The coronavirus disease pandemic was declared in March 2020, as the southern hemisphere’s winter approached. Australia expected co-circulation of severe acute respiratory syndrome coronavirus 2, influenza and other seasonal respiratory viruses. However, influenza notifications were 7,029 (March–September) compared with an average 149,832 for the same period in 2015–2109, despite substantial testing. Restrictions on movement within and into Australia may have temporarily eliminated influenza. Other respiratory pathogens also showed remarkably changed activity in 2020.

Brachiopods are (perhaps all too) familiar to any geology student who has taken an invertebrate paleontology course; they may well be less familiar to biology students. Even though brachiopods are among the most significant components of the marine fossil record by virtue of their considerable diversity, abundance, and long evolutionary history, fewer than 500 species are extant. Reconciling the geological and biological perspectives is necessary in order to test hypotheses, not only about phylogenetic relationships among brachiopods but also about their spectacular decline in diversity in the end-Permian mass extinction, which permanently reset their evolutionary trajectory. Studying brachiopod ontogeny and development, population genetics, ecology, physiology, and biogeography, as well as molecular systematics and phylogenomics, enables us to better understand the context of evolutionary processes over the short term. Investigating brachiopod morphological, taxonomic, and stratigraphic records over the Phanerozoic Eon reveals historical patterns of long-term macroevolutionary change, patterns that are simply unknowable from a biological perspective alone.

Members of the neogastropod muricid subfamily Rapaninae are abundant, shallow-water predators whose phylogeny was previously investigated by Kool (1993b), who used mainly anatomical characters. In order to deepen understanding of the evolution of this important clade and to incorporate functional, ecological fossil evidence, we performed a phylogenetic analysis based on 34 shell characters in 45 genus-level taxa, including five muricid outgroups. Cladograms based on shell characters alone differed from those founded on anatomical features these analyses differed from the phylogenetic reconstruction combining all available morphological evidence. The preferred cladogram incorporates all evidence and reveals a “Thais group” and an “Ergalatax clade” that both emerge from the derived portion of a more primitive, paraphyletic group of other rapanines. The Ocenebrinae, the other four outgroup taxa three ergalataxine taxa all lie outside the rapanine clade that includes the remaining ergalataxines as a derived subclade.We used the phylogenetic results to probe aspects of the ecological history of the Rapaninae. Our data imply that antipredatory shell defenses (elongated aperture, denticles on the inner side of the outer lip robust external spines and tubercles) evolved multiple times, mainly in post–early Miocene clades in the Indo–West Pacific region. These results support earlier nonphylogenetic inferences.We compared known prey types and methods of predation of living rapanines with their distribution on our phylogenetic tree. The plesiomorphic mode of feeding in the Rapaninae is drilling of hard-shelled prey. Feeding by other means and on such soft-bodied prey as sipunculan and polychaete worms evolved several times independently among post–early Miocene rapanines in the Indo–West Pacific. Methods of predation on hard-shelled prey that involve edge-drilling or attack by way of the aperture also evolved independently several times, but did so throughout the geographical range of the subfamily.Specialization for life on the upper shore occurred in at least eight lineages, all but two of which are confined to the Indo–West Pacific. Ecological diversification of the Rapaninae was therefore most common in the tropical Indo–West Pacific during and after early Miocene time. This diversification occurred in a setting of already high biological diversity and intense competition and predation.

