Trunk Appendages Research Articles

Early post-embryonic polyxenidan millipedes from Saxonian amber (Eocene)

Early post-embryonic stages of arthropods are rarely found in the fossil record. They seem even more rarely reported in literature, possibly as it is more difficult to establish new species based on these specimens. Here we report two immature specimens of polyxenidan myriapods (Diplopoda, Polyxenida) preserved in Saxonian amber (Eocene). Specimen 1 represents a stage I individual with only three pairs of trunk appendages, the collum and three additional tergites. It is interpreted as a representative of the polyxenidan ingroup Polyxenidae. Specimen 2 represents a stage II individual with four pairs of trunk appendages, the collum and three well-sclerotised tergites. It is interpreted as a representative of the polyxenidan ingroup Synxenidae. We additionally discuss the challenges of further systematically interpreting immatures and fossil polyxenidans in general. We also discuss how the further exploration of data of immature polyxenidans, extant and fossil, has the potential to improve our understanding of their evolution and biology.

Untangling the Echiniscus Gordian knot: Stellariscus gen. nov. (Heterotardigrada : Echiniscidae) from Far East Asia

Species constituting the family Echiniscidae are highly derived, armoured and inhabit terrestrial habitats, in contrast to other heterotardigrades that are predominantly marine. The genus Echiniscus C.A.S. Schultze, 1840, nominal for the family Echiniscidae, is currently the most speciose tardigrade genus. However, the great morphological variability, in comparison with other heterotardigrade genera, suggests the polyphyletic character of the genus. Here, we analyse new specimens of Echiniscus pseudelegans Séméria, 1994 collected in Japan and conclude that the species as well as two other related taxa, E. elegans Richters, 1907 and E. latifasciatus Dudichev and Biserov, 2000, represent a new genus, Stellariscus, gen. nov. The new genus is characterised by a mixture of peculiar morphological apomorphies: black eyes, star-like dorsal plate sculpturing, no trunk appendages (only cephalic cirri present), two types of ventral plates, and striking sexual dimorphism in both qualitative and quantitative traits. Morphological phylogeny of the family Echiniscidae suggests a close affinity between Stellariscus, Hypechiniscus Thulin, 1928 and Pseudechiniscus Thulin, 1911. The polyphyletic status of both Echiniscus and Testechiniscus Kristensen, 1987 is also inferred. The taxonomic significance of ventral armature in echiniscid phylogeny and taxonomy is discussed.

The gnathobasic spine microstructure of recent and Silurian chelicerates and the Cambrian artiopodan Sidneyia: Functional and evolutionary implications

Gnathobasic spines are located on the protopodal segments of the appendages of various euarthropod taxa, notably chelicerates. Although they are used to crush shells and masticate soft food items, the microstructure of these spines are relatively poorly known in both extant and extinct forms. Here we compare the gnathobasic spine microstructures of the Silurian eurypterid Eurypterus tetragonophthalmus from Estonia and the Cambrian artiopodan Sidneyiainexpectans from Canada with those of the Recent xiphosuran chelicerate Limulus polyphemus to infer potential variations in functional morphology through time. The thickened fibrous exocuticle in L. polyphemus spine tips enables effective prey mastication and shell crushing, while also reducing pressure on nerve endings that fill the spine cavities. The spine cuticle of E. tetragonophthalmus has a laminate structure and lacks the fibrous layers seen in L. polyphemus spines, suggesting that E. tetragonophthalmus may not have been capable of crushing thick shells, but a durophagous habit cannot be precluded. Conversely, the cuticle of S. inexpectans spines has a similar fibrous microstructure to L. polyphemus, suggesting that S. inexpectans was a competent shell crusher. This conclusion is consistent with specimens showing preserved gut contents containing various shelly fragments. The shape and arrangement of the gnathobasic spines is similar for both L. polyphemus and S. inexpectans, with stouter spines in the posterior cephalothoracic or trunk appendages, respectively. This differentiation indicates that crushing occurs posteriorly, while the gnathobases on anterior appendages continue mastication and push food towards and into the mouth. The results of recent phylogenetic analyses that considered both modern and fossil euarthropod clades show that xiphosurans and eurypterids are united as crown-group euchelicerates, with S. inexpectans placed within more basal artiopodan clades. These relationships suggest that gnathobases with thickened fibrous exocuticle, if not homoplasious, may be plesiomorphic for chelicerates and deeper relatives within Arachnomorpha. This study shows that the gnathobasic spine microstructure best adapted for durophagy has remained remarkably constant since the Cambrian.

