Evolutionary and biogeographical shifts in response to the Late Ordovician mass extinction

The Late Ordovician mass extinction was an interval of high extinction with inferred low ecological selectivity, resulting in little change in community structure after the event. In contrast, the mass extinction may have fundamentally changed evolutionary dynamics in the surviving groups. We investigated the phylogenetic relationships among strophomenoid brachiopods, a diverse brachiopod superfamily that was a primary component of Ordovician ecosystems. Four Ordovician families/subfamilies sampled in the analysis (Rafinesquinidae, Strophomeninae, Glyptomenidae and Furcitellinae) were reconstructed as monophyletic groups, and the base of the strophomenoid clade that dominated the Silurian recovery was reconstructed as diversifying alongside these families during the Middle Ordovician. We time‐calibrated the phylogeny and used geographical occurrences to investigate biogeographical changes in the strophomenoids through time with the R package BiogeoBEARS . Our results indicate that extinction was higher in taxa whose ranges were constrained to tropical or subtropical regions. Furthermore, our results suggest important shifts in the diversification patterns of these brachiopods after the mass extinction. While most of the strophomenoid families survived the Late Ordovician event, ecologically abundant taxonomic groups during the Ordovician were either driven to extinction, reduced in diversity, or slowly died off during the Silurian. The new abundant strophomenoid taxa derived from one clade (consisting of Silurian–Devonian groups such as Douvillinidae, Strophodontidae and Amphistrophiidae) that diversified during the post‐extinction radiation. Our results suggest the selective diversification during the Silurian radiation, rather than selective extinction in the Late Ordovician, had a greater impact on the evolutionary history of strophomenoid brachiopods.     

Oldest known Dicoelosia and Epitomyonia,deep water brachiopods from the Beiguoshan Formation (Middle Katian,Upper Ordovician), Shaanxi,north China

Abstract: A diverse brachiopod fauna from a relatively deep water carbonate facies of the Upper Ordovician Beiguoshan Formation (uppermost Caradoc – lower Ashgill, middle Katian) is characterized by small shells and contains the oldest known Dicoelosia and Epitomyonia, two diagnostic taxa of deep water brachiopod palaeocommunities during the Late Ordovician and Silurian. Three new species are recognized: Dicoelosia cordiformis sp. nov., Dicoelosia perbrevis sp. nov. and Epitomyonia fui sp. nov. These pioneer forms of the family Dicoelosiidae show a relatively high degree of morphological plasticity. The shells of Dicoelosia from the Beiguoshan Formation range from the typical slender‐lobed form with a concavoconvex profile to the strongly equibiconvex, fat‐lobed morphotype that was not known previously until the late Silurian. The Beiguoshan dicoelosiids point to an important attribute of the deep water brachiopods: small generalists with high morphological plasticity, which make them ideal candidates as progenitors for the evolution of shallow water brachiopod faunas in shelf and platform depositional environments.     

Influence of valve geometry, ornamentation, and microstructure on fractures in Late Ordovician brachiopods

Among Late Ordovician brachiopods from southeastern Indiana. strophomenids display a ratio of 4:1 parabolic to linear repaired fractures in contrast to the 1:2 ratio found for orthids and rhynchonellids. Additionally, only strophomenids display repaired elliptical fractures. The weakest parts of strophomenid and orthid-rhynchonellid shells are the regions of the adductor muscle scars and the sulcus, respectively. Fractured biplanate shells of strophomenids are commonly cleaved anteriorly to posteriorly, whereas fractures are localized on the anterior of strongly curved to geniculate conspecific specimens. Rugae on leptaenids, thickened anterior margins of the brachial valve of rafinesquinids, and the dense distribution of pseudopunctate in all strophomenids, functioned to localize anteriorly the (un)repaired linear and parabolic fractures. No sublethal fracture occurs on any biconvex shell where the height is greater than 14 mm, despite the fact that numerous specimens of certain species attained shell heights of 20 mm or more, an observation suggesting the upper limit in the gape of the crushing elements of the predator. Crushing experiments on valve 'models' reveal that the inflated equibiconvex, plicate shape of the shells of Plarysfrophia is the strongest. However, the architecture of the concave strophomenid valves is relatively stronger than the corresponding valves of many orthids and rhynchonellids when normalized for valve thickness.     

