 and Archeoentomichnus Hasiotis & Dubiel 1995a.

Bromley (1990, 1996) mentioned diagenetic haloes as a particular category of wall lacking ichnotaxonomical value. Hasiotis et al. (1993) interpreted the external coating of Scaphichnium hamatum as diagenetic cement, but no particular wall was described for this ichnofossil (Bown & Kraus 1983; Hasiotis et al. 1993). The last category mentioned by Bromley (1990, 1996) - wall ornamentation - seems to be more probably a wall feature than a type of wall in itself. Orna- mentation may, potentially, be present in several different types of wall. Edwards et al. (1998) described unnamed trace fossils from the Bem- bridge Limestone Formation (England), which

are generally preserved as ovoid internal casts, whose complex surface sculpture can be interpreted as scratch marks (cast in positive relief) from depressions in the chamber wall. Similar casts preserving the original sculpture of the chamber wall are known in Teisseirei barattinia (Melchor et al. 2002, fig. 12 I). The constructed wall of both ichnospecies of Eatonichnus shows a helical pattern that is present on both wall surfaces, although more clearly in the outer surface. E. utahensis also exhibits a distinct superimposed bioglyph (Bown et al. 1997) (Fig. 2d). External bioglyphs are more pronounced when a space is present between the constructed wall and the excavated chamber. The external ornamentation of bee (Apidae, Emphorini) cells shows flattened pellets because the constructed wall is built against the cavity boundary (e.g. Hazeldine 1997; Genise & Poir+ 2000) (Fig. 1). In contrast, the wall of Apicotermitinae (Termitidae) nests - which are separated by a space from the bearing excavated chambers - show a very pronounced, complex sculpture (e.g. Grass~ 1984).

Fil l ings

Filling material and structure is another important taxobase in invertebrate ichnotaxonomy. Passive fill enters a burrow gravitationally, whereas active fillings are emplaced by the trace-maker (Bromley 1990, 1996). Both kinds of fill occur in insect trace fossils. As described previously, adult insects provision their nests with different kinds of organic matter to rear their larvae. Some dung-beetles are known to arrange the provision of their brood masses in a meniscate pattern (Halffter & Matthews 1966; Halffter & Edmonds 1982) (Fig. 3a). Such meniscate fills are seen in Coprinisphaera, Monesichnus, Eatonichnus and Scaphichnium (Bown & Kraus 1983; Hasiotis et al. 1993; Bown et al. 1997; Genise & Laza, 1998; Duringer et al. 2000a, 2000b) (Figs 2d, 3a). These authors attributed their material to dung-beetles based on the meniscate structure of the fills. The absence of an active fill in Rebuffoichnus, Pallichnus, Ficto- vichnus and Teisseirei is concomitantly taken to suggest that these ichnotaxa represent coleop- reran pupation cells (Retallack 1984; Johnston et al. 1996; Genise et al. 2002a; Melchor et al. 2002). Despite the fact that bees, ants and termites also provision their nests actively (e.g. Stephen et al. 1969; Grass6 1984; H611dobler & Wilson 1990), their trace fossils show no recognizable active fill (e.g. Laza 1982; Genise & Bown 1994b; Hasiotis & Dubiel 1995a; Bown et al. 1997; Genise 2000).




Burrow system

Burrows associated with fossil bee cells are uncommon (e.g. Cellicalichnus and Ellipsoideich- nus) (Genise 2000), and the same is true for coleopteran trace fossils. The coleopteran trace fossil Pallichnus is the single exception: it shows an associated burrow system, albeit only obser- vable microscopically (Retallack 1984). The poor preservation of burrows in fossil nests of solitary insects may be a result of the absence of constructed or lined burrow walls, in contrast to those of brood and pupation cells (Genise & Bown 1994a).

The burrows within nests of solitary insects have a constant diameter that corresponds to the body diameter of their constructor, as in other invertebrate trace fossils (Ekdale et al. 1984). The situation is different in ant and termite nests, whose burrow systems result from the cooperative work of many. Burrows are commonly very complex, and burrow diameter is variable. In many cases, larger burrows (first order) are connected with medium-sized ones (second order) and smaller ones (third order), the latter representing individual passages that match the size of the workers (Grass6 1984; Sands 1987; Genise & Bown 1994b) (Fig. 6c, d). Even when individual chambers are similar to those constructed by coleopterans, the associated burrow systems clearly distinguish the constructions of social from solitary insects. The burrow systems may:

�9 arise from the base of isolated chambers (e.g. Tacuruichnus);

�9 be present in the wall or in the interior of chambers (e.g. Tacuruichnus, Termitichnus qatranii); or

�9 connect different chambers (e.g. Krausichnus, Termitichnus, Vondrichnus, Fleaglellius, A rcheoentomichnus) (Genise & Bown 1994b; Hasio- tis & Dubiel 1995a; Genise 1997).

This ichnotaxobase distinguishes ant and termite nests from others and enables them to be classi- fied together in a suprageneric group.

Ichnofamilies of palaeosol trace fossils attributed to insects

Insect trace fossils in palaeosols can be grouped into four morphological groups or ichnofamilies based on the ichnotaxobases described above. The first ichnogeneric group comprises the ichnofamily Celliformidae, erected by Genise (2000), and includes the following ichnogenera, all of which are attributed to bees: Celliforma

Brown 1934; Uruguay Roselli 1939; Palmiraich- nus Roselli 1987; Ellipsoideichnus Roselli 1987; Rosellichnus Genise & Bown 1996; Corimbatich- nus Genise & Verde 2000; and Cellicalichnus Genise 2000. These ichnogenera can be recognized by the presence of cells having rounded bases and fiat tops, which commonly bear a spiral closure. Cells may be isolated or clustered, and may be connected by tunnels of similar diameters.

The second group comprises the new ichnofamily Coprinisphaeridae introduced herein, and includes unnamed trace fossils and spherical, pear-shaped or ovoid ichnogenera with active or passive fill and constructed walls. Ichnogenera included are: Fontanai Roselli 1939; Teisseirei Roselli 1939; Coprinisphaera Sauer 1955; Mone- sichnus Roselli 1987; Rebuffoichnus Roselli 1987; and Eatonichnus Bown et al. 1997. These ichnogenera are interpreted as the brood masses of dung-beetles or pupation chambers of various coleopteran taxa.

The third group, also representing trace fossils attributed to pupation chambers and brood masses of beetles, consists of ichnofossils lacking a constructed wall. Such ichnotaxa are included in the new ichnofamily Pallichnidae, and include: Scaphichnium Bown & Kraus 1983; Pallichnus Retallack 1984; and Fictovichnus Johnston et at. 1996. Pallichnus shows two features that distinguish it from somewhat similar coprinisphaerid ichnogenera. The Pallichnidae lack a discrete constructed wall. This fact suggested to Retal- lack (1984) that these trace fossils would probably represent pupation cells of Geotrupinae or Scarabaeinae rather than brood masses. Also, the chambers are connected to lateral tunnels that radiate from a vertical shaft, another character that is absent in Coprinisphaeridae. Fictovich- nus resembles Pallichnus in having a lined wall defined on its inner surface by a clay layer (Retal- lack 1984; Johnston et al. 1996). Fictovichnus has also been attributed to pupation chambers of beetles (Johnston et al. 1996). Scaphichnium lacks a lined or constructed wall, and its characteristic hamate or lunate shape with a bulbous termination is very different from both Pallichnus and Fictovichnus. Its inclusion in Pallichnidae is tentative. The meniscate filling of Scaphichnium probably reflects provisioning organic matter inside the excavated chambers for rearing larvae, as seen in modern examples of the Geo- trupinae (Hasiotis et al. 1993).

Some internal casts of Coprinisphaeridae may be indistinguishable from Pallichnidae. Such is the case for internal casts of Rebuffoichnus resembling Fictovichnus (Fig. 3e), as well as internal casts of Coprinisphaera, which may resemble



426 J. F. GENISE

Key to separate ichnofamilies of chambered trace fossils in palaeosols

1 Trace fossils composed of cells having rounded bases and flat tops that commonly bear a spiral closure Celliformidae Genise Trace fossils showing another combination of characters 2

2 Trace fossils composed of: (a) chambers of different shapes interconnected by a boxwork showing burrows of different diameters; or (b) isolated chambers bearing an internal boxwork; and/or (c) chambers from which a burrow system radiates. Intersecting grooves, scratch and scrape markings absent Krausichnidae ifam. nov. Trace fossils showing another combination of characters 3

3 Trace fossils composed of isolated or clustered, spherical, pear-shaped or ovoid chambers, surrounded by a discrete, constructed wall Coprinisphaeridae ifam. nov. Trace fossils composed of spherical, ovoid, hamate, or lunate chambers lacking a constructed wall Pallichnidae if am. nov.

Pallichnus. However, internal casts are commonly associated with complete specimens in outcrop, leaving no doubt as to the identity of the trace fossils (e.g. Genise et al. 2002a). Ovoid trace fossils included in the Coprini- sphaeridae and Pallichnidae may commonly show a broken end, or may be perforated or flattened following the emergence of the adult insect. In these cases, specimens may take a celliformid-like shape (e.g. Edwards et al. 1998). However, the flattened end lacks the spiral design as of well-preserved Celliformidae.

The fourth group, the new ichnofamily Krau- sichnidae, consists of trace fossils composed either of chambers of different shapes interconnected by burrow systems of different diameters, or of isolated chambers associated with burrow systems of different diameters. Ichnogenera include: Attaichnus Laza 1982; Termitichnus Bown 1982; Syntermesichnus Bown & Laza 1990; Krausichnus Genise & Bown 1994b; Flea- glellius Genise & Bown 1994b; Vondrichnus Genise & Bown 1994b; Archeoentomichnus Hasiotis & Dubiel 1995a; Parowanichnus Bown et al. 1997; and Tacuruichnus Genise 1997. These ichnogenera are attributed to the work of social insects, namely ants and termites. The presence of tunnels of different diameters in the same structure is the distinctive feature for recog- nition of traces constructed by the cooperative work of specialized types of individual (of differing sizes) within a society (Genise 1997).

A recently described but unnamed trace fossil from the Pleistocene of Chad (Duringer et al. 2000a, 2000b) deserves further comment. The trace fossil represents several specimens of Copri- nisphaera connected by a tunnel system. As such, it could be seen as a link between Coprinisphaer- idae and Krausichnidae that would almost preclude their separation. Such structures are unknown among dung-beetle traces, a fact that originally suggested to Duringer et al. (2000a, 2000b) that specimens of Coprinisphaera and

the interconnecting tunnels were constructed by different trace-makers (Duringer et al. 2000b). Later, the same authors recognized termites as possible constructors of these tunnel systems (Duringer et al. in press). The traces are thus a composite trace fossil (sensu Pickerill 1994) in which the termites exploit food reserves intended for larvae of Coleoptera.

Systematic ichnology

I c h n o f a m i l y C o p r i n i s p h a e r i d a e ifam. nov.

Type ichnogenus. Coprinisphaera Sauer 1955. Diagnosis. Trace fossils consist of spherical,

subspherical, pear-shaped, ovoid, or sub-ovoid chambers, generally isolated, rarely clustered. Chambers are surrounded by a discrete constructed wall, which may show a circular or ovoid hole. Some ichnogenera show empty or passively filled chambers, whereas in others active infill is the norm.