Gene expression reveals evidence for EGFR-dependent proximal-distal limb patterning in a myriapod.

Evolution of segmented limbs is one of the key innovations of Arthropoda, allowing development of functionally specific specialized head and trunk appendages, a major factor behind their unmatched evolutionary success. Proximodistal limb patterning is controlled by two regulatory networks in the vinegar fly Drosophila melanogaster, and other insects. The first is represented by the function of the morphogens Wingless (Wg) and Decapentaplegic (Dpp); the second by the EGFR-signaling cascade. While the role of Wg and Dpp has been studied in a wide range of arthropods representing all main branches, that is, Pancrustacea (= Hexapoda + Crustacea), Myriapoda and Chelicerata, investigation of the potential role of EGFR-signaling is restricted to insects (Hexapoda). Gene expression analysis of Egfr, its potential ligands, and putative downstream factors in the pill millipede Glomeris marginata (Myriapoda: Diplopoda), reveals that-in at least mandibulate arthropods-EGFR-signaling is likely a conserved regulatory mechanism in proximodistal limb patterning.

First report of the branchiobdellidan Holtodrilus truncatus (Annelida: Clitellata) found on the freshwater atyid shrimp Neocaridina sp. from Korea

The branchiobdellidan species Holtodrilus truncatus (Liang, 1963) is reported for the first time in Korea. The genus Holtodrilus Gelder and Brinkhurst, 1990 is monotypic within the family Xironodrilidae, and the genus and family of this species are also new to the Korean fauna. The branchiobdellidan specimens were collected from the exoskeletons of the freshwater atyid shrimp, Neocaridina sp., sampled from the 'Andeok Valley', Jeju-do Province, South Korea. Holtodrilus truncatus is characterized by its terete body form, 7/7 dental formula (with a large medial tooth and three pairs of smaller lateral teeth), and no trunk appendages. In addition, we determined a partial sequence of the mitochondrial cytochrome c oxidase subunit 1 (CO1) gene as a DNA barcode marker for H. truncatus. The present study is the first record of the branchiobdellidan occurrence on a non-crayfish host (shrimp) in Korea. To date, seven species (four genera in three families) of branchiobdellidans have been reported in the Korean fauna.

A restudy of Utahcaris orion (Euarthropoda) from the Spence Shale (Middle Cambrian, Utah, USA)

AbstractUtahcaris orion Conway Morris & Robison, 1988, from the lower middle Cambrian (Series 3, Stage 5) Spence Shale Member situated in Utah, USA, is redescribed based on a restudy of the original material. Newly recognized features, including ventral lateral eyes, trunk appendages, and a bulbous cephalic shield with subtriangular extension, reinforce similarities with Sanctacaris uncata from the middle Cambrian Burgess Shale. Both are assigned to the new family, Sanctacarididae. Sanctacaridids represent the oldest chelicerates. Their ecology and that of their nearest non-chelicerate relatives indicate that Chelicerata were plesiomorphically predatory.

