Baring it all- undressing Cambrian ´Orsten´phosphatocopine arthropods using synchrotron X-ray tomographic microscopy

Synchrotron radiation X-ray tomographic microscopy (SRXTM) was used to virtually dissect and peel the shields off of the microscopic, bivalved phosphatocopine crustaceans in the Cambrian ‘Orsten’ type of preservation of Sweden. Doing so opened up for an array of concealed internal structures to be observed in a fully enclosed specimen of Hesslandona ventrospinata and a semi-enclosed specimen of Hesslandona angustata. For comparison, also a head-larva stage specimen of H. angustata, with shields in ‘butterfly position’, was analysed. The X-ray tomographic data sets revealed excellently preserved structures, such as labrum, sternum, antennae, mandibular and post-mandibular limbs with their minute setae, all of which were more or less dis-guised by the enclosing shields. This, moreover, allowed assignment to growth stages of the specimens, which is impossible based solely on external morphology and size.Microspherules observed inside the shields of the semi-enclosed H. angustata specimen may represent remains of food particles, and the feeding biology of phosphatocopines is discussed in detail. Our analyses suggest that phosphatocopines were particle feeders. The SRXTM technique offers the ability to three-dimensionally reconstruct the morphology in high resolution, construct virtual serial sections and study concealed structures. The resulting data allow for new structures to be revealed for previously known taxa and for new taxa to be identified, with the added benefit of not destroying the specimens in the process. Hence, we do not longer have to rely on serendipitous finds of broken and/or open phosphatocopine specimens to elucidate their diagnostic ventral morphology

Naupliar and Metanaupliar development of Thysanoessa raschii (Malacostraca, Euphausiacea) from Godthåbsfjord, Greenland, with a reinstatement of the ancestral status of the free-living Nauplius in Malacostracan evolution

The presence of a characteristic crustacean larval type, the nauplius, in many crustacean taxa has often been considered one of the few uniting characters of the Crustacea. Within Malacostraca, the largest crustacean group, nauplii are only present in two taxa, Euphauciacea (krill) and Decapoda Dendrobranchiata. The presence of nauplii in these two taxa has traditionally been considered a retained primitive characteristic, but free-living nauplii have also been suggested to have reappeared a couple of times from direct developing ancestors during malacostracan evolution. Based on a re-study of Thysanoessa raschii (Euphausiacea) using preserved material collected in Greenland, we readdress this important controversy in crustacean evolution, and, in the process, redescribe the naupliar and metanaupliar development of T. raschii. In contrast to most previous studies of euphausiid development, we recognize three (not two) naupliar (= ortho-naupliar) stages (N1-N3) followed by a metanauplius (MN). While there are many morphological changes between nauplius 1 and 2 (e.g., appearance of long caudal setae), the changes between nauplius 2 and 3 are few but distinct. They involve the size of some caudal spines (largest in N3) and the setation of the antennal endopod (an extra seta in N3). A wider comparison between free-living nauplii of both Malacostraca and non-Malacostraca revealed similarities between nauplii in many taxa both at the general level (e.g., the gradual development and number of appendages) and at the more detailed level (e.g., unclear segmentation of naupliar appendages, caudal setation, presence of frontal filaments). We recognize these similarities as homologies and therefore suggest that free-living nauplii were part of the ancestral malacostracan type of development. The derived morphology (e.g., lack of feeding structures, no fully formed gut, high content of yolk) of both euphausiid and dendrobranchiate nauplii is evidently related to their non-feeding (lecithotrophic) status

The implications of a Silurian and other thylacocephalan crustaceans for the functional morphology and systematic affinities of the group

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

Exceptionally Preserved Cambrian Trilobite Digestive System Revealed in 3D by Synchrotron-Radiation X-Ray Tomographic Microscopy

The Cambrian ‘Orsten’ fauna comprises exceptionally preserved and phosphatised microscopic arthropods. The external morphology of these fossils is well known, but their internal soft-tissue anatomy has remained virtually unknown. Here, we report the first non-biomineralised tissues from a juvenile polymerid trilobite, represented by digestive structures, glands, and connective strands harboured in a hypostome from the Swedish ‘Orsten’ fauna. Synchrotron-radiation X-ray tomographic microscopy enabled three-dimensional internal recordings at sub-micrometre resolution. The specimen provides the first unambiguous evidence for a J-shaped anterior gut and the presence of a crop with a constricted alimentary tract in the Trilobita. Moreover, the gut is Y-shaped in cross section, probably due to a collapsed lumen of that shape, another feature which has not previously been observed in trilobites. The combination of anatomical features suggests that the trilobite hypostome is functionally analogous to the labrum of euarthropods and that it was a sophisticated element closely integrated with the digestive system. This study also briefly addresses the preservational bias of the ‘Orsten’ fauna, particularly the near-absence of polymerid trilobites, and the taphonomy of the soft-tissue-harbouring hypostome

Functional morphology of giant mole crab larvae: a possible case of defensive enrollment

BACKGROUND: Mole crabs (Hippidae) are morphologically distinct animals within Meiura, the “short-tailed” crustaceans. More precisely, Hippidae is an ingroup of Anomala, the group which includes squat lobsters, hermit crabs, and numerous “false” crabs. Within Meiura, Anomala is the sister group to Brachyura, which includes all true crabs. Most meiuran crustaceans develop through two specific larval phases. The first, pelagic one is the zoea phase, which is followed by the transitory megalopa phase (only one stage). Zoea larvae are rather small, usually having a total size of only a few millimeters. Zoea larvae of some hippidan species grow significantly larger, up to 15 mm in size, making them the largest known zoea larvae of all anomalan, and probably all meiuran, crustaceans. It has been suggested that such giant larvae may be adapted to a specific defensive strategy; i.e., enrollment. However, to date such giant larvae represent a rarity. METHODS: Eight specimens of large-sized hippidan larvae from museum collections were photographed with a Canon Rebel T3i digital camera under cross-polarized light. Additionally, one of the specimens was documented with a Keyence BZ-9000 fluorescence microscope. The specimen was subsequently dissected to document all appendages in detail. UV light (377 nm) was used for illumination, consistent with the specimen’s autofluorescence capacities. For high-resolution images, composite imaging was applied. RESULTS: All specimens differ in important aspects from all other known hippidan zoea larvae, and thus probably represent either previously unreported larvae or stages of known species, or larvae of unknown species. The sixth pleon segment articulates off the telson, a condition not previously reported in hippidan zoea larvae, but only for the next larva phase (megalopa). The larvae described here thus most likely represent the ultimate pelagic larval stages, or rare cases of ‘early megalopae’. The morphological features indicate that giant hippidan larvae perform defensive enrollment. CONCLUSIONS: Our investigation indicates a larger morphological diversity of hippidan larvae than was known previously. Moreover, their assumed functional morphology, similar to the condition in certain stomatopod larvae, indicates a not yet directly observable behavior by these larvae, namely defensive enrollment. In a wider context, we are only just beginning to understand the ecological roles of many crustacean larvae. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1186/s40851-016-0052-5) contains supplementary material, which is available to authorized users