Ichnogenus Fontanai Roselli 1939 (Fig. 2c)

v*1939 Fontanai Roselli p. 79, figs 23, 24, 31 (8). p. 1975 'Nidos f6siles pOtreos de Cole6pteros"

Francis p. 553. 1976 Fontanaichnus Roselli p. 167 (junior

synonym). p. 1981 'Nidos f6siles de Cole6pteros'

Sprechmann, Bossi and Da Silva p. 266. 1982 Fontanaichnus Roselli; Martinez p. 58. v1987 Fontanaichnus Roselli; Roselli p. 34, pl. II,

fig. 6; pl. III, fig. 5. p. 1988 'Nidos de insectos f6siles (cole6pteros) '

Ford p. 47. 1990a Fontanaichnus Roselli; Retallack, p. 219,

figs 201 A,B. 1993 Fontanai Roselli; Genise p. 53. 1994 Fontanaichnus Roselli; Donovan p. 209, figs

8.5 A, B. 1994a Fontanai Roselli; Genise & Bown p. 112.




Key to ichnogenera within Coprinisphaeridae

1 Spherical, subspherical or pear-shaped chambers Ovoid or sub-ovoid chambers

2 Spherical chambers showing a raised rim surrounding the emergence hole Spherical, subspherical or pear-shaped chambers lacking a raised rim to the aperture

3 Ovoid structures, external wall showing a helical design Ovoid structures showing smooth or rugose outer wall

4 Chambers showing meniscate fill Empty or passively filled chambers

5 Chambers having a depressed outline in cross-section; inner surface may show scratch marks; some specimens show a short tunnel at the entrance or antechamber Chambers having rounded outline in cross-section

1996 Fontanai Roselli; Johnston, Eberth & Anderson p. 522.

1998 Fontanai Roselli; Genise & Laza p. 213. 1998 Fontanaichnus Roselli; Buatois, M~ingano,

Genise & Taylor p. 226. 2000 Fontanai Roselli; Genise, Mfingano,

Buatois, Laza & Verde p. 54. 2000 Fontanai Roselli; Krell p. 891. 2002 Fontanaichnus Roselli; Buatois, Mfi.ngano

& Acefiolaza p. 189. Type and only known ichnospecies. Fontanai

kragtievichi Roselli 1939. Diagnosis. Spherical chambers having a thick

constructed wall and an emergence hole surrounded by a raised rim or neck.

Remarks. This ichnogenus is pending the ichnotaxonomic revision along with the closely related Coprinisphaera (Laza, personal communication). Possible trace-makers are dung- beetles (Scarabaeinae). The presence of a neck is the only feature that allows the separation of Fontanai from Coprinisphaera. The neck may be interpreted as the remains of a separate egg chamber that some dung-beetles construct over the provision chamber in their brood masses (Halffter & Matthews 1966). However, if the neck is interpreted as the remains of a former egg chamber, it would be possible to trace a continuous morphological series between both ichnogenera.

Ichnogenus Coprinisphaera Sauer 1955 (Fig. 2a, b)

v1938a 'Bolas de escarabeidos' Frenguelli p. 348. v1938b 'Pallottole di Scarabeidi' Frenguelli p. 77,

fig. 5; pl. VII, figs 1-8. v1939 Devincenzia Roselli p. 81, figs 26, 27, 28

and 31 (5-6) (non Kraglievich 1932). v1939a 'Nidos f6siles de Escarabeidos" Frenguelli

p. 270. v1939b 'Nidos f6siles de Escarabeidos' Frenguelli

p. 379, figs 4-9, pls I-II. v 1940 'Nidos f6siles de Escarabeidos" Frenguelli

p. 70.

2 3

Fontanai Roselli

CoprO~isphaera Sauer Eatonichnus Bown et al.

4 Monesichnus Roselli

5

Teisseirei Roselli Rebuff oichnus Roselli

v1941 'Nidos de escarabeidos' Frenguelli p. 87. 1950 'Bolas de Cangagua' Bruet p. 280, pl. I, figs

2,3. * 1955 Coprinisphaera Sauer p. 123, figs 1-5. 1955 Cangabola Lengerken p. 937, figs 6-8. 1956 Coprinisphaera Sauer; Sauer p. 550, figs 1~,. 1959 'Nidos de Scarabaeidae' Halffter p. 174. 1959 Coprinisphaera Sauer; Sauer p. 119. 1962 Coprinisphaera Sauer; H~intzschel W189. 1966 'Fossil scarab balls' Halffter & Matthews

p. 154. 1966 'Nidos f6siles de escarabeidos' Camacho

p. 490, pl. XVI figs n, o. 1972 'Nidos de escarabajos' Andreis p. 91. p. 1975 'Nidos f6siles pOtreos de Cole6pteros'

Francis p. 553. 1975 Coprinisphaera Sauer; H~intzschel p. W52. 1976 Devincenzichnus Roselli p. 167 (junior

synonym). 1977 'Paleonidos de escarabeidos' Spalletti &

Mazzoni p. 267. 1981 'Nidos de escarabeidos' Pascual & Bondesio

p. 125. 1981 'Nidos de escarabajos' Andreis p. 34. p. 1981 'Nidos f6siles de Cole6pteros'

Sprechmann, Bossi & Da Silva p. 266. 1982 'Nidos de escarabeidos' Alonso, Gonzf.lez &

Pelayes p. 2. 1982 Coprinisphaera Sauer; Martinez p. 48. 1982 Devincenzichnus Roselli; Martinez p. 48. 1986a 'Icnof6siles de Scarabaeinae' Laza p. 13. 1986b 'Nidos de Scarabaeinae' Laza p. 19. 1987 'Heliocopris dung ball' Sands p. 423. v1987 Devizenzichnus Roselli, Roselli p. 21, pl. I,

fig. 1. *v1987 Martinezichnus Roselli p. 22, pl. I, figs 2,

2a (new synonym). *v1987 Madinaichnus Roselli p. 23, pl. I, fig. 3

(new synonym). *v1987 Microicoichnus Roselli p. 49, pl. I, fig. 8

(new synonym). p. 1988 'Nidos de cole6pteros' Ford p. 47. v 1990 'Nidos de escarabeidos' Laza & Reguero

p. 245.



428 J .F . GENISE

Fig. 2. Coprinisphaeridae. (a) Copr&isphaera ecuadoriensis Sauer 1955: thick constructed wall, empty chamber, and emergence hole. Eocene-Miocene Sarmiento Formation, Argentina; MACN-LI1802. Bar: 1 cm. (MACN- LI, Museo Argentino de Ciencias Naturales, Laboratorio de Icnologia.) (b) Holotypes of Devincenzichnus murguiai Roselli 1939 (left) (MFLR 479), Martinezichnusfrancisi Roselli 1987 (centre) (MFLR607), and Madinaichnus larranagai Roselli 1987 (right) (MFLR 641). Note the almost continuous range of ball and hole sizes. Late Cretaceous-Early Tertiary Asencio Formation, Uruguay. Bar: 1 cm. (MFLR: Museo Francisco Lucas Roselli, Nueva Palmira, Uruguay.) (c) Fontanai kraglievichi Roselli 1939: raised rim around the emergence hole. Late Cretaceous-early Tertiary Asencio Formation, Uruguay; MACN-LI 1786. Bar: 1 cm. (d) Eatonichnus utahensis (Guilliland and La Rocque, 1952). Holotype: external bioglyph and internal chamber with meniscate filling. Palaeocene Colter Formation, USA, OM20201. Bar: I cm. (Photograph courtesy of T. Bown.) (OM: Ohio State University Orton Geological Museum.)

1990a Coprinisphaera Sauer; Retallack p. 219, fig. 201 G-I .

1990a Devincenzichnus Roselli; Retallack p. 219, fig. 201 C, D.

1990b 'Dung beetle trace fossils' Retallack p. 436.

1991 Coprinisphaera Sauer; Retallack p. 182. 1993 Coprinisphaera Sauer; Genise p. 50.




1993 Devincenzichnus Roselli; Genise p. 50. 1994 Coprinisphaera Sauer; Donovan p. 209, fig.

8.5 G-I. 1994 Devincenzichnus Roselli; Donovan p. 209,

fig. 8.5C, D. 1994a Coprinisphaera Sauer; Genise & Bown,

p. 109, figs 3, 4. 1995 Coprinisphaera Sauer; Genise & Cladera

p. 78, figs 1E, Fig. IF, left and right. 1995 Martinezichnus Roselli; Genise & Cladera

p. 78, figs 1E, Fig. IF, centre. 1995 'Nidos de escarabajos' Fontaine; Ballesteros

& Powell p. 12. 1996 'Nidos de escarabajos' Iriondo & Kr6hling

p. 43. 1996 Devincenzichnus Roselli; Veroslavsky &

Martinez p. 41, pl. I, fig. 5. 1996 Devincenzichnus Roselli; Johnston, Eberth

& Anderson p. 522. 1996 Coprinisphaera Sauer; Johnston, Eberth &

Anderson p. 522. 1998 Devincenzichnus Roselli; Buatois,

M~ngano, Genise & Taylor p. 226. 1998 Martinezichnus Roselli; Buatois, M~ngano,

Genise & Taylor p. 226. 1998 Madinaichnus Roselli; Buatois, Mhngano,

Genise & Taylor p. 226. 1998 Microicoichnus Roselli; Buatois, M~ngano,

Genise & Taylor p. 226. 1998 Coprinisphaera Sauer; Buatois, Mhngano,

Genise & Taylor p. 227. 1998 Microicoichnus Roselli; Genise & Laza

p. 213. 1998 Madinaichnus RoseUi; Genise & Laza p. 213. 1998 Martinezichnus Roselli; Genise & Laza

p. 213. 1998 Devincenzichnus Roselli; Genise & Laza

p. 213. 1998 Coprinisphaera Sauer; Genise & Laza p. 220. 1999 Coprinisphaera Sauer; Genise p. 110. 1999 Martinezichnus Roselli; Genise p. 110. 1999 Madinaichnus Roselli; Genise p. 110. 1999 Microicoichnus Roselli; Genise p. 110. 2000 Coprinisphaera Sauer; Genise, M~ngano,

Buatois, Laza & Verde p. 54. 2000 Martinezichnus Roselli; Genise, MS.ngano,

Buatois, Laza & Verde p. 54. 2000 Madinaichnus Roselli; Genise, MS.ngano,

Buatois, Laza & Verde p. 54. 2000 Microicoichnus Roselli; Genise, Mhngano,

Buatois, Laza & Verde p. 54. 2000 Coprinisphaera Sauer; Krell p. 890. 2000 Microicoichnus Roselli; Krell p. 891. 2000 Madinaichnus Roselli; Krell p. 891. 2000 Martinezichnus Roselli; Krell p. 891. 2000a 'Fossil dung-beetle brood ball' Duringer,

Brunet, Cambefort, Likius, Mackaye, Schuster & Vignaud p. 277, figs 3-6.

2000b 'Boules de bousiers fossiles' Duringer, Brunet, Cambefort, Beauvilain, Mackaye, Vignaud & Schuster p. 259, figs 1-10.

2000 'Boules-nids fossiles de bousiers'Schuster, Duringer, Nel, Brunet, Vignaud & Mackaye p. 17.

2001a Coprinisphaera Sauer; Retallack p. 142. 2001 a Devincenzichnus Roselli; Retallack p. 142. 2001a Martinezichnus Roselli; Retallack p. 142. 2001 a Madinaichnus Roselli; Retallack p. 142. 2002 Martinezichnus Roselli; Buatois, M~ngano

& Acefiolaza p. 22. 2002 Microicoichnus Roselli; Buatois, M~mgano

& Acefiolaza p. 189. 2002 Madinaichnus Roselli; Buatois, M~ngano &

Acefiolaza p. 189. 2002 Devincenzichnus Roselli; Buatois, M~ngano

& Acefiolaza p. 189. 2002 Coprinisphaera Sauer. Buatois, M~ingano &

Acefiolaza p. 189. Type ichnospecies. Coprinisphaera ecuadoriensis

Sauer 1955. Diagnosis. Spherical, subspherical and pear-

shaped chambers having a constructed wall. Some specimens may show a hole (possibly an emergence hole). Internal cavities empty or containing a meniscate or passive fill.

Included ichnospecies. Coprinisphaera murgiai Roselli 1939; C. francisi Roselli 1987; C. larranagai Roselli 1987; C. lafurcadai Roselli 1987; C. ecuadoriensis Sauer 1955; C. frenguellii Genise & Bown 1994a.

Remarks. Coprinisphaera is one of the most common trace fossils in the Tertiary palaeosols of South America, and it was appropriately one of the first recorded insect trace fossils (Fren- guelli 1938a; Roselli 1939). It has subsequently been described from different localities and ages of South and Central America, Asia, Antarctica and Africa (Genise et al. 2000; Krell 2000 and references therein). The mention of 'nidos de escarabajos o escarabeidos" [nests of dung-beetles or scarabs] is common in the Argentinean sedimentological and palaeontological literature (e.g. Andreis 1972, 1981; Spalletti & Mazzoni 1977; Pascual & Bondesio 1981; Alonso et al. 1982; Laza 1986a, 1986b; Laza & Reguero 1990; Fontaine et al. 1995; Iriondo & Kr6hling 1996). In these cases, it is clear that the authors recognized the typical spherical or pear-shaped forms described by Frenguelli (1938b, 1939b), and as such these occurrences are attributed herein to Coprinisphaera ispp. The trace fossils described by Frenguelli (1938b, 1939b) are the same as those recorded by Halffter (1959), Cama- cho (1966) and Halffter & Matthews (1966). In turn, Uruguayan sedimentologists (e.g. Francis 1975; Sprechmann et al. 1981; Ford 1988)



430 J. F. GENISE

mentioned coleopteran nests described by Roselli (1939) which may be assigned indeterminately to Coprinisphaera or Fontanai.

