Megascopic eukaryotic algae from the 2.1-billion-year-old negaunee iron-formation, Michigan.

Hundreds of specimens of spirally coiled, megascopic, carbonaceous fossils resembling Grypania spiralis (Walcott), have been found in the 2.1-billion-year-old Negaunee Iron-Formation at the Empire Mine, near Marquette, Michigan. This occurrence of Grypania is 700 million to 1000 million years older than fossils from previously known sites in Montana, China, and India. As Grypania appears to have been a photosynthetic alga, this discovery places the origin of organelle-bearing eukaryotic cells prior to 2.1 billion years ago.

Nucleic acid and protein clocks.

The use of pairwise comparisons of correctly aligned DNA and protein sequences for the measurement of time in historical biology remains a contentious matter. However, the limited success of some molecular evolutionary clocks provides a stimulus to attempt to improve their resolution by the judicious selection of sequences for ease of alignment, commonality of function, taxonomic breadth and appropriate rates of evolution. Existing algorithms for correcting observed distances for superimposed nucleotide substitutions or amino acid replacements appear adequate for the task, given the noise that results from the inherent variability of the process. Some possible approaches are illustrated through the use of gene and protein sequences of the large subunit of ribulose 1,5-bisphosphate carboxylase/oxygenase: an enzyme that is demonstrably homologous from purple bacteria to flowering plants.

Collagen gene construction and evolution.

Collagen genes appear to have been assembled by the tandem repetition of homologous primary (9 base pair), secondary (54 base pair), and tertiary (702 base pair) modules. In vertebrate interstitial collagen genes many of the secondary modules are separated by introns, but in invertebrate collagen genes the non-coding sequences lie near the ends of supposed tertiary modules and are therefore about 702 (54 X 13) base pairs apart. The genes for vertebrate interstitial collagens (types I-III) seem to have been constructed by the tandem repetition of five tertiary modules, three of which were subsequently shortened by internal deletions. This shortening of the gene resulted in the non-integral relationship between the period of the fibrils and the length of the molecules of vertebrate collagens, and was therefore responsible for the mechanical properties of the completed product. Comparisons of the amino acid sequences of various collagens indicate that the main types of collagen evolved about 800-900 million years ago, a date that agrees well with the fossil record of primitive Metazoa.

Derivation of the globins from type b cytochromes.

Similarities in the amino acid sequences of vertebrate and invertebrate globins, b5 and b2 cytochromes and chicken sulfite oxidase point to a common ancestry for all of these proteins. The distal heme ligand (histidine or its equivalent) is common to both sets of proteins, but the proximal histidine ligand of the cytochromes is replaced by another histidine residue in the globins. This explains why the heme is reversed between globins and b5 cytochromes. It seems likely that the genes for primitive globins contained three exons, the first two of which were derived from a cytochromelike DNA sequence. A model is presented to show how globins may have evolved from a pre-existing type b cytochrome; the complexity of the required changes is an indication that all globins are monophyletic.

Molluscan phylogeny: the paleontological viewpoint.

Stasek (1) theorized that the extant mollusks are the progeny of three separate lineages that separated before the phylum was well established. He wrote that no known intermediate forms, fossil or living, bridge the "enormous gaps between any two of the three lineages," and therefore treated each as a separate subphylum. These subphyla are (i) the subphylum Aculifera Hatscheck 1891, containing only the class Aplacophora, derived from the most primitive ancestors of the Mollusca; (ii) the subphylum Placophora von Jhering 1876, containing only the class Polyplacophora, and emphasizing the pseudometamerism of its more advanced premollusk ancestor; and (iii) the subphylum Conchifera Gegenbaur 1878, containing the Monoplacophora and the other classes derived from it. We point out that the Polyplacophora may be derived from the Monoplacophora instead of a more primitive ancestral stock. We also suggest that the Conchifera can be separated into two major lineages, each worthy of the rank of subphylum. The fossil record indicates that the Monoplacophora gave rise to the Gastropoda, Cephalopoda, Rostroconchia, and possibly Polyplacophora, and that the Pelecypoda and Scaphopoda are derived from the Rostroconchia. These last three classes thus form a lineage that diverged from the Monoplacophora in the Early Cambrian. They emphasized a shell form that in all groups is primitively open at both ends, allowing the gut to remain relatively straight, with an anterior mouth and posterior anus. They became burrowing (infaunal) deposit or filter feeders. We coin the term Diasoma (through-body) for the subphylum containing these three classes (Rostroconchia, Pelecypoda, and Scaphopoda). The remaining three classes (Monoplacophora, Gastropoda, and Cephalopoda) emphasize a conical univalved shell, usually twisted into a spiral. The relatively small single aperture forces the anus to lie close to the mouth, and the gut is bent into a "U." Most are surface-dwelling (epifaunal) grazers or carnivores. We coin the name Cyrtosoma (hunchback-body) for the subphylum containing these three classes. Strictly speaking, the cyrtosomes are the ancestors of the diasomes but, in fact, both subphyla appeared and began to diversify within a few million years in the Early Cambrian. Note added in proof: After proofs were corrected we were informed that the new genus Opikella (40) is preoccupied by (Opikella = Oepikella) Thorslund 1940, an Ordovican ostracod. We rename the mollusk genus Oepikila.

Fordilla troyensis Barrande: The Oldest Known Pelecypod.

Specimens of the small bivalved animal Fordilla troyensis Barrande from New York State show that this fossil is the oldest known pelecypod mollusk and not a conchostracan arthropod. This finding extends the range of the class Pelecypoda backward in time from the Early Ordovician (about 495 million years ago) to the Early Cambrian (about 540 to 570 million years ago). The morphology of Fordilla troyensis suggests that it lived infaunally and that it was ancestral to the pelecypod subclasses Heteroconchia and Isofilibranchia.

Rostroconchia: a new class of bivalved mollusks.

Four Paleozoic bivalved genera are assigned to the new molluscan class Rostroconchia: Eopteria, Euchasma, Conocardium, and Pseudoconocardium. These mollusks have ani uncoiled univalved larval shell; an untorted bivalved adult shell; no hinge teeth, ligament, or adductor muscles; and a fused, almost inflexible. hinge. Rostroconchianis developed separately from the pelecypods through the ribeirioids, but are regarded as more closely related to the Pelecypoda and Scaphopoda than to other known classes of mollusks.