Andreas Maas

An epipodite-bearing crown-group crustacean from the Lower Cambrian.

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. Finds in Lower Cambrian rocks of the Phosphatocopina, the fossil sister group to eucrustaceans, 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. 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: they occur in a set of three, derive from setae and are a ground-pattern feature of Eucrustacea.

Cambrian Derivatives of the Early Arthropod Stem Lineage, Pentastomids, Tardigrades and Lobopodians An ‘Orsten’ Perspective

Abstract ‘Orsten’-type preservation is the phosphatisation of cuticular surfaces without any further deformation and has yielded completely three-dimensional fossils, mainly arthropods at scale of 100 μm—2 mm. Records of such exceptional fossils are now reported from several continents and from the Early Cambrian (approx. 520 Mi. years BC) to the Early Cretaceous (approx. 100 Mi. years BC). ‘Orsten’-type konservat - Lagerstatten from the Middle and Upper Cambrian have brought up mainly crustaceans and representatives of the Euarthropoda, but also derivatives of their early stem lineage. These are larval stages of Upper Cambrian pentastomids, tongue worms, which today are parasites of various tetrapods. A Middle Cambrian representative of the minute tardigrades is currently under description; it possibly represents the adelphotaxon of extant Tardigrada. New to science are fragments of a small tubular, finely annulated organism with similarly annulated segmental tubular limbs. This Upper Cambrian form, currently under investigation, may represent the first lobopodian in an ‘Orsten’-type preservation and expands their record to the late Cambrian. It shares with several of the Lower to Middle Cambrian Chengjiang and Burgess Shale fauna lobopodians the lobopodian design of its limbs, paired segmental dorsal outgrowths, and the finely annulated tubular body, which is smooth in the limb regions. New information presented by this ‘Orsten’ fossil concerns cuticular details undetectable on flattened fossils. Pentastomids, tardigrades and the onychophorans/lobopodians have been called “pro-” or “prot-arthropods” because they still lack, or partly lack, characteristic features known from later derivatives of the evolutionary line of Arthropoda. Among the three taxa, the Pentastomida are assumed to be the latest offshoot because they possess segmented limbs with pivoted joints between the articles. The now established record of all of the “pro-arthropods” and even true crustaceans in the earliest Palaeozoic implies, in our view, that the ancestry of Arthropoda lies even further back well in the Pre-Cambrian.

Phylogeny and life habits of Early Arthropods——Predation in the Early Cambrian Sea

Abstract We investigated two new arthropods from the Maotianshan-Shale fauna of southern China in the course of our research on life strategies, particularly predation, in Early Cambrian marine macrofaunal biota. One form clearly belongs to the so-called “great-appendage” arthropods, animals that were, most likely, active predators catching prey with their first pair of large, specialized frontoventral appendages. Based on this, we hypothesize that the new species and many others, if not all of the “great-appendage” arthropods were derivatives of the chelicerate stem lineage and not forms having branched off at different nodes along the evolutionary lineage of the Arthropoda. Rather, we consider the “great-appendage” arthropods as belonging to a monophyletic clade, which modified autapomorphically their first pair of appendages (antennae in general arthropod terminology) into raptorial organs for food capture. The second new form resembles another Maotianshan-Shale arthropod. Fuxianhuia protensa, in shari...

Autofluorescence imaging, an excellent tool for comparative morphology

Here we present a set of methods for documenting (exo‐)morphology by applying autofluorescence imaging. For arthropods, but also for other taxa, autofluorescence imaging combined with composite imaging is a fast documentation method with high‐resolution capacities. Compared to conventional micro‐ and macrophotography, the illumination is much more homogenous, and structures are often better contrasted. Applying different wavelengths to the same object can additionally be used to enhance distinct structures. Autofluorescence imaging can be applied to dried and embedded specimens, but also directly on specimens within their storage liquid. This has an enormous potential for the documentation of rare specimens and especially type specimens without the need of preparation. Also for various fossils, autofluorescence can be used to enhance the contrast between the fossil and the matrix significantly, making even smallest details visible. ‘Life‐colour’ fluorescence especially is identified as a technique with great potential. It provides additional information for which otherwise more complex methods would have to be applied. The complete range of differences and variations between fluorescence macrophotography and different types of fluorescence microscopy techniques are here explored and evaluated in detail. Also future improvements are suggested. In summary, autofluorescence imaging is a powerful, easy and fast‐to‐apply tool for morphological studies.

MOUTHPARTS OF THE PONTO-CASPIAN INVADER DIKEROGAMMARUS VILLOSUS (AMPHIPODA: PONTOGAMMARIDAE)

Abstract The pontogammarid amphipod Dikerogammarus villosus, originally a Ponto-Caspian faunal element, has, in the recent 15-20 years, successfully invaded various aquatic systems in Europe including Lake Constance. In these rivers and lakes it had and still has severe ecological impact on native macro-invertebrates, often eliminating the native and earlier established gammaridean species. In order to test the hypothesis that the mode of food acquisition of D. villosus is of significance for this phenomenon, we focused on the mouthparts of D. villosus, i.e., mandibles, the two pairs of maxillae and the maxillipeds using SEM. Contrary to expectations, provoked by field and laboratory observations, the results of this study show that the mouthparts of D. villosus are not highly specialized just for carnivory and predation. Indeed, the stout mandibles, with their well-developed incisors enable to kill even prey with robust integument, but other modes of feeding are possible. On the maxillulae, maxillae, and maxillipeds we found setae that can be used, together with the gnathopods and the antennae, for filtering suspended algae and other small particles from the respiration current. The same structures are involved in collecting detritus. In contrast, D. villosus does not possess any specific tools for scraping periphyton from the substrate. Feeding on macrophytes may be possibly but not very effective because the surfaces of the molars are not well suited for grinding such plant material. It is shown that D. villosus is neither a shredder, as traditionally predicated for most gammarideans, nor is it a specialized carnivore, as predation experiments proposed, but rather unspecialized. Its ability to be carnivorous and to use a wide spectrum of other food may be an important reason for the success of this invader, being an advantage compared to mainly herbivorous gammarideans, which have been eliminated in many places by D. villosus.

