Bigeneric nomina: An historical and evolutionary perspective

It is suggested that the addition of bigeneric or bispecific nomina to designate fossils which are intermediate between two chronogenera or chronospecies is inadvisable, because the use of either of these categories to describe cases involving a single phyletic line, consisting of a continuous chain of time-sequential populations, misrepresents the nature of the evolutionary processes involved; and that in such cases fossils might best be referred to by specimen numbers.     

Genomic signatures of evolution in Nautilus—An endangered living fossil

Living fossils are survivors of previously more diverse lineages that originated millions of years ago and persisted with little morphological change. Therefore, living fossils are model organisms to study both long‐term and ongoing adaptation and speciation processes. However, many aspects of living fossil evolution and their persistence in the modern world remain unclear. Here, we investigate three major aspects of the evolutionary history of living fossils: cryptic speciation, population genetics and effective population sizes, using members of the genera Nautilus and Allonautilus as classic examples of true living fossils. For this, we analysed genomewide ddRAD‐Seq data for all six currently recognized nautiloid species throughout their distribution range. Our analyses identified three major allopatric Nautilus clades: a South Pacific clade, subdivided into three subclades with no signs of admixture between them; a Coral Sea clade, consisting of two genetically distinct populations with significant admixture; and a widespread Indo‐Pacific clade, devoid of significant genetic substructure. Within these major clades, we detected five Nautilus groups, which likely correspond to five distinct species. With the exception of Nautilus macromphalus, all previously described species are at odds with genomewide data, testifying to the prevalence of cryptic species among living fossils. Detailed F_ST analyses further revealed significant genome‐wide and locus‐specific signatures of selection between species and differentiated populations, which is demonstrated here for the first time in a living fossil. Finally, approximate Bayesian computation (ABC) simulations suggest large effective population sizes, which may explain the low levels of population differentiation commonly observed in living fossils.     

Stable carbon isotopes and the metabolism of the terrestrial Devonian organism Spongiophyton

Devonian fossils of Spongiophyton have been identified as a terrestrial evolutionary intermediate between algae and vascular land plants on the basis of their dichotomously branched, tubular morphology, thick cuticles, and the scattered distribution of surface pores. Our understanding of their physiology is, however, severely limited, but may be increased through the use of stable carbon isotope measurements. One such study led to the hypothesis that Spongiophyton carried a carbon isotope (δ¹³C) signature characteristic of lichens ( Jahren et al., 2003 Geology 31 , 99–102). Here, we outline three difficulties with accepting this idea and report independent isotopic measurements of Spongiophyton fossils from Canada and Ghana. Our results show that the isotopic discrimination of analogous tissues of Spongiophyton, extant lichens, liverworts and mosses are statistically indistinguishable. We suggest therefore that claims to have definitively identified lichen metabolism are premature, and cannot be sustained.     

EXTINCT TRANSITIONAL FAGACEAE FROM THE OLIGOCENE AND THEIR PHYLOGENETIC IMPLICATIONS

Fruits, catkins, and associated leaves of at least two extinct trigonobalanoid taxa have been discovered at an Oligocene fossil plant locality rich in fagaceous remains. These fossils exhibit a mosaic of fruit and pollen characters found in the two extant subfamilies Castaneoideae and Fagoideae of Fagaceae. Comparison with cladograms based on modern taxa suggests that these extinct taxa were similar to the ancestors of subfamily Fagoideae and may have been intermediate between Fagus and the modern trigonobalanoid genera. Pollen types isolated from the fossil staminate catkins provide unique character states that are transitional between modern pollen types in Fagaceae and are important in understanding the evolution of exine micromorphology within the family. This analysis provides a striking example of the use of character data from fossils to determine character-state adjacency prior to polarization of characters using outgroup comparison. Because of the mosaic nature of their character complexes, these fossils support monophyly in both the family Fagaceae and the subfamily Fagoideae. In addition, the occurrence of trigonobalanoid fossils in the Oligocene of North America has interesting biogeographic implications and provides insights into the nature of North American Fagaceae during the Tertiary.     