Roselli (1987) created Martinezichnus, Mad# naichnus and Microicoichnus based on the different sizes of the chambers and the sizes of emergence holes, but these ichnotaxobases show significant overlap, precluding any clear- cut distinction between the different ichnogenera (Fig. 2b). They are formally considered herein as synonyms following Genise (1999). The probable trace-makers are dung-beetles (Scarabaeinae).

Ichnogenus Eatonichnus Bown et al. 1997 (Fig. 2d)

v* 1997 Eatonichnus Bown, Hasiotis, Genise, Maldonado & Browers p. 52, figs 8C-F, 9.

1998 Eatonichnus Bown, Hasiotis, Genise, Maldonado and Browers; Genise & Laza p. 214.

1999 Eatonichnus Bown, Hasiotis, Genise, Maldonado & Browers; Genise p. 111.

2000 Eatonichnus Bown, Hasiotis, Genise, Maldonado & Browers; Genise, Mfingano, Buatois, Laza & Verde p. 54.

2000 Eatonichnus Bown, Hasiotis, Genise, Maldonado & Browers; Krell p. 891.

v2001 Eatonichnus Bown, Hasiotis, Genise, Maldonado & Browers; Genise, Cladera & Tancoff p. 45.

Type ichnospecies. Eatonichnus utahensis (Gilliland & La Rocque 1952). Diagnosis. Trace fossils composed of closely

appressed whorls, tightly spiralled around a central cylindrical cavity and converging terminally, thereby forming a closed, spindle-shaped helix. Helices may be either sinistral or dextral, with whorls inclined away from the transverse axis. Whorl diameter constant along the helix and invariably greater than the diameter of the central cavity. Central cavity fill closely packed, with or without meniscae (Bown et al. 1997).

Included ichnospecies. E. utahensis (Gilliland & La Rocque 1952); E. claronensis Bown et al. 1997.

Remarks. The helical design of the outer wall is unique in Coprinisphaeridae, but its ovoid shape and meniscate fillings relate it to the unpatterned Monesichnus. The distinction between the two named ichnospecies of Eatonichnus is based on their very dissimilar sizes, whorl inclination, cavity fill and external ornamentation. A third, unnamed, ichnospecies has also been described based on its larger size and external bioglyph (Bown et al. 1997). Later, Genise et al. (2001) recorded Argentinean specimens of E. claronensis that show intermediate sizes, suggesting that size may be a misleading ichnotaxobase for this

group. Possible trace-makers are dung-beetles (Scarabaeinae) (Bown et al. 1997). The affinity with dung-beetle brood masses is suggested by its shape and meniscate fillings, similar to the ichnogenus Monesichnus. In addition, some modern dung-beetles are known to excavate helical tunnels (Bown et al. 1997). However, the producer of this trace fossil is unknown because there are no modern analogues for this structure.

Ichnogenus Monesichnus Roselli 1987 (Fig. 3a)

v* 1987 Monesichnus Roselli p. 39, pl. I, fig. 7. v1994 Monesichnus Roselli; Laza, Genise &

Bown p. 397. 1997 Monesichnus Roselli; Bown, Hasiotis,

Genise, Maldonado & Browers p. 52. 1998 Monesichnus Roselli; Buatois, Mfingano,

Genise & Taylor p. 226. v1998 Monesichnus Roselli; Genise and Laza

p. 213, figs 3-5. 1999 Monesichnus Roselli; Genise p. 110. 2000 Monesichnus Roselli; Genise, M~mgano,

Buatois, Laza & Verde p. 54. 2000 Monesichnus Roselli; Krell p. 891. 2001 Monesichnus Roselli; Retallack p. 142. 2002 Monesichnus Roselli; Buatois, Mfingano &

Acefiolaza p. 189. Type and only known ichnospecies. Monesichnus ameghinoi Roselli 1987.

Diagnosis. Discrete structures, fusiform to ovate, composed of a constructed, unpatterned wall (sometimes showing a longitudinal groove), and an internal cavity that in some cases is empty and in others exhibits a meniscate fill (Genise & Laza 1998).

Remarks. Monesichnus is morphologically similar to Eatonichnus, but its unpatterned wall clearly distinguishes it from the latter. Its meniscate fill, interpreted as the provisions of a dung-beetle brood mass (Fig. 3a), distinguishes it from other ichnogenera such as Teisseirei and Rebuffoichnus, which are interpreted as pupation chambers. Some broken specimens of Monesichnus may also be empty, their fills probably lost by weather- ing. Even in these cases, the rounded cross-section of the chamber distinguishes it from Teisseirei, and the absence of a rounded emergence hole precludes its assignment to Rebuffoichnus. These structures are similar to brood masses of the representatives of the modern dung-beetle genera Dichotomius, Grornphas and Oruscatus (Scarabaeinae) (Genise & Laza 1998).

Ichnogenus Teisseirei Roselli 1939 (Fig. 3b, c)

v* 1939 Teisseirei Roselli p. 82, figs. 29, 30, 31(7). 1976 Tesseirichnus Roselti p. 167 (junior

synonym). 1982 Tesseirichnus Roselli; Martlnez p. 61.




Fig. 3. Coprinisphaeridae. (a) Monesichnus ameghinoi Roselli 1987 (left): wall and meniscate filling. Late Cretaceous-Early Tertiary Asencio Formation, Uruguay; MACN-LI233. A sectioned sample of a brood mass of the dung-beetle Oruscatus davus (Scarabaeinae) (right) showing the same type of wall and fillings. Bar: 1 cm. (b) Teisseirei barattinia Roselli 1939: antechamber and bioglyph in the chamber wall; included in a piece of rock matrix. Late Cretaceous-Early Tertiary Asencio Formation, Uruguay; MACN-LI876. Bar: 1 cm. (e) T. barattinia Roselli 1939: cross-section showing the constructed wall and the depressed outline of the chamber. Eocene-Miocene Sarmiento Formation, Argentina; MPEF-IC 253. Bar: 1 cm. (MPEF-IC: Museo Paleontol6gico Egidio Feruglio-Icnologia). (d) Rebuffoichnus casamiquelai Roselli 1987. Late Cretaceous Laguna Palacios Formation; Argentina; MACN-LII202. Bar: 1 cm. (e) R. casamiquelai: internal cast with remains of the outer wall. Late Cretaceous Laguna Palacios Formation, Argentina; MACN-LI 1181. Bar: 1 cm.

v1987 Teisserichnus Roselli p. 24, pl. I, fig. 5 (lapsus).

v1987 lsociesichnus Roselli p. 38, pl. II, fig. 5 (new synonym).

1990a Teisseirichnus Roselli; Retal lack p. 219. 1993 Teisseirei Roselli; Genise p. 53.

1996 Teisseirei Roselli; Veroslavsky and Mart inez p. 41, pl. I, fig. 4.

1998 Teisseirei Roselli; Buatois, M~mgano, Genise & Taylor p. 226.

1998 Isociesichnus Roselli; Buatois, M~ingano, Genise & Taylor p. 226.



432 J.F. GENISE

1998 Teisseirei Roselli; Genise & Laza p. 213. 1998 Isociesichnus Roselli; Genise & Laza p. 213. 2000 Teisseirei Roselli; Genise, Mfingano,

Buatois, Laza & Verde p. 54. v2001 Teisseirei Roselli; Genise & Zelich p. 44. 2001 Teisseirei Roselli; Retallack p. 142. v2002 Teisseirei Roselli; Melchor, Genise &

Miquel p. 35, fig. 12 A-E, I. Type and only known ichnospecies. Teisseirei

barattinia Roselli 1939. Diagnosis. Depressed chambers slightly

arched downwards and having constructed walls. Some specimens may show a small, rounded antechamber. Inner surface of the wall displays, in well-preserved specimens, a distinct lining bearing small elliptical scratches oriented mostly longitudinally (modified from Melchor et al. 2002).

Remarks. The internal bioglyph (Fig. 3b) and the depressed cross-section (Fig. 3c) distinguish this ichnogenus from other Coprinisphaeridae. These trace fossils commonly show three preser- vational morphologies: (1) empty or passively filled chambers found in situ in palaeosols; (2) isolated, detached clasts composed of the chamber fillings; or (3) empty or passively filled chambers surrounded by a thick wall. Specimens found in situ in palaeosols of the Asencio Forma- tion suggested that these chambers were merely excavated structures (Genise 1999). However, the recent finding of hundreds of specimens bearing constructed walls in the Tertiary of Patagonia and re-examination of previous discoveries sug- gests that Teisseirei barattinia has a constructed wall. The holotype (and only documented specimen) of lsociesichnus diplocamara RoseUi 1987 is actually a structure composed of two attached specimens of Teisseirei barattinia. Thus it is regarded herein as a junior synonym of Teisseirei. Despite the excellent preservation of many specimens that display an internal bioglyph showing the scratches of the constructors, the origin of Teisseirei is unknown. Its shape and the absence of active fill indicate the lack of original provisions, and suggest that it is probably a coleopteran pupation chamber.

Ichnogenus Rebuffoichnus Roselli 1987 (Fig. 3d, e)

1925 'Calcareous insect puparia' Lea p. 35, pl. I, figs 1-20.

v* 1987 Rebuffoichnus Roselli p. 24, pl. I, fig. 4; pl. III, fig. 4.

p. 1996 Fictovichnus Johnston, Eberth and Anderson p. 516, figs 2, 3C-D.

1998 Rebuffoichnus Roselli; Genise and Laza p. 213.

1998 Rebuffoichnus Roselli; Buatois, M~ingano, Genise & Taylor p. 226.

v1999 Rebuffoichnus Roselli; Genise, Sciutto, Laza, Gonzfilez & Bellosi p. 29.

2000 Rebuffoichnus Roselli; Krell p. 892. 2000 Rebuffoichnus Rose|li; Genise, M~ngano,

Buatois, Laza & Verde p. 54. 2001 a Rebuffoichnus Roselli; Retallack p. 142. v2002 Rebuffoichnus Roselli; Genise, Sciutto,

Laza, Gonzfilez & Bellosi p. 230, figs 5D, 7. 2002 Rebuffoichnus Roselli; Buatois, Mfingano &

Acefiolaza p. 189. v2002 Rebuffoichnus Roselli: Genise, Laza,

Fernandez & Frogoni p. 160. Type and only known ichnospecies.

Rebuffoichnus casamiquelai Roselli 1987. Diagnosis. Sub-ovoid to subcylindrical struc-

tures composed of a wall, whose exterior aspect is rugose and lumpy, whereas the interior is smooth or showing a faint bioglyph. The internal chamber is ovoid and has a circular cross-section. The wall may be perforated by a rounded hole (Genise et al. 2002b).

Remarks. Rebuffoichnus differs from other more or less ovoid insect trace fossils, such as Monesichnus, in lacking any active fill and in the presence of a rounded hole. Trace fossils from the Quaternary of Australia described by Lea (1925) and Read (1974) were assigned to an unnamed ichnospecies of Fictovichnus by Johnston et al. (1996). However, the type ichnospecies of Fictovichnus lacks a constructed wall as seen in the material of Lea (1925) and Read (1974). These specimens have a general aspect and similar features to those of Rebuffoichnus casamiquelai (Genise et al. 2002b). R. casamiquelai is attributed to pupation chambers of Curculionidae, Scarabaeidae, or Tenebrionidae (Johnston et al. 1996). However, species of the Curculionidae are the most likely constructors because of the circular body outline, which would accord with the circular holes in the trace fossils (Genise et al. 2002b). This assump- tion is also confirmed by the discovery of the constructor within a Rebuffoichnus from Australia (Lea 1925).

Ichnofamily Pal l ichnidae if am. nov.