Notch signaling induces cell proliferation in the labrum in a regulatory network different from the thoracic legs

The insect head is composed of several segments and an anterior non-segmental region. While patterning of the segmental region relies – at least in part – on the known trunk patterning mechanisms, development of the anterior most region remains poorly understood. The labrum is an enigmatic structure of the anterior median region (AMR) of the insect head. Based on similar development and gene expression patterns it has been suggested to be a serial homolog of trunk appendages. However, its position in the non-segmental region indicated an independent origin. In order to learn more about development of the AMR including the labrum, we screened the results of the large scale RNAi screen iBeetle to identify novel genes. We found the Notch ligand Tc-Serrate and the ubiquitin ligase Tc-mind bomb1 to be required for labrum formation. Further studies showed that Notch signaling is acting upstream of the genetic hierarchy and is required for regulating cell proliferation. We combined our work with previous data to compare the regulatory gene networks of labrum and trunk appendage formation. This reveals that despite the involvement of a similar set of genes, the genetic interactions are quite different.

Enalikter aphson is an arthropod: a reply to Struck et al . (2014)

<i>Enalikter aphson</i> is an arthropod: a reply to Struck <i>et al</i> . (2014)

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.

Multi-Segmented Arthropods from the Middle Cambrian of British Columbia (Canada)

A new arthropod,Kootenichela deppin. gen. n. sp., is described from the Stanley Glacier exposure of the middle Cambrian (Series 3, Stage 5) Stephen Formation in Kootenay National Park (British Columbia, Canada). This taxon possesses a number of primitive arthropod features such as an elongate, homonomous trunk (consisting of at least 29 segments), poorly sclerotised trunk appendages, and large pedunculate eyes associated with an anterior (ocular) sclerite. The cephalon encompasses a possible antenna-like appendage and enlarged raptorial appendages with a bipartite peduncle and three spinose distal podomeres, indicative of megacheiran (“great-appendage” arthropod) affinities. The relationships of megacheirans are controversial, with them generally considered as either stem-euarthropods or a paraphyletic stem-lineage of chelicerates. An extensive cladistic analysis resolvedKootenichelaas sister-taxon to the enigmaticWorthenella cambriafrom the middle Cambrian (Series 3, Stage 5), Burgess Shale Formation in Yoho National Park (British Columbia), which is herein reinterpreted as a megacheiran arthropod. Based on their sister-group relationship, both taxa were placed in the new family Kootenichelidae, to whichPseudoiuliafrom the Chengjiang biota is also tentatively assigned. All of these taxa possess an elongate, multi-segmented body and subtriangular exopods. This family occupies a basal position within a paraphyletic Megacheira, the immediate outgroup of Euarthropoda (crown-group arthropods). The resultant topology indicates that analyses that have resolved megacheirans as stem-chelicerates have done so because they have rooted on inappropriate taxa, e.g., trilobitomorphs and marrellomorphs.

Arthroaspis n. gen., a common element of the Sirius Passet Lagerstätte (Cambrian, North Greenland), sheds light on trilobite ancestry.

BackgroundExceptionally preserved Palaeozoic faunas have yielded a plethora of trilobite-like arthropods, often referred to as lamellipedians. Among these, Artiopoda is supposed to contain taxa united by a distinctive appendage structure. This includes several well supported groups, Helmetiida, Nektaspida, and Trilobita, as well as a number of problematic taxa. Interrelationships remain unclear, and the position of the lamellipedian arthropods as a whole also remains the subject of debate.ResultsArthroaspis bergstroemi n. gen. n. sp., a new arthropod from the early Cambrian Sirius Passet Lagerstätte of North Greenland shows a striking combination of both dorsal and ventral characters of Helmetiida, Nektaspida, and Trilobita. Cladistic analysis with a broad taxon sampling of predominantly early Palaeozoic arthropods yields a monophyletic Lamellipedia as sister taxon to the Crustacea or Tetraconata. Artiopoda is resolved as paraphyletic, giving rise to the Marrellomorpha. Within Lamellipedia, a clade of pygidium bearing taxa is resolved that can be shown to have a broadly helmetiid-like tergite morphology in its ground pattern. This morphology is plesiomorphically retained in Helmetiida and in Arthroaspis, which falls basally into a clade containing Trilobita. The trilobite appendages, though similar to those of other lamellipedians in gross morphology, have a unique outward rotation of the anterior trunk appendages, resulting in a ‘hard wired’ lateral splay, different to that observed in other Lamellipedia.ConclusionsThe combination of helmetiid, trilobite, and nektaspid characters in Arthroaspis gives important hints concerning character polarisation within the trilobite-like arthropods. The distinctive tergite morphology of trilobites, with its sophisticated articulating devices, is derived from flanged edge-to-edge articulating tergites forming a shield similar to the helmetiids, previously considered autapomorphic for that group. The stereotypical lateral splay of the appendages of lamellipedians is a homoplastic character shown to be achieved by several groups independently.

