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A detailed account of the morphology and ontogeny of the late Middle Cambrian crustacean †Henningsmoenicaris scutula is presented. Ten successive ontogenetic stages could be recognised in the material collected from various localities in Sweden. Morphogenetic changes include the development of a pair of stalked lateral eyes and the increase in the number and size of appendages and their setal armature. Notably, early stages lack ‘proximal endites’ on all post-antennular appendages; such a spine-bearing endite has previously been thought to appear simultaneously on these limbs. In †H. scutula a single functional endite appears on the third limb in an advanced stage; an additional endite appears on the second limb and, subsequently, further endites appear on more posterior limbs. Furthermore, a single specimen of †Sandtorpia vestrogothiensis gen. et sp. nov. is described. Based on this new information and data of other ‘Orsten’ taxa, particularly those assigned already to the early evolutionary lineage of Crustacea, a small-scale computer-based phylogenetic analysis was performed. This resolved the basal branchings of Crustacea s. l. as follows: †Oelandocarididae (=†Oelandocaris oelandica+†H. scutula+†S. vestrogothiensis)+(†Cambropachycopidae (=†Goticaris longispinosa+†Cambropachycope clarksoni)+ (†Martinssonia elongata+Labrophora (=†Phosphatocopina+Eucrustacea))). Plotting ontogenetic data on the phylogram and comparing the ground pattern at every node led to the detection of three peramorphic heterochronic events in the evolutionary lineage towards Eucrustacea.

Background Thylacocephala is a group of enigmatic extinct arthropods. Here we provide a full description of the oldest unequivocal thylacocephalan, a new genus and species Thylacares brandonensis , which is present in the Silurian Waukesha fauna from Wisconsin, USA. We also present details of younger, Jurassic specimens, from the Solnhofen lithographic limestones, which are crucial to our interpretation of the systematic position of Thylacocephala. In the past, Thylacocephala has been interpreted as a crustacean ingroup and as closely related to various groups such as cirripeds, decapods or remipeds. Results The Waukesha thylacocephalan, Thylacares brandonensis n. gen. n. sp., bears compound eyes and raptorial appendages that are relatively small compared to those of other representatives of the group. As in other thylacocephalans the large bivalved shield encloses much of the entire body. The shield lacks a marked optical notch. The eyes, which project just beyond the shield margin, appear to be stalked. Head appendages, which may represent antennulae, antennae and mandibles, appear to be present. The trunk is comprised of up to 22 segments. New details observed on thylacocephalans from the Jurassic Solnhofen lithographic limestones include antennulae and antennae of Mayrocaris bucculata , and endites on the raptorial appendages and an elongate last trunk appendage in Clausocaris lithographica . Preserved features of the internal morphology in C. lithographica include the muscles of the raptorial appendage and trunk. Conclusions Our results indicate that some ‘typical’ thylacocephalan characters are unique to the group; these autapomorphies contribute to the difficulty of determining thylacocephalan affinities. While the new features reported here are consistent with a eucrustacean affinity, most previous hypotheses for the position of Thylacocephala within Eucrustacea (as Stomatopoda, Thecostraca or Decapoda) are shown to be unlikely. A sister group relationship to Remipedia appears compatible with the observed features of Thylacocephala but more fossil evidence is required to test this assertion. The raptorial appendages of Thylacocephala most likely projected 45 degrees abaxially instead of directly forward as previously reconstructed. The overall morphology of thylacocephalans supports a predatory mode of life.