Can the Lilliput Effect be detected in the brachiopod faunas of South China following the terminal Ordovician mass extinction?

In the immediate aftermath of global extinctions, organisms were normally much smaller than those prior to these events. This ‘Lilliput Effect’ can be subdivided into two types: 1) a specific type, following the original definition of the effect which targets species-level taxa associated with inhospitable environments, and 2) a more general type, related to the reactions of higher-rank taxa above the species-level. The body sizes of brachiopods from South China through the Ordovician and Silurian transition (Late Katian, Hirnantian, and earliest Rhuddanian) are compared at generic, superfamilial, ordinal, and class levels. The results indicate that the body sizes of the taxa of lower rank (e.g. genus-level) are highly variable within these different intervals. The type of evidence for the Lilliput Effect through the end Ordovician mass extinction is thus quite different from that of the end Permian mass extinction probably reflecting differences in the intensity of these two major bioevents. However, the relationships between the contrasting trends in body-size change of some taxa of higher rank (e.g. at the ordinal-level) and the relative dominance of these taxa in the latest Ordovician and earliest Silurian suggest that the brachiopods of the two major Ordovician groups, the strophomenoids and orthoids, adopted different survival strategies during and immediately after the crisis from those of the pentamerides and rhynchonellides, that were common in Silurian assemblages.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

Decline in diversity of early Palaeozoic loosely coiled gastropod protoconchs

Quantitative data on molluscan larval conch fossil assemblages of ages ranging from the Ordovician (Argentina and the Baltic region), through Silurian (Austria), Devonian (Poland) to Carboniferous (Texas) supplement knowledge of early planktonic gastropods communities transformations. They show that larval shells of the bilaterally symmetrical bellerophontids and dextrally coiled gastropods with a hook-like straight apical portion of the first whorl initially dominated. Their relative frequency, as well as that of the sinistrally coiled ‘paragastropods’, diminished during the Ordovician and Silurian to virtually disappear in the Late Devonian and Early Carboniferous. Already during the Ordovician, diversity of larvae with gently loosely coiled first whorl increased, to be replaced then with more and more tightly coiled forms. Both the aperture constrictions and mortality peaks, probably connected with hatching and metamorphosis, indicate that the Ordovician protoconchs with hook-like first coil represent both the stage of an embryo developing within the egg envelope and a planktonic larva. The similarity of the straight apex to larval conchs of hyoliths and advanced thecosome pteropods is superficial, as these were not homologous stages in early development.     

Predation in the Ordovician and Silurian of Baltica

Signs of predation appear in the Middle Ordovician of Baltica. Shell repair dominates over the predatory borings in the Ordovician and Silurian. Predators attacked molluscs, brachiopods and tentaculitoids in the Ordovician and molluscs, tentaculitoids, brachiopods and ostracods in the Silurian. There is an increase in the number of prey species in the Late Ordovician, which could be related to the Great Ordovician Biodiversification Event. Molluscs are the favourite prey taxon in the Ordovician, but in the Silurian, molluscs became less dominant as the prey. This is probably not an artefact of preservation as Ordovician and Silurian molluscs are equally well preserved.     

Did the Late Ordovician mass extinction event trigger the earliest evolution of ‘strophodontoid’ brachiopods?

‘Strophodontoid’ brachiopods represented the majority of strophomenide brachiopods in the Silurian and Devonian periods. They are characterized by denticles developed along the hinge line. The evolution of denticles correlated with the disappearance of dental plates and teeth and were already present when the clade originated in the Late Ordovician. Specimens of Eostropheodonta parvicostellata from the Kuanyinchiao Bed (early–middle Hirnantian, uppermost Ordovician) in the Hetaoba Section, Meitan, Guizhou Province, South China, display clear fossil population variation, during a process of loss of dental plates and the development of denticles. Three phenotypes of E. parvicostellata are recognized in a single fossil bed, likely heralding a speciation process. Non-metric multidimensional scaling (NMDS) based on five key characters of genera of the Family Leptostrophiidae shows a much wider morphospace for Silurian genera than for those in the Devonian. Phylogenetic analysis of the Family Leptostrophiidae supports the NMDS analysis and mostly tracks their geological history. The fossil population differentiation in E. parvicostellata discovered between the two phases of the Late Ordovician mass extinction event (LOME) linked to a major glaciation, suggests a Hirnantian origination of the ‘strophodontoid’ morphology, and links microevolutionary change to a macroevolutionary event.     

