An assemblage of 50 species of small shelly fossils is described from Cambrian Series 2 (Stage 4) strata in North Greenland, the present day northernmost part of the paleocontinent of Laurentia. The fossils are derived from the basal member of the Aftenstjernesø Formation at Navarana Fjord, northern Lauge Koch Land, a condensed unit that accumulated in a sediment-starved outer ramp setting in the transarctic Franklinian Basin, on the Innuitian margin of Laurentia. Most other small shelly fossil assemblages of similar age and composition from North America are described from the Iapetan margin of Laurentia, from North-East Greenland south to Pennsylvania. Trilobites are uncommon, but include Serrodiscus. The Australian bradoriid Spinospitella is represented by a complete shield. Obolella crassa is the only common brachiopod. Hyoliths, including Cassitella, Conotheca, Neogloborilus, and Triplicatella, are abundant and diverse, but most are represented just by opercula. Sclerites interpreted as stem-group aculiferans (sachitids) are conspicuous, including Qaleruaqia, the oldest described paleoloricate, Ocruranus?, Inughuitoconus n. gen., and Hippopharangites. Helcionelloid mollusks are diverse, but not common; they are associated with numerous specimens of the bivalve Pojetaia runnegari. The fauna compares best with that of the upper Bastion Formation of North-East Greenland, the Forteau Formation of western Newfoundland, and the Browns Pond Formation of New York, but several taxa have a world-wide distribution. Many specimens are encrusted with crystals of authigenic albite. New species: Anabarella? navaranae, Stenotheca? higginsi, Figurina? polaris, Hippopharangites groenlandicus, Inughuitoconus borealis, and Ocruranus? kangerluk.

UUID: http://zoobank.org/160a17b1-3166-4fcf-9849-a3cabd1e04a3

In this study, middle to late Mississippian microfossil assemblages from the Maritimes Basin of eastern Canada (Nova Scotia, SW Newfoundland, and New Brunswick) are closely compared to those from Western Paleotethys basins. The comparison is focused mainly on foraminifers and calcareous algae. Most foraminifers and algae described from the Maritimes Basin are considered cosmopolitan, and the occurrence in western Europe and northern Africa of taxa previously considered endemic to the North America Realm suggests a close paleobiogeographic relationship. This European/African correlation is further supported by other foraminiferal/algal taxa, the importance of which were previously overlooked, including: Plectogyranopsis ex gr. P. hirosei (Okimura, 1965), Mikhailovella Ganelina, 1956, Koktjubina windsorensis (Mamet, 1970), Polysphaerinella bulla Mamet, 1973, Mstinia Dain in Dain and Grozdilova, 1953, Haplophragmina Reitlinger, 1950, Omphalotis Shlykova, 1969, Pseudolituotuba Vdovenko, 1971, Pseudoendothyra Mikhailov, 1939, Saccamminopsis (Sollas, 1921) Vachard and Cózar, 2003, Kamaenella Mamet and Roux, 1974, and Anthracoporellopsis Maslov, 1956. Some species recorded in the Maritimes Basin have been typically recorded in Britain and Ireland in the southern platform of Laurussia. This implies a connection via the Rhenohercynian Ocean, whereas statistical analyses suggest that Maritimes Basin assemblages are closer to those of the Gondwana platform, which could have been established via the Paleotethys Ocean, and also with terranes northwest of the Variscan Front, in which its most logical connection should be with a still-open Rheic Ocean during the Visean and early Serpukhovian. Those taxa demonstrate a more-or-less continuous faunal and microfloral interchange between the Maritimes Basin and the Western Paleotethys paleobiogeographic realm. Furthermore, the width of the Paleotethys and Rheic oceans separating these regions is not considered excessive, particularly during the late Visean and early Serpukhovian.

