How diverse were ferns in the Baltic amber forest?

Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopteris” humboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopteris” humboldtiana in the new genus Berendtiopteris. “Alethopteris” serrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.     

Effects of alternate reef states on coral reef fish habitat associations

The present study describes ontogenetic shifts in habitat use for 15 species of coral reef fish at Rangiroa Atoll, French Polynesia. The distribution of fish in different habitats at three ontogenetic stages (new settler, juvenile, and adult) was investigated in coral-dominated and algal-dominated sites at two reefs (fringing reef and inner reef of motu). Three main ontogenetic patterns in habitat use were identified: (1) species that did not change habitats between new settler and juvenile life stages (60% of species) or between juvenile and adult stages (55% of species—no ontogenetic shift); (2) species that changed habitats at different ontogenetic stages (for the transition “new settler to juvenile stage”: 15% of species; for the transition “juvenile to adult stage”: 20% of species); and (3) species that increased the number of habitats they used over ontogeny (for the transition “new settler to juvenile stage”: 25% of species; for the transition “juvenile to adult stage”: 25% of species). Moreover, the majority of studied species (53%) showed a spatial variability in their ontogenetic pattern of habitat use according to alternate reef states (coral reef vs algal reef), suggesting that reef state can influence the dynamics of habitat associations in coral reef fish.     

Further observations on a method for estimating hominoid dental developmental patterns

Several recent studies have concluded that early hominines resembled apes in patterns of dental ontogeny and have inferred from this conclusion that they were ape-like in other aspects of growth and development as well. Prior to their employment on fossil junveniles, the comparative methods used in these studies were not first tested for their predicitive accuracy in distinguishing modern humans from apes on the basis of proposed patterns of dental ontogeny. We find that 92% of a smaple of 48 modern children are classified as ape-like or nonhuman by the criteria employed in one of these studies and overlap entirely with fossil juvenile dental patterns. The use of these methods to characterize early hominines as “ape-like” or distinctly divergent from humans is thus shown to be unwarranted. © 1993 Wiley-Liss, Inc.     

Impact of increased character sampling on the phylogeny of Cetartiodactyla (Mammalia): combined analysis including fossils

The phylogenetic position of Cetacea (whales, dolphins and porpoises) is an important exemplar problem for combined data parsimony analyses because the clade is ancient and includes many well‐known and relatively complete fossil species. We combined data for 71 terminal taxa (43 extinct/28 extant) to test where Cetacea fits within Cetartiodactyla, and where various fossil hoofed mammals (e.g., ?entelodonts, “?anthracotheriids” and ?mesonychians) are positioned. We scored 635 phenotypic characters (osteology, dentition, soft tissue, behavior), approximately three times the number of characters in the last major analysis of this clade, and combined these with > 40 000 molecular characters, including new data from 10 genes. The analysis supported a topology consistent with the majority of recently published molecular studies. Cetacea was the extant sister taxon of Hippopotamidae, followed successively by Ruminantia, Suina and Camelidae. Several extinct taxa were phylogenetically unstable, upsetting resolution of the strict consensus and limiting branch support, but the positions of several key fossils were consistently resolved. The wholly extinct ?Mesonychia was more closely related to Cetacea than was any “artiodactylan.”“?Anthracotheriids” were paraphyletic, and, with the exception of one species, were more closely related to Hippopotamidae than to any other living taxon. The total evidence analysis overturned a highly nested position for Moschus supported by molecular data alone. The character partition that could be scored for the fossil taxa (osteological and dental characters) included more informative characters than most molecular partitions in our analysis, and had the fewest missing data. The osteological–dental data alone, however, did not support inclusion of cetaceans within crown “Artiodactyla.” Recently discovered ankle bones from fossil whales reinforced the monophyly of Cetartiodactyla but provided no particular evidence of derived similarities between hippopotamids and fossil cetaceans that were not shared with other “artiodactylans”. © The Willi Hennig Society 2007.     

Ontogeny and phylogeny in temnospondyls: a new method of analysis

A new method has been devised to compare the ontogeny and phylogeny of some of the better documented tetrapods from the Stephanian to the Trias. This approach is based on global parsimony analysis of several temnospondyl amphibians, in which some ontogenetic sequences have been highlighted. Forty-one homologous morphoanatomical character states have been separated into larval, juvenile and adult stages of each of six tetrapod species. The taxonomic congruence (TC) approach involves comparing trees based on larval, juvenile and adult character states. These so-called 'ontotrees' are not congruent, either in their topologies or in the distribution of the character states. The total evidence (TE) approach involves a combined analysis of all the character states observed in the various growth stages of the taxa, and is secondarily used in order to test this taxonomic incongruence. In this case, the TE result corroborates the TC analysis: the TE tree is robust and reveals a few homoplasies which cause the taxonomic incongruence. This is interpreted as either the result of heterochronic events in temnospondyl evolution, or as the product of inaccurate identification of larval and metamorphic fossil forms.     

