Two new eurypterids, a pterygotid Pterygotus wanggaii n. sp. and an adelophthalmoid Parahughmilleria fuea n. sp., are described from the Early Devonian (Lochkovian) Xiaxishancun Formation of Yunnan province, southwest China. This discovery represents the first record of Parahughmilleria from Gondwana and the first Pterygotus from China. Pterygotus wanggaii n. sp. is characterized by the first primary denticles (d1,d1′) being located on the middle part of the cheliceral ramus and third primary denticles (d3,d3′) elongate, even longer than the first primary denticles. Parahughmilleria fuea n. sp. is differentiated by being a large Parahughmilleria with strongly developed lateral epimera from tergites T4 to T12. These discoveries not only extend the geographical extent of the genera Pterygotus and Parahughmilleria from Euramerica to SW China, but also give insight into the similarity of ecosystem structures across the Early Devonian world. In addition, based on previous studies, the new discoveries further support the hypothesis that eurypterids underwent a crisis during the Silurian–Devonian boundary interval.

A new hibbertopterid eurypterid, Cyrtoctenus bambachi n. sp., is described from the Early Mississippian (Tournaisian) Price Formation of western Virginia. The same unit yields an unidentifiable stylonurine eurypterid. These are the first eurypterids documented from the Mississippian of North America, and only the fourth locality of this age anywhere in the world to yield eurypterids.

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Horseshoe crabs as a group are renowned for their morphological conservatism punctuated by marked shifts in morphology associated with the occupation of non-marine environments and have been suggested to exhibit a consistent developmental trajectory throughout their evolutionary history. Here, we report a new species of horseshoe crab from the Ordovician (Late Sandbian) of Kingston, Ontario, Canada, from juvenile and adult material. This new species provides critical insight into the ontogeny and morphology of the earliest horseshoe crabs, indicating that at least some Palaeozoic forms had freely articulating tergites anterior to the fused thoracetron and an opisthosoma comprising 13 segments.

Two new eurypterids, a pterygotid Erettopterus qujingensis n. sp., and a slimoniid, Slimonia sp., are described from the upper Silurian (Pridolian) Yulongsi Formation of Yunnan Province, China. Erettopterus qujingensis n. sp. is characterized by several inversely curved ramus denticles and a metastoma with a deep notch in the center. The discovery not only extends the geographic extent of the genus Erettopterus and Slimonia from Euramerica to southwest China, but also gives insight into the similarity of ecosystem structures across the Silurian world.

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One of the oldest fossil horseshoe crabs figured in the literature is Entomolithus lunatus Martin, 1809, a Carboniferous species included in his Petrificata Derbiensia. While the species has generally been included within the genus Belinurus Bronn, 1839, it was recently used as the type species of the new genus Parabelinurus Lamsdell, 2020. However, recent investigation as to the appropriate authority for Belinurus (see Lamsdell and Clapham, 2021) revealed that all the names in Petrificata Derbiensia were suppressed in Opinion 231 of the International Commission on Zoological Nomenclature (1954) for being consistently nonbinomial under Article 11.4 of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature, 1999). Despite the validation of several species names for anthozoans, brachiopods, and cephalopods described in Petrificata Derbiensia in subsequent rulings (International Commission on Zoological Nomenclature, 1956a, b), Belinurus lunatus has not been the subject of any subsequent Commission ruling or opinion, and so its use in Petrificata Derbiensia remains suppressed. The Belinurus lunatus species name was used in several subsequent publications during the 1800s, none of which made the name available under ICZN article 11.5; Parkinson (1811) is also suppressed for being nonbinomial, while Woodward (1830), Buckland (1837), Bronn (1839), and Baily (1859) refer to the species only as a synonym of Belinurus trilobitoides (Buckland, 1837) through citation to the suppressed Pretificata Derbiensia. The first author to make Belinurus lunatus an available name was Baldwin (1905), who used the name in reference to a new figured specimen from Sparth Bottoms, Rochdale, UK, but again as an explicit junior synonym of Belinurus trilobitoides (Buckland, 1837). Therefore, it was not until Eller (1938) treated B. lunatus as a distinct species from B. trilobitoides that B. lunatus became an available name as per ICZN Article 11.6.1 under the authorship of Baldwin (1905) following ICZN Article 50.7.

