The ichnofamily Celliformidae for Celliforma and allied ichnogenera

This contribution undertakes a comprehensive revision, lacking until now, of all the ichnotaxa attributed to fossil bee cells, one of the most common traces in paleosols. These ichnotaxa are morphologically related to Celliforma and consequently grouped herein in the new ichnofamily Celliformidae. Two new ichnogenera are formulated, Cellical‐ichnns igen. nov., for several ichnospecies previously included in Celliforma, and a new ichnospecies, C. chubutensis isp. nov., belonging to this ichnofamily, and Brownichnus igen. nov., for the ichnospecies favosites, which is not morphologically related to Celliforma. The ichnogenera included in this ichnofamily involve traces in which Celliforma, the most simple trace of the group, is part of their structure. Paltniraichnus is akin to Celliforma with antechambers and discrete walls. Uruguay, Corim‐batichnus and Rosellichnus are clusters composed of adjacent rows of Celliforma, or Palmiraichnus‐like cells. Ellipsoideichnus and Cellicalichnus are different arrangements of Celliforma‐like cells attached to tunnels.

Celliformidae are based on the morphology of the traces, all of them comprising cells, groups of cells and cells attached to tunnels. As such, its component ichnotaxa are based exclusively on morphological ichnotaxobases, which, in turn, were evaluated and selected with respect to the nest architecture of the trace makers, the bees. This paper analyzes some procedures and clues (i.e. behavioral homologies) that may be used to select the proper taxobases to erect ichnotaxa when the identity of the trace makers is known, as in this study case.     

Curved fossil bee cells as tools for reconstructing the evolutionary history and palaeogeographical distribution of Diphaglossinae (Apoidea,Colletidae)

The new ichnospecies Celliforma curvata is described to include curved fossil bee cells from Argentina, Uruguay and the USA. The upper part of the cell (neck) of the new ichnospecies is curved, and accordingly, it can be attributed to bees of the subfamily Diphaglossinae (Colletidae). The oldest record of C. curvata, from the early Eocene of North America (52–49 Ma), provides a minimum age for the appearance of this subfamily, in accordance with an already proposed calibrated phylogeny. It is also proposed that these fossil cells could be used for future calibrations of molecular clocks. C. curvata indicates that Diphaglossinae had a widespread distribution, from southern Utah to extra‐Andean Patagonia at 42°S. In contrast, extant representatives reach only 38°S in this region. Diphaglossinae were more extended southwards in the past thanks to better environmental conditions in extra‐Andean Patagonia.     

Bioerosion in the Miocene Reefs of the northwest Red Sea,Egypt

Macroborings provide detailed information on the bioerosion, accretion and palaeoenvironment of both modern and fossil reefs. Dolomitized reefal carbonates in the Um Mahara Formation exhibit an outstanding example of spatially distributed, well‐preserved bioerosion structures in tropical to subtropical syn‐rift Miocene reefs. Ten ichnospecies belonging to five ichnogenera are identified; three belonging to the bivalve‐boring ichnogenus Gastrochaenolites, three attributed to the sponge‐boring ichnogenus Entobia, and four ichnospecies assigned to three worm‐boring ichnogenera Trypanites, Maeandropolydora and Caulostrepsis. The distribution of the reported borings is strongly linked to the palaeo‐reef zones. Two distinctive ichnological boring assemblages are recognized. The Gastrochaenolites‐dominated assemblage reflects shallower‐marine conditions, under water depths of a few metres, mostly in back‐reef to patch‐reef zones of a back‐reef lagoon. The Entobia‐dominated assemblage signifies relatively deeper marine conditions, mostly in reef core of the fringing Miocene reefs. These ichnological assemblages are attributed herein to the Entobia sub‐ichnofacies of the Trypanites ichnofacies. This ichnofacies indicates boring in hard carbonate substrates (such as corals, rhodoliths, carbonate cements and hardgrounds) during periods of non‐sedimentation or reduced sediment input.     

Scolicia shirahamensis isp. nov.: a triple-corded scolicia and its ichnological implications

Abstract

Peculiar meniscate burrows with three sediment cords occur in early to middle Miocene tidal-flat deposits of southwestern Japan. Two of the cords are situated at the bottom and the other is at its center. Detailed observations of the burrow structures and comparative neoichnological studies of modern spatangoid burrows in a tidal flat revealed that the former two were true drainage tubes and the latter was fecal in origin. The trace fossil was thus assigned to the ichnogenus Scolicia. Based on these findings, a new ichnospecies Scolicia shirahamensis isp. nov. has been described here. The central sediment cord is seemingly identical to the drainage tube of the ichnogenus Bichordites, another ichnogenus that has been commonly ascribed to a fossil spatangoid burrow, similar to Scolicia. Careless ichnogeneric identification of a spatangoid burrow, based only on the central sediment cord, therefore, may produce an incorrect identification.     