New bethylid and chrysidid wasps (Hymenoptera: Chrysidoidea) from Canadian Late Cretaceous amber

A new bethylid species, Celonophamia granama, and two new chrysidid species, Procleptes eoliami, and P. hopejohnsonae, are described from Late Cretaceous (Campanian) amber collected at the Grassy Lake locality in Alberta, Canada. Within the deposit these taxa constitute the first bethylid, and the second and third chrysidid species to be described, respectively. The new taxa expand the sparse fossil record of Chrysidoidea, particularly that of Chrysididae—a group that was previously represented by only three described species in the Mesozoic. The presence of Celonophamia species in both Canadian amber and Siberian (Taimyr) amber further emphasizes faunal similarities between these two northern Late Cretaceous amber deposits. Given the prevalence of metallic coloration in Chrysididae, the specimens described here also provide evidence for the taphonomic alteration of perceived insect colors in Cretaceous amber inclusions.     

Species concepts and the recognition of ancestors

Whether or not ancestral species can be recognised depends on the species concept adopted. A “metaspecies”; is a species that completely lacks autapomorphies, and which might (or might not) be ancestral to other species. Such taxa have been identified among both living and fossil organisms. Under the most commonly‐used species concepts (biological, evolutionary, phenetic, phylogenetic, ecological, recognition and cohesion), “metaspecies”; can be assumed to be ancestral. Even if the known members of a metaspecies are not ancestral to anything, parsimony dictates that the (as yet unknown) ancestral lineage is identical to the metaspecies and, under these species concepts, assignable to the same species. Only the cladistic and monophyletic species concepts would deny “metaspecies”; ancestral status, but these species concepts are problematical and have never been used by practising systematists.     

An ant-associated mesostigmatid mite in Baltic amber

Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first—and so far only—fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.     

Early Oligocene Callitris and Fitzroya (Cupressaceae) from Tasmania

This paper documents Early Oligocene fossilized foliage and ovulate cones from Lea River, Tasmania and identifies them as belonging to two extant southern hemisphere Cupressaceae genera, Callitris and Fitzroya. Most importantly, it sheds some light on evolutionary trends within Callitris, a genus with numerous extant Australian species and two extant New Caledonian species. Callitris has a very poor fossil record and, because of the present absence of a molecular data set that includes all species, its phylogeny remains somewhat ambiguous. Although Fitzroya foliage has previously been described from a number of Tasmanian sites, this is the first recording of fertile material. The ovulate cones of the Callitris and Fitzroya macrofossils are characterized by bract-scale complexes in two whorls of three and are subtended by scale-like leaves in whorls of three. The fossilized foliage specimens consist of scale-like leaves in whorls of three. These morphological characteristics are only exhibited by three extant southern hemisphere cupressaceous genera, Callitris, Actinostrobus, and Fitzroya. The assignment of the fossils to extinct Callitris and Fitzroya species is made by comparisons with species from these three extant genera. Although much of the Lea River flora are wet rainforest taxa, the Callitris fossils have characteristics of both wet- and dry-adapted extant species.     

On Occurrence and Significance about the Fossil Plants from the Yuntaiguan Formation Located between Hunan and Hubei

This paper deals with the fossil plants from the Yuntaiguan formation located between Hunan and Hubei province. There are six genera and ten species (containing a new genus and five new species) to be discussed here, such as Protolepidodendron scha-ryanum Krejci, Barrandeina dusliana (Krejci) Stur, B. contigua sp. nov., B. lixianensis sp. nov., B. multistriata sp. nov., B. laxa sp. nov., Drepanophycus spinaeformis Goepp., “Protopteridium”? minutum Halle, Psilophyti4es sp. and Longostachys latisporophyllus gen. et sp. nov., etc. The appearance of this fossil flora concerned here can be correlated to the Protolepidodendron scharyanum-Barrandeina dusliana Assemblage. Among these fossil plants from Yuntaiguan formation, both the Protolepidodendron scharyanum and Barrandeina dusliana are also the index fossils, the prior was wide spread during the Middle Devonian in whole world and the other was only found at the late Middle Devonian in Europe. In the Yuntaiguan formation the genus Barrandeina containing no less than five species and, the other taxa were also found from the Middle or the late Middle Devonian. So that this formation should be referred to the late Middle Devonian in age and this conclusion corresponds with the result of spore-pollen analysis in the Xiushuigou formation, Xiongzi county, which is similar to the Yuntaiguan formation and has been referred to the Middle Devonian. In this paper, only four species of Barrandeina and one species of Longostachys have been described, those of the former have not been found before in China. The authors think; this will be significant on studying fossil plants and in comparison with the age of the strata.     