Morphometric variation in Plio-Pleistocene hominid distal humeri

The magnitude and meaning of morphological variation among Plio-Pleistocene hominid distal humeri have been recurrent points of disagreement among paleoanthropologists. Some researchers have found noteworthy differences among fossil humeri that they believe merit taxonomic separation, while others question the possiblity of accurately sorting these fossils into different species and/or functional groups. Size and shape differences among fossil distal humeri are evaluated here to determine whether the magnitude and patterns of these differences can be observed within large-bodied, living hominoids. Specimens analyzed in this study have been assigned to various taxa (Australopithecus afarensis, A. africanus, A. anamensis, Paranthropus, and early Homo) and include AL 288-1m, AL 288-1s, AL 137-48a, AL 322-1, Gomboré IB 7594, TM 1517, KNM-ER 739, KNM-ER 1504, KMN-KP 271 (Kanapoi), and Stw 431. Five extant hominoid populations are sampled to provide a standard by which to consider differences found between the fossils, including two modern human groups (Native American and African American), one group of Pan troglodytes, and two subspecies of Gorilla gorilla (G. g. beringei, G. g. gorilla). All possible pairwise d values (average Euclidean distances) are calculated within each of the reference populations using an exact randomization procedure. This is done using both raw linear measurements as well as scale-free shape data created as ratios of each measurement to the geometric mean. Differences between each pair of fossil humeri are evaluated by comparing their d values to the distribution of d values found within each of the reference populations. Principal coordinate analysis and an unweighted pair group method with arithmetic averages (UPGMA) cluster analysis are utilized to further assess similarities and differences among the fossils. Finally, canonical variates analysis and discriminant analysis are employed using all hominoid samples in order to control for correlations among variables and to identify those variables that discriminate among groups; possible affinities of individual fossils with specific extant species are also examined. The largest size differences, those between the small Hadar specimens and the two largest fossils (KNM-ER 739, IB 7594), can be accommodated easily within the ranges of variation of the two Gorilla samples, but are extreme relative to the other reference samples. The d values between most of the fossils based on shape data, with the notable exception of those associated with KNM-ER 739 and KNM-ER 1504, can be sampled safely within all five reference samples. Subsequent analyses further support the inference that KNM-ER 739 and KNM-ER 1504 are different from the other hominid humeri and possess a unique total morphometric pattern. In overall shape, the distal humeri of the other fossils (non-Koobi Fora) are most similar to living chimpanzees. The distal humerus of Paranthropus from Kromdraai (TM 1517e) is most similar to one of the Hadar specimens of A. afarensis (AL 137-48a), whereas the first specimen of A. africanus from Sterkfontein (Stw 431) is not closely linked to any of the other australopithecines. The A. anamensis humerus from Kanapoi exhibits no special affinities to A. afarensis or to modern humans. © 1996 Wiley-Liss, Inc.     

Insights on the Behavior of Late Paleozoic Aquatic Crustaceans (Pygocephalomorpha?): Compound Trace Fossils from Western Argentina

We describe intergradations between the arthropod repichnial trace fossils Diplichnites gouldi (Gevers et al., 1971), Dendroidichnites Demathieu et al., 1992, and Monomorphichnus Crimes, 1970 with two distinct cubichnial traces, Gluckstadtella cooperi Savage, 1971 and Huilmuichnus santracruzensis new ichnogenus and ichnospecies. These compound ichnofossils are part of high ichnodiversity arthropod-dominated trace fossil associations from the Late Carboniferous Agua Escondida Formation of the San Rafael Basin (Argentina). The complex ichnotaxonomy of Diplichnites and the ichnospecies D. gouldi are discussed and some possible working solutions are proposed. The most likely producer of the compound trace fossils are the Pygocephalomorpha, a group of Late Paleozoic crustaceans that are recorded in nearby basins from Uruguay and Brazil. The compound trace fossils are used to infer a number of complex behaviors of pygocephalomorphs in a subaqueous setting, including caridoid (tail flip) or more primitive escape reactions, landing followed by resting and transient resting during normal walking. In addition, different gaits can be inferred from well-preserved D. gouldi, which can be correlated with contrasting trackway morphology.     