More Orsten-type fossils In recent years, 'Orsten-type' fossils, characterized by remarkable preservation of the soft tissue of putative embryos and arthropod larvae, have transformed our view of the early evolution of metazoans. They are termed 'Orsten' fossils from the name of the oily alum shale deposits in Sweden where they were first found; similarly fine preservation has since been found elsewhere. Now, importantly, elsewhere includes the Lower Cambrian in China. Fossils of 'eucrustacea' — crustaceans of modern aspect — are common in the fossil record from around 500 million years ago. Newly unearthed Orsten-type fossils from China include the earliest known eucrustacean in exquisite three-dimensional detail, significantly extending the fossil record of this group. Fossils of 'eucrustacea' (crustaceans of modern aspect) are common in the fossil record from around 500 million years ago. A remarkable new 'Orsten' type Konservat-Lagerstäette from the Lower Cambrian of China reveals the earliest known eucrustacean in exquisite three-dimensional detail, significantly extending the fossil record of this important group. Crown-group crustaceans (Eucrustacea) are common in the fossil record of the past 500 million years back to the early Ordovician period, and very rare representatives are also known from the late Middle and Late Cambrian periods 1 . Finds in Lower Cambrian rocks of the Phosphatocopina, the fossil sister group to eucrustaceans 2 , imply that members of the eucrustacean stem lineage co-occurred, but it remained unclear whether crown-group members were also present at that time. ‘Orsten’-type fossils are typically tiny embryos and cuticle-bearing animals, of which the cuticle is phosphatized and the material is three-dimensional and complete with soft parts. Such fossils are found predominantly in the Cambrian and Ordovician and provide detailed morphological and phylogenetic information on the early evolution of metazoans. Here we report an Orsten-type Konservat-Lagerstätte from the Lower Cambrian of China that contains at least three new arthropod species, of which we describe the most abundant form on the basis of exceptionally well preserved material of several growth stages. The limb morphology and other details of this new species are markedly similar to those of living cephalocarids, branchiopods and copepods and it is assigned to the Eucrustacea, thus representing the first undoubted crown-group crustacean from the early Cambrian. Its stratigraphical position provides substantial support to the proposition that the main cladogenic event that gave rise to the Arthropoda was before the Cambrian 3 . Small leaf-shaped structures on the outer limb base of the new species provide evidence on the long-debated issue of the origin of epipodites 4 , 5 : they occur in a set of three, derive from setae and are a ground-pattern feature of Eucrustacea.

Background Calmanostraca is a group of branchiopod eucrustaceans, with Triops cancriformis and Lepidurus apus as most prominent representatives. Both are regularly addressed with the inaccurate tag “living fossil”, suggesting that the morphology has remained stable for several millions of years. Yet, T. cancriformis and L. apus represent only a fraction of the morphological diversity occurring in Calmanostraca, comprising the two groups Notostraca and Kazacharthra. Notostracans, commonly called tadpole shrimps, comprise the two groups Lepidurus and Triops with their elongated and rather narrow (in dorsal view) head shields . Kazacharthrans are exclusively fossil calmanostracans with broad and rather short shields, known from the Jurassic and Triassic period. One formation where fossil calmanostracans have been found is the Yixian Formation of northeastern China (Lower Cretaceous, 125–121 million years). It is part of the Jehol Group, an ecosystem known for its exceptionally well-preserved fossils, including vertebrates and plants, but also diverse arthropods. Two calmanostracan species have to date been described from the Yixian Formation, Jeholops hongi and Chenops yixianensis . Results We describe here a new calmanostracan crustacean from the Yixian Formation, Notostraca oleseni , and additionally a simple tool using a morphospace analysis to delineate different species. Measurements characterising the shield and trunk proportions of different calmanostracan species were performed, data were size-corrected, and used for this morphospace analysis to compare the different morphologies. As sclerotised body parts are more likely to be preserved in fossils than soft tissue, shields and parts of the trunk are in many cases the only morphological structures available for study. Therefore, the present analysis represents a simple tool for distinguishing between different species, as well as allowing the inclusion of specimens that are only preserved fragmentarily. Additionally, it provides a tool to demarcate the kazacharthran-like specimen described, but not formally named, by Wagner et al. (Paleontol Res. 22:57–63, 2018). Hence, we amended the description and name the species Calmanostraca hassbergella . Conclusion Our results indicate a large diversity in shield and trunk morphology in calmanostracans, in contrast to their often claimed highly conserved and uniform morphology. Especially extinct forms such as Notostraca oleseni add up to this result and point to the species richness and morphological diversity within Calmanostraca.