The ontogenetical and supposed phylogenetical fate of the parietal muscles in the Ctenostomata (Bryozoa)

A new term, “trophon”, has been introduced for the feeding and sexually propagating unit in the stolonate ctenostomes; the “stolon” and the “trophon” represent “sukzooids”, because they are homologous only to parts of the autozooids in the uniserially arranged ctenostomes. – The ontogeny of the muscles, especially the parietal muscles, has been studied in 17 species of ctenostomatous bryozoans belonging to different sub-groups of this order; the main effort was directed toward the details of the differentiations in living animals (14 of the investigated species). In various species of the Ctenostomata two separate sets of parietal muscles develop – probably as a consequence of the elongation of the cystid; the functional parietal muscles in the trophons of the stolonial forms are homologous to the secondary parietal muscles found in several uniserial ctenostomes. Contrary to the supposition by Soule (1954) in the different families of the serially arranged forms the muscles appear in different sequences during the development of the zooid and partly have different ontogenetical fates. Neither the “Carnosa” nor the “Paludicellea” (s. 1.) nor the “Stolonifera” can be maintained as taxa with the argument of a similar ontogeny of the muscles. A detailed comparison regarding the fate of the parietal muscles (and the polypide anlage) showed similarities between certain of the uniserial to either some of the stolonial, the sheet like, or the cystid-fusing forms suggesting a separate evolutionary origin of the advanced forms in those groups of the serially growing forms. – A new family, Pottsiellidae fam. nov., has been introduced and defined.     

ONTOGENY AND TAPHONOMY: AN EXPERIMENTAL TAPHONOMY STUDY OF THE DEVELOPMENT OF THE BRINE SHRIMP ARTEMIA SALINA

Abstract: Although the relationship between ontogeny and phylogeny has been of long‐standing interest to palaeontologists, the fossil record has provided little insight into the development of long extinct organisms. This has changed with the discovery of numerous assemblages of fossilized invertebrate embryos and larvae, but realising their evolutionary significance is hampered by a paucity of data on the relationship between ontogeny and taphonomy. We describe the results of an experimental taphonomy study of the development of the anostracan brine shrimp Artemia salina, which show that in conditions of aqueous aerobic and anaerobic autolysis and microbial decay, the developmental stages exhibit differential preservation potential. The most decay resistant developmental stage is the diapause cyst, encapulsating the gastrula, in which the gross morphology of the embryo can be maintained for 18 months or more in simple anaerobic conditions. Otherwise, the embryo shrinks within the cyst and cellular and tissue detail of breaks down as lipid droplets coalesce. Postembryonic excysted larvae decay more rapidly. The rate of decay is similar among all larval stages with the exception of the L₄ larva, which resists cuticle failure for longer than later developmental stages. The larvae decay leading to liquefaction of the muscles and viscera, leaving an intact but empty and progressively shrunken and distorted cuticle that eventually loses structural integrity and collapses. Our experimental results provide an explanatory model for the phenomenal abundance of putative diapause stage embryos, in the absence of postembryonic stages, as seen in the Ediacaran Doushantuo Formation of South China and the incompleteness of fossilized developmental sequences of embryos and larvae more generally. It also cautions against the association of developmental stages in fossil deposits without additional evidence. Finally, the pattern of decay seen in larvae provides an explanation for the preservation style of Orsten‐type Lagerstätten where preservation of cuticular detail can be astonishingly fine, but extends internally to muscles and viscera only rarely.     