Scolicia and its producer in shallow-marine deposits of the Miocene Chenque Formation (Patagonia,Argentina): functional morphology and implications for understanding burrowing behavior

Abstract

Scolicia is one of the most conspicuous trace fossils in lower shoreface deposits of the Miocene Chenque Formation of Patagonia, Argentina. This ichnotaxon consists of horizontal, sinuous or meandering trace fossils with a laminated backfill and two parallel strings located at the base. Abundant body fossils attributed to Brisaster iheringi occur in close association to these trace fossils. The echinoids are very well preserved, and most of the specimens have some areas with their spines attached in life position. In particular, preservation of the subanal tufts of spines, associated to isopores (which are interpreted as being associated to funnel building tube feet), supports the interpretation that these organisms are the producers of Scolicia isp. A shaft connecting the burrow with the sediment-water interface was not observed in vertical sections of the excavations, although on bedding plane surfaces, some small circular tube outlets are recognized. However, the great diversity and abundance of ichnofossils co-occurring in these deposits may preclude adequate identification of vertical structures. These deposits provide an excellent opportunity to integrate trace fossil data, body fossil information and observations from modern analogues in order to perform a morpho-functional analysis of Scolicia.     

An aquatic microfossil assemblage from Cenomanian amber of France

The ‘osseous’ amber from the Cenomanian of northwestern France contains numerous microscopic inclusions, some of which are fairly well preserved and identifiable as protists. This paper describes three cyanobacteria similar to modern Plectonema. Lyngbya, and Coelosphaeriunr, fungus-like fossils of uncertain affinities (cf. ?Candida); a colorless chrysomonad similar to Monas; a desmid identical with Closterium; and naked ciliates of uncertain affinities (cf. Cyrtolophosis). All of these fossils are in a single sample of amber from Bretagnolles (Eure Département). This assemblage is comparable to modern limnetic microbial communities. It is typical of shallow freshwater environments in which productivity and respiration are both high. This interpretation fits paleoecological reconstructions drawn from the arthropod fossils from French amber. ***Chrysomonads, ciliates, Cretaceous, cyanobacteria, desmids, fungi, micropaleontology.     

Middle Jurassic fossils of the genus Sharasargus from Inner Mongolia, China (Diptera: Archisargidae)

To date, only two species of the genus Sharasargus are known. These Upper Jurassic species were found in Mongolia and Kazakhstan, respectively. We herein describe two new species from the Middle Jurassic, which were found in Inner Mongolia, China. They are the oldest known fossils of this genus. A key to Sharasargus species is given.     

The female reproductive unit of Ephedra (Gnetales): comparative morphology and evolutionary perspectives

Morphological variation in Ephedra (Gnetales) is limited and confusing from an evolutionary perspective, with parallelisms and intraspecific variation. However, recent analyses of molecular data provide a phylogenetic framework for investigations of morphological traits, albeit with few informative characters in the investigated gene regions. We document morphological, anatomical and histological variation patterns in the female reproductive unit and test the hypothesis that some Early Cretaceous fossils, which share synapomorphies with Ephedra, are members of the extant clade. Results indicate that some morphological features are evolutionarily informative although intraspecific variation is evident. Histology and anatomy of cone bracts and seed envelopes show clade‐specific variation patterns. There is little evidence for an inclusion of the Cretaceous fossils in the extant clade. Rather, a hypothesized general pattern of reduction of the vasculature in the ephedran seed envelope, probably from four vascular bundles in the fossils, to ancestrally three in the living clade, and later to two, is consistent with phylogenetic and temporal analyses, which indicate that extant diversity evolved after the Cretaceous–Tertiary boundary. Notwithstanding striking similarities between living and Cretaceous Ephedra, available data indicate that the Mesozoic diversity went almost entirely extinct in the late Cretaceous causing a bottleneck effect in Ephedra, still reflected today by an extraordinarily low level of genetic and structural diversity. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 387–430.     

Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca,Spain

Gibert, J.M. de, Mas, G. & Ekdale, A.A. 2012: Architectural complexity of marine crustacean burrows: unusual helical trace fossils from the Miocene of Mallorca, Spain. Lethaia, Vol. 45, pp. 574–585. Unusual helical trace fossils occur in Tortonian shelfal calcarenites in the island of Mallorca. Their morphology may be either simple (ichnogenus Gyrolithes) or, more commonly, consist of two concentric helical burrows (ichnogenus Lapispira). They bear a very characteristic pelleted lining and are associated and probably connected to Thalassinoides and Ophiomorpha burrow systems very abundant in the same unit. These features allow the interpretation that the tracemaker was a thalassinidean shrimp. The complex and compound nature of these trace fossils is comparable to that seen in other modern and fossil crustacean burrow systems, and it reflects the behavioural plasticity of the architects. □Trace fossils, ichnology, ichnofabrics, crustacean burrows, Miocene, shallow marine, Lapispira.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

Ladinian carbonates of the Cabo Cope Unit (Betic Cordillera,SE Spain): a tethys-maláguide palaeogeographic gateway

Summary The Cabo Cope Unit, which outcrops east of Aguilas (Murcia), belongs to the Maláguide tectonic Complex (Betic Internal Zone) and displays stratigraphic characteristics of particular interest, including Triassic bioclastic carbonate beds which are not common in the Maláguide units. Biostratigrafic fossils have been found in these beds and may correlate with Triassic alpine biofacies. Alpine fauna fossils only appeared in those palaeogeographic units of the Internal Zone of the Cordillera referred to as Alpujárride units, while the influence of the Sephardic faunal province is evident in almost all the cordillera. For these reasons it is noteworthy that new alpine fauna fossils have been found in an Internal Zone unit in which relevant fossils rarely appear. The Triassic succession of the unit studied in this paper can be subdivided into two members: a lower one, which is clastic and contains thick gypsum beds, and an upper one, consisting of carbonate rocks. The lower member has been interpreted as a fluvial-coastal deposit. The upper member is interpreted as a sequence of carbonate ramp deposits. This ramp evolved into a shallow platform with tidal flats typical of a coastal zone. The bivalve fossilsDaonella cf.lommeli (Wissmann) and “Posidonia” sp. have been found in the carbonate member, along with the conodontSephardiella mungoensis (Diebel). These fossils are of the Late Ladinian age and have been found only in this outcroup of the Betic Cordillera. The presence of this fossil assemblage, which belongs to the alpine faunal province, indicates a connection during the Late Ladinian between the Tethys sea and this area of the Maláguide palaeogeographic domain. The palaeogeographic location of the Cabo Cope Unit during the Middle Triassic was at the south-easternmost part of the Betic Basin, implying that the connection between the Tethys and the Betic Basin was established in the easternmost domains of the basin.     

Natural assemblages of Hindeodus conodonts from a Permian–Triassic boundary sequence,Japan

Hindeodus parvus and Hindeodus typicalis occur in a deep‐water chert and claystone section in the Mino Terrane, Japan, which has been identified as a Jurassic accretionary complex. Conodont fossils are preserved as natural assemblages of impression fossils on bedding planes in claystone. In this study, 13 assemblages of Hindeodus species were recognized, comprising at most 13 elements which generally maintain the original composition and structure of an apparatus. We discriminated pairs of carminiscaphate P₁, angulate P₂ and makellate M elements, as well as a single alate S₀ element and two digyrate and four bipennate elements constituting the S array. Although the digyrate and bipennate elements are preserved in the S₂ and S_3–4 positions, respectively, a pair of S₁ elements was not found due to incompleteness in the natural assemblages. The conodont biostratigraphy indicates that the lithological boundary between chert and claystone units in the study section corresponds exactly to the Permian–Triassic boundary.     