The middle Eocene Washakie Formation of Wyoming, USA, provides a rare window, within a single depositional basin, into the faunal transition that followed the early Eocene warming events. Based on extensive examination, we report a minimum of 27 species of carnivorous mammals from this formation, more than doubling the previous taxic count. Included in this revised list are a new species of carnivoraform, Neovulpavus mccarrolli n. sp., and up to ten other possibly new taxa. Our cladistic analysis of early Carnivoraformes incorporating new data clarified the array of middle Eocene taxa that are closely related to crown-group Carnivora. These anatomically relatively derived carnivoraforms collectively had an intercontinental distribution in North America and east Asia, exhibiting notable variations in body size and dental adaptation. This time period also saw parallel trends of increase in body size and dental sectoriality in distantly related lineages of carnivores spanning a wide range of body sizes. A new, model-based Bayesian analysis of diversity dynamics accounting for imperfect detection revealed a high probability of substantial loss of carnivore species between the late Bridgerian and early Uintan North American Land Mammal ‘Ages’, coinciding with the disappearance of formerly common mammals such as hyopsodontids and adapiform primates. Concomitant with this decline in carnivore diversity, the Washakie vertebrate fauna underwent significant disintegration, as measured by patterns of coordinated detection of taxa at the locality level. These observations are consistent with a major biomic transition in the region in response to climatically induced opening-up of forested habitats.

UUID: http://zoobank.org/9162f1a6-a12c-4d55-ba1d-dc66e8cda261

New, distinctive, well-preserved and previously undescribed constituents of a Middle Ordovician (middle Darriwilian, Dw2) radiolarian assemblage from the Table Cove Formation in Newfoundland are described. Three-dimensional X-ray micro-computed tomography (μ-CT) facilitates detailed examination of key specimens revealing hitherto unknown details of the internal morphologies of key lower Paleozoic taxonomic groups, among which a lack of knowledge has previously impeded resolution of higher taxonomic rankings.

Twenty-seven archaeospiculid and entactinarian taxa are described and illustrated including six new species: Westernbrookia polygonata n. sp., Neopalaeospiculum piccadilliensis n. sp., Ramuspiculum laxum n. sp. Spongentactinia nazarovi n. sp., Aspiculum irregulare n. sp., and Nyfrieslandia ramosissima n. sp. The investigation extends the known ranges of the species: Pararcheoentactinia reedae Won and Iams, 2002; Sphaeroentactinia robusta Won and Iams, 2015; Varispiculum ectospiculatum Won and Iams, 2015; and Svalbardospiculum multifurcatum (Won, Iams, and Reed, 2005), together with the genus Echidnina to the mid-Darriwilian.

UUID: http://zoobank.org/74826b7b-bb86-45d5-ad23-e6e65e0706df.

This is the first of two comprehensive taxonomic works on the early Miocene (ca. 23–20 Ma) bryozoan fauna associated with coral reefs from the Siamaná Formation, in the remote region of Cocinetas Basin in the La Guajira Peninsula, northern Colombia, southern Caribbean. Fifteen bryozoan species in 11 families are described, comprising two cyclostomes and 13 cheilostomes. Two cheilostome genera and seven species are new: Antropora guajirensis n. sp., Calpensia caribensis n. sp., Atoichos magnus n. gen. n. sp., Gymnophorella hadra n. gen. n. sp., Cribrilaria multicostata n. sp., Cribrilaria nixor n. sp., and Figularia bragai n. sp. Eight species are identified only at genus level and remain in open nomenclature. Of the species found, 27% have erect colonies and 73% encrusting colonies. Both types contributed to the reef framework and produced sediment. The observed bryozoan diversity was higher in the barrier reefs than in the lagoonal patch reefs.

UUID: http://zoobank.org/5c8468ef-31b0-4e7e-ba93-60a2e2f30b76.