In the first half of the nineteenth century, a marked shift occurred in our understanding and treatment of the chelicerate fossil record, with the differentiation and recognition of entirely extinct genera for the first time. At the heart of this taxonomic revolution were the Eurypterida (sea scorpions) and Xiphosura (horseshoe crabs), although both groups were in fact considered crustaceans until Lankester's (1881) seminal comparative anatomical study of the extant xiphosuran Limulus Müller, 1785 and modern scorpions. The oldest available eurypterid genus is Eurypterus deKay, 1825; the oldest available fossil arachnid genus name is that of the scorpion Cyclophthalmus Corda, 1835. However, there has been considerable historical confusion over the oldest available fossil xiphosuran genus name, which has been recognized alternately as Belinurus König (with a publication date of either 1820 or 1851) or the synonymous Bellinurus Pictet, 1846. Most recent treatments (e.g., Selden and Siveter, 1987; Anderson and Selden, 1997; Anderson et al., 1997; Lamsdell, 2016, 2021; Bicknell and Pates, 2020) have favored Bellinurus Pictet, 1846 as the available name; however, Haug and Haug (2020) recently argued that Belinurus König, 1820 is valid and has priority, a position then followed by Lamsdell (2020), prompting a reinvestigation of the taxonomic history of the genus. Upon review, it is clear that neither of the previously recognized authorities for Belinurus are accurate and that the two candidate type species for each genus are, in fact, synonyms. Given the convoluted and at times almost illogical history of the competing names, along with the most recent controversy as to which has priority, we present a complete history of the treatment of the genus to resolve the issue.

The burgeoning field of phylogenetic paleoecology (Lamsdell et al. 2017) represents a synthesis of the related but differently focused fields of macroecology (Brown 1995) and macroevolution (Stanley 1975). Through a combination of the data and methods of both disciplines, phylogenetic paleoecology leverages phylogenetic theory and quantitative paleoecology to explain the temporal and spatial variation in species diversity, distribution, and disparity. Phylogenetic paleoecology is ideally situated to elucidate many fundamental issues in evolutionary biology, including the generation of new phenotypes and occupation of previously unexploited environments; the nature of relationships among character change, ecology, and evolutionary rates; determinants of the geographic distribution of species and clades; and the underlying phylogenetic signal of ecological selectivity in extinctions and radiations. This is because phylogenetic paleoecology explicitly recognizes and incorporates the quasi-independent nature of evolutionary and ecological data as expressed in the dual biological hierarchies (Eldredge and Salthe 1984; Congreve et al. 2018; Fig. 1), incorporating both as covarying factors rather than focusing on one and treating the other as error within the dataset.

Principal component analysis has been used to test for similarities in ecology and life habit between modern and fossil birds; however, the two main portions of the hind limb—the foot and the long bone elements—have not been examined separately. We examine the potential links between morphology, ecology, and phylogeny through a synthesis of phylogenetic paleoecological methods and morphospace analysis. Both hind limb morphologies and species’ ecologies exhibit extreme phylogenetic clumping, although these patterns are at least partially explainable by a Brownian motion style of evolution. Some morphologies are strongly correlated with particular ecologies, while some ecologies are occupied by a variety of morphologies. Within the morphospace analyses, the length of the hallux (toe I) is the most defining characteristic of the entire hind limb. The foot and hind limb are represented on different axes when all measurements are considered in an analysis, suggesting that these structures undergo morphological change separately from each other. Early birds tend to cluster together, representing an unspecialized basal foot morphotype and a hind limb reliant on hip-driven, not knee-driven, locomotion. Direct links between morphology, ecology, and phylogeny are unclear and complicated and may be biased due to sample size (~60 species). This study should be treated as a preliminary analysis that further studies, especially those examining the vast diversity of modern birds, can build upon.