Fossil eggs of clam shrimps (Spinicaudata) are rare and little attention has been paid to the study of their shape and microstructures. Here, we report the discovery of exceptionally preserved three-dimensional eggs from numerous specimens of Eosestheria elliptica Chen, 1976 from the lacustrine Early Cretaceous Yixian Formation in western Liaoning, China. These three-dimensionally preserved fossil eggs display a spherical shape with smooth surface, part of the tertiary envelope, and possibly the first embryonic cuticle, which were previously unknown or ambiguous. The eggs are abundant and assumed to be attached to the exopod as in extant Spinicaudata. Moreover, the exceptional three-dimensional preservation and delicate preparation of slices of the eggs allowed us to document the microstructures and elemental composition of fossil eggs of clam shrimps from the Jehol Biota. Energy dispersive spectroscopy of the fossilized envelope revealed a calcium phosphate composition. However, the egg contents display two completely different elemental compositions. Some exhibit the same elemental composition as the envelope, whereas others has been replaced by alumino-silicate. The taphonomic process is also briefly discussed in this paper.

Here we describe a phosphatocopid arthropod with preserved soft anatomy from Lower Cambrian rocks of Shropshire, England, which provides evidence for the occurrence of Crustacea, including Eucrustacea, in the Early Cambrian. The find identifies an important, stratigraphically early source of well-preserved fossils (Konservat-Lagerstätte).

The Phosphatocopina (Crustacea, Arthropoda) is an important group of ‘Orsten-type’ preserved fossils. It is resolved as the sister-group of Eucrustacea. Here we study the labrum, the median eyes, and the inner lamella of Phosphatocopina based on the selected specimens respectively from western Hunan, South China, and Sweden. The labrum characters of different species exhibit interspecific difference. The ratio of labrum length/shield length is significant to the reconstruction of the ontogeny of phosphatocopines. The labrum characters should be introduced to the diagnosis of Phosphatocopina. In the earliest growth stage, the median eyes were inconspicuous, almost as flat as the other area of the hypstome. However, with the growth of the individuals they became conspicuous. There were two obviously concave structures on the ventral side of the hypostome in later stages, which were the positions where the median eyes should have been located. In addition, the two modes of preservation of the inner lamella, i.e., that of shrinking and that of bulging up, are interpreted as the preservational artifact herein.

The Phosphatocopina were long considered as the oldest, Cambrian, record of ostracode Crustacea. However, our detailed analysis of more than 2,500 specimens from the Upper Cambrian 'Orsten' of Sweden reveals that Phosphatocopina are neither Ostracoda nor Eucrustacea. The antenna and mandible of the phosphatocopines investigated consist of a prominent limb stem which carries a two-segmented endopod and multi-annulated exopod. This stem portion is now recognised as the fusion product of the coxa and basipod during ontogeny. Phosphatocopina share features, such as the coxa and basipod on antennae and mandibles, as well as ventral body structures such as the prominent pre-oral labrum and a single post-oral cephalic plate, the sternum (with paragnaths on the mandibular sternal portion), exclusively with the Eucrustacea. As a plesiomorphy, the ontogeny of Phosphatocopina starts with a 'head larva' with four pairs of limbs, a larva type found in the ground pattern of the Euarthropoda as well as the Crustacea. In contrast, eucrustacean ontogeny begins with a nauplius with three pairs of limbs, a 'short-head larva' or orthonauplius. Again, the post-mandibular limbs of phosphatocopines retain the plesiomorphic limb design of a basipod with a setiferous 'proximal endite', whereas Eucrustacea, including the Ostracoda, have their first post-mandibular limb differentiated into a 'mouthpart', the maxillula. Autapomorphies of Phosphatocopina include the small antennula with few terminal setae, a bivalved shield with interdorsum, and the fused coxa and basipod on antenna and mandible. We therefore consider the Phosphatocopina to be the sister group of the Eucrustacea. The respective phosphatocopine species of the Upper Cambrian of southern Sweden are restricted to a particular time zone and may be useful as stratigraphic markers.