No abstract

Functional consequences of the variation in geometry and morphology of the articulate brachiopod hinge mechanism are poorly understood, despite the fact that hinge structures have considerable importance in brachiopod taxonomy. Jaanusson (1971) proposed that the ability to resorb shell material during growth, particularly in the hinge structures, can be used to distinguish two groups within the articulates, the deltidiodonts and the cyrtomatodonts. He considered the two groups to be morphologically distinct, “natural” phylogenetic groups, separated by a “functional discontinuity.” In order to test the morphological, functional, and phylogenetic implications of shell resorption, comparisons of the hinge-system geometry and diductor muscle moment are made here between deltidiodont and cyrtomatodont brachiopods. A truss network composed of landmarks relevant to the valve opening mechanism is constructed to characterize hinge-system geometry. The function of the hinge mechanism is analyzed in the context of valve opening, and diductor muscle force, effort lever arm, and moment are compared between deltidiodonts and cyrtomatodonts. The distribution of resorption among brachiopods is investigated with respect to a phylogenetic hypothesis proposed by Williams and Rowell (1965a).Deltidiodont brachiopods are morphologically more variable than cyrtomatodonts, and a greater proportion of the variability is correlated with size. Deltidiodonts and cyrtomatodonts employ different strategies to open the valves; deltidiodont lever arms are relatively longer, whereas cyrtomatodont diductor muscles have relatively larger cross-sectional areas. The greatest muscle moment in deltidiodont hinge systems is realized in the maintenance of a gape angle; in the cyrtomatodont system, it is achieved at the initiation of a gape. Although they are morphologically and functionally distinct, it is doubtful that the two groups are separated by a “functional discontinuity.” Because the phylogenetic relationships among brachiopod orders are not yet resolved, the status of shell resorption as a homologue is still unclear. Resorption is manifest in at least some members of each major group of articulates (except the pentameraceans); it is likely that resorption has evolved independently several times in brachiopod evolution, in part because of the increased morphological flexibility it confers.

A new classification of the Brachiopoda is proposed to take into account recent advances in our understanding of the anatomy, shell morphology, ontogeny and phylogeny of the phylum. The use of phylogenetic analysis to help rationalize this new information did not obviate the dilemma facing all previous classifications of how best to reconcile fossil and living data. Over 95% of all recognized genera are founded on extinct species, with the greatest diversity occurring in Cambro-Ordovician times when all but two of the 26 major groups constituting the phylum first appeared. Only five of these groups survive to the present day, albeit as well dispersed representatives of the early diversity. To com pare phylogenies extrapolated from these data, phylogenetic analyses of Recent and Cambro-Ordovician groups were conducted independently by using 55 biological characters for the former group and 69 morphological (and inferred anatomical) features for the latter; only 12 characters were common to both exercises. The cladogram derived for seven Recent suprafamilial taxa, with Phoronis and cyclostome and ctenostome bryozoans as outgroups, is virtually the same as that being obtained by studies of the brachiopod genome. It is also largely compatible with the cladogram for 33 Cambro-Ordovician suprafamilial taxa with Phoronis as outgroup. This cladogram has, in turn, been subjected to stratocladistic tests and has been shown to be consistent with the stratigraphic records of the taxa analysed. A reconciliation of the genealogies derived from the Recent and Cambro-Ordovician data, represented by 14 taxa and clades (with Phoronis as outgroup), was effected by using the 19 synapomorphies characterizing these groups. The resultant cladogram shows living organophosphatic-shelled lingulids (and discinids) as a sister group to a clade of all other living brachiopods. This clade, however, includes the extinct organophosphatic-shelled paterinids and the organocalcitic-shelled craniids. The inclusion of the craniids, in particular, is a cladistic compromise that is inconsistent with genetic and some anatomical and morphological evidence. It was therefore decided to accommodate these inconsistencies by dividing the Brachiopoda into three subphyla, each typified by Recent species with early Palaeozoic ancestors and defined by easily identifiable synapomorphies. The inarticulated Linguliformea, consisting of two classes (Lingulata and Paterinata), is characterized by an organophosphatic shell with a stratiform secondary layer and by planktotrophic larvae. Its modern representatives are the lingulids and discinids. The inarticulated Craniiformea is primarily distinguished by an organocarbonate shell with a laminar secondary layer and the absence of a pedicle throughout ontogeny. The craniids are the sole Recent descendants. The mainly articulated Rhynchonelliformea is the largest subphylum as it embraces five Classes (Chileata, Obolellata, Kutorginata, Strophomenata and Rhynchonellata). Its synapomorphies include an organocarbonate shell with a fibrous secondary layer, the presence of a pedicle without a coelomic core and the development of a recognizable diductor muscle system controlling the opening of the valves about a hinge axis defined by interareas. All Recent brachiopod species articulating with cyrtomatodont teeth and sockets are rhynchonelliforms.