Eurypterids are generally considered to comprise a mixture of active nektonic to nektobenthic predators and benthic scavenger-predators exhibiting a mode of life similar to modern horseshoe crabs. However, two groups of benthic stylonurine eurypterids, the Stylonuroidea and Mycteropoidea, independently evolved modifications to the armature of their anterior appendages that have been considered adaptations toward a sweep-feeding life habit, and it has been suggested the evolution toward sweep-feeding may have permitted stylonurines to capture smaller prey species and may have been critical for the survival of mycteropoids during the Late Devonian mass extinction. There is a linear correlation between the average spacing of feeding structures and prey sizes among extant suspension feeders. Here, we extrapolate this relationship to sweep-feeding eurypterids in order to estimate the range of prey sizes that they could capture and examine prey size in a phylogenetic context to determine what role prey size played in determining survivorship during the Late Devonian. The mycteropoid Cyrtoctenus was the most specialized sweep-feeder, with comblike appendage armature capable of capturing mesoplankton out of suspension, while the majority of stylonurines possess armature corresponding to a prey size range of 1.6–52 mm, suggesting they were suited for capturing small benthic macroinvertebrates such as crustaceans, mollusks, and wormlike organisms. There is no clear phylogenetic signal to prey size distribution and no evolutionary trend toward decreasing prey sizes among Stylonurina. Rather than prey size, species survivorship during the Late Devonian was likely mediated by geographic distribution and ability to capitalize on the expanding freshwater benthos.

The occupation of new environments by evolutionary lineages is frequently associated with morphological changes. This covariation of ecotype and phenotype is expected due to the process of natural selection, whereby environmental pressures lead to the proliferation of morphological variants that are a better fit for the prevailing abiotic conditions. One primary mechanism by which phenotypic variants are known to arise is through changes in the timing or duration of organismal development resulting in alterations to adult morphology, a process known as heterochrony. While numerous studies have demonstrated heterochronic trends in association with environmental gradients, few have done so within a phylogenetic context. Understanding species interrelationships is necessary to determine whether morphological change is due to heterochronic processes; however, research is hampered by the lack of a quantitative metric with which to assess the degree of heterochronic traits expressed within and among species. Here I present a new metric for quantifying heterochronic change, expressed as a heterochronic weighting, and apply it to xiphosuran chelicerates within a phylogenetic context to reveal concerted independent heterochronic trends. These trends correlate with shifts in environmental occupation from marine to nonmarine habitats, resulting in a macroevolutionary ratchet. Critically, the distribution of heterochronic weightings among species shows evidence of being influenced by both historical, phylogenetic processes and external ecological pressures. Heterochronic weighting proves to be an effective method to quantify heterochronic trends within a phylogenetic framework and is readily applicable to any group of organisms that have well-defined morphological characteristics, ontogenetic information, and resolved internal relationships.

The Upper Famennian (Upper Devonian) Strud locality has yielded very abundant and diversified flora as well as vertebrate and arthropod faunas. The arthropod fauna, mostly recovered from fine shales deposited in a calm, confined floodplain habitat including temporary pools, has delivered a putative insect and various crustaceans including eumalacostracans and notostracan, spinicaudatan and anostracan branchiopods. Here we present the Strud eurypterids, consisting of semi-articulated juvenile specimens assigned to Hardieopteridae recovered from the pool and floodplain deposits, as well as larger isolated fragments of potential adults recovered from stratigraphically lower, coarser dark sandy layers indicative of a higher-energy fluvial environment. The Strud fossils strongly suggest that, as proposed for some Carboniferous eurypterids, juvenile freshwater eurypterids inhabited sheltered nursery pools and migrated to higher-energy river systems as they matured.