Type ichnogenus. Pallichnus Retallack 1984. Diagnosis. Group of ichnogenera comprising

subspherical, ovoid, hamate or lunate chambers lacking a constructed wall. Chambers are surrounded by linings or diagenetic haloes. Fillings may be passive or meniscate.

Remarks. Scaphichnium, the first named ichnogenus of the ichnofamily (Bown & Kraus 1983), shows peculiar characters that make its inclusion in the Pallichnidae tentative. Pallichnus Retallack




Key to identify the ichnogenera of Pallichnidae

1 Hamate to lunate with bulbous termination, unlined walls, meniscate filling Scaphichnium Bown and Kraus Spherical or ovoid shape, lined walls, empty chambers 2

2 Spherical shape Pallichnus Retallack Ovoid shape Fictovichnus Johnston et al.

1984 is herein designated as the type ichnogenus. The presence of associated tunnels as found in Pallichnus is an unusual trait for this ichnofamily and is therefore not considered as diagnostic. Apart from the known ichnogenera, many unnamed ovoid-shaped trace fossils that are devoid of constructed walls may be included in this ichnofamily, including cocoons (e.g. Bown et al. 1997), pseudoeggs (e.g. Hirsch 1994a), misidentified eggs (e.g. Johnston et al. 1996) and pupation chambers (e.g. Edwards et al. 1998).

Ichnogenus Scaphichnium Bown & Kraus 1983 (Fig. 4a)

* 1983 Scaphichnium Bown & Kraus p. 106, figs 4C, 5E-G, 6C, E, 9B, D.

1993 Scaphichnium Bown & Kraus; Hasiotis, Asian & Bown p. 2, figs 2, 4.

1998 Scaphichnium Bown & Kraus; Genise & Laza p. 214.

1998 Scaphichnium Bown & Kraus; Buatois, M~ngano, Genise & Taylor p. 227.

1999 Seaphichnium Bown & Kraus; Genise p. 110. 2000 Scaphichnium Bown & Kraus; Genise,

M~ngano, Buatois, Laza & Verde p. 54. 2000 Scaphichnium Bown & Kraus; Krell p. 892. Type and only known ichnospecies.

Scaphichnium hamatum Bown & Kraus 1983. Diagnosis. Discrete, hook-shaped to lunate,

meniscate endostratal burrow fills, oriented with long axis vertical to concave upward and with rounded, bulbous, lower (distal) termina- tions (Bown & Kraus 1983).

Remarks. Scaphichnium hamatum is an unusual insect trace fossil in many aspects. Its particular shape, meniscate fill and lack of a lined or constructed wall confer to this structure a very distinctive architecture unknown in other trace fossils. Consequently, its inclusion in Pallichni- dae is tentative. These trace fossils are attributed to brood masses of Scarabaeinae beetles, possibly Geotrupinae (Hasiotis et al. 1993).

Ichnogenus Fictovichnus Johnston et al. 1996 (Fig. 4c)

?1982 'Ovoid vesicles' Freytet & Plaziat p. 65, pl. 49, figs G, H.

?1987 'Cocoons' Ritchie p. 435, pl. 11.14, figs 9, 10.

?1994 'Cocoons' Thackray p. 796.

? 1994 'Egglike concretions' '? lizard eggs' Mikhailov, Sabath & Kurzanov p. 106, figs 7.16D-F, 7.20.

?1994a 'Pseudoeggs' Hirsch p. 281, fig. 11.3A. ?1994b 'Pseudoeggs' Hirsch p. 145, fig. 10-6F. ?1996 Celliforma sp. Veroslavsky & Martinez

p. 41, pl. I, fig. 2. v1996 'N6dulos ovoidales" Sciutto & Martinez

p. 74. * 1996 Fictovichnus Johnston, Eberth & Anderson

p. 521, figs 1, 3A-B and 4. v1997 'Wasp traces' Bown, Hasiotis, Genise,

Maldonado & Browers p. 48, figs 6A, C-E, 8A-B.

v1998 'Cocoon-like trace fossils' Edwards, Jarzembowski, Pain & Daley p. 25, figs 3-6.

1999 Fictoviehnus Johnston, Eberth & Anderson; Genise, Sciutto, Laza, Gonz/dez & Bellosi p. 29.

2000 Fictovichnus Johnston, Eberth & Anderson; Genise, M/mgano, Buatois, Laza & Verde p. 54.

2002 Fictovichnus Johnston, Eberth & Anderson; Genise, Sciutto, Laza, Gonz~.lez & Bellosi p. 230.

v2002 'Cocoons' Melchor, Genise & Miquel p. 24, fig. 12G-H.

Type ichnospecies. Fictovichnus gobiensis Johnston et al. 1996. Diagnosis. Ellipsoid chambers enveloped by

thin clay-rich zone, the inner surface of which is clearly defined and smooth. The outer surface of the clay-rich zone is gradational with the surrounding matrix. The long axis of the chamber is parallel to bedding, with a terminal exit hole that may be subterminal or medial (on the upper surface of the chamber relative to bedding). Where the ichnogenus occurs in calcretized soils, the chamber is commonly surrounded by variably developed calcite halo. Trace fossils occur as hollow structures with a wall formed of a clay-rich zone and calcite halo, or as egg- shaped internal moulds, depending on local calcretization and induration of host sediment (Johnston et al. 1996). Included ichnospecies. F. gobiensis Johnston et al. 1996 (=parvus Johnston et al. 1996 syn. nov.).

Remarks. Both named ichnospecies of Ficto- vichnus - F. gobiensis and F. parvus - were diag- nosed on the basis of differing size, probably reflecting the fact that they were made by different species of insect. However, the potential



434 J. F. GENISE

Fig. 4. Schematic drawings of Pallichnidae and Krausichnidae described by Bown & Kraus (1983), Retallack (1984), Johnston et al. (1996), Bown et al. (1997), and Hasiotis & Dubiel (1995a). (a) Scaphichnium hamatum Bown & Kraus 1983: Lower Eocene Willwood Formation, USA. (b) Pallichnus dakotensis Retallack 1984: Oligocene Brule Formation, USA. (c) Fictovichnus gobiensis Johnston et al. 1996: Late Cretaceous Djadokhta Formation, Mongolia. (d) Parowanichnusformicoides Bown et al. 1997: Palaeocene-Eocene Claron Formation, USA. (e) Idealized reconstruction of Archeoentomichnus metapolypholeos Hasiotis & Dubiel 1995a: Late Triassic Chinle Formation, USA.

existence of trace fossils of intermediate size would confuse the ichnospecific taxonomy: thus the two ichnospecies are considered herein to be synonymous. A third, unnamed ichnospecies from the Quaternary of Australia, included in Fictovichnus by Johnston et al. (1996), is trans- ferred to Rebuf fo ichnus owing to the presence of a constructed wall in the Australian material (Genise et al. 2002a).

Many ovoid casts were described or mentioned as 'cocoons', 'ovoid structures' and/or illustrated

showing an ovoid shape (e.g. Freytet & Plaziat 1982; Ritchie 1987; Thackray 1994; Veroslavsky & Martinez 1996; Sciutto & Martinez 1996; Bown et al. 1997; Melchor et al. 2002). They are attributed herein tentatively to Fictovichnus ispp. In addition, Johnston et al. (1996 and references therein) mentioned a long list of doubtful vertebrate fossil eggs or pseudoeggs that are also probably attributable to Fictovichnus (e.g. Hirsch 1994a, 1994b; Mikhailov et al. 1994). However, most of these cocoons or pseudoeggs




lack any evidence of the emergence hole, an important diagnostic character to definitively associate these dubiofossils with this ichnogenus.

In contrast, the cocoon-like trace fossils described by Edwards et al. (1998) show evidence of an emergence hole, a complex bioglyph and, in some cases, an associated burrow, features important enough to suggest a new ichnospecies of Fictovichnus. This ichnogenus is attributed to pupation chambers of Coleoptera, probably Curculionidae, Tenebrionidae or Scarabaeidae (Johnston et al. 1996).

Ichnogenus Pallichnus Retallack 1984 (Fig. 4b)

* 1984 Pallichnus Retatlack p. 580, figs 7, 9, 10. 1990b Pallichnus Retallack; Retallack p. 221,

figs 204, 206. 1993 Pallichnus Retallack; Genise p. 50. 1994b Pallichnus Retallack; Genise & Bown

109. 1996 Pallichnus Retallack; Johnston, Eberth &

Anderson p. 522. 1998 Pallichnus Retallack; Buatois, Mfingano,

Genise & Taylor p. 227. 1999 Pallichnus Retallack; Genise p. 110. 2000 Pallichnus Retallack; Retallack, Bestland &

Fremd p. 178. 2000 Pallichnus Retallack; Genise, Mfingano,

Buatois, Laza & Verde p. 54. 2000 Pallichnus Retallack; Krell p. 891. 2000 Pallichnus Retallack; Genise & Poir6 p. 7. 2002 Pallichnus Retallack; Melchor, Genise &

Miquel p. 29. Type and only known ichnospecies. Pallichnus

dakotensis Retallack 1984. Diagnosis. Nearly spherical chambers, defined

by thin wall of dark clay and organic matter; inner boundary of wall sharp and smooth, such that the internal cast is easily separated from rock matrix. The outer boundary of the wall grades outward into the surrounding matrix. One side of the chamber is disrupted to form large, irregularly circular, exit cavity, usually about half the diameter of the main chamber. Each nearly spherical chamber is arranged at end of short branches from the vertical burrow, so that the exit cavity faces into the branch burrow; both vertical and branch burrows of slightly lesser diameter than the nearly spherical chamber (Retallack 1984).

Remarks. The presence of a lined wall clearly distinguishes this trace fossil from those included in Coprinisphaeridae. The preservation of tunnels is unusual among those ichnogenera attributed to the work of solitary insects, grouped in Celliformidae, Coprinisphaeridae and Pallichnidae. Accordingly, tunnel remains in Pallichnus are detected only in thin sections

(Retallack 1984). These trace fossils are interpreted as pupation cells of scarabaeid beetles, particularly Geotrupinae and Scarabaeinae (Retallack 1984).

Ichnofamily Kraus ichnidae ifam. nov.

Type ichnogenus. Krausichnus Genise & Bown 1994b.

Diagnosis. Group of ichnogenera showing chambers associated with burrow systems, composed in most cases of burrows of very different diameters. Burrows are devoid of scratch marks and/or intersecting grooves. Chambers have no radiating tunnels from their upper parts and are commonly linked to a burrow system that nor- mally interconnects them with other chambers. Chambers may be empty, passively or actively filled, and/or they may contain secondary systems of burrows of different diameters and smaller chambers on or within their inner walls.

Remarks. The first described and most representative ichnogenera of this ichnofamily, A ttaichnus Laza 1982 and Termitichnus Bown 1982, bear names relating to ants and termites respectively. However, names such as Termitich- nidae or Attaichnidae should be avoided because Krausichnidae comprises trace fossils that can be attributed indistinctly to ants or termites, and also because trace fossil names should not make reference to possible producers (e.g. Brom- ley 1990). In contrast, Krausichnus shows one of the most complex morphologies of the group, and its name does not refer to the presumed constructor. Among the ichnotaxa included within the Krausichnidae, Termitichnus, Vondrichnus and Fleaglellius constitute a well-defined morphological group that may deserve the creation of an ichnofamily distinct from the Krausichni- dae once they are better known.

In addition to named ichnogenera, there is an extensive pedologic literature in relating to the origin of laterites, dealing with unnamed trace fossils attributed to termites, mostly from the Pliocene and Pleistocene of tropical areas (e.g. Machado 1983; Grass6 1986; Schaefer 2001). It is almost impossible to deal with the ichnotaxonomy of this material because in most cases the supposed nests were only studied micromorpho- logically and show no clear boundaries. Meaning that structures were only studied micromor- phologically (not macromorphologically) macro- morphological studies are lacking. Conversely, this kind of study is needed in already named trace fossils, and could usefully be conducted in order to evaluate the degree of reliability of the attribution of these ichnogenera to termites



Jaqueline

Realce

436 J. F. GENISE

Key to the ichnogenera of Krausichnidae

1 Chamber walls neither constructed nor lined 2 Chamber walls constructed or lined 3

2 Spherical chambers having only one main vertical burrow connected to the lower part or, rarely, another one at the top of the chamber. Secondary burrows connecting the main ones and chambers at different points. The distribution of chambers is similar throughout the structure Attaichnus Laza Oblate to subspherical chambers interconnected by burrows of similar diameter arising from top, bottom and sides of chambers. Chambers and burrows more numerous in the central part of the nest Parowanichnus Bown et al.