New fossil Pachygastrinae from Mexican amber (Diptera: Stratiomyidae)

We describe the new genus Elektrothopomyia n. gen., with two new species E. saltensis n. sp. and E. tzotzili n. sp. as second and third fossil Pachygastrinae from the Mexican amber. These fossils from the Totolapa amber deposit confirm the presence of this group in Central America during the late Oligocene to middle Miocene period. Their exact affinities remain uncertain because of the current problems in the phylogenetic classification of these flies. These fossils are new elements of comparison between the entomofaunas of the Dominican and Mexican amber.     

Middle Jurassic trace fossils from the Ridang Formation in Sajia County,South Tibet,and their palaeoenvironmental significance

An ichnofauna consisting of 18 ichnospecies (one of them new) in 14 ichnogenera are described for the first time from the Middle Jurassic Ridang Formation in Sajia County, South Tibet. The ichnofauna can be subdivided into two ichnoassemblages in ascending stratigraphic order: the Palaeophycus–Megagrapton ichnoassemblage in the lower and middle parts of the Ridang Formation, followed by the Cosmorhaphe–Nereites–Paleodictyon ichnoassemblage in the upper Ridang Formation. Overall, the trace fossils occur in a middle–distal turbidite fan sequence, as evidenced by both sedimentological analysis and the composition of the trace fossils. Several subenvironments of the middle–distal fan system have been recognized on the basis of the spatial distribution of the trace fossils. Typically, the channel-fill deposits in the middle part of the turbidite fan lack trace fossils, the interchannel and upper channel-fill (levee) subenvironments of the middle turbidite fan contain abundant facies-crossing trace fossils, in contrast to the distal part of the turbidite fan where deep-water trace fossils are dominant. The ichnofauna is similar to typical flysch ichnofaunas from Europe and North America in characteristics, and is interpreted to represent a typical deep-sea Nereites ichnofacies. The presence of these deep-sea trace fossils therefore would suggest that a continental slope environment existed in southern Tibet during the Middle Jurassic and the study area was located in a slope-abyssal plain setting.     

How big were early hominids?

The recent discovery of new postcranial fossils, particularly associated body parts, of several Plio-Pleistocene hominids provides a new opportunity to assess body size in human evolution.¹ Body size plays a central role in the biology of animals because of its relationship to brain size, feeding behavior, habitat preference, social behavior, and much more. Unfortunately, the prediction of body weight from fossils is inherently inaccurate because skeletal size does not reflect body size exactly and because the fossils are from species having body proportions for which there are no analogues among modern species. The approach here is to find the relationship between body size and skeletal size in ape and human specimens of known body weight at death and to apply this knowledge to the hominid fossils, using a variety of statistical methods, knowledge of the associated partial skeletons of the of early hominids, formulae derived from a modern human sample, and, finally, common sense. The following modal weights for males and females emerge: Australopithecus afarensis, 45 and 29 kg; A. africanus, 41 and 30 kg; A. robustus, 40 and 32 kg; A. boisei, 49 and 34 kg; H. habilis, 52 and 32 kg. The best known African early H. erectus were much larger with weights ranging from 55 kg on up. These estimates imply that (1) in the earliest hominid species and the “robust” australopithecines body sizes remained small relative to modern standards, but between 2.0 and 1.7 m.y.a. there was a rapid increase to essentially modern body size with the appearance of Homo erectus; (2) the earliest species had a degree of body size sexual dimorphism well above that seen in modern humans but below that seen in modern gorillas and orangs which implies (along with other evidence) a social organization characterized by kin-related, multi-male groups with females who were not kin-related; (3) relative brain sizes increased through time; (4) there were two divergent trends in relative cheek-tooth size—a steady increase through time from A. afarensis to A. africanus to the “robust” australopithecines, and a decrease beginning with H. habilis to H. erectus to H. sapiens.     

Silurian trace fossils in carbonate turbidites from the Alexander arc of southeastern Alaska

Early to Late Silurian (Llandovery‐Ludlow) body and trace fossils from the Heceta Formation of southeastern Alaska are preserved in the oldest widespread carbonates in the Alexander terrane. These fossils represent the earliest benthos to inhabit diverse shallow and deep subtidal environments in the region and are important indicators of early stages in benthic community development within the evolving Alexander arc. The ichnofossils are particularly significant because they add to a small but growing body of knowledge about trace fossils in deep‐water carbonates of Paleozoic age.