The Embryonic and Larval Development of Polypterus senegalusCuvier, 1829: its Staging with Reference to External and Skeletal Features, Behaviour and Locomotory Habits

The embryonic and larval development of the Polypteriformes, the presumed sister group of all other living actinopterygians, is poorly known. The main reason is the scarcity of successful breedings in captivity and therefore the lack of developmental series of any one polypterid species. A series of five successful breedings of P. senegalusnow makes it possible to define developmental stages of this species based on numerous closely timed specimens. The staging given here focuses on external embryonic and larval features: epidermal surface structures documented by SEM, colour pattern, development of fins and squamation, larval feeding and locomotory behaviour. The development of P. senegalusis characterized by a long free embryonic phase. Suction feeding is performed from the beginning of larval life (apterolarval phase). The pectoral fins start to become employed for slow locomotion and as supportive structures at around the same time. Olfactorily guided prey capture, however, is observed later in the pterolarval phase. Quantitative kinematic data also demonstrate a change in the mode of undulatory locomotion during this phase. Sustained axial undulation becomes confined to the posterior abdominal and caudal region of the body. At about the same time the paraxial high frequency undulation of the pectoral fin fold is replaced by the characteristic propeller-like movement of much greater amplitude and wavelength. Surfacing for aerial breathing is not seen before a marked change in colouration has taken place at the beginning of the juvenile period. The external gills slowly become reduced during this period. The definitions of larval and juvenile stages given here may advance understanding of developmental processes in the ontogeny of these primitive actinopterygians, and may serve as a tool for comparison with the ontogeny of Tetrapoda and Dipnoi, as well as to that of some “primitive” groups of Actinopterygii. Judging from its distribution among extant taxa, embryonic and larval ciliation is a character that most probably belongs to the grundplan? of Osteognathostomata. Phylogenetic evaluation is not so clear for the two other prominent embryonic and larval specializations found in Polypterus: upper labial attachment glands and opercular external gills. © 1997 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd.     

The anatomy, affinity, and phylogenetic significance of Markuelia

The fossil record provides a paucity of data on the development of extinct organisms, particularly for their embryology. The recovery of fossilized embryos heralds new insight into the evolution of development but advances are limited by an almost complete absence of phylogenetic constraint. Markuelia is an exception to this, known from cleavage and pre-hatchling stages as a vermiform and profusely annulated direct-developing bilaterian with terminal circumoral and posterior radial arrays of spines. Phylogenetic analyses have hitherto suggested assignment to stem-Scalidophora (phyla Kinorhyncha, Loricifera, Priapulida). We test this assumption with additional data and through the inclusion of additional taxa. The available evidence supports stem-Scalidophora affinity, leading to the conclusion that scalidophorans, cyclonerualians, and ecdysozoans are primitive direct developers, and the likelihood that scalidophorans are primitively metameric.     

Utilizing auxology to understand ontogeny of extinct hominins: A case study on Homo naledi

The methods used to study human growth and development (auxology) have not previously been applied within the setting of hominin maturation (ontogeny). Ontogeny is defined here as the pattern of biological change into an adult form, both at the individual and species level. The hominin fossil record has a lack of recovered immature materials, due to such factors as taphonomic processes that destroy pre-adults; the fragility of immature compared to adult bone; and the lower mortality rates of juveniles compared to adults. The recent discovery of pre-adult hominin skeletal material from a single, homogeneous Homo naledi species from the Rising Star cave system in South Africa provides the opportunity for a broader application of auxology methods and thus the need to understand their use in a modern context. Human auxology studies benefit from a robust database, across multiple populations, and with longitudinal studies in order to assess the patterns and variations in typical growth, development and life history stages. Here, we review the approach, vocabulary, and methods of these human studies, investigate commonalities in data with the fossil record, and then advance the reconstruction of ontogeny for the extinct hominin species H. naledi. To this end, we apply an auxology model into the paleontological context to broadly predict H. naledi birthweight of the offspring at 2.06 kg with a range (±1 SD) of 1.89 to 2.24 kg, with a length at birth 45.5 cm. We estimate a H. naledi juvenile partial skeleton DH7 to be a height of 111–125 cm at death.     

Diverse fossil amoebae in German Mesozoic amber

Fossil amoebae are very rare, although their evolutionary history extends back into the Proterozoic. The Cenomanian amber of Schliersee (southern Germany) is very rich in micro-organisms and contains the highest diversity of fossil freshwater rhizopods (Gymnamoebia and Testacealobosia) yet discovered. Altogether seven testate amoebae and one gymnamoebian species are recorded from this Mesozoic amber. The four newly discovered taxa described in this paper can be assigned to the extant species Centropyxis delicatula , Centropyxis hirsuta , Phryganella acropodia and Phryganella paradoxa . Over 200 individuals of Phryganella paradoxa are preserved. Together with their syninclusions, the amoebae are species of limnetic or limnetic–terrestrial microcoenoses. The presence of 100-myr-old fossils with extant representatives suggests evolutionary stasis of these freshwater amoebae. However, not all modern testacean families have been recorded from Mesozoic limnetic habitats. Our experimental studies verify that naked and testate amoebae can be embedded in resins.     