The development of the nervous system in Laevicaudata (Crustacea, Branchiopoda): insights into the evolution and homologies of branchiopod limbs and ‘frontal organs’

We investigated the development of the external morphology and of the nervous system in Lynceus biformis and Lynceus brachyurus (Laevicaudata, Branchiopoda), by using immunohistochemical methods in combination with a confocal laser scanning analysis. In both Lynceus species, a free-swimming nauplius larva, equipped with three appendages, hatches from resting eggs. Despite their close phylogenetic relationship to each other, considerable differences are present in their external morphology. Hatching L. brachyurus larvae are equipped with a large and flattened labrum, where in contrast, the L. biformis larvae possess a smaller labrum with four conspicuous posteriorly directed spines at its margin. Despite these differences, the development of the nervous system is quite similar in both species. The hatching larvae are equipped with a naupliar nervous system, and only in the more advanced stages, the development of the ventral nerve cord starts. Furthermore, our investigation into the nervous system provided insights into architecture and evolution of protocerebral sensory organs, the dorsal setae field and the dorsal frontal organ, only present in Laevicaudata. The identification of frontal filaments with an associated frontal filament organ in Lynceus revealed—after a comprehensive comparison with other branchiopods—that these organs exist throughout Branchiopoda and are comparable to those in other crustaceans. Additionally, our results of the peripheral nervous system analysis showed that the innervation pattern of the naupliar appendages (antenna and mandible) and the trunk appendages could be serially homologized, despite much difference in gross morphology of these. Based on the innervation pattern of limbs, we suggest that the larval uniramous mandibular palp, found in the larvae of all ‘large’ branchiopods, is largely exopodal of nature (contrary to most earlier statements) and that the endopodite of the trunk limbs consists of only one distal endite-like segment (confirming some earlier statements) and not of three as proposed by others.

Captopodus poschmanni gen. et sp. nov. a new stem-group arthropod from the Lower Devonian Hunsrück Slate (Germany)

A new arthropod from the Lower Devonian Hunsrück Slate is described on the basis of four specimens. The body of Captopodus poschmanni comprises a head, a trunk with an anal portion. The high number of trunk appendages (≥66 segments) is unusual. The function of one pair of cupola-like structures of the head shield is unclear. The presence of large grasping appendages in the head superficially resembles the ‘short great appendages’ of other euarthropods and grasping appendages of thylacocephalans. The phylogenetic position of the arthropod cannot be determined in detail, though several morphological aspects indicate a phylogenetic position as a stem lineage representative of the Euarthropoda, the morphology of the trunk appendages seem to indicate a more advanced phylogenetic position. This new taxon underlines the exceptional diversity of arthropods within the Hunsrück Slate in comparison to other Devonian fossil sites and highlights the significance of the Hunsrück Slate for the evolution of early arthropods.

Tardigrada from Santo Antão Island (Archipelago of Cape Verde, West Africa) with the description of a new species

Three tardigrade species from Santo Antão Island, Archipelago of Cape Verde are firstly reported from Republic of Cape Verde: Echiniscus scabrospinosus Fontoura, 1982, Echiniscus clavispinosus sp. nov. and Milnesium tardigradum Doyère, 1840. E. clavispinosus sp. nov. belongs to the ‘viridis group’ of species characterized by the green colour and plate ornamentation comprised of tubercles, fine dots and light spots, by lacking dorsal and lateral trunk appendages (cirrus A excluded) and well developed claws. E. clavispinosus sp. nov. differs from all the known species of the ‘viridis group’ in having the area between the paired plate III and the terminal plate unsculptured, in details of the cuticular ornamentation, pointed clavae instead of papillate apices, and stronger spur on internal claws. A dichotomous key to the species of the Echiniscus viridis group and a list of Macaronesian species are also given.