Heterochrony is considered to be an important and ubiquitous mechanism of evolutionary change. Three components are necessary to describe heterochrony: phylogenetic relationships, size and shape change, and timing of developmental events. Patterns and processes of heterochrony are all too often invoked before all three components have been investigated. Phylogenetic hypotheses affect the interpretation of heterochrony in three ways: rooting of a clade, topology of a clade, and character polarity. To study these effects we examined the distribution of shell microstructure, lophophore support structures, and body size in four different phylogenetic hypotheses of thecideide brachiopods (Triassic to Recent), a group of minute, cryptic, benthic marine invertebrates.Thecideides are consistently monophyletic in experiments using terebratulide, strophomenate, and spire-bearing outgroups together and separately, varying ingroup membership, and experimentally withholding certain character complexes. Thecideide monophyly is also supported by bootstrap analysis. Hypotheses of heterochrony in thecideide origins and evolution are therefore not merely artifacts of classification and can be pursued further. Using either strophomenate or spire-bearing outgroups, Triassic Thecospira is the most primitive thecideide. Trees constructed using terebratulide outgroups are rooted instead at Eudesella, a taxon derived in every other phylogenetic reconstruction, and the Triassic thecideides occupy derived rather than primitive positions.Our phylogenetic results support the traditional interpretation of the reduction or loss of the secondary fibrous shell layer as a paedomorphic pattern, whereas the evolution of lophophore support structures suggests a peramorphic pattern. Reduction in thecideide adult body size is gradual, phylogenetically, and results in an overall paedomorphic pattern. Heterochrony in these three character suites may play a role in the subsequent evolution of the clade, but apparently not in the origin of the clade, as is commonly thought. Heterotopy, rather than—or in addition to—heterochrony, may account for both the origin and evolution of the lophophore support structures and in the reduction and loss of the secondary shell layer. These phylogenetic hypotheses suggest that heterochrony can result from a complex mosaic of processes and provide specific, testable predictions about the processes responsible for producing the patterns, whether heterochronic or not. Categorizing an entire clade (such as thecideides), rather than individual characters, as globally paedomorphic may allow interesting peramorphic patterns in individual characters to be overlooked.

Abstract— The monophyletic status of the Brachiopoda and phylogenetic relationships within the phylum have long been contentious issues for brachiopod systematists. The relationship of brachiopods to other lophophore‐bearing taxa is also uncertain; results from recent morphological and molecular studies are in conflict. To test current hypotheses of relationship, a phylogenetic analysis was completed (using PAUP 3.1.1) with 112 morphological and embryological characters that vary among extant representatives of seven brachiopod superfamilies, using bryozoans, phoronids, pterobranchs and sipunculids as outgroups. In the range of analyses performed, brachiopod monophyly is well supported, particularly by characters of soft anatomy. Arguments concerning single or multiple origins of a bivalved shell are not relevant to recognizing brachiopods as a clade. Articulate monophyly is very strongly supported, but inarticulate monophyly receives relatively weak support. Unlike previous studies, the nature of uncertainties about the clade status of Inarticulata are detailed explicitly here, making them easier to test in the future. Calcareous inarticulates appear to share derived characters with the other inarticulates, while sharing many primitive characters with other calcareous brachiopods (the articulates). Experimental manipulation of the data matrix reveals potential sources of bias in previous hypotheses of brachiopod phylogeny. Although not tested explicitly, lophophorate monophyly is very tentatively supported. Molecular systematic studies of a diverse group of brachiopods and other lophophorates will be particularly welcome in providing a test of the conclusions presented here.