The Stem Crustacean Oelandocaris oelandica Re-Visited

The arthropod Oelandocaris oelandica from the upper Middle Cambrian “Orsten” of Sweden was recently recognized as a member of the early phase of crustacean evolution based on additional morphological detail from new specimens. Here we present a detailed investigation of all available material. It includes the description of a 400 µm long specimen probably representing an early developmental stage. Variation in size correlated with variation of trunk-segment numbers allowed recognition of different instars. The largest specimens do not exceed an estimated length of about 1 mm, indicating that our material may consist only of immature specimens. The characteristic, extremely long antennula of O. oelandica branches into three long rods. It may have served as the major structure to sweep in food, aided by the two subsequent appendages. These and the more posterior limbs were also responsible for locomotion. Minute pores on the outer edges of the posterior limbs and on the trunk tergites possibly contained sensilla originally, which may have served as water-current detectors. The presence of a minute proximal endite only on the third head appendage suggests a rather basal position of this species within Crustacea, because comparable developmental stages of other known stem crustaceans have such an endite on more of their appendages. Reconstruction of O. oelandica and its life attitudes (referred to the largest instar known) benefited from the application of 3D modelling. These helped, e.g., in identifying the combination of the plesiomorphic feeding function of the antennulae and the specialisation of the exopods of the next two appendages as a step toward the development of a sweep-net mode of feeding, one of the key novelties in the evolution of Crustacea. Such a mode of feeding coupled with locomotion of the three anterior appendages is still practiced in the naupliar and metanaupliar phases of many extant eucrustaceans, and even some adults.

The evolutionary history of crustacean segmentation: a fossil-based perspective.

Summary The evolution of segmentation in Crustacea, that is, the formation of sclerotized and jointed body somites and arrangement of somites into tagmata, is viewed in light of historical traits and functional constraints. The set of Early to Late Cambrian ‘Orsten’ arthropods have informed our current views of crustacean evolution considerably. These three‐dimensionally preserved fossils document ancient morphologies, as opposed to purely hypothetical models and, because of the unusual preservation of larval stages, provide us with unparalleled insight into the morphogenesis of body somites and their structural equipment. The variety of evolutionary levels represented in the ‘Orsten’ including lobopodians, tardigrades, and pentastomids also allows phylogenetic interpretations far beyond the Crustacea. The ‘Orsten’ evidence and data from representatives of the Lower Cambrian Chengjiang biota in southwestern China, including phylogenetically earlier forms, form the major source of our morphology‐based review of structural and functional developments that led toward the Crustacea. The principal strategy of arthropods is the simultaneous development of head somites, as expressed in a basal “head larva,” and a successive addition of postcephalic somites from a preterminal budding zone with progressive maturation of metameric structures. This can be recognized in the developmental patterns of extant and fossil representatives of several euarthropod taxa, particularly crustaceans, trilobites, and chelicerates (at least basally). The development of these taxa points to an early somite‐poor and free‐living hatching stage. Embryonic development to a late stage within an egg, as occurring in recent onychophorans and certain in‐group euarthropods, is regarded as achieved several times convergently.

A Eucrustacean Metanauplius from the Lower Cambrian

A new eucrustacean arthropod, Wujicaris muelleri gen. et sp. nov, is represented by a Lower Cambrian early metanauplius of strikingly modern morphology despite being the oldest known fossil of such an early immature crustacean larva. The morphology of the metanauplius closely mirrors that of corresponding developmental stages of living barnacles and copepods, and it is likely that its appendages had a similar function for feeding and locomotion. The metanauplius larva demonstrates remarkable stasis in morphology, life history, and lifestyle of (small) eucrustaceans over 525 million years, probably as a result of adaptation to a long-lasting physical niche and regime involving low Reynolds numbers and laminar current flow.

A new Late Cambrian pentastomid and a review of the relationships of this parasitic group

Pentastomida, tongue worms, are a taxon of about 130 species of parasites, living exclusively in the respiratory tracts of vertebrates. Three-dimensionally preserved Upper Cambrian larvae already demonstrate a high degree of adaptation to parasitism, striking morphological conservatism, and a high diversification by the Late Cambrian, thereby suggesting a likewise diversified host group. Not least due to their highly modified morphology, the systematic affinities of pentastomids remain controversial. The two major alternatives place the group as either close to branchiuran crustaceans or as stem-lineage derivatives of the Euarthropoda. To this set of Cambrian fossil representatives of the pentastomids we can add a new form from Lower Ordovician boundary beds from Sweden, most likely reworked from Upper Cambrian horizons. Based on this new species, named Aengapentastomum andresi gen. et sp. nov., and the available information about fossil and Recent pentastomids, we review the diverging ideas on the systematic position of this fully parasitic taxon.

Phosphatocopina - ostracode-like sister group of Eucrustacea

AbstractThe 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.