Carbonate turbidites that originated along a deep marine slope within the arc yield a low‐diversity suite of trace fossils consisting of five distinct biogenic forms. Simple burrows (Planolites, two forms), ramifying tunnels (Chondrites), and tiny cylindrical burrows (?Chondrites) represent the feeding activities (fodinichnia) of pre‐turbidite animals that burrowed in the lime mud before the influx of coarser sediment deposited by turbidity currents. These trace fossils are associated locally with cross‐cutting burrows created as domichnia (Palaeophycus). Rarer hypichnial burrows and endichnial traces were created by post‐turbidite animals that fed soon after the deposition of coarse detritus from turbidity flows.

Trace fossils in these deposits reflect much lower diversity levels than in Paleozoic siliciclastic turbidites. This difference may represent unfavorable environmental conditions for infaunas, differential preservation, or significant paleogeographic isolation of the Alexander terrane during the Silurian. Greater utilization of trace fossils in terrane analysis may help to resolve this issue and provide new data for reconstructing the paleogeography of circum‐Pacific terranes.     

Evidence for targeted elasmobranch predation on thalassinidean shrimp in the Miocene Taliao Formation,NE Taiwan

Terrestrial and marine invertebrate organisms both leave records of their activities in the sediment in the form of trace fossils, at least during certain stages of their ontogeny. In contrast, trace fossils produced by vertebrate organisms are scarce, although terrestrial trace fossils provide exclusive insights into the social behaviour of their producers. In the marine realm, vertebrate trace fossils are relatively rare, difficult to identify and problematic to interpret. However, in certain settings, observations on serendipitously preserved and exposed trace fossils can shed light on the predatory behaviour of marine vertebrates. In Miocene outer shelf to nearshore sandstones of the Taliao Formation in NE Taiwan, large numbers of bowl‐shaped trace fossils can be observed. Morphology and size range (diameter typically 10–30 cm, average depth around 10 cm) of these trace fossils agree well with feeding traces of modern stingrays, and the trace fossil Piscichnus waitemata, which has been attributed to bottom feeding rays. Stingrays direct a jet of water from their mouths to excavate a bowl‐shaped pit to expose their prey. In the material filling the excavated bowl, broken pieces of two other common trace fossils, Ophiomorpha and Schaubcylindrichnus, are often found, and in a number of cases, vertical shafts of Ophiomorpha surrounded by dispersed pieces of wall material have been observed. In contrast, surrounding sediment rarely contains this kind of broken pieces of wall material. These observations clearly indicate that stingrays specifically targeted the producers of the trace fossils: thalassinoid crustaceans and worms, respectively. The targeted predation of these relatively deep burrowers furthermore suggests that the rays used their electroreceptive organs to locate the prey; as such, direct targeting of buried prey only based on olfactory senses has been shown to be ineffective in experiments with extant myliobatiform rays.     

Late-Pleistocene Anthelia (Hepaticae), an arctic-alpine,snow-bed indicator at a low elevation site in Massachusetts,U.S.A.

Abstract

A fossil of Anthelia from late-Pleistocene sediments at Tom Swamp, Massachusetts, consisted of a portion of a plant bearing the characteristic three rows of isomorphic, deeply bifid leaves. The fossil Anthelia indicates the existence of areas of late-lying snow in an essentially treeless, late-Pleistocene landscape. Associated bryophyte (mainly moss) and tracheophyte fossils establish the presence of additional hygric and mesic habitat types.

The occurrence of Anthelia and other leafy liverwort fossils in the basal inorganic sediments at Tom Swamp is unusual. Features of these fossils suggest that the translucent quality of some of the fragments and their small size (1 mm or smaller), which relates to fragmentation during transport, diagenesis and extraction (the breakage resulting from a weak middle lamella between the cells), may be reasons for the rarity of Pleistocene and Holocene fossils of liverworts.

Published as contribution number 589 of the New York State Science Service.