Palaeoecology and diversity of endosymbionts in Palaeozoic marine invertebrates: Trace fossil evidence

Endosymbionts are organisms that live within the growing skeleton of a live host organism, producing a cavity called a bioclaustration. The endosymbiont lives inside the bioclaustration, which it forms by locally inhibiting the normal skeletal growth of the host, a behaviour given the new ethological category, impedichnia. As trace fossils, bioclaustrations are direct evidence of past symbioses and are first recognized from the Late Ordovician (Caradoc). Bioclaustrations have a wide geographic distribution and occur in various skeletal marine invertebrates, including tabulate and rugose corals, calcareous sponges, bryozoans, brachiopods, and crinoids. Ten bioclaustration ichnogenera are recognized and occur preferentially in particular host taxa, suggesting host-specificity among Palaeozoic endosymbionts. The diversity of bioclaustrations increased during the Silurian and reached a climax by the late Middle Devonian (Givetian). A collapse in bioclaustration diversity and abundance during the Late Devonian is most significant among endosymbionts of host coral and calcareous sponge taxa that were in decline leading up to the Frasnian-Famennian mass extinction.     

Did incumbency play a role in maintaining boundaries between Late Ordovician brachiopod realms?

Studies of ecosystem level changes in the geological record have found that the major extinction events eliminated many incumbent clades that had been ecologically dominant for long intervals. Surviving clades that had not been able to compete with the extinct incumbents were then able to evolve adaptations that allowed them to move into the niches vacated by the incumbents. Underlying this pattern is the inability of clades that do not occupy a particular niche to evolve adaptations that would permit them to compete with incumbent clades that are already successfully occupying that niche. The zoogeographic distributions of brachiopods in the Late Ordovician of Laurentia may also have been maintained by incumbency, which was disrupted by the end-Ordovician extinction event. Following the extinction event, an Early Silurian zoogeographic reorganization occurred, during which surviving clades evolved into the vacated epeiric sea niches in the Early Silurian. Just as incumbency plays a role in long-term evolutionary patterns, zoogeographic realms and provinces are also partially maintained by incumbency.     

Facies distribution patterns of some marine benthic faunas in early paleozoic platform environments

Worldwide Late Cambrian—Silurian lithofacies patterns indicate that the platforms of that time were sites of accumulation of two essentially different rocks suites: the platform carbonate rocks and the platform terrigenous rocks. Most of the platform rocks accumulated as sediments in shallow marine environments similar to those of the present but far more widely spread.Present-day marine benthic faunas are distributed in depth zones which are primarily controlled by temperature. Faunas tend to occur in substrate-related discrete clusters (communities) within each life zone; similar substrates in different depth zones commonly have different faunal associations. Individual phyletic stocks may encounter environmental optimum or near-optimum conditions in certain areas, that commonly are revealed by an abundance of species and individuals within species in each stock. Environmental optimum conditions depend upon availability of food that may be utilized, modes of feeding of the animals present, water motion, and substrate, among other factors. Organisms in past seas were distributed in patterns similar to those of the present.Carbonate platforms were particularly widespread during the latest Cambrian—Early Ordovician. Intertidal environments spread widely across those platforms during that time and characteristic faunal associations developed in them. Saukiid and related tribolites dominated latest Cambrian carbonate platform intertidal faunas. The Early Ordovician carbonate platform intertidal was dominated by archeogastropod-nautiloid cephalopod faunas. These animals were joined by tabulate corals and certain brachiopods during the latter part of the Ordovician and Silurian as prominent faunal elements in the carbonate platform intertidal—shallow subtidal. Cruziana and related trace fossils, bivalves, and certain tribolites (notably homalonotids and dalmanitids) dominated most terrigenous platform intertidal—shallow subtidal faunas of the Ordovician and Silurian.Articulate brachiopods (primarily orthoids, strophomenoids, and rhynchonelloids) appear to have been relatively prominent during the Early Ordovician in shallow subtidal environments on both carbonate and terrigenous platforms and to have spread down the bathymetric gradient into increasingly deeper subtidal areas of both platforms during the latter part of the Ordovician. Tribolites dominated faunas in relatively moderate to deep subtidal environments on both platforms during the early part of the Ordovician. They were gradually replaced by brachiopods in first the shallower, and later the deeper subtidal as dominant members of the faunas. Brachiopods (primarily pentameroids and spiriferoids) dominated nearly all Silurian warm-water subtidal environments from the shallow subtidal to the edges of the platforms.Platform uplifts in the Middle Ordovician and glacio-eustatic sea-level fluctuations in the Late Ordovician caused environmental changes across the platforms that were accompanied by marked replacements among marine benthic faunas in all environments. The distribution of Ordovician carbonate platforms and glacial deposits suggests that an Ordovician polar region may have been close to present-day equatorial Africa and that Ordovician warm temperate-tropical regions lay close to the present-day North Pole.     