Diverse and abundant fossil taxa have been described in the lower Cambrian Shipai Formation in the Three Gorges area of Hubei Province, South China, but the taxonomy and diversity of the co-occurring brachiopod fauna are still far from clear. Here we describe the brachiopod fauna recovered from the Shipai Formation in the Three Gorges area of South China, including representatives of the subphylum Linguliformea: linguloids (Lingulellotreta ergalievi, Eoobolus malongensis, and Neobolidae gen. indet. sp. indet.), and an acrotretoid (Linnarssonia sapushanensis); and representatives from the subphylum Rhynchonelliformea: the calcareous-shelled Kutorginates (Kutorgina sinensis, Kutorgina sp., and Nisusia liantuoensis). This brachiopod assemblage and the first occurrence of Linnarssonia sapushanensis shell beds permit correlation of the Shipai Formation in the Three Gorges area of Hubei Province with the Stage 4 Wulongqing Formation in the Wuding area of eastern Yunnan. This correlation is further strengthened by the first appearance datum (FAD) of the rhynchonelliform brachiopod Nisusia in the upper silty mudstone of both the Shipai and Wulongqing formations. The new well-preserved material, derived from siliciclastic rocks, also gives critical new insights into the fine shell structure of L. sapushanensis. Microstructural studies on micromorphic acrotretoids (like Linnarssonia) have previously been restricted to fossils that were acid-etched from limestones. This is the first study to carry out detailed comparative ultrastructural studies on acrotretoid shells preserved in siliciclastic rocks. This work reveals a hollow tube and solid column microstructure in the acrotretoid shells from the Shipai Formation, which is likely to be equivalent of traditional column and central canal observed in shells dissolved from limestones.

In this study, a new assemblage of Ediacaran metazoan fossils is reported from the basal Stáhpogieddi Formation on the Digermulen Peninsula of Arctic Norway, including Anulitubus n. gen. Moczydłowska in Moczydłowska et al., Anulitubus formosus n. gen. n. sp. Moczydłowska in Moczydłowska et al., Coniculus n. gen. Moczydłowska in Moczydłowska et al., Coniculus elegantis n. gen. n. sp. Moczydłowska in Moczydłowska et al., Fistula n. gen. Moczydłowska in Moczydłowska et al., and Fistula crenulata n. gen. n. sp. Moczydłowska in Moczydłowska et al. The specimens are three-dimensionally preserved and include tubular and conical skeletons that are morphologically distinguished by their body-wall constructions, radial symmetry, polarity, segmentation, and annulation. The skeletons are interpreted to be biomineralized by primary silica based on computed micro-tomographic, petrographic, geochemical, and spectroscopic evidence of originally rigid body wall with layers of constant thicknesses, composed of opal, microcrystalline quartz, and an admixture of carbonaceous material, which differ from the host sediment mineralogy and do not show replacement or encrustation. The fossil-bearing interval immediately overlies strata of Gaskiers age and can be bracketed within 580–541 Ma, but it is estimated to be ca. 575 Ma on the basis of averaged sedimentation rates and biostratigraphic correlations with Ediacaran biota found in up-section deposits of ca. 558–555 Ma. Future new findings of such fossils in different preservation modes and further multi-collector inductively coupled plasma mass spectrometry, which shows the silicon fractionation and traces its biogenic origin versus inorganic mineralization, may corroborate the interpretation of biogenic silicification of these earliest skeletal fossils.

UUID: https://zoobank.org/6bccada1-870e-47b0-b819-82685152ea54

The filter-feeding organ of some extinct brachiopods is supported by a skeletal apparatus called the brachidium. Although relatively well studied in Atrypida and Athyridida, the brachidial morphology is usually neglected in Spiriferida. To investigate the variations of brachidial morphology in Spiriferida, 65 species belonging to eight superfamilies were analyzed. Based on the presence/absence of the jugal processes and normal/modified primary lamellae of the spiralia, four types of brachidium are recognized. Type-I (with jugal processes) and Type-II (without jugal processes), both having normal primary lamellae, could give rise to each other by losing/re-evolving the jugal processes. Type-III, without jugal processes, originated from Type-II through evolution of the modified lateral-convex primary lamellae, and it subsequently gave rise to Type-IV by evolving the modified medial-convex primary lamellae. The evolution of brachidia within individual evolutionary lineages must be clarified because two or more types can be present within a single family. Type-III and Type-IV are closely associated with the prolongation of the crura, representing innovative modifications of the feeding apparatus in response to possible shift in the position of the mouth towards the anterior, allowing for more efficient feeding on particles entering the mantle cavity from the anterior gape. Meanwhile, the modified primary lamellae adjusted/regulated the feeding currents. The absence of spires in some taxa with Type-IV brachidium might suggest that they developed a similar lophophore to that in some extant brachiopods, which can extend out of the shell.

