Carolin Haug

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

BackgroundThylacocephala 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.ResultsThe 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.ConclusionsOur 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.

Evolution of insect wings and development - new details from Palaeozoic nymphs

The nymphal stages of Palaeozoic insects differ significantly in morphology from those of their modern counterparts. Morphological details for some previously reported species have recently been called into question. Palaeozoic insect nymphs are important, however – their study could provide key insights into the evolution of wings, and complete metamorphosis. Here we review past work on these topics and juvenile insects in the fossil record, and then present both novel and previously described nymphs, documented using new imaging methods. Our results demonstrate that some Carboniferous nymphs – those of Palaeodictyopteroidea – possessed movable wing pads and appear to have been able to perform simple flapping flight. It remains unclear whether this feature is ancestral for Pterygota or an autapomorphy of Palaeodictyopteroidea. Further characters of nymphal development which were probably in the ground pattern of Pterygota can be reconstructed. Wing development was very gradual (archimetaboly). Wing pads did not protrude from the tergum postero‐laterally as in most modern nymphs, but laterally, and had well‐developed venation. The modern orientation of wing pads and the delay of wing development into later developmental stages (condensation) appears to have evolved several times independently within Pterygota: in Ephemeroptera, Odonatoptera, Eumetabola, and probably several times within Polyneoptera. Selective pressure appears to have favoured a more pronounced metamorphosis between the last nymphal and adult stage, ultimately reducing exploitation competition between the two. We caution, however, that the results presented herein remain preliminary, and the reconstructed evolutionary scenario contains gaps and uncertainties. Additional comparative data need to be collected. The present study is thus seen as a starting point for this enterprise.

Life habits, hox genes, and affinities of a 311 million-year-old holometabolan larva

BackgroundHolometabolous insects are the most diverse, speciose and ubiquitous group of multicellular organisms in terrestrial and freshwater ecosystems. The enormous evolutionary and ecological success of Holometabola has been attributed to their unique postembryonic life phases in which nonreproductive and wingless larvae differ significantly in morphology and life habits from their reproductive and mostly winged adults, separated by a resting stage, the pupa. Little is known of the evolutionary developmental mechanisms that produced the holometabolous larval condition and their Paleozoic origin based on fossils and phylogeny.ResultsWe provide a detailed anatomic description of a 311 million-year-old specimen, the oldest known holometabolous larva, from the Mazon Creek deposits of Illinois, U.S.A. The head is ovoidal, downwardly oriented, broadly attached to the anterior thorax, and bears possible simple eyes and antennae with insertions encircled by molting sutures; other sutures are present but often indistinct. Mouthparts are generalized, consisting of five recognizable segments: a clypeo-labral complex, mandibles, possible hypopharynx, a maxilla bearing indistinct palp-like appendages, and labium. Distinctive mandibles are robust, triangular, and dicondylic. The thorax is delineated into three, nonoverlapping regions of distinctive surface texture, each with legs of seven elements, the terminal-most bearing paired claws. The abdomen has ten segments deployed in register with overlapping tergites; the penultimate segment bears a paired, cercus-like structure. The anterior eight segments bear clawless leglets more diminutive than the thoracic legs in length and cross-sectional diameter, and inserted more ventrolaterally than ventrally on the abdominal sidewall.ConclusionsSrokalarva berthei occurred in an evolutionary developmental context likely responsible for the early macroevolutionary success of holometabolous insects. Srokalarva berthei bore head and prothoracic structures, leglet series on successive abdominal segments – in addition to comparable features on a second taxon eight million-years-younger – that indicates Hox-gene regulation of segmental and appendage patterning among earliest Holometabola. Srokalarva berthei body features suggest a caterpillar-like body plan and head structures indicating herbivory consistent with known, contemporaneous insect feeding damage on seed plants. Taxonomic resolution places Srokalarva berthei as an extinct lineage, apparently possessing features closer to neuropteroid than other holometabolous lineages.