Taxonomic and functional implications of mandibular scaling in early hominins

Body mass estimates for fossil hominin taxa can be obtained from suitable postcranial and cranial variables. However, the nature of the taphonomic processes that winnow the mammalian fossil record are such that these data are usually only available for the minority of the specimens that comprise the hypodigm of a species. This study has investigated the link between species mean body mass and the height and width of the mandibular corpus in a core sample of 23 species of extant simians. The slopes of the least-squares regressions for the whole sample and for the hominoid subset are similar. However, the intercepts differ so that for a given body mass, a hominoid will generally have a smaller mandible than a generalized simian. The same mandibular measurements were taken on 75 early hominin mandibles assigned to eight species groups. When mandibular corpus height- and width-derived estimates of body mass for the fossil taxa were compared with available postcranial and cranial-derived body mass estimates, the eight early hominin species sort into four groups. The first, which includes A. afarensis and A. africanus, has mandibles which follow a “generalized simian” scaling relationship. The second group, which comprises the two “robust” australopithecine species, P. boisei and P. robustus, has mandibles which scale with body mass as if they are “super-simians,” for they have substantially larger mandibles than a simian with the same body mass. The two “early Homo” species, H. habilis sensu stricto and H. rudolfensis, make up the third group. It has mandibular scaling relationships that are intermediate between that of the comparative simian sample and that of the hominoid subsample. The last of the four groups comprises H. ergaster and H. erectus; their mandibles scale with body mass as if they were hominoids, so that of the four groups they have the smallest mandibles per unit body mass. These results are related to comparable information about relative tooth size. Their relevance for attempts to interpret the dietary adaptations of early hominins are explored. Am J Phys Anthropol 105:523–538, 1998. © 1998 Wiley-Liss, Inc.     

Fossils and seed plant phylogeny reanalyzed

In a cladistic analysis of Recent seed plants, Loconte and Stevenson (1990) obtained results that conflict with our 1986 analysis of both extant and fossil groups and argued that fossil data had led us to incorrect conclusions. To explore this result and the general influence of fossils on phylogeny reconstruction, we assembled new “Recent” and “Complete” (extant plus fossil) data sets incorporating new data, advances in treatment of characters, and those changes of Loconte and Stevenson that we consider valid. Our Recent analysis yields only one most parsimonious tree, that of Loconte and Stevenson, in which conifers are linked with Gnetales and angiosperms (anthophytes), rather than with Ginkgo, as in our earlier Recent and Complete analyses. However, the shortest trees derived from our Complete analysis show five arrangements of extant groups, including that of Loconte and Stevenson and our previous arrangements, suggesting that the result obtained from extant taxa alone may be misleading. This increased ambiguity occurs because features that appear to unite extant conifers and anthophytes are seen as convergences when fossil taxa are interpolated between them. All trees found in the Complete analysis lead to inferences on character evolution that conflict with those that would be drawn from Recent taxa alone (e.g., origin of anthophytes from plants with a “seed fern” morphology). These results imply that conclusions on many aspects of seed plant phylogeny are premature; new evidence, which is most likely to come from the fossil record, is needed to resolve the uncertainties.     

Plant Traces Resembling Skolithos

The ichnogenus Skolithos Haldeman 1840 is a simple tubular trace fossil that was initially described as a “fucoid” or seaweed impression, i.e., a fossilized marine plant. Today the tracemaker is commonly considered to have been a vermiform marine invertebrate, although this interpretation is not acceptable to all. It is the name giver to the archetypal, Seilacherian Skolithos ichnofacies, which is a widely acknowledged indicator of relatively high energy, shallow water, nearshore to marginal marine environments. Here we record and describe analogous cylindrical structures, unquestionably reflecting plant tracemakers. These have been recognized in Quaternary consolidated “coffee” sandrock and Recent coastal dune, marsh or swamp, and salt meadow settings from northern New Zealand and Sapelo Island, USA.

We conclude that Skolithos is not an unequivocal indicator of shallow marine settings. Our observations confirm the opinions of a number of previous workers who have expressed the need for caution when using this ichnotaxon as a shallow-water, higher energy palaeoenvironmental indicator, especially when supporting evidence is otherwise lacking. In particular we note that in high intertidal and non-aquatic settings, Skolithos may be a plant trace fossil rather than serve as evidence for invertebrate activities. At the present time there is pressing need for a thorough systematic revision of the Skolithos ichnotaxon.