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Concepts of the family Hyolithidae Nicholson fide Fisher and the genera Hyolithes Eichwald and Orthotheca Novak have been expanded through time to encompass a variety of morphologically dissimilar shells. The Hyolithidae is here considered to include only those hyolithid species which have a rounded (convex) dorsum; slopes on the dorsum are inflated, and the venter may be flat or slightly inflated. Hyolithes encompasses species which possess a low dorsum and a prominent longitudinal sulcus along each edge of the dorsum; the ligula is short and the apertural rim is flared. The emended concept of Orthotheca includes only those species of orthothecid hyoliths which have a subtriangular transverse outline and longitudinal lirae covering the shell on both dorsum and venter.Eighteen species of Hyolithes and one species of Orthotheca from the Appalachian region and Western Interior were reexamined in light of more modern taxonomic concepts and standards of quality for type material. Reexamination of type specimens of H. similis Walcott from the Lower Cambrian of Newfoundland, H. whitei Resser from the Lower Cambrian of Nevada, H. billingsi Walcott from the Lower Cambrian of Nevada, H. gallatinensis Resser from the Upper Cambrian of Wyoming, and H. partitus Resser from the Middle Cambrian of Alabama indicates that none of these species represents Hyolithes. Hyolithes similis is here included under the new genus Similotheca, in the new family Similothecidae. Hyolithes whitei is designated as the type species of the new genus Nevadotheca, to which H. billingsi may also belong. Hyolithes gallatinensis is referred to Burithes Missarzhevsky with question, and H. partitus may represent Joachimilites Marek. The type or types of H. attenuatus Walcott, H. cecrops Walcott, H. comptus Howell, H. cowanensis Resser, H. curticei Resser, H. idahoensis Resser, H. prolixus Resser, H. resseri Howell, H. shaleri Walcott, H. terranovicus Walcott, and H. wanneri Resser and Howell lack shells and/or other taxonomically important features such as a complete aperture, rendering the diagnoses of these species incomplete. Their names should only be used for the type specimens until better preserved topotypes become available for study. Morphology of the types of H.? corrugatus Walcott and “Orthotheca” sola Resser does not support placement in the Hyolitha; the affinities of these species are uncertain.

Hyolithids are a group of Paleozoic lophotrochozoans with a four-pieced skeleton consisting of a conch, an operculum, and a pair of lateral ‘spines' named helens. Both the conch and operculum are relatively well known and, to a certain extent, have modern analogues in other lophotrochozoan groups. The helens, on the other hand, are less well known and do not have clear modern analogues. This has hindered the knowledge of the complete morphology of the hyolithid skeleton, as well as other aspects of hyolithid biology, such as the organization of soft parts, and their ability to move. The material studied herein, consisting of disarticulated skeletal elements from the Silurian of Gotland, Sweden, illustrates a complete developmental sequence of a hyolithid species and includes the first complete, three-dimensionally preserved helens. Our material confirms that helens were massive skeletal elements, whose growth started proximally with the deposition of a central, coherent lamella. Further shell accretion took place around this lamella, but followed a particular accretion pattern probably constrained by the presence of marginal muscle attachment sites on the proximal-most portion of the helens. These muscle attachment sites were ideally located to allow a wide range of movements for the helens, suggesting that hyolithids may have been relatively mobile organisms.

ABSTRACT. Re-examination of type specimens of Early Cambrian and early Mid Cambrian hyoliths from Sweden confirms placement of Hyolithes teretiusculus Linnarsson in Hexitheca Syssoiev, and reassignment to the order Hyolithida rather than Orthothecida. Inclusion of Hyolithes affinis Holm within Decoritheca Syssoiev is re-confirmed, and Hyolithes socialis Linnarsson is now referred to Nevadotheca Malinky and Slapylites Marek. An operculum included under Hyolithes oelandicus Holm is transferred provisionally to Maxilites Marek, and conchs of that species are placed under Tulenicornus Val'kov with question. No other hyolith species of Holm can be definitely assigned to genus. The rarity of well preserved opercula is a major impediment to recognizing hyolith taxa confidently, and we follow Holm in attributing the comparative rarity of opercula to the fact that in some hyolith taxa the operculum may have been composed of a different, and perhaps less stable, material than the rest of the conch.

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