3 Structures composed of tiered, tabular and flat chambers supported by pillars and ramps 4 Structures composed of subspherical, ovoid or oblate chambers 5

4 Tiered, tabular and flat chambers composing spindle-shaped to columnar compound chambers lacking an external, discrete, wall. Compound chambers interconnected by a burrow system and/or isolated burrows Krausichnus Genise & Bown Tiered, tabular and flat chambers composing a columnar, central structure surrounded by a discrete wall. A burrow system connects this structure to peripheral oblate chambers and possibly other columns Archeoentomichnus Hasiotis & Dubiel

5 Structures composed of a single, large, cup-shaped chamber surrounded by a thick constructed wall bearing a boxwork of secondary burrows and chambers and a peripheral system of radiating burrows with differing diameters Tacuruiehnus Genise Structures composed of a system of chambers having similar sizes 6

6 Chambers connected by primary burrows of similar sizes 7 Chambers connected by primary and secondary burrows 8

7 At most three apposed, obovate chambers. Commonly having a rind of anastomosing burrows at top and sides of chambers. Diffuse arrangement of chambers Two or more apposed chambers commonly forming towers. Bases and tops of chambers convex upwards without a rind of anastomosing burrows

8 Spherical to subspherical chambers. Burrows, simple or compound and well differentiated from chambers, having much smaller diameters Elongate to oblate chambers. Burrows consistently simple, in some cases comparable in size to chambers

Vondrichnus Genise & Bown

Fleaglellius Genise & Bown

Termitichnus Bown

Syntermesichnus Bown & Laza

or ants. For instance, whereas Krausichnus, Termitichnus and Attaichnus leave few doubts about their termite and ant origin respectively, Taeuruichnus is known from a single specimen and requires further confirmation of its structure using more material. Vondrichnus, Fleaglellius and Parowanichnus are comparatively simple structures lacking the distinctive boxwork (sensu Ekdale et al. 1984) composed of tunnels of different diameters. Finally, Syntermesichnus and Archeoentomichnus were described from fragmentary material, from which the whole structure and ichnogeneric diagnoses were inferred. Micromorphological confirmation of the biological affinities of these trace fossils is critical, moreover, as the Triassic Archeoentomich- nus predates the oldest (Cretaceous) termite body fossils by about 150 million years (Hasiotis & Dubiel 1995a) (Table 1).

The taxonomy of this ichnofamily is complex because the trace fossils were described using different criteria to select taxobases and particularly because the diagnoses were strongly influenced by the architecture of modern nests of the sup-

posed constructors (e.g. Bown & Laza 1990; Laza 1995, 1997; Hasiotis & Dubiel 1995a).

Ichnogenus Attaichnus Laza 1982 (Fig. 5a)

v* 1982 A ttaichnus Laza p. 112, pls II, III. 1993 Attaichnus Laza; Genise p. 53. 1997 A ttaichnus Laza; Bown, Hasiotis, Genise,

Maldonado & Browers p. 45. 1998 Attaichnus Laza; Buatois, Mfingano,

Genise & Taylor p. 227. 1999 Attaichnus Laza; Genise p. 111. 2000 A ttaichnus Laza; Genise, Buatois,

M~mgano, Laza 7 Verde p. 55. 2001a Attaichnus Laza; Retallack p. 143. Type and only known ichnospecies. Attaichnus

kuenzelii Laza 1982. Diagnosis. System of spherical chambers

interconnected by primary and secondary burrows. Primary burrows connected to the chambers vertically to the lower part, forming inside a folded rim in some cases. A second primary burrow can be connected at the opposite side of the chamber. The secondary burrows inter- connect chambers with primary ones. The




Fig. 5. Krausichnidae. (a) Attaichnus kuenzelii Laza 1982: chamber and burrow casts. Late Miocene Cerro Azul Formation, Argentina (MACN-LI1787, 1788, 1791, 1792). Bar: I cm. (b) Krausichnus trompitus Genise & Bown 1994b, close-up of the holotype: flat chambers with parallel roofs and floors, vertical pillars (centre), and dark linings and fillings. Late Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 1 cm. (c) Tacuruichnus farinai Genise 1997: holotype. Close to the knife it is possible to see the external wall bearing a system of burrows and chambers; on the left, large burrows radiating from the cup-shaped nest. Late Pliocene Barranca de Los Lobos Formation, Argentina. Bar: 10cm.

chamber system occupies a conical area up to 7 m in diameter and 3 m in height, in which the chambers are regularly distributed (modified from Laza 1982).

Remarks. This trace fossil is known only from the type locality. However, its morphological features are clearly distinguishable from those

of the other representatives of Krausichnidae. The large (140-170 mm) spherical chambers connected by one or at most two primary burrows at the base and at the top are unmistakable traits. This trace fossil is preserved as detached casts of chambers and burrows having no indication of the presence of a lined or constructed wall



438 J. F. GENISE

(Fig. 5a). Laza (1982) attributed this trace fossil to ants of the genus Atta because of the gross morphology of the structure, the size and shape of chambers and burrows, the folded rim of primary burrows entering to the chambers, and the secondary burrows.

Ichnogenus Parowanichnus Bown et al., 1997 (Fig. 4d)

* 1997 Parowanichnus Bown, Hasiotis, Genise, Maldonado & Brouwers p. 45, figs 4, 5.

1999 Parowanichnus Bown, Hasiotis, Genise, Maldonado & Brouwers; Genise p. 111.

2000 Parowanichnus Bown, Hasiotis, Genise, Maldonado & Brouwers; Genise, Buatois, M~ngano, Laza & Verde p. 55.

Type and only known ichnospecies. Parowanichnus formicoides Bown et al. 1997. Diagnosis. System composed of oblate to

subspherical chambers interconnected by burrows of similar diameter. Burrows connected at tops, sides and bottoms of chambers. The gallery network is crudely trellate in plan, forming a grid-like lattice with descending shaft galleries and lateral tunnel galleries set more or less perpendicular to one another. Lined or constructed walls are lacking. Burrows, and the chambers they connect, radiate away from the centre of the structure and gradually decline in number. The chamber system occupies an area up to approximately 1 m in height and 3.3 m in width (modified from Bown et al. 1997).

Remarks. Parowanichnus differs from A ttaieh- nus in having: (1) a much smaller total nest volume; (2) much smaller and less densely packed chambers; (3) chambers oblate to hemi- spherical rather than globular; (4) galleries of one basic size (smaller than in Attaichnus); (5) galleries providing access equally to top, sides and bottom of chambers; and (6) chambers and burrows more densely grouped in the central part of the structure (Bown et al. 1997). As with Attaichnus, chambers and burrows lack linings or constructed walls. This trace fossil is attributed to ants because its structure is composed of burrows connected with chambers, and it lacks lined or constructed walls (Bown et al. 1997).

Ichnogenus Krausichnus Genise & Bown, 1994 (Fig. 5b)

?1981 'Fossilized nests of Hodotermitidae' Coaton p. 79, fig. 1.

1993 'Chevron-shaped chambers' Bown & Genise p. A58.

v* 1994b Krausichnus Genise & Bown p. 169, figs 6H-I, 7, 8D, 9-11, 12A.

1998 Krausichnus Genise & Bown; Buatois, M~mgano, Genise & Taylor p. 227.

V?1998 Krausichnus Genise & Bown; Genise, Pazos, Gonz~lez, T6falo & Verde p. 12.

1999 Krausichnus Genise & Bown; Genise p. 112.

2000 Krausichnus Genise & Bown; Genise, Buatois, M~ingano, Laza & Verde p. 55.

2000 Krausichnus Genise & Bown; Miller & Mason p. 210.

?2000 'Termitibresfossiles" Schuster, Duringer, Nel, Brunet, Vignaud & Mackaye p. 15, figs 3-5.

2002 Krausichnus Genise & Bown; Pazos, T6falo & S~nchez-Bettuci p. 34.

2002 Krausichnus Genise & Bown; Buatois, M~ingano & Acefiolaza p. 189.

Type ichnospecies. Krausichnus trompitus Genise & Bown 1994b. Diagnosis. Tiered arrangement of tabular, flat

chambers with roofs and floors flat and parallel. Chambers exhibit vertical pillars, partition walls and conspicuous linings. Successive chambers are connected by tiny passages. Arrangement of tiered chambers may take various forms, such as spindles or columns, resulting in compound chambers without surrounding walls. These compound chambers may be interconnected by isolated or interconnecting burrows (modified from Genise & Bown 1994b).

Included ichnospecies. K. trompitus Genise & Bown 1994b; K. altus Genise & Bown 1994b.

Remarks. The original ichnogeneric diagnosis included details of the shape and arrangement of compound chambers, but it is now clear that such taxobases are more useful at ichnospecific level. A third, unnamed ichnospecies (Coaton 1981; Schuster et al. 2000) demonstrates that the morphological features that compose a recurrent architecture of ichnogeneric rank include tabular and tiered chambers showing pillars and forming compound chambers without external walls. The shape of the compound chambers and their interconnections are useful to distinguish ichnospecies. The whole structure may consist of a single compound chamber or a system of related compound chambers. Another ichnogenus that shows tiered tabular chambers is Archeoentomichnus, in which these structures are connected with simple elongate oblate chambers.

In K. trompitus and K. altus, plus the Chadian material (Schuster et al. 2000), the combination of chambers and burrows indicates a social insect. The conspicuous linings and the constructed pillars and vertical walls specifically suggest termites (Genise & Bown 1994b; Schuster et al. 2000). Accordingly, Coaton (1981) attributed his unnamed fossil nests to the Hodotermitidae.




Ichnogenus Archeoentomichnus Hasiotis & Dubiel 1995a (Fig. 4e)

1993 Archeoentomoichnos Hasiotis & Dubiel p. 177 (nomen nudum).

1994 Archeoentomichnus Hasiotis & Dubiel p. 8 (nomen nudum).

* 1995a Archeoentomichnus Hasiotis & Dubiel; Hasiotis & Dubiel p. 121, figs 3-5.

1995b Archeoentomoichnus Hasiotis & Dubiel; Hasiotis & Dubiel p. 86, fig. 1.6 (lapsus).

1998 Archeoentomiehnus Hasiotis & Dubiel; Buatois, M~ngano, Genise & Taylor p. 225.

1999 Archeoentomichnus Hasiotis & Dubiel; Genise p. 112.

2000 Archeoentomichnus Hasiotis & Dubiel; Genise, Buatois, M/mgano, Laza & Verde 2000 p. 54.

2000 Archeoentomichnus Hasiotis & Dubiel; Hasiotis p. 154.

2001 a Archaeoentomonichnus Retallack p. 143 (lapsus).

Type and only known ichnospecies. Archeoentomichnus metapolypholeos Hasiotis & Dubiel 1995a. Diagnosis. Peripheral part of the structure

composed of large and small anostomosing galleries and elongate and oblate chambers. Central part columnar and partially subterranean, internally with stacked floor levels and slender, steeply inclined, connecting ramps. Galleries connect the central part to elongate oblate chambers and possibly to other columnar parts (modified from Hasiotis & Dubiel 1995a).

Remarks. The original diagnosis is slightly modified to avoid interpretative terms such as periecie or endoecie that correspond specifically to termite architecture. This ichnogenus is described from fragmentary material. The diagnosis is based on an idealized reconstruction, rather than the holotype's morphology as illustrated by Hasiotis & Dubiel (1995a). The reconstructed morphology is similar to that of Krausichnus, but in Archeoentomichnus the tiered chambers are connected with elongate, oblate, simple chambers, which are lacking in Krausichnus. Moreover, the tabular chambers are thicker, and the central part of the nest less organized in general aspect than in Krausichnus.

Ichnogenus Tacuruichnus Genise 1997 (Fig. 5c)

v* 1997 Tacuruichnus Genise p. 140, figs 2, 3. 1999 Tacuruichnus Genise; Genise p. 112. 2000 Tacuruichnus Genise; Genise, Buatois,

M~mgano, Laza & Verde p. 55. Type and only known ichnospecies. Tacuruich- nus farinai Genise 1997.