Members of the echinoid order Spatangoida, a highly diverse and abundant marine invertebrate clade, were important denizens of the Cretaceous Western Interior Seaway (WIS), an epicontinental seaway that divided North America in two during an interval of greenhouse conditions between roughly 100 and 65 million years ago. A phylogenetic analysis of spatangoids was conducted using a character matrix of 32 characters from 21 species. Species that occur in the WIS were considered comprehensively, and species from other regions such as South America, Europe, and North Africa were also incorporated into the analysis. Phylogenetic patterns retrieved are largely congruent with preexisting family-level classifications; however, species within several genera, especially Hemiaster and Heteraster, need to be reassigned so that classification better reflects phylogeny. The genera Washitaster and Heteraster are closely related, as are Mecaster, Palhemiaster, and Proraster; Pliotoxaster, Macraster, and Hemiaster; and Micraster and Diplodetus. Biogeographic patterns were also considered using the phylogeny, and several episodes of vicariance and range expansion were identified. These were possibly related to some of the various major episodes of sea-level rise and fall during the Cretaceous. In particular, Valangian–mid-Aptian regressions may have caused vicariance within Heteraster and Washitaster while other early spatangoid vicariance may be related to regressions during the late Aptian–early Cenomanian. Further, vicariance caused by regressions during the mid-Cenomanian–Maastrichtian may have driven diversification within Micraster and Diplodetus. Last, transgressions during the late Aptian–early Cenomanian seem to have spurred prominent range expansions in Mecaster and Hemiaster.

Permian millipedes are rare, especially so considering the relative abundance of millipedes in Carboniferous rocks. We report an early Permian millipede fauna containing three new genera and species of millipedes (Oklahomasoma richardsspurense new genus new species, Karstiulus fortsillensis new genus new species, and Dolesea subtila new genus new species) found in fossil-producing pockets of the Fort Sill fissures exposed in the Dolese Quarry near Richards Spur, southwest Oklahoma, USA. These are the first new genera of invertebrates to be described from this site, one of the most prolific fossil-vertebrate sites in the world. We also comment on taxa with morphological similarities and note previously described occurrences of Permian millipedes as well as occurrences of fossil myriapods (millipedes and centipedes) in karst deposits (caves and fissure fills) in Europe, Africa, Asia, North America, and the Caribbean. In contrast with the forms found at Richards Spur, most of these previous accounts of millipedes found in caves and fissure fills are of Pleistocene forms that are closely allied to modern taxa. The taxa from Richards Spur bear some similarities to Pennsylvanian forms. Karst (cave and fissure) faunas should be ranked with concretion faunas, cannel coals, and amber faunas as a major source of fossil myriapods.

UUID: http://zoobank.org/5d58e1fb-4e5b-4597-9cd9-5cc9e2096b4d

Cribrilinid bryozoans originating from Pleistocene deep-water sediments from two localities near Messina (Sicily, Italy)—Capo Milazzo (Gelasian) and Scoppo (Calabrian)—were examined. Five cribrilinid species were found, three in each locality and time interval, with only one species shared. Three species, Cribrilaria profunda n. sp., Glabrilaria transversocarinata n. sp., and Figularia spectabilis n. sp., are new to science. Of the two remaining species, Figularia figularis was already known from local fossil associations, whereas Glabrilaria pedunculata, a present-day Mediterranean species, is recorded for the first time as a fossil. New combinations are suggested for two species previously assigned to Puellina, Cribrilaria saldanhai (Harmelin, 2001) n. comb. and Cribrilaria mikelae (Harmelin, 2006) n. comb. The diagnosis of the genus Figularia was amended to include an erect growth morphology in addition to the encrusting form, and the occurrence of ooecia formed by the distal kenozooid. Following a literature revision of all species currently assigned to Figularia, the new combinations Vitrimurella capitifera (Canu and Bassler, 1929) n. comb. and Hayamiellina quaylei (Powell, 1967a) n. comb. are suggested, and problematic species are listed and briefly discussed.