Soft-part anatomy of the Early Cambrian bivalved arthropodsKunyangellaandKunmingella: significance for the phylogenetic relationships of Bradoriida

Bradoriids are small bivalved marine arthropods that are widespread in rocks of Cambrian to Early Ordovician age. They comprise seven families and about 70 genera based on shield ('carapace') morphology. New bradoriid specimens with preserved soft-part anatomy of Kunmingella douvillei (Kunmingellidae) are reported from the Early Cambrian Chengjiang Lagerstätte of China together with, for the first time to our knowledge, a second bradoriid species with preserved soft parts, Kunyangella cheni (Comptalutidae). Kunmingella douvillei has a 10-segmented limb-bearing body with uniramous ninth and tenth appendages and a series of homogeneous, apparently (proximal parts not preserved) unspecialized post-antennal biramous limbs with setose leaf-shaped exopods. Each endopod consists of five podomeres. A presumed penultimate instar of Ky. cheni preserves remnants of three head and two trunk appendages, and the adult is reconstructed as having four head appendages. This material allows testing of the affinity of the Bradoriida. Kunmingella is identified as a stem crustacean in character-based analyses, through both morphological comparisons and cladistic reconstructions. Global parsimony analysis recovers a monophyletic Bradoriida as the sister group to crown crustaceans.

The bivalved arthropods Isoxys and Tuzoia with soft‐part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island, Australia)

Abstract: Abundant material from a new quarry excavated in the lower Cambrian Emu Bay Shale (Kangaroo Island, South Australia) and, particularly, the preservation of soft‐bodied features previously unknown from this Burgess Shale‐type locality, permit the revision of two bivalved arthropod taxa described in the late 1970s, Isoxys communis and Tuzoia australis. The collections have also produced fossils belonging to two new species: Isoxys glaessneri and Tuzoia sp. Among the soft parts preserved in these taxa are stalked eyes, digestive structures and cephalic and trunk appendages, rivalling in quality and quantity those described from better‐known Lagerstätten, notably the lower Cambrian Chengjiang fauna of China and the middle Cambrian Burgess Shale of Canada.

The insect upper lip (labrum) is a nonsegmental appendage‐like structure

The insect upper lip--the labrum--is a lobe-like structure anterior to the mouth opening. Whether the labrum represents a fused pair of segmental appendages or evolved independently is heavily debated. Here, we identify additional similarities of the regulatory gene network active in labrum and trunk appendages. However, we do not find a labral parasegment boundary and we show that labral Tc-Dll expression is independent of Tc-wg and Tc-hh signals. In contrast, Tc-Dll expression in all trunk appendages does require these signals. Finally, we identify crucial differences between the location of the labrum and trunk appendages: the labrum develops in median rather than lateral tissues and is part of an anterior nonsegmental tissue marked by and dependent on Tc-six3 activity. To reconcile these seeming contradictory results, we propose that the genetic network evolved in either labrum or trunk appendages and became redeployed at a novel location to form the other structure.

The mesoparasitic copepod Trifur tortuosus Wilson, 1917 (Pennellidae): Redescription with notes on its congeners

AbstractThe genus TrifurWilson, 1917 (Siphonostomatoida, Pennellidae) comprises relatively large and conspicuous parasitic copepods infecting the body surface and gills of several marine fishes. Variations in the gross morphology have been used to discriminate between the existing species of Trifur, however all previous descriptions of these species are brief, lacking many important characteristics, especially those of the head and trunk appendages, which usually were lost in