We describe a new stylonurid eurypterid from the evaporitic potassium-salt deposits of the Upper Devonian (Famennian) Soligorsk Formation in the Pripyat Trough of Belarus. All specimens are assigned to Soligorskopterus tchepeliensis new genus new species, which represents the first formally described eurypterid species from Belarus. The occurrence of well-preserved eurypterids in these unusual evaporite deposits is most likely due to transport from freshwater stream habitats into a hypersaline setting following death. Soligorskopterus tchepeliensis n. gen. n. sp. appears to be intermediate between the traditionally considered parastylonurids and stylonurids and thus extends our understanding of stylonurid evolution in the mid-Paleozoic. Soligorskopterus n. gen. extends the occurrence of Famennian eurypterids into eastern Laurussia and the Stylonuridae into the Upper Devonian, and this taxon could be part of a global eurypterid habitat shift that took place in the Late Devonian.

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A single specimen of a new species of the chasmataspidid Diploaspis Størmer, 1972 is described from the upper Silurian (Pridoli) Phelps Member of the Fiddlers Green Formation (Bertie Group) in Herkimer County, New York State, USA. Diploaspis praecursor sp. nov. is distinguished by the shape of the posterolateral margins of the buckler, which are drawn out into angular epimera, and by the lack of elongate tubercles on the postabdomen. This discovery increases the taxonomic diversity of the Bertie Group by extending the geographic extent of Diploaspididae into North America. D. praecursor pre-dates previously known species of Diploaspis by more than 10 million years.

An isolated ramus of the pterygotid eurypterid Jaekelopterus cf. howelli from the Early Devonian (Pragian) Beartooth Butte Formation (Cottonwood Canyon, north-western Wyoming) is described. Pterygotid taxonomy and synonymy is briefly discussed with the genera Pterygotus, Acutiramus and Jaekelopterus shown to be potential synonyms. The use of cheliceral denticulation patterns as a generic-level character is discouraged in light of variations within genera and its unsuitability as a major characteristic in the other eurypterid families.

A single specimen of a new species of the synziphosurine Kasibelinurus Pickett, 1993 is described from the Lower Devonian (Lochkovian) Xiaxishancun Formation of Yunnan Province, China. The new species, K. yueya sp. nov., extends the geographic extent of the family Kasibelinuridae from the Australian palaeocontinent to the South China palaeocontinent, and the stratigraphic range back some 50 Ma from Late to Early Devonian.

Drepanopterus pentlandicus Laurie, 1892 is redescribed from the original type material along with previously unfigured specimens. A cleft metastoma is confirmed as a characteristic of the genus, along with the armature of the second and third prosomal appendages being modified into flattened blades, while the species is shown to possess a somewhat enlarged second tergite and lateral prosomal margins that overlap the first opisthosomal tergite. Different ontogenetic stages of D. pentlandicus are described, and reveal that these latter characters develop only later in ontogeny, suggesting that described specimens of Drepanopterus abonensis Simpson, 1951 may represent juveniles. Cladistic analysis of Stylonurina shows the genus Drepanopterus to be monophyletic consisting of D. pentlandicus, D. abonensis and D. odontospathus sp. nov.: it forms a basal clade of mycteropoids. Hibbertopteroidea Kjellesvig-Waering, 1959 is shown to be a junior subjective synonym of Mycteropoidea Cope, 1886, with the synonymy of many of the hibbertopterid genera hypothesised and Hibbertopterus Kjellesvig-Waering, 1959 suggested to represent juvenile specimens of Cyrtoctenus Størmer & Waterston, 1968. Hibbertopterus permianus Ponomarenko, 1985 is transferred to Campylocephalus Eichwald, 1860. The role of heterochrony in the morphological development of the mycteropoid lineage is discussed, with both hibbertopterids and mycteropids suggested to be hypertrophic and pre-displacement peramorphs respectively.