UUID: http://zoobank.org/b7b36152-bf7b-4e00-b6ec-2614b2a58f1b

Eight bryozoan species are described from the Hanchiatien Formation (lower Silurian, Telychian) of southern Chongqing, South China. Four species are new: the trepostomes Asperopora sinensis n. sp., Trematopora jiebeiensis n. sp., and Trematopora tenuis n. sp., and the fenestrate Moorephylloporina parvula n. sp. One species, the cystoporate Hennigopora sp. indet., is described in open nomenclature. Moorephylloporina parvula n. sp. is eurytopic, occurring in all types of facies within the bioherms. Erect Moorephylloporina Bassler, 1952, Trematopora Hall, 1852, and Leioclema Ulrich, 1882 formed pioneering communities on weakly cemented substrata, whereas encrusting Fistulipora M‘Coy, 1849, Hennigopora Bassler, 1952, and Asperopora Owen, 1969 occurred on hardgrounds and formed densely compact framestones. Robust branched Trematopora and Leioclema tend to occur out of the reef core (framework) where they could have formed reef-flank thickets in more agitated conditions. The generic composition of the studied fauna correlates with other localities in South China, and they show general paleobiogeographic relations to Siberia and Indiana, USA.

UUID http://zoobank.org/3326dd2f-7c9e-43bc-9dab-84047b274f89

In order to maximize the utility of future studies of trilobite ontogeny, we propose a set of standard practices that relate to the collection, nomenclature, description, depiction, and interpretation of ontogenetic series inferred from articulated specimens belonging to individual species. In some cases, these suggestions may also apply to ontogenetic studies of other fossilized taxa.

A new sinistrally coiled univalved mollusk Catalanispira n. gen. is described with two species; Catalanispira reinwaldti (Öpik, 1930) from the Middle Ordovician Kõgekallas Formation (Darriwilian) of Estonia and Catalanispira plattevillensis n. gen. n. sp. from the Upper Ordovician Platteville Formation (Sandbian) of northern Illinois, USA. Morphological features include a large, low-trochiform shell, a narrow lenticular aperture, a deep funnel-like umbilicus, a falcate inner lip and a large (1.4 mm wide) protoconch. Ornamentation consists of fine commarginal growth lines or ribs but superimposed on a slightly irregular shell surface. Catalanispira n. gen. is placed within the sinistrally coiled order Mimospirida and the family Onychochilidae, and Catalanispirinae n. subfam. is proposed. The large Lower Ordovician (Tremadocian) Pelecyogyra Ebbestad and Lefebvre, 2015 from Morocco and France is transferred to this new subfamily. The well-preserved initial growth stage of Catalanispira plattevillensis n. gen. n. sp. is cap-shaped, slightly asymmetrical, unusually large, and smooth, and represents either an unusually large embryonic shell (protoconch 1) or a larval shell (protoconch 2). It differs from the smaller protoconch described for the clisospirine Mimospira Koken in Koken and Perner, 1925, which might include a multiwhorled larval shell (protoconch 2). Mimospirids are dominantly Ordovician, and have been classified as untorted mollusks (only distantly related to gastropods), dextral hyperstrophic gastropods, or sinistral orthostrophic gastropods. Sinistral asymmetry already in the embryonic shell and lack of conclusive evidence for coiling direction, e.g., an operculum, could suggest that Catalanispira n. gen. or similar mimosprids were sinistral orthostrophic gastropods. Currently the group is therefore classified as a group of sinistral orthostrophic gastropods, unranked within the Gastropoda.