Phanerozoic changes in the high-rank suprageneric diversity structure of brachiopods: Linear and non-linear effects

Phanerozoic evolution of brachiopods produced many linear (established by a comparison of successive geologic time units) and non-linear (established by a comparison of non-successive geologic time units) effects, which can be examined quantitatively by using the similarity coefficients (Czekanowski's Quantified Coefficient and Gower Index) and correlation tools. The high-rank suprageneric diversity structure accounts for a number of superfamilies in each of 26 orders for every epoch of geological time. The intensity of turnovers in this structure was generally low during the entire Phanerozoic. It was slightly stronger during the Early Paleozoic, but close to zero during the Cenozoic, when the high-rank suprageneric diversity structure of brachiopods stabilized finally. Significant turnovers took place at the Middle Cambrian–Early Ordovician, the Late Ordovician–Early Silurian, the Late Silurian–Early Devonian, the Middle Devonian–Mississippian, and the Permian–Triassic transitions. Influences of mass extinctions, both major like those End Ordovician or Permian/Triassic and minor like Early Jurassic or Jurassic/Cretaceous, on the high-rank suprageneric diversity structure of brachiopods is registered. The strongest was the consequences of the Permian/Triassic catastrophe, which perhaps even reset the brachiopod evolution. No evident direct relationships are established between intensity of turnovers and eustatic fluctuations. However, the changes in the diversity structure recorded with the Gower Index provide evidence that eustatic lowstands were more favorable for intensification in these changes.     

Reef reconstruction after extinction events of the latest ordovician in the Yangtze platform,South China

Summary Early Silurian reef reconstruction on the Yangtze Platform, in the northern part of the South China Block, is preceded by a combination of regional and global processes. During most of Ashgill time (Late Ordovician), the area was dominated by Wufeng Formation deep water graptolitic black shales. Reefs largely disappeard in the middle of the Ashgill Stage, from the northwestern margin of Cathaysian Land (southeastern South China Block), in advance of the Late Ordovician glaciation and mass extinction, due to regional sea-level changes and regional uplift, unrelated to the mass extinction itselt. Late Ordovician microbial mudmound occurrence is also found in the western margin of the Yangtze Platform, its age corresponding to theDicellograptus complexus graptolite biozone of pre-extinction time. On the Yangtze Platform, a thin, non-reef-bearing carbonate, the Kuanyinchiao Formation (=Nancheng Formation in some sites), thickness generally no more than 1m, occurs near several landmasses as a result of Hirnantian regression. Reappearance of the earliest Silurian carbonates consisting of rare skeletal lenses in the upper part of Lungmachi Formation, are correlated to theacensus graptolite biozone, early Rhuddanian of Shiqian, northeastern Guizhou, near Qianzhong Land. Carbonate sediments gradually developed into beds rich in brachiopods and crinoids in the lower part of Xiangshuyuan Formation, middle Rhuddanian. In the middle part of Xiangshuyan Formation, biostromes, containing abundant and high diversity benthic faunas such as corals, crinoids and brachiopods, show beginnings of reconstruction of reef facies. Substantial reef recovery occurred in the upper part of Xiangshuyuan Formation, lower Aeronian, as small patch reefs and biostromes. During the late Aeronian, carbonate sediments, especially reefs and reef-related facies, expanded on the upper Yangtze Platform, and radiation of reefs occurred in Ningqiang Formation, upper Telychian. The long period of reef recovery, taking several million years, remains difficult to explain, because redistribution of any refugia faunas would be expected to take place soon after the extinction. Reefs and reef-related facies subsequently declined after Telychian time due to regional uplift of the major portion of the Yangtze Platform. Carbonate facies are therefore uncommon in South China during the rest of Silurian time.     