Diagnosis. Cup-shaped structure composed of a wall bearing a net of anostomosing burrows surrounding a chamber. Exteriorly the wall is connected to a gallery system composed of burrows of different diameters (Genise 1997).

Remarks. Tacuruichnus looks like a large specimen of Termitiehnus qatranii; however, in the latter ichnogenus the chambers are spherical and closed, because they would have been located completely below ground (Genise & Bown 1994b; Genise 1997). Tacuruichnus is the only krausichnid represented by a structure composed of a single large chamber. However, the characteristic burrow and chamber systems of the ichnofamily are present in the chamber's external wall and periphery (Fig. 5c).

Its architecture closely resembles the hypo- geous part of the nest of Cornitermes cumulans (Nasutitermitinae) (Genise 1997). The presence of a single chamber, interpreted as a nest, is also compatible with nests of C. cumulans. It would be important to find more specimens to determine the complete morphology and the taxonomic affinity of this trace fossil.

Ichnogenus Vondrichnus Genise & Bown, 1994 (Fig. 6a)

*1994b Vondrichnus Genise & Bown p. 165, figs 5J; 6A, B.

1998 Vondrichnus Genise & Bown; Buatois, M/mgano, Genise & Taylor p. 227.

1999 Vondrichnus Genise & Bown; Genise p. 112.

2000 Vondrichnus Genise & Bown; Genise, M~ngano, Buatois, Laza & Verde p. 54.

2002 Vondrichnus Genise and Bown; Buatois, M~ngano & Acefiolaza p. 189.

Type and only known ichnospecies. Vondrichnus obovatus Genise & Bown 1994b. Diagnosis. Diffuse, polychambered, excavated

subterranean systems. Obovate chambers occur in dense swarms of near 300 in cross-section. Burrows simple, branched or unbranched, exit- ing from one or more points on periphery of chamber and comprising a dense mass of anastomosing burrows that may connect chambers. Sediment in the centre of the chambers is alveolar and commonly arranged in concentric bands. Chambers expanded by apposition of 1-3 chambers against one another (Genise and Bown 1994b).

Remarks. Vondrichnus differs from Termitich- nus in: (1) consistently smaller mean chamber size; (2) obovate form of chamber; (3) tighter packing of associated chambers; (4) absence of isolated chambers; (5) larger number of galleries in each cluster; (6) lack of gallery ornamentation; (7) greater density of galleries between chambers;



440 J .F . GENISE

Fig. 6. Krausichnidae. (a) Vondrichnus obovatus Genise & Bown 1994b. Late Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 1 cm. (b) Fleaglellius pagodus Genise & Bown 1994b: three-chambered tower. Late Eocene~)ligocene Jebel Qatrani Formation, Egypt. Bar: 1 cm. (e) Syntermesichnusfontanae Bown & Laza 1990: interconnected chambers lined with a lighter fine material; cross-section of an individual thin passage (on the left). Miocene Pinturas Formation, Argentina; MACN-LI81. Bar: 1 cm. (d) Termitichnus simplicidens Genise & Bown 1994b: main burrows (particularly visible on the right) radiate from the base of the chamber. Late Eocene-Oligocene Jebel Qatrani Formation, Egypt. Bar: 10cm.

and (8) absence of compound galleries. Vondrich- nus differs from Fleaglellius in: (1) never having more than two apposed chambers; (2) rarely having only vertically apposed chambers; (3) not having convex-upward chambers; (4) commonly possessing a rind of anastomosed peripheral galleries at the tops and sides of chambers; and (5) occurring as diffuse but interconnected groups of chambers (Genise & Bown 1994b). Some of the characters stated by the

authors, such as size, packing of chambers and number of burrows, are undoubtedly very useful for dividing the morphological complex of Termitichnus-Fleaglellius-Vondrichnus; however, they are not very satisfactory as ichnotaxobases in a more general context. Vondrichnus and Fleaglellius show a primary burrow system devoid of secondary tunnels that distinguish them from Termitichnus. Vondrichnus can be clearly separated from Fleaglellius because in




the latter the chambers are convex upwards and form 'towers' by vertical apposition (Genise & Bown 1994b) (Fig. 6a, b). Possible trace-makers are termites, possibly Macrotermitinae (Genise & Bown 1994b). The very simple architecture of this trace fossil necessitates further examination of micromorphological characters to confirm its affinities.

Ichnogenus Fleaglellius Genise & Bown, 1994 (Fig. 6b)

1994b Fleaglellius Genise & Bown p. 167, figs 6C-G, 8E-F.

1998 Fleaglellius Genise & Bown; Buatois, M~ngano, Genise & Taylor p. 227.

1999 Fleaglellius Genise & Bown; Genise p. 112. 2000 Fleaglellius Genise & Bown; Genise,

M~ngano, Buatois, Laza & Verde p. 54. 2002 Fleaglellius Genise & Bown; Buatois,

M~ingano & Acefiolaza p. 189. Type and only known ichnospecies. Fleaglellius

pagodus Genise & Bown 1994b. Diagnosis. Diffuse, polychambered, excavated

subterranean system. Chambers oblate, with bases and tops of chambers almost invariably convex-upward and generally apposed. Succes- sive chambers are added vertically or near vertically, with the top of the lower completely overlapped by the base of the upper chamber, such that completed structures make up towers of 2-35 chambers. Towers are widely dispersed in groupings of up to 40 specimens and are connected by dense masses of simple galleries. Gal- leries, both branched and unbranched, connect different towers at all levels (Genise & Bown 1994b).

Remarks. This ichnogenus differs from its closest morphological counterpart, Vondrichnus, in that Fleaglellius: (1) always has more than one and up to 35 apposed chambers, always added vertically; (2) invariably has individual chambers that are noticeably convex-upwards; (3) lacks a peripheral rind of anastomosed galleries at tops and/or sides of chambers; and (4) lacks a diffuse distribution of individual chambers (Genise & Bown 1994b). Possible trace-makers are termites of unknown affinities (Genise & Bown 1994b). The particular kind of enlargement of nest by vertical apposition of chambers is unknown in subterranean termites, but resembles that of certain subaerial nests. This ichnogenus requires micromorphological analysis of its affinities.

Ichnogenus Termitichnus Bown, 1982 (Fig. 6d)

v* 1982 Termitichnus Bown p. 259, figs 2-7. ? 1984 Termitichnus Bown; Tandon & Naug

p. 285, figs 7B, 9B-E. 1993 Termitichnus Bown; Genise p. 54.

?1993 Termitichnus Bown; Smith, Mason & Ward p. 592, fig. 16.

V1994b Termitichnus Bown; Genise & Bown p. 160, figs 4, 5, 8A-C.

1997 Termitichnus Bown; Genise p. 140. 1998 Termitichnus Bown; Buatois, M~ngano,

Genise & Taylor p. 227. 1998 Termitichnus Bown; Smith & Mason p. 555,

figs 12, 13, 14A. 1999 Termitichnus Bown; Genise p. 112. 2000 Termitichnus Bown; Genise, M~ngano,

Buatois, Laza & Verde p. 54. 2000 Termitichnus Bown; Miller & Mason p. 208,

figs 12-16. 2001a Termitichnus Bown; Retallack p. 143. 2002 Termitichnus Bown; Buatois, M~ingano &

Acefiolaza p. 189. Type ichnospecies. Termitichnus qatranii Bown,

1982. Diagnosis. Diffuse, polychambered excavated

subterranean systems with spherical to subspherical chambers connected to one another by a network of simple and/or compound galleries of different diameters. Primary burrows arise from the base of chambers. Chambers may be isolated, as expanded globular clusters of chambers or in associated constellate aggrega- tions of tens to hundreds of chambers (modified from Genise & Bown 1994b).

Included ichnospecies. T. qatranii Bown 1982; T. simplicidens Genise & Bown 1994b; T. namibiensis Miller & Mason 2000.

Remarks. The diagnosis has been modified to exclude characters such as size and fill of chambers, which are more useful for ichnospecific diagnosis. Also excluded are those characters referred to 'nest expansion', which is a concept linked to the supposed constructors rather than to documented morphology. The basal position of the primary burrows is an important character added to the diagnosis, because it is constant and very representative in both ichnospecies. Differ- ences from Fleaglellius and Vondrichnus were commented on in previous sections. Attaichnus has consistently spherical chambers always, and a single primary burrow arises from the base of chambers, and in some cases other from the top. The architecture of Syntermesichnus is clearly different from that of Termitichnus; however, in terms of ichnotaxobases it is difficult to translate these differences avoiding measure- ments and other taxobases of dubious merit. In addition the commonest type of Termitichnus comprises spherical or subspherical chambers from which one to five primary burrows arise, some of which may connect another distant chamber (Fig. 6d). In contrast, Syntermesichnus takes on the aspect of a boxwork of burrows



442 J. F. GENISE

and chambers of rather similar diameters (Fig. 6c).

Not all of the trace fossils attributed to Termi- tichnus after its original description (Bown 1982) and redescription (Genise & Bown 1994b) belong to that ichnogenus. Smith et al. (1993) reported supposed Termitichnus from the late Pleistocene of Namibia, and proposed a Termitichnus ichnofacies, but their figured specimen does not closely resemble Termitichnus. Probable Termitichnus have subsequently been illustrated from the Tertiary of Namibia by Smith & Mason (1998): these are discussed below with unnamed Krau- sichnidae.

An additional ichnospecies of Termitichnus from early Pleistocene deposits of South Africa, T. namibiensis, is composed of isolated or interconnected chambers, surrounded by a thick wall and a network of simple tunnels (Miller & Mason 2000). Chambers of T. namibiensis are filled with meniscate, tiered, galleries. A thick wall and peripheral tunnel system surround isolated or interconnected subspherical chambers typical of Termitichnus. The attribution to this ichnogenus to fossil termite nests is also supported by the tiered arrangement of shelves, as in other fossil and modern termite nests (e.g. Sands 1987). However, some characters raise doubts about the ichnotaxonomical placement of this trace fossil. Chambers apparently lack the characteristic primary tunnels of Termitich- nus, and the interpretation of the internal structure is confusing. It is not clear how to interpret a gallery that extends horizontally and also shows a meniscate filling, which commonly results from the backfilling of burrows of similar diameter from that of the trace-maker (e.g. D'Alessandro & Bromley 1987; Keighley & Pickerill 1994). Also, the turning points at the end of galleries are unusual, in that the thin layers between two successive shelves commonly become thicker towards the sides before joining the external wall (e.g. Sands 1987). These thickenings help to reinforce the whole structure. In T. namibiensis the thickenings are disconnected from the outer wall at the turning points. In addition, T. namibiensis also lacks the ramps, pillars and openings that are common in Krau- sichnus, as well as the thin-layered ovoids and the hive of modern termite nests (e.g. Sands 1987; Genise & Bown 1994b). In sum, the internal structure of the chambers of T. namibiensis (Miller & Mason 2000, fig. 12) plus the above- mentioned characters leave some doubts about the trace-makers of this internal structure, which may be different from those of the chambers. These characters, plus the absence of primary burrows arising from the base of

the chambers, suggest that the inclusion of this trace fossil in Termitichnus would require further work. On the other hand, although the absence of external wall makes some specimens comparable to Krausichnus, these specimens are incomplete and eroded (Miller & Mason 2000). Possible trace-makers include termites belonging to Macrotermitinae (Genise & Bown 1994b).

Ichnogenus Syntermesichnus Bown & Laza 1990 (Fig. 6c)

v* 1990 Syntermesichnus Bown & Laza p. 74, figs 2-4.

1993 Syntermesichnus Bown & Laza; Genise p. 54.

1995 Syntermesichnus Bown & Laza; Genise and Cladera p. 80, fig. 2C-D.

1995 Syntermesichnus Bown & Laza; Constantino p. 460.

?1997 Syntermesichnus Bown & Laza; Smith & Kitching p. 41, figs 16, 17.