UUID: http://zoobank.org/affc8dcf-4c0f-493d-bee5-75a457996e84

The Lakeview Limestone is one of the westernmost Cambrian exposures in the northwestern United States and occurs on the western edge of the Montania paleotopographic high. These deposits occur between the deeper water deposits to the west and carbonate banks and intracratonic basins to the east and provide critical link between the regions. A re-investigation of the Cambrian trilobite faunas from the lower portion of the Lakeview Limestone, Pend Oreille Lake, Idaho, is undertaken due to the inadequate illustrations and descriptions provided by Resser (1938a). Resser's type specimens and additional material are figured and described. The trilobite assemblages represent the Ptychagnostus praecurrens Zone, Wuliuan Stage, Miaolingian Series and including two new taxa: Itagnostus idahoensis n. sp., and Utia debra n. sp. Because of the similarity between some species of Amecephalus from the Lakeview Limestone to specimens from the Chisholm Shale, Nevada, the type specimens of Amecephalus piochensis (Walcott, 1886) and Am. packi (Resser, 1935), Walcott's and Resser's type specimens are re-illustrate and their taxonomic problems are discussed. Utia curio Walcott, 1924 from the Spence Shale, Utah, have never been re-illustrated since Walcott (1925), these type specimens are also re-illustrated and compared to Utia debra n. sp.

UUID: http://zoobank.org/5ef6ee5e-bddc-4395-becf-9bcea54c679c

Bituminous limestone of the Ediacaran Shibantan Member of the Dengying Formation (551–539 Ma) in the Yangtze Gorges area contains a rare carbonate-hosted Ediacara-type macrofossil assemblage. This assemblage is dominated by the tubular fossil Wutubus Chen et al., 2014 and discoidal fossils, e.g., Hiemalora Fedonkin, 1982 and Aspidella Billings, 1872, but frondose organisms such as Charnia Ford, 1958, Rangea Gürich, 1929, and Arborea Glaessner and Wade, 1966 are also present. Herein, we report four species of Arborea from the Shibantan assemblage, including the type species Arborea arborea (Glaessner in Glaessner and Daily, 1959) Glaessner and Wade, 1966, Arborea denticulata new species, and two unnamed species, Arborea sp. A and Arborea sp. B. Arborea arborea is the most abundant frond in the Shibantan assemblage. Arborea denticulata n. sp. resembles Arborea arborea in general morphology but differs in its fewer primary branches and lower length/width ratio of primary branches. Arborea sp. A and Arborea sp. B are fronds with a Hiemalora-type basal attachment. Sealing by microbial mats and authigenic cementation may have played an important role in the preservation of Arborea in the Shibantan assemblage. The Shibantan material of Arborea extends the stratigraphic, ecological, and taphonomic ranges of this genus.

UUID: http://zoobank.org/554f21da-5f09-4891-9deb-cbc00c41e5f1

Dilophosaurus wetherilli was the largest animal known to have lived on land in North America during the Early Jurassic. Despite its charismatic presence in pop culture and dinosaurian phylogenetic analyses, major aspects of the skeletal anatomy, taxonomy, ontogeny, and evolutionary relationships of this dinosaur remain unknown. Skeletons of this species were collected from the middle and lower part of the Kayenta Formation in the Navajo Nation in northern Arizona. Redescription of the holotype, referred, and previously undescribed specimens of Dilophosaurus wetherilli supports the existence of a single species of crested, large-bodied theropod in the Kayenta Formation. The parasagittal nasolacrimal crests are uniquely constructed by a small ridge on the nasal process of the premaxilla, dorsoventrally expanded nasal, and tall lacrimal that includes a posterior process behind the eye. The cervical vertebrae exhibit serial variation within the posterior centrodiapophyseal lamina, which bifurcates and reunites down the neck. Iterative specimen-based phylogenetic analyses result in each of the additional specimens recovered as the sister taxon to the holotype. When all five specimens are included in an analysis, they form a monophyletic clade that supports the monotypy of the genus. Dilophosaurus wetherilli is not recovered as a ceratosaur or coelophysoid, but is instead a non-averostran neotheropod in a grade with other stem-averostrans such as Cryolophosaurus ellioti and Zupaysaurus rougieri. We did not recover a monophyletic ‘Dilophosauridae.’ Instead of being apomorphic for a small clade of early theropods, it is more likely that elaboration of the nasals and lacrimals of stem-averostrans is plesiomorphically present in early ceratosaurs and tetanurans that share those features. Many characters of the axial skeleton of Dilophosaurus wetherilli are derived compared to Late Triassic theropods and may be associated with macropredation and an increase in body size in Theropoda across the Triassic-Jurassic boundary.

