Desmond Collins

TUZOIA: MORPHOLOGY AND LIFESTYLE OF A LARGE BIVALVED ARTHROPOD OF THE CAMBRIAN SEAS

Abstract For almost 30 years, paleontologists have analyzed evolutionary sequences in terms of simple null models, most commonly random walks. Despite this long history, there has been little discussion of how model parameters may be estimated from real paleontological data. In this paper, I outline a likelihood-based framework for fitting and comparing models of phyletic evolution. Because of its usefulness and historical importance, I focus on a general form of the random walk model. The long-term dynamics of this model depend on just two parameters: the mean (μstep) and variance (σ2step) of the distribution of evolutionary transitions (or “steps”). The value of μstep determines the directionality of a sequence, and σ2step governs its volatility. Simulations show that these two parameters can be inferred reliably from paleontological data regardless of how completely the evolving lineage is sampled. In addition to random walk models, suitable modification of the likelihood function permits consideration of a wide range of alternative evolutionary models. Candidate evolutionary models may be compared on equal footing using information statistics such as the Akaike Information Criterion (AIC). Two extensions to this method are developed: modeling stasis as an evolutionary mode, and assessing the homogeneity of dynamics across multiple evolutionary sequences. Within this framework, I reanalyze two well-known published data sets: tooth measurements from the Eocene mammal Cantius, and shell shape in the planktonic foraminifera Contusotruncana. These analyses support previous interpretations about evolutionary mode in size and shape variables in Cantius, and confirm the significantly directional nature of shell shape evolution in Contusotruncana. In addition, this model-fitting approach leads to a further insight about the geographic structure of evolutionary change in this foraminiferan lineage.

The Burgess Shale Anomalocaridid Hurdia and Its Significance for Early Euarthropod Evolution

As the largest predators of the Cambrian seas, the anomalocaridids had an important impact in structuring the first complex marine animal communities, but many aspects of anomalocaridid morphology, diversity, ecology, and affinity remain unclear owing to a paucity of specimens. Here we describe the anomalocaridid Hurdia, based on several hundred specimens from the Burgess Shale in Canada. Hurdia possesses a general body architecture similar to those of Anomalocaris and Laggania, including the presence of exceptionally well-preserved gills, but differs from those anomalocaridids by possessing a prominent anterior carapace structure. These features amplify and clarify the diversity of known anomalocaridid morphology and provide insight into the origins of important arthropod features, such as the head shield and respiratory exites.

The "evolution" of Anomalocaris and its classification in the arthropod class Dinocarida (nov.) and order Radiodonta (nov.)

The remarkable “evolution” of the reconstructions of Anomalocaris, the extraordinary predator from the 515 million year old Middle Cambrian Burgess Shale of British Columbia, reflects the dramatic changes in our interpretation of early animal life on Earth over the past 100 years. Beginning in 1892 with a claw identified as the abdomen and tail of a phyllocarid crustacean, parts of Anomalocaris have been described variously as a jellyfish, a sea-cucumber, a polychaete worm, a composite of a jellyfish and sponge, or have been attached to other arthropods as appendages. Charles D. Walcott collected complete specimens of Anomalocaris nathorsti between 1911 and 1917, and a Geological Survey of Canada party collected an almost complete specimen of Anomalocaris canadensis in 1966 or 1967, but neither species was adequately described until 1985. At that time they were interpreted by Whittington and Briggs to be representatives of “a hitherto unknown phylum.” Here, using recently collected specimens, the two species are newly reconstructed and described in the genera Anomalocaris and Laggania, and interpreted to be members of an extinct arthropod class, Dinocarida, and order Radiodonta, new to science. The long history of inaccurate reconstruction and mistaken identification of Anomalocaris and Laggania exemplifies our great difficulty in visualizing and classifying, from fossil remains, the many Cambrian animals with no apparent living descendants.

New Burgess Shale Fossil Sites Reveal Middle Cambrian Faunal Complex

Soft-bodied and lightly sclerotized Burgess shale fossils have been found at more than a dozen new localities in an area extending for 20 kilometers along the front of the Cathedral Escarpment in the Middle Cambrian Stephen Formation of the Canadian Rockies. Five different fossil assemblages from four stratigraphic levels have been recognized. These assemblages represent distinct penecontemporaneous marine communities that together make up a normal fore-reef faunal complex.