Three-dimensionally preserved minute larva of a great-appendage arthropod from the early Cambrian Chengjiang biota

Significance Understanding the nature of the Cambrian radiation involves knowing not only the morphologies of adult animals but also their developmental pathways. However, fossil evidence of early larvae is rare. Here we describe a well-preserved 2-mm-long larva of the short-great-appendage arthropod Leanchoilia illecebrosa from the early Cambrian Chengjiang biota. The exceptional 3D preservation has allowed our microcomputed tomography analyses to resolve a series of rudimentary limb Anlagen in the posterior portion of the larva—an arrangement resembling that in late-stage eucrustacean metanauplii. L. illecebrosa is considered as an early representative of either chelicerates or of euarthropods as a whole. Therefore, this discovery provides fossil evidence that posthatching segment addition is a feature rooted in the ancestor of Euarthropoda. A three-dimensionally preserved 2-mm-long larva of the arthropod Leanchoilia illecebrosa from the 520-million-year-old early Cambrian Chengjiang biota of China represents the first evidence, to our knowledge, of such an early developmental stage in a short-great-appendage (SGA) arthropod. The larva possesses a pair of three-fingered great appendages, a hypostome, and four pairs of well-developed biramous appendages. More posteriorly, a series of rudimentary limb Anlagen revealed by X-ray microcomputed tomography shows a gradient of decreasing differentiation toward the rear. This, and postembryonic segment addition at the putative growth zone, are features of late-stage metanauplii of eucrustaceans. L. illecebrosa and other SGA arthropods, however, are considered representative of early chelicerates or part of the stem lineage of all euarthropods. The larva of an early Cambrian SGA arthropod with a small number of anterior segments and their respective appendages suggests that posthatching segment addition occurred in the ancestor of Euarthropoda.

The evolution of centipede venom claws - open questions and possible answers.

The maxilliped venom claw is an intriguing structure in centipedes. We address open questions concerning this structure. The maxillipeds of fossil centipedes from the Carboniferous (about 300 million years old) have been described, but not been depicted previously. Re-investigation demonstrates that they resemble their modern counterparts. A Jurassic geophilomorph centipede (about 150 million years old) was originally described as possessing a rather leg-like maxilliped. Our re-investigation shows that the maxilliped is, in fact, highly specialized as in modern Geophilomorpha. A scenario for the evolution of the centipede maxilliped is presented. It supports one of the two supposed hypotheses of centipede phylogeny, the Pleurostigmophora hypothesis. Although this hypothesis appears now well established, many aspects of character evolution resulting from this phylogeny remain to be told in detail. One such aspect is the special joint of the maxilliped in some species of Cryptops. Cryptops is an in-group of Scolopendromorpha, but its maxilliped joint can resemble that of Lithobiomorpha or even possess a mixture of characters between the both. Detailed investigation of fossils, larger sample sizes of extant species, and developmental data will be necessary to allow further improvements of the reconstruction of the evolutionary history of centipedes.

The presumed oldest flying insect: more likely a myriapod?

The early fossil record of insects is scarce, with only few finds in the Devonian. All these finds appear problematic and controversial, partly due to incomplete preservation and challenging interpretation of many structures. We provide details of one of these important forms, Rhyniognatha hirsti from the famous Rhynie Chert Lagerstätte with up-to-date 3D imaging techniques. The fossil has been interpreted as the remains of one of the earliest flying insects. The specimen mainly preserves the remains of the head. The structures of the mandibles have been used as a main argument for an interpretation as an insect, but these are in fact less easy to interpret. New observed structures include the remains of a head capsule and an additional pair of mouth parts. Structures formerly suggested to represent remains of the head capsule or apodemes are more likely to be representing glands of ectodermal origin. The newly observed structures do not support an interpretation as an insect. Instead they make the interpretation as a myriapod more likely, possibly as a centipede. Centipede remains from the Rhynie Chert are known from scutigeromorphs. We therefore point out that R. hirsti could be interpreted as an early centipede.

What nymphal morphology can tell us about parental investment – a group of cockroach hatchlings in Baltic Amber documented by a multi-method approach

We describe a piece of Baltic amber, about 50 million years old, which contains a group of 13 small cockroach nymphs. These specimens were documented with different methods to explore the advantages and limitations of certain imaging techniques: (1) light-based methods, such as stereo-macro photography, composite imaging under polarised light, combined with virtual surface reconstruction, and (2) X-ray micro-computed tomography, processed as volume renderings and surface reconstructions. All nymphs within the amber piece are of the same size and do not exhibit any noticeable morphological variance. Their developmental state and the way in which they are arranged indicate that these nymphs represent hatchlings. Dictyopterans (including Mantodea and Blattodea with Isoptera as ingroup) exhibit a wide range of different types of social and brood care behaviour. The evolution of this complex set of characters has been addressed repeatedly in extant-based approaches, yet deep-time aspects of this evolutionary process have rarely been addressed. The specimens described here could represent a case of a group of blattodean nymphs hatching from an ootheca, which would represent the first fossil record of such a process, or even possibly provide the first indirect evidence of social behaviour in fossil non-termite dictyopterans, indicating that it was already developed 50 million years ago.