A new species of microconchid tubeworm, Microconchus cravenensis is described from the Mississippian Cracoean reefs of North Yorkshire, United Kingdom. Despite the fact that some other microconchid species could have attained large tube length, the new species possesses the largest recorded diameter (to 7.7 mm) of the planispirally-coiled (attachment) tube and the largest recorded aperture diameter (8.3 mm) in the helically uncoiled portion. Thus, with respect to these features, Microconchus cravenensis n. sp. is the largest and most robust microconchid species recognized so far. At present, it is only known from the Craven Reef Belt of North Yorkshire, where it attached to corals and possibly bivalve shells, and was preyed upon by small durophagous animals, as indicated by repaired injuries preserved on one of the tubes.

UUID: http://zoobank.org/2ba8f87b-ec1c-4bb3-8615-115e7a527376

Lovén's Law described the position of larger basicoronal ambulacral plates in echinoids. The smaller basicoronal plates form first in ontogeny. We restate Lovén's Law to describe the position of first ambulacral plates using Carpenter's ambulacra and left (L) or right (R) as: AR, BL, CR, DL, ER, with EA the pair of ambulacra both identical and adjacent. First ambulacral plates of the Cambrian edrioasteroid, Walcottidiscus, code identically. The transition from a tri-radiate 1-1-1 pattern to the 2-1-2 ambulacral pattern of Walcottidiscus and other pentaradiate Paleozoic echinoderms results in Lovén's Law. This provides an hypothesis for the origin of Lovén's Law and predicts its widespread occurrence among echinoderm classes.

The ‘BD different’ pattern of primary brachioles in pentaradiate glyptocystitoid Rhombifera results from subterminal branching of the ambulacra. The ontogenetic sequence was triradiate, then lateral ambulacra bifurcated, and finally second brachioles developed. Positions of second brachioles of pentaradiate glyptocystitoids code as AR, BL, CR, DL, ER. Other examples of Lovén's Law occur in Diploporita, Ophiuroidea, Edrioasteroidea, Edrioblastoidea, Holothuroidea, Ophiocistioidea, and Bothriocidaris. Our working hypothesis is that Lovén's Law arose with pentaradiality. An objective test requires a reliable independent landmark across all classes, which does not exist currently. Our hypothesis is only parsimonious if Lovén's Law arose just once in echinoderm evolution and is widespread within the phylum. So far, both conditions appear to be met, but edrioasteroid first ambulacral cover plates commonly exhibit an alternative Lovén-like pattern of AL, BL, CR, DL, ER, with AB the identical adjacent ambulacra.

UUID: http://zoobank.org/4c40d338-8bfa-4515-9037-934f3bbab153

A locality in the Upper Cretaceous Two Medicine Formation of Montana preserves abundant and variable horseshoe crab tracks and trails of the ichnotaxon Kouphichnium isp. These specimens span six morphologies differing in track form and trail configuration. These differences likely reflect variations in track-maker locomotion and behavior, substrate consistency, epichnial versus hypichnial preservation, and undertrack versus true tracks. Several tracks preserve the first clear appendage impressions for an extinct horseshoe crab. This discovery adds new information to the fossil horseshoe crab diversity in the Cretaceous Period. Trackway dimensions, such as the external width across the pusher legs or of the prosomal drag mark, provide information on the track-maker size. Most trackways correspond with crabs 9–14 cm wide; the abundance but limited size range of the traces suggests the large assemblage corresponds to a mating aggregation. The trace fossil record of xiphosurids indicates that throughout their history, horseshoe crabs inhabited both marine and nonmarine settings. They were definitively present in freshwater habitats from the lower Carboniferous through at least the Paleogene. Horseshoe crab trace abundance is highest from the upper Carboniferous through the Jurassic and likely reflects two factors: true upper Carboniferous taxonomic diversity and a preponderance of suitable sites for trackway preservation in the Late Triassic and Jurassic. Cretaceous traces are uncommon, and this Two Medicine locality is the first occurrence of horseshoe crab traces in the Late Cretaceous worldwide. Overall, track abundance and diversity would seem to correspond well with the reported horseshoe crab body fossil record.