1997 Syntermesichnus Bown & Laza; Bown, Hasiotis, Genise, Maldonado & Browers p. 46.

1998 Syntermesichnus Bown & Laza; Buatois, Mfingano, Genise & Taylor p. 227.

1999 Syntermesichnus Bown & Laza; Genise p. 112.

2000 Syntermesichnus Bown & Laza; Genise, Mfingano, Buatois, Laza & Verde p. 54.

2002 Syntermesichnus Bown & Laza; Buatois, Mfingano & Acefiolaza p. 189.

Type and only known ichnospecies. Syntermesichnusfontanae Bown & Laza 1990. Diagnosis. Peripheral part of the structure,

tabular, with large and small anastomosing burrows and elongate, oblate chambers. Large burrows arising from chambers, but small burrows branch from larger ones. Systems of small passages at one level extend the structure in the horizontal plane and permit communication to other levels. The same style of branching is present at all levels. Walls of burrows and chambers lined with compacted sediment (finer than host matrix) (modified from Bown & Laza 1990).

Remarks. Modifications of the original diagnosis were made to avoid any reference to modern termite nests. Syntermesichnus shows one of the simplest morphologies of Krausichni- dae and, consequently, one of the most difficult to define and separate from the other Krausich- nidae. The whole structure lacks definite limits and, in the field, resembles a diffuse boxwork of tunnels and chambers of more or less similar diameter occupying entire beds (Genise & Bown, unpublished data). The abundance of similar Syntermesichnus-like traces in pyroclastic




deposits from the Cretaceous and Tertiary of southern South America (Genise, unpublished data) gives this trace fossil a particular importance. The chamber and burrow systems, lined with fine material, suggested to Bown & Laza (1990) that Syntermesichnus was a fossil termite nest. However, the lack of a definite architecture (e.g. nest limits) and its pervasive presence in beds preclude a definite attribution without micromorphological analysis of the lined walls. Trace fossils comparable to Syntermesichnus are described from the Jurassic Elliot Formation of South Africa (Smith & Kitching 1997), although this attribution needs further analysis.

Possible trace-makers are termites of the genus Syntermes (Bown & Laza 1990). However, as discussed above, this attribution requires micromorphological study and the examination of further material to look for well-delimited nests. In addition, as Constantino (1995) noted, the descriptions of Syntermes nests in the literature are inadequate, so that the affinities of Syntermesichnus should be considered doubtful.

Unnamed trace fossils attributable to Krausichnidae Various trace fossils from the Pliocene Laetoli Formation of Tanzania, all attributable to Krausichnidae, were described by Sands (1987). He recognized seven basic types.

The first type is composed of systems of anastomosing burrows of different diameters, which may show vertical shafts associated with flattened chambers. These systems are comparable to the structures attributed by Smith et al. (1993) to Termitichnus. However, these structures are not clearly compatible with the ichnogeneric diagnosis. The second type with thick-layered ovoids (=chambers) shows particular features, such as the presence of a surrounding cavity, that are unknown from other fossil termite nests. These ovoids lack an external wall and associated burrow systems. They somewhat resemble very roughly ichnospecies of Krausichnus in having a layered structure, but the layers are unusually thick. A third type of trace fossil with shafted chambers is not comparable to any other Krausichnidae. This type is composed of a bell-shaped chamber from which vertical shafts arise, which are capped by smaller shafts, chambers and pores. The fourth type, composed of a single ovoid, is similar to the second type, but has an external wall and ramps and passages from one floor to the next. The fourth type can be attributed to neither Krausichnus nor Termitichnus because Krausichnus does not have an outer wall and Termitichnus does not have a comparable layered

structure. The fifth type is represented by three small ovoids showing pits, protrusions, ramps and openings respectively. These resemble similar structures from the Tertiary of Namibia attributed to Termitichnus by Smith and Mason (1998), and also unnamed trace fossils from the early Miocene of Ethiopia (Bown & Genise 1993). They superficially resemble Termitichnus, but lack the associated burrow system, which is diagnostic. The sixth type is similar to the first type but also has chambers and pores arranged in anastomosing columnar and alveolar structures. No other trace fossils are comparable to the sixth type. The last type, from the upper part of the Laetoli beds, consists of thin-layered ovoids that show a layered internal structure having ramps and connecting openings surrounded by a sculptured wall. This type is very similar to unnamed trace fossils described by Coaton (1981) and Schuster et al. (2000) from the Pleistocene of South Africa and the Pliocene of Chad respectively, and is attributable to a distinct ichnospecies of Krausichnus. However, the ovoids from Laetoli have a constructed wall, an unusual trait for Krausichnus that is lacking in the other African material. Miller & Mason (2000) considered these structures attributable to their ichnospecies Termitichnus namibiensis.

Sands (1987) related most of these types of trace fossil to different parts of Macrotermitinae nests at different stages of development and/or constructed in different types of soil, with two exceptions: the vertical shaft with flattened chambers, which more probably relates to ant nests of Camponotus, and the thin-layered ovoids, which are attributable to neither the Macrotermitinae nor the Hodotermitidae (Sands 1987).

In summary, the Laetoli material (Sands 1987) represents a spectrum of samples of krausichnids that share common features with other representatives of the ichnofamily from Africa (e.g. Coaton 1981; Bown & Genise 1993; Smith et al. 1993; Genise & Bown 1994b; Smith & Mason 1998; Schuster et al. 2000). This similarity reflects the origin of these trace fossils in common lineages of African termites. Ichnotaxo- nomically, the Laetoli material presents some problems. Considering the architecture of Macrotermitinae nests, each type described is more likely a part of a more complex structure than a complete structure in itself. This fact reiter- ates a common trend in evolution of behaviour in insects, in which complex architectures and behaviours are the result of the addition of simple behaviours and architectures respectively. One possibility is to include the types one and six



444 J .F . GENISE

(tunnel systems) in Syntermesichnus, interpreting this ichnogenus as diffuse boxworks of tunnels and chambers. It would also be possible to include the thick- and thin-layered ovoids in Krausichnus or Termitichnus, but important features distinguish these ovoids from the known ichnospecies. Another standpoint might be to include each Laetolian type in a new ichnotaxon, even if each is part of a more complex structure; this procedure would be supported by the fact that some of these types are recurrent as individual trace fossils in other localities and ages in Africa (e.g. Coaton 1981; Bown & Genise 1993; Smith & Mason 1998; Schuster et al. 2000).

Other unnamed Krausichnidae were described by Laza (1995, 1997) from Pliocene and Pleisto- cene deposits of Argentina. They include seven types of trace fossil: two attributed to termites, and five to ants. One type of termite nest is currently being redescribed by Laza (personal communication) and two ant nests described in 1995 were redescribed by Laza (1997). Hasiotis & Demko (1996) also described a supposed ant nest from the Jurassic Morrison Formation. The importance of all this material for our knowledge of the diversity of Krausichnidae is unquestionable. However, in all cases the diagnoses and descriptions are influenced by the supposed modern analogue, which, in combination

with the fragmentary nature of the material, makes any ichnotaxonomical consideration very difficult. There are still other unnamed trace fossils, cited as termite or ant nests (e.g. Tandon & Naug 1984; Iriondo & Kr6hling 1996; Tauber 1996; Andreis & Cladera 1998) that are only mentioned and will require analysis and proper description.

A sound description of this unnamed material, in the light new understanding of krausichnid morphology, will aid incorporation of this material into the emerging ichnotaxonomic framework.

I c h n o s t r a t i g r a p h y

There are 25 described ichnogenera attributed to insect trace fossils in palaeosols, 19 of which are reviewed herein, and the remaining six in a previous contribution (Genise 2000) (Table 1). Almost all of them show comparable stratigraphic ranges that, in turn, accord with our present knowledge of the evolutionary history of their trace-makers: bees, beetles, ants and termites (e.g. Crowson 1981; Kuschel 1983; Krishna 1990; Jarzembowski & Ross 1996; Grimaldi et al. 1997; Grimaldi 1999; Engel 2000; Grimaldi & Agosti 2000; Krell 2000; Schaefer 2001; Nel et al. in press).

Table 1. Stratigraphic ranges and abundance of insect ichnotaxa in palaeosols

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Pleistocene 4 1 1 3 1 1 Pliocene 4 1 1 1 1 2 Miocene 3 O 1 O 1 1 1 1 2 4 2 1 Oligocene 2 1 1 1 1 1 1 3 1 Eocene 1 5 2 1 3 1 4 1 1 1 1 1 1 1 3 1 1 1 Palaeocene 1 1 2 Upper Cretaceous @ ~ 0 �9 1 3 ~ @ G @ @ @ 0 Middle Cretaceous Lower Cretaceous Jurassic @ 1 Triassic Permian Carboniferous Devonian Silurian Ordovician Cambrian

O O O O O O O O O O ?

1

Coprinisphaeridae (1-6), Pallichnidae (7-9), Krausichnidae (10-18) and Celliformidae (19-25). 1, Fontanai; 2, Coprinisphaera; 3, Eatonichnus; 4, Monesichnus; 5, Teisseirei; 6, Rebuffoichnus; 7, Pallichnus; 8, Fictovichnus; 9, Scaphichnium; 10, A ttaichnus; 11, Parowanichnus; 12, Krausichnus; 13, A rcheoentomichnus; 14, Tacuruichnus; 15, Vondrichnus; 16, Fleaglellius; 17, Termitichnus; 18, Syntermesichnus; 19, Palmiraichnus; 20, Celliforma; 21, Corimbatichnus; 22, Uruguay; 23, Rosellichnus; 24, Ellipsoideichnus; 25, Cellicalichnus. Numbers in cells represent formations in which the ichnotaxon is recorded. Circles indicate the oldest body fossil of the potential trace-maker.




The Coprin isphaer idae

Among the Coprinisphaeridae, Fontanai has been recorded only from the Late Cretaceous- Early Tertiary Asencio Formation of Uruguay (Roselli 1939). These redbed deposits could not be dated precisely until now because of the complete absence of datable rocks or fossil remains other than the fossil nests (Genise et al. 2004). Genise et al. (2002a) suggested a possible Eocene age for this unit, owing to the presence of an important unconformity with the under- lying Cretaceous Yapeyfi Formation (Pazos et al. 2002), comparison of the ichnofauna with other Palaeogene deposits in South America, and the abundance and diversity of dung-beetle nests. This abundance and diversity could reflect the increased availability of herbivore faeces corresponding to the Eocene diversification and abundance of South American herbivores. In Table 1 all the records from this formation are considered to be Eocene. Because this formation is one of the richest in fossil insect nests - it includes 10 of the 25 described ichnogenera - it produces a peak of diversity for this age that may be an artefact, not simply because the age of this formation is unknown, but also because intensive research on other formations and continental deposits has just begun. Never- theless, it cannot be overlooked that, during the early Eocene, a climatic optimum occurred (e.g. Zachos et al. 2001), which would have favoured the abundance and diversification of some groups of insects (e.g. Wilf & Labandeira 1999).

Coprinisphaera and Celliforma are the most widely recorded ichnogenera in palaeosols. Coprinisphaera has been recorded from the aforementioned Asencio Formation in Uruguay (Roselli 1939, 1987; Genise et al. 2004), the early Eocene Casamayor Formation, Argentina (Frenguelli 1938b; Laza 1986a), the late Eocene Musters Formation, Argentina (Andreis 1972; Laza 1986a), the late Eocene La Meseta Forma- tion, Antarctica (Laza & Reguero 1990), the Eocene-Miocene Sarmiento Formation (Laza 1986a; Bellosi et al. 2001), the Oligocene Deseado Formation, Argentina (Frenguelli 1938b; Laza 1986a), the early Miocene Pinturas Formation, Argentina (Genise & Bown 1994a), the late Miocene Coll6n-Curfi Formation, Argentina (Frenguelli 1939; Laza 1986b), the late Miocene Paso de las Carretas Formation, Argentina (Pascual & Bondesio 1981), the Plio- cene Monte Hermoso Formation, Argentina (Laza 1986b), the Pliocene Piquete Formation, Argentina (Alonso et al. 1982), the Pliocene Laetoli Formation, Kenya (Sands 1987), the

Pliocene Chad Formation, Chad (Duringer et al. 2000a, 2000b), the Pleistocene Ensenada Formation, Argentina (Frenguelli 1938a), the Pleistocene Tezanos Pinto Formation, Argentina (Iriondo & Kr6hling 1996), the Pleistocene Taft del Valle Formation, Argentina (Fontaine et al. 1995), and from an unnamed Pleistocene formation, from Ecuador (Sauer 1955). Eatonichnus has been recorded from the Palaeocene Colter Formation, USA (Gilliland & La Rocque 1952), the late Palaeocene-Eocene Claron Formation, USA (Bown et al. 1997), and the early Palaeocene Pefias Coloradas Formation, Argentina (Genise et al. 2001). Monesichnus is another ichnogenus that has been recorded exclusively from the Late Cretaceous-Early Tertiary Asencio Formation of Uruguay (Roselli 1987).