Corimbatichnus fernandezi: A cluster of fossil bee cells from the late cretaceous‐early tertiary of Uruguay

Corimbatichnus fernandezi n.igen, n. isp. is a cluster of fossil bee cells from the Uruguayan Late Cretaceous‐early Tertiary Asencio Formation. It is composed of rows of small excavated cells contained in paleosol peds. Cells have inner surfaces somewhat rough and are closed with loose material. Separation among cells is very thin, and the cells are orientated uniformly, showing a great economy of space as in some halictine constructions. Corimbatichnus can be distinguished from Rosellichnus and Uruguay by its convex face, excavated cells and the spatial arrangement of them.     

Scorpionid Resting Trace from The Lower Permian of Southern New Mexico,USA

We describe an arthropod body impression associated with arthropod trackways of the ichnogenus Stiaria from the Lower Permian (upper Wolfcampian) Robledo Mountains Formation (Hueco Group) in the Prehistoric Trackways National Monument of southern New Mexico. The probable producer of these traces was a scorpion, and we name the likely scorpionid resting trace Alacranichnus braddyi, new ichnogenus and ichnospecies. There are no prior reports of scorpionid body impressions from terrestrial settings in the fossil record.     

New Miocene scarabeid and hymenopterous nests and early miocene (santacrucian) paleoenvironments,patagonian Argentina

Three new trace fossils are described from Miocene paleosols of southern Argentina. Celliforma pinturensis, n. ichnosp. and Celliforma rosellii, n. ichnosp. are interpreted as cells of digging bees, possibly Anthophoridae, and Coprinisphaerafrenguellii, n. ichnosp. are brood balls of dung‐beetles. Both burrowing bees and dung‐beetles are common nesters in relatively open areas, confirming previous reconstructions of the paleoenvironment of the Pinturas Formation. A brief review of scarabeid and bee fossil nests from South America is presented, and we propose that constructed nests have a higher preservation potential than excavated nests. This fact explains their more common occurrences as trace fossils in paleosols. A new ethological category, calichnia, is proposed for hymenopterous and coleopterous traces, in which adult individuals make nests exclusively for larvae.     

A New Ichnospecies of Gyrolithes from the Austin Chalk,Upper Cretaceous,Texas, USA

Gyrolithes, a helical marine trace fossil, occurs in beds of the Austin Chalk near Waco, McLennan County, Texas. The new ichnospecies Gyrolithes texanus is characterized by unique morphology and wall construction. This discovery expands the current geographic and environmental range of Gyrolithes, extending this ichnogenus into the chalk-dominated beds of the Cretaceous of Texas. Irregular bedding features from the Gyrolithes locality indicate this section of the Austin Chalk was deposited within a storm-dominated depositional regime, interspersed with periods of quiescence which allow for firm ground formation and colonization by the trace maker. Occurrences of Gyrolithes have been associated by previous publications with marginal marine settings but not associated with storm deposits; therefore, this discovery constitutes an expansion of associated environment for this ichnogenus.     

Parataenidium,a new taenidium‐like ichnogenus from the carboniferous of Ireland

The new ichnogenus Parataenidium is erected for backfilled tubular trace fossils that can appear superficially similar to Taenidium, but are divided horizontally into two distinct levels. Two ichnospecies are recognised: Parataenidium mullaghmorensis isp. nov. and Parataenidium moniliformis (Tate 1859). The latter ichnospecies is transferred from Eione Tate 1859, which is a junior homonym of Eione Rafinesque 1814, and therefore unavailable for Tate's ichnotaxon. The ichnogenus is an important component of late Paleozoic shallow‐water siliciclastic sediments, and can be considered as a “guide”; indicator for the Carboniferous.     

The New Ichnogenus Flagrichnus—A Paleoenvironmental Indicator for Cold-Water Settings?