The Arthropod Alalcomenaeus cambricus Simonetta, from the Middle Cambrian Burgess Shale of British Columbia

More than 300 specimens of the previously rare arthropod Alalcomenaeus cambricus Simonetta have been collected from a new Burgess Shale locality in the Glossopleura Zone on Mount Stephen, British Columbia. This new material provides much more complete information on its morphology. The cephalon was covered by a shield. A pair of pedunculate eyes and three median eyes were followed by a large anterior appendage, the ‘great appendage’, bearing three long flagella. The two posterior head appendages, like those of the trunk, were biramous. They consisted of a segmented, inner branch, and a flap-like outer branch, fringed with long filaments. The trunk consisted of 11 somites, each protected by a tergite and bearing a pair of biramous limbs. The telson was paddle-like and fringed posteriorly with wide flat spines. Alalcomenaeus was probably a predator, moving mainly by swimming. It is now known to be one of the more abundant, widely distributed and longest ranging of Burgess Shale arthropod genera. Its affinities lie with the Arachnomorpha.

Adolescent Growth and Maturity in Nautilus

The shell of Nautilus is commonly thought to be fully mature when it has certain characteristics, namely, a white venter, a thickened apertural edge, a black band along the inside of the aperture, a pronounced ocular sinus on each side, a marked thickening of the final septum, and approximation of the final septa (Stenzel, 1964). This chapter will examine and evaluate these and other putative indicators of maturity and will endeavor to show how mature modifications develop in the ontogeny of the animal.

First Report of Sphenothallus Hall, 1847 in the Middle Cambrian

Sphenothallus Hall, 1847 is a widespread Paleozoic marine taxon that has been interpreted most recently as a tubiculous annelid or other ‘worm’ or as a thecate hydrozoan or scyphozoan cnidarian (e.g., Mason and Yochelson, 1985; Feldmann et al., 1986; Van Iten et al., 1992, 1996; Neal and Hannibal, 2000; Zhu et al., 2000). Members of this genus are characterized by a very gently tapered, finely lamellar apatitic tube bearing a closed subconical holdfast and a pair of robust longitudinal thickenings situated at the ends of the tubes greatest diameter (Zhu et al., 2000). Rarely, the tube exhibits an internal transverse wall that extends adaperturally along the inner surface of the tube proper and may also exhibit a subcylindrical terminal protuberance (Van Iten et al., 1992, figs. 5, 6). The transverse wall of Sphenothallus is essentially identical in gross morphology and microstructure to the schott (apical wall) of conulariids, an extinct group of thecate cnidarians (Van Iten, 1991, 1992a, 1992b; Jerre, 1994; Van Iten et al., 1996; Hughes et al., 2000) that may have been closely related to Sphenothallus (Van Iten et al., 1992). Clarke (1913, pl. 26, figs. 16–18; see also Van Iten et al., 1992, fig. 3) illustrated a compound specimen of S. sica consisting of numerous “daughter” tubes arranged in a highly regular manner (in opposition and evenly spaced) along a single, relatively large “parent” tube whose test wall appears to be confluent with the base of the “daughter” tubes. This compound specimen is distinctly different from the more common associations of epibiontic, holdfast-bearing Sphenothallus arranged in a less orderly fashion on Sphenothallus tubes or other shells (see for example Feldmann et al., 1986, fig. 2), and probably is best interpreted as a clonal …

Soft−part preservation in two species of the arthropod Isoxys from the middle Cambrian Burgess Shale of British Columbia, Canada

More than forty specimens from the middle Cambrian Burgess Shale reveal the detailed anatomy of Isoxys, a worldwide distributed bivalved arthropod represented here by two species, namely Isoxys acutangulus and Isoxys longissimus. I. acutangulus had a non-mineralized headshield with lateral pleural folds (= “valves” of previous authors) that covered the animals body almost entirely, large frontal spherical eyes and a pair of uniramous prehensile appendages bearing stout spiny outgrowths along their anterior margins. The 13 following appendages had a uniform biramous design—i.e., a short endopod and a paddle-like exopod fringed with marginal setae with a probable natatory function. The trunk ended with a flap-like telson that protruded beyond the posterior margin of the headshield. The gut of I. acutangulus was tube-like, running from mouth to telson, and was flanked with numerous 3D-preserved bulbous, paired features interpreted as digestive glands. The appendage design of I. acutangulus indicates that the animal was a swimmer and a visual predator living off-bottom. The general anatomy of Isoxys longissimus was similar to that of I. acutangulus although less information is available on the exact shape of its appendages and visual organs. I. longissimus is characterized by extremely long anterior and posterior spines. There are now seven Isoxys species known with soft-part preservation, I. acutangulus, I. longissimus from the Burgess Shale, I. auritus and I. curvirostratus from the Maotianshan Shale of China, I. communis and I. glaessneri from the Emu Bay Shale of Australia and I. volucris from Sirius Passet in Greenland. The frontal appendages of Isoxys strongly resemble those of other Cambrian arthropods, characterized by a single pair of “great appendages” with a shared prehensile function yet some variability in length and shape.

Misadventures in the Burgess Shale

One hundred years after Charles Doolittle Walcott found a wealth of Cambrian fossils in the Rocky Mountains of British Columbia, Desmond Collins reflects on the bumpy road of their classification.