Brachiopod morphology,life mode,and crushing predation in the Ordovician Arnheim Formation,Kentucky, USA

Predation on ancient shelled prey is an often-studied topic in paleoecology, but the early Paleozoic and the brachiopods that dominated the seafloor at that time are relatively underrepresented in the predation literature. We assessed predatory repair scar frequencies among the brachiopod genera from the Early Richmondian (Late Ordovician) Oregonia Member (Arnheim Formation) near Flemingsburg, Kentucky. We found higher repair frequencies on the concavo-convex Rafinesquina and Leptaena relative to the bi-convex genera. There were no trends in repair frequency through the stratigraphic section and no relationships between repair frequency and community diversity metrics. It is possible that concavo-convex brachiopods’ flat shape, thin shell profile, and free-lying (no pedicle attachment) lifestyle made them more likely or appealing targets of Ordovician crushing predators. It is also possible that concavo-convex brachiopods were better suited to survive crushing attacks than biconvex taxa. We also found differences in shell ornament that may influence the visibility of repair scars.     

The first report of epipunctae in non-plaesiomyid brachiopods from the lowest Silurian of southeastern China

Epipunctae are microscopic perforations that do not penetrate the shell but are confined to the outer layers of the shell. They have been known previously only in the orthidine family Plaesiomyidae. The striated walls of epipunctae are considered unique to brachiopods and thus of significance for the study of stem-group brachiopod. In this study, we report the first occurrence of epipunctae in a non-plaesiomyid brachiopod based on a well-preserved external mould of Cathaysiorthis yushanensis from the lower Silurian Shiyang Formation of Jiangxi, southeastern China. The species belongs to the Family Cathaysiorthidae which differs from the Family Plaesiomyidae in many aspects especially in dorsal internal structures. The new data imply that epipunctae may be more widespread in orthide brachiopods than previously thought, strengthening the notion that this is a plesiomorphic character of the brachiopod shell. The discovery of epipunctae, however, depends on excellent preservation of the outer surface of the shells as well as careful cleaning and observation.     

华南与缅甸奥陶纪末赫南特贝动物群中的无铰类腕足动物

在奥陶纪末生物大灭绝的两幕之间,海洋底域繁盛着赫南特贝腕足动物群（Hirnantia Fauna）。它数量丰富,分布广泛,历程短暂。以往国内外学者研究这个动物群时,常把有铰类腕足动物作为重点,而无铰类只被简单描述或列出名单,整体面貌不明。无铰类化石尽管材料有限、研究基础薄弱,但只就属的数目而言,占据了全球赫南特贝动物群总属数的近1/5,其群落、演化和环境意义不可小觑。文中专门记述上扬子区（华南古板块）观音桥层（赫南特早中期）与缅甸曼德勒地区（滇缅马苏古地体）Hwe Mawng紫色页岩段（赫南特中期）所产赫南特贝动物群的无铰类化石,计有3目、4超科[Linguloidea（舌形贝超科）、Discinoidea（平圆贝超科）、Craniopsoidea（似髑髅贝超科）和Cranioidea（髑髅贝超科）]的10属、12种,包括5个命名属种[Plectoglossa cf. davidsoni (Barrande),Schizotretinia cf. euxina (Havlí?ek), Pseudopholidops partibilis (Rong),Petrocrania cribrum (Temple),Xenocrania haimei (Reed)]和7个未定种（Trematis sp.,Paracraniops sp.,Acanthocrania sp.,Petrocrania? sp. 1,Petrocrania? sp. 2, Pseudolingula? sp. 和Orbiculoidea sp.）,其中,Pseudolingula? sp.和Orbiculoidea sp.两个未定名种因标本不佳,未正式描述。其中,以Pseudopholidops最为常见,其次是Xenocrania和Petrocrania。前两属是华南、滇缅马苏、波罗的卡、阿瓦隆尼亚、佩鲁尼卡等古板块或地体赫南特贝动物群的常见分子。它们已知限于南、北纬30°之间的低纬度地区,这可能与化石采集和研究程度有关,但更可能反映奥陶纪末全球气候仍存在分异现象。研究识别了奥陶纪晚期无铰类腕足动物7个目,可归为3个类群：灭绝目（仅神父贝目Paterinida）、消减目（多样性与丰度大幅衰减,尤其是三分贝目Trimerellida和乳孔贝目Acrotretida在Hirnantian销声匿迹）和延续目（适应能力强、忍耐阈值高,成功穿越大灾难的首幕）。这些分类单元对研究奥陶纪-志留纪交界期腕足动物群的多样性、群落生态、生物地理及宏演化等有重要意义。     