The new marine microboring ichnogenus Flagrichnus is described from Pleistocene mollusc shell beds of Rhodes (Greece) and the Costa Brava (Spain). Recent equivalents are studied in detail from the cold-temperate setting of the Swedish Kosterfjord area. Two ichnospecies are recognized: Flagrichnus profundus consists solely of a deeply penetrating gallery with a basal swelling while Flagrichnus baiulus comprises a single to multiple sack-shaped cavity at the base of a branching filamentous gallery that is penetrating deeply into the substrate. The traces are distributed from the euphotic zone down to aphotic depths suggesting (chemo)heterotrophic microendoliths and more specifically marine fungi as the trace makers. For Flagrichnus profundus, the thraustochytrid fungus Schizochytrium is recognized as its trace maker; for Flagrichnus baiulus, no definite candidate has been identified. While Flagrichnus profundus is distributed from tropical to arctic waters, Flagrichnus baiulus is only known from fossil and Recent non-tropical settings suggesting an applicability of this ichnospecies as an indicator for low paleotemperatures.     

A Foraminiferal Parasite on the Sea Urchin Echinocorys: Ichnological Evidence from the Late Cretaceous (Lower Maastrichtian,Northern Germany)

Archaeonassa is a poorly known ichnogenus, originally described from the Cambrian of North America, which is a member of the Scolicia ’group’, but can be regarded as distinct from Scolicia. Archaeonassa was originally based on modern material, and therefore falls into a taxonomic grey zone concerning its availability, but as A. fossulata was erected on fossil material the ichnogenus is here retained. Although exhibiting a degree of morphological plasticity in its style of ornament, Archaeonassa is considered to be monospecific, with only the ichnospecies A. fossulata recognised. The ichnogenus appears to be restricted to the Paleozoic, although similar material is well documented from modern environments. Thus the ichnogenus is expected to be geographically and stratigraphically more widespread than it otherwise appears. Archaeonassa is a valuable tool for environmental reconstruction within the Paleozoic as it occurs within intertidal deposits. The ichnogenus is interpreted chiefly as the work of gastropods, although in many cases may have been produced by trilobites or even echinoids, and represents the exogenic expression of either surface or shallow subsurface locomotion.     

A New Ichnospecies of Arthrophycus from the Late Carboniferous (Pennsylvanian) of Michigan,USA

A unique ichnofossil assemblage from Pennsylvanian-age sandstones near Eaton Rapids, Michigan, USA, comprises straight-to-curved traces preserved in convex hyporelief, with the transverse ridges and median grove associated with the ichnogenus Arthrophycus. The Michigan traces show some branching or pseudo-branching (also known from other Arthrophycus specimens) and are among the smallest structures (millimeters in diameter) attributed to this ichnogenus. The orientation of the Michigan Arthrophycus burrows is distinct from other ichnospecies of this taxon in the pronounced co-planar alignment of the burrows, as opposed to the multiplanar, interweaving, “bundled” nature typical of the ichnogenus. On this basis we assign the Michigan specimens to a new ichnospecies of Arthrophycus. The “paralleling behavior” of the new taxon may reflect a strategy of the tracemaker to avoid previous burrows and reflect differences in resource availability or current energy. This new taxon supports previous records of the occurrence of this ichnogenus in Upper Carboniferous strata.     

Selection for barriers between honey bees and a devastating parasite

Rivaling pesticides and a dearth of flowers, the parasitic mite Varroa destructor presents a tremendous threat to western honey bees, Apis mellifera. A longstanding, but minor, pest for the Asian honey bee Apis cerana, these obligate bee parasites feast on developing and adult A. mellifera across several continents. Varroa reproduction is limited to a short window when developing bee pupae are concealed in wax cells. Mated females target developing bees just before pupation and then have about one day to initiate reproduction, eventually laying one male and up to several female offspring. Female mites often fail to reproduce at all, instead waiting in cells until their bee host finishes development and then hitching dangerous rides on a succession of adult bees for up to several weeks, before scouting for a new host pupa. In this issue of Molecular Ecology, Conlon et al. (2019) have explored mite reproductive success via a clever and thought‐provoking association study. In so doing, they have identified a protein whose actions could be integral to the dance between bees and their mite parasites.     

Vagorichnus,a new ichnogenus for feeding burrow systems and its occurrence as discrete and compound ichnotaxa in Jurassic lacustrine turbidites of Central China

A new ichnogenus, Vagorichnus, is figured and described from Jurassic lacustrine turbidites of the Anyao Formation, Henan Province, Central China. Vagorichnus consists of complex burrow systems composed of discontinuous curved to meandering segments in which irregular networks are formed. The ichnospecies V. anyao covers burrow systems characterized by constricted and irregularly annulated segments, and ridge‐like knobs. Although traces are preserved as hypichnial ridges on the sole of turbidite sandstones, several lines of evidence indicate that these ichnofossils record the activity of a post‐event infauna. Vagorichnus occurs as discrete and compound ichnotaxa. In the latter case, it grades into Gordia and Tuberculichnus. Vagorichnus is interpreted as invertebrate feeding structures (Fodinichnia).     