The previously mentioned ichnogenera of Coprinisphaeridae are attributed to Scarabaei- nae (Frenguelli 1938; Laza 1986b; Bown et al. 1997; Genise & Laza 1998; Genise 1999; Krell 2000), whose oldest body fossils come from the Late Cretaceous strata (Krell 2000). In this case the body fossils shortly predate the trace fossils of the group, as there is no ichnological record for the Cretaceous. Moreover, the presence of Coprinisphaera is taken as indicative of Cenozoic deposits in South America (Genise et al. 2000). Halffter & Edmonds (1982) and Cambefort (1991) suggested that dung-beetles might have diversified by the end of the Cretaceous, helped by the radiation of the herbivorous dinosaurs and/or the increase of mammal excrement. The only Cretaceous trace fossils attributable to dung-beetles (non-Scarabaeinae) are those described by Chin & Gill (1996) from dinosaur coprolites.

The other two ichnogenera of Coprinisphaeri- dae deserve particular attention. Teisseirei has been recorded from the Late Cretaceous-Early Tertiary Asencio Formation, Uruguay (Roselli 1939), the Eocene Gran Salitral Formation, Argentina (Melchor et al. 2002), and the Eocene- Miocene Sarmiento Formation, Argentina (Bellosi et al. 2001). Despite the well-preserved macro- and micromorphological characters of this trace fossil, as well as its abundance in Tertiary deposits of South America, the trace-maker cannot yet be identified more precisely than as probably being a coleopteran. Hence it is impossible to compare the stratigraphic range of Teisseirei with that of the body fossil record of any particular coleopteran. Rebuffoichnus has been recorded from the Late Cretaceous Laguna Palacios Formation, Argentina (Genise et al. 200219), the Late Cretaceous-Early Tertiary Asencio Formation, Uruguay (Roselli 1987), and the



446 J.F. GENISE

Pleistocene of Australia (Lea 1925; Johnston et al. 1996). This ichnogenus, probably attributable to the work of weevils (Genise et al. 2002b), is one of only two that crosses the K-T boundary, being recorded from the Late Cretaceous to the Pleisto- cene. As such, it is one of the oldest trace fossils that can be definitely attributed to insects (Genise et al. 2002b). The body fossil record of their probable constructors, the curculionids, extends back to Upper Jurassic-Lower Cretaceous deposits (Crowson 1981; Kuschel 1983; Jarzem- bowski & Ross 1996). As a whole, the body fossil record of possible trace-makers predates the Coprinisphaeridae, even though the thick constructed walls confer to these trace fossils a high preservation potential.

The Pal l ichnidae

The Pallichnidae pose a different problem from that of the Coprinisphaeridae. Scaphichnium has been recorded exclusively from the Eocene Willwood Formation, USA (Bown & Kraus 1983), whereas its possible producers, the Geo- trupinae (Hasiotis et al. 1993), are recorded since the late Oligocene (Krell 2000). Pallichnus has been recorded only from the Oligocene Brule Formation in the USA (Retallack 1984), whereas its possible producers, the Geotrupinae or Scarabaeinae (Retallack 1984), are recorded from the late Oligocene and Late Cretaceous respectively (Krell 2000). Fictovichnus, in a broad sense, may include very simple trace fossils showing a morphology that has been recorded from the Late Jurassic Morrison Formation, USA (Hirsch 1994b), the Late Cretaceous Barun Goyot Formation, Mongolia (Mikhailov et al. 1994), the Late Cretaceous Djadokhta Formation, Mongolia (Johnston et al. 1996), the Late Cretaceous Bajo Barreal Formation, Argentina (Sciutto & Martinez 1996), the Palaeo- cene of Uruguay (Veroslavsky & Martinez 1996), the Eocene Claron Formation, USA (Bown et al. 1997), the Eocene of France (Freytet & Plaziat 1982; Hirsch 1994a), the Eocene Gran Salitral Formation, Argentina (Melchor et al. 2002), the late Eocene Bembridge Limestone Forma- tion, England (Edwards et al. 1998), the Miocene Higewi Formation, Kenya (Thackray 1994), and the Pliocene of Tanzania (Ritchie 1987). It is impossible to determine a unique trace-maker for Fictovichnus, whose makers may have included the Tenebrionidae, Curculionidae, Scarabaeidae (Johnston et al. 1996), or even other families of Coleoptera. This precludes any precise comparison with the body fossil record of the coleopteran families.

The Kraus ichnidae

Representatives of Krausichnidae show, in most cases, similar characteristics: body fossils predating trace fossils and scarcity of data, with some exceptions. Attaichnus and Parowanichnus, the two ichnogenera attributed to ants, have been recorded exclusively from the Miocene Epecu~n Formation, Argentina (Laza 1982) and from the Eocene Claron Formation, USA (Bown et al. 1997) respectively. The oldest body fossils of ants come from the Cretaceous of the USA and France (Grimaldi et al. 1997; Grimaldi & Agosti 2000; Nel et al. in press).

Krausichnus has been recorded from the Late Cretaceous-Early Tertiary Asencio Formation, Uruguay (Genise et al. 1998), the late Eocene Qasr el Sagha Formation and late Eocene-Oligo- cene Jebel Qatrani Formation, Egypt (Genise & Bown 1994b), the late Miocene Baynunah For- mation, Abu Dhabi Emirate (Bown & Genise 1993), the Pliocene of Chad (Schuster et al. 2000), and the Pleistocene of South Africa (Coaton 1981). Tacuruichnus has been recorded exclusively from the late Pliocene Barranca de los Lobos Formation, Argentina (Genise 1997), whereas Vondrichnus and Fleaglellius from the late Eocene-Oligocene Jebel Qatrani Formation, Egypt (Genise & Bown 1994b). Termitichnus shows a broader distribution, being recorded from the late Eocene-Oligocene Jebel Qatrani Formation, and the Late Eocene Qasr el Sagha Formation, Egypt (Bown 1982), the early Mio- cene Kashab Formation, Egypt (Genise & Bown 1994b), the Plio-Pleistocene Boulder For- mation, India (Tandon & Naug 1984), the late Pleistocene Homeb Silts, Namibia (Smith et al. 1993), the late Pleistocene Sossus Sand, Namibia (Smith & Mason 1998), and the Pleistocene of South Africa (Miller & Mason 2000). Finally, Syntermesichnus has been recorded from the Miocene Pinturas Formation, Argentina (Bown & Laza 1990), and there is a doubtful record from the Early Jurassic Elliot Formation, South Africa (Smith & Kitching 1997). With the exception of this last record, all other occurrences are predated by the oldest fossil termites from the Lower Cretaceous (Krishna 1990).

The case of Archeoentomichnus from the Triassic Chinle Formation, USA (Hasiotis & Dubiel 1995a) deserves particular attention because it is the only datum that does not fit the general picture shown in Table 1. These authors attributed this trace fossil to termites despite the fact that it would predate the oldest body fossils by 150 million years (Hasiotis & Dubiel 1995a). Many invertebrate trace fossils are more preservable than their producers:




accordingly, fossil bee nests predate the oldest bees (Elliott & Nations 1998; Genise 2000). However, the few million years involved in this difference between the oldest bee trace fossils and the oldest fossils of the probable constructors is an expected one. It is consistent with our previous knowledge of the evolutionary history of bees and their relationship with the coevolving plant groups (Grimaldi 1999; Engel 2001). In the case of Archeoentomichnus, the palaeoentomolo- gical significance that would result from the discover of a Triassic termite nest predating the oldest termites by 150 million years would require sound proof, such as micromorphological studies, to be accepted as such. At present, pending further work, this occurrence is not accepted.

The whole interpretation of Mesozoic evolution of palaeosol ichnofaunas attributed to insects by Hasiotis (2000) is mostly unsupported, in that the attribution of these early purported trace fossils to insects is not well documented or well founded. The conclusions are based largely on poorly supported interpretations of Triassic and Jurassic trace fossils from the Chinle and Morrison Formations. These interpretations are at odds with the strong evidence that indicate that ants, termites, bees and dung- beetles arose and diversified during the Cretac- eous (Krishna 1990; Labandeira 1998; Grimaldi 1999; Krell 2000; Engel 2001; Schaefer 2001). This conclusion is also supported by the ichnological record of insect nests in palaeosols (Table 1). Accordingly, Labandeira (1998), Grimaldi (1999), Genise (2000) and Engel (2001) objected to such interpretations of Chinle and Morrison trace fossils because of inadequate documentation.

Triassic and Jurassic trace fossils are unknown from other studied deposits of similar age, which adds significance to the Chinle and Morrison trace fossils, as increasing the necessity for sound descriptions and interpretations of the fossils. Other Triassic and Jurassic palaeosols studied are of low ichnodiversity, composed mainly of simple trace fossils such as Skolithos, Macanopsis, Taenidium and Edaphichnium, none of which can be certainly attributed to insects (Retallack 1976, 1980; Smith & Kitching 1997; Melchor et al. 2001; Genise et al. 2004). The morphology of trace fossils in palaeosols ranges from very simple to very complex. The attribution of traces of very simple morphology to modern taxa is a misleading procedure because they can commonly be attributed to several different groups of organisms (Ratcliffe & Fagerstrom 1980; Retallack 1990b; Genise et al. 2004). Trace fossils in Mesozoic palaeosols that

can unequivocally be attributed to insects are few, and are recorded from only three Late Cretaceous formations in the USA, Mongolia and Argentina (Johnston et al. 1996; Elliott & Nations 1998; Genise et al. 2002b). The record of insect fossil nests from rocks of this age accords with the body fossil record of their probable constructors and/or reflects their supposed evolutionary history. A scenario proposed by Genise & Bown (1994a) stated that the diversity of insect fossil nests in palaeosols increased significantly after the Cretaceous and not earlier. This hypothesis is based on the fact that most common trace fossils in palaeosols are constructions belonging to termites, bees and dung- beetles, groups that arose during that period based on body fossil evidence. This hypothesis was later corroborated by the discovery of Late Cretaceous bee nests and coleopteran pupal chambers (Johnston et al. 1996; Elliot & Nations 1998; Genise 2000; Genise et al. 2002b). The Genise and Bown hypothesis is also supported by the increase in abundance of records of insect fossil nests in Tertiary palaeosols, in contrast to the general absence of records from pre-Cretaceous deposits.

Although Labandeira & Sepkoski (1993) stated that the appearance and expansion of angiosperms had no influence on insect familial diversification, their quantitative analysis did not evaluate the ecological importance of the families involved. The origin of termites, ants, bees and dung-beetles during the Cretaceous was probably related to the origin and diversification of angiosperms and to the broad-scale ecological changes that resulted in the K-T mass extinction event. These insect trace- makers would have played a major role in this event as the new soil colonizers of the emergent ecosystems. The record of latosols, from the Cre- taceous onwards in the stratigraphic column, has been strongly related to the radiation of termites and angiosperms (Schaefer 2001). In contrast to other groups that became extinct or less domi- nant following the K-T event, insect ichnofossils in palaeosols show that their producers were part of the new flourishing ecosystems.

This evolutionary scenario is reflected in the stratigraphic range and abundance of insect trace fossils in palaeosols reflected in Table 1, which shows the absence of Celliformidae, Coprinisphaeridae, Pallichnidae and Krausichni- dae in pre-Cretaceous rocks. It also shows that the oldest record of two of these ichnofamilies came from Cretaceous rocks, which contain a lower diversity and abundance of nests than Tertiary ones. Even so, Cretaceous nests are very important ones, in relation to the origin of



448 J. F. GENISE

important groups of insects and their building behaviour. Tertiary rocks have the most diverse, abundant and well-preserved assemblages of insect fossil nests, in accordance with the diversification of insects and their building behaviour.

The initial manuscript benefited from comments by A. Uchman and A. Rindsberg. I thank also D. McIlroy, M. Verde and A. Rindsberg for improving the final version. This research was partially supported by a grant from the National Agency of Scientific and Technical Promotion of Argentina (FONCYT-PICT 6156/99) to the author.

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