The evolution of a key character, or how to evolve a slipper lobster

A new fossil lobster from the Cretaceous of Lebanon, Charbelicaris maronites gen. et sp. nov., is presented here, while the former species Cancrinos libanensis is re-described as Paracancrinos libanensis comb. nov. P. libanensis is shown to be closer related to the contemporary slipper lobsters than to Cancrinos claviger (lithographic limestones, Jurassic, southern Germany). A finely-graded evolutionary scenario for the slipper-lobster morphotype is reconstructed based on these fossil species and extant forms. The evolutionary changes that gave rise to the current plate-like antennae of Scyllaridae, a key apomorphy of this group, are traced back through time. The antenna of what is considered the oldest slipper lobster became petaloid and consisted of about 20 fully articulated elements. For this group the name Scyllarida sensu lato tax. nov. is introduced. In a next evolutionary step, the proximal articles became conjoined and a lateral extension appeared on peduncle element 3. The entire distal petaloid region is conjoined already at the node of Verscyllarida tax. nov. In modern slipper lobsters, Neoscyllarida tax nov., the distal region is no longer petaloid in shape but asymmetrical. The study also emphasizes that exceptionally preserved fossils need to be documented with optimal documentation techniques to obtain all available information.

Intermetamorphic developmental stages in 150 million-year-old achelatan lobsters--The case of the species tenera Oppel, 1862.

We re-investigated the fossil species tenera Oppel, 1862, an achelatan lobster (traditionally named Palinurina tenera) found in 150 million years old limestones of southern Germany. All known specimens attributed to this species show a mixture of characters, which in modern forms occur either in larvae or post-larval juveniles. Hence these specimens provide insight into a phase in ontogeny that is no longer present in the developmental sequence of any modern achelatan lobster, as the latter ones skip this phase and replace it by a drastic metamorphosis. Comparable cases have been described earlier, yet did only comprise single stages or two successive ones at most. In the here described case four developmental stages are preserved. The reconstructed ontogeny of tenera therefore represents the currently best known sequence of an early achelatan lobster that covers this specific intermediate phase. The largest known stage most likely still represents an immature of a yet undiscovered adult. These findings support the interpretation that early achelatan lobsters developed in a more gradual ontogenetic sequence than modern forms. It furthermore demonstrates that it was even more gradual than anticipated previously.

The ontogeny of Limulus polyphemus (Xiphosura s. str., Euchelicerata) revised: looking “under the skin”

In recent years, methods for investigating the exo-morphology of zoological specimens have seen large improvements. Among new approaches, auto-fluorescence imaging offers possibilities to document specimens under high resolution without introducing additional artifacts as, for example, seen in scanning electron microscopy (SEM) imaging. Additionally, while SEM imaging is restricted to the outer morphology of the current instar, auto-fluorescence imaging can be used to document changes of the outer morphology of the next instar underneath the cuticle of the current instar. Thus, reinvestigating seemingly well known species with these methods may lead to interesting new insights. Here we reinvestigate the late embryonic development of the xiphosuran (“sword tail”) Limulus polyphemus, which is often treated as a proxy for early eucheliceratan evolution. In addition to entire specimens, the appendages of the embryos were dissected off and documented separately with composite-autofluorescence microscopy. Based on these data, we can distinguish six developmental stages. These stages do not match exactly the formerly described stages, as these were largely based on SEM investigation. Our stages appear to represent earlier or later phases within what has in other studies been identified as one stage. This finer subdivision is visible as we can see the developing cuticle under the outer cuticle. In comparison to data from fossil xiphosurans, our results and those of other studies on the ontogeny of L. polyphemus point to a derived mode of development in this species, which argues against the idea of L. polyphemus as a “living fossil.”