The earliest evidence of holometabolan insect pupation in conifer wood

Background

The pre-Jurassic record of terrestrial wood borings is poorly resolved, despite body fossil evidence of insect diversification among xylophilic clades starting in the late Paleozoic. Detailed analysis of borings in petrified wood provides direct evidence of wood utilization by invertebrate animals, which typically comprises feeding behaviors.

Methodology/Principal Findings

We describe a U-shaped boring in petrified wood from the Late Triassic Chinle Formation of southern Utah that demonstrates a strong linkage between insect ontogeny and conifer wood resources. Xylokrypta durossi new ichnogenus and ichnospecies is a large excavation in wood that is backfilled with partially digested xylem, creating a secluded chamber. The tracemaker exited the chamber by way of a small vertical shaft. This sequence of behaviors is most consistent with the entrance of a larva followed by pupal quiescence and adult emergence — hallmarks of holometabolous insect ontogeny. Among the known body fossil record of Triassic insects, cupedid beetles (Coleoptera: Archostemata) are deemed the most plausible tracemakers of Xylokrypta, based on their body size and modern xylobiotic lifestyle.

Conclusions/Significance

This oldest record of pupation in fossil wood provides an alternative interpretation to borings once regarded as evidence for Triassic bees. Instead Xylokrypta suggests that early archostematan beetles were leaders in exploiting wood substrates well before modern clades of xylophages arose in the late Mesozoic.     

Taxonomic reassessment ofBolonia Meunier, 1886 (trace fossil) based on new material from the type area in Boulonnais, northern France

The ichnospeciesBolonia lata Meunier, 1886 is a straight, bilobate trace fossil with a somewhat heart-shaped outline in cross-section. It has previously been reported from various localities mainly from the Jurassic by different authors under various names such asGyrochorte, Scolicia, orProtovirgularia. Because none of these previous determinations fully agrees with the trace fossils’ morphology, the ‘forgotten name’ is reintroduced, and due to a lack of type material a neotype from the type area of Boulonnais, France is proposed. The ichnogenus is currently represented only by one ichnospecies,B. lata Meunier, 1886. Diagnostic criteria ofBolonia and morphologically related trace fossils such asScolicia, Gyrochorte, Bichordites, “Taphrhelminthopsis nelsoni”, andProtovirgularia are discussed, the synonymy ofB. lata is provided and finally a discussion on the possible producer ofBolonia is given.      

Micro‐ and nano‐scale ultrastructure of cell walls in Cryogenian microfossils: revealing their biological affinity

Moczyd?owska, M., Schopf, J.W. & Willman, S. 2009: Micro‐ and nano‐scale ultrastructure of cell walls in Cryogenian microfossils: revealing their biological affinity. Lethaia, Vol. 43, pp. 129–136. Recently established protocols and methods in advanced microscopy and spectrometry applied to studies of ancient unicellular organic‐walled microfossils of uncertain biological affinities (acritarchs) provide new evidence of the fine ultrastructure of cell walls and their biochemistry that support the interpretation of some such microfossils as photosynthesizing microalgae. The micro‐scale and nanoscale ultrastructure of the cell walls of late Cryogenian sphaeromorphic acritarchs from the Chichkan Formation (Kazakhstan) revealed by the advanced techniques and studied originally by Kempe et al. (2005) is here further analysed and compared with that of modern microalgal analogues. On the basis of such comparison, we interpret the preserved cell wall ultrastructure to reflect original layering and lamination within sub‐layers of the fossil wall, rather than being a result of taphonomic and diagenetic alteration. The outer thick layer represents the primary wall and the inner layer the secondary wall of the cell, whereas the laminated amorphous sub‐layers, 10–20 nm in thickness and revealed by transmission electron and atomic force microscopy, are recognized as trilaminar sheath structure. Because two‐layered cell walls, trilaminar sheaths and the position of the TLS within the fossil cell wall are characteristic of the mature developmental state in cyst morphogenesis in modern microalgae, we infer that the Chichkan sphaeromorphs are probably resting cells (aplanospores) of chlorophyceaen green microalgae from the order Volvocales. □Biological affinity, cell wall, Cryogenian, microfossils, ultrastructure.     

Ichnologic note

Palaxius salataensis, an anomuran microcoprolite, is described from Cretaceous deposits of the MIT Guyot in the northwest Pacific, about 1500 km southeast of Japan. This is the first report of the ichnogenus in this area and the first report of the ichnospecies in Cretaceous deposits. The range of P. salataensis is thus very long, from Permian to Cretaceous.