Euryhalinity of Palaeozoic articulate brachiopods

OMonotypic and very low diversity virgianid shell beds from the Upper Ordovician to Lower Silurian dolomites of North Greenland were formed in marginal marine quiet-water hypersaline environments. In the light of this evidence the salinity tolerances of other Palaeozoic articulate brachiopods is evaluated. There are only a small number of species apparently invading hypersaline or brackish environments, but it is significant considering that previously all articulate brachiopods were thought to be fully marine. Two types of occurrence are noted, those species specifically related to marginally marine environments, disappearing with the introduction of fully marine faunas, and the majority of species which extend their normal marine range into marginal conditions. No brachiopod species appears to have invaded very hypersaline or truly brackish conditions. No single group of articulate brachiopods specifically specialised in colonising marginal marine environments, apart from possibly the virgianid pentamerids. Palaeozoic, Upper Ordovician, Lower Silurian, Brachiopoda. Pentamerida, Virgianidae, Greenland, palaeoecology. hypersaline environments, brackish environments .     

Intraspecific variability in rhynchonellid brachiopods: test of a competition hypothesis

The variability in the plicae of the central fold of eight rhynchonellid brachiopods – Lepidocyclus capax (Ordovician), Stegerhynchus whitii (Silurian), Megalopterorhynchus baldwini (Devonian), 'Camarotoechia' purduei (Mississippian), Wellerella osagensis (Pennsylvanian), Leiorhynchus weeksi (Permian), 'Pugnoides'; Iriassicus (Triassic) and Tetrarhynchia sp. (Jurassic) – is inversely related to the diversity of brachiopods within the faunal assemblage. Explanations for the phenotypic variability due to taxonomic splitting, sexual polymorphism or ontogenetic development are precluded or unsupported by the morphologic analysis, although a small percentage of the morphologic variability can be ascribed to ecophenotypic differentiation by environmental stimuli.
After considerations of the abiotic influences of time, geographic location and isolation, and environmental stability and homogeneity, most of the morphologic variability in these brachiopods is attributed to the biotic influence, namely competition. Other proposed relationships, i.e. population abundance, sample size, shell size or ribbing pattern and intraspecific variability are not statistically significant.     

Lipanorthis Benedetto from the Tremadocian of NW Argentina reidentified as a dalmanellidine: Significance for the origin and early radiation of the punctate orthide brachiopods

The dalmanellidine brachiopods formed a diverse, abundant and long-lived group of Palaeozoic benthos, reaching their acme during the later Ordovician. The oldest members of the suborder occur in Arenig rocks, where both superfamilies, the Enteletoidea and Dalmanelloidea, are represented by relatively simple morphotypes. The reidentification of Lipanorthis Benedetto from the Tremadocian rocks of Argentina as a linoporellid significantly extends the record of the dalmanellidines into older strata. The relatively complex morphology of the recently described taxon and its marked morphological divergence from other Arenig dalmanellidine taxa suggest a significant pre-Ordovician record for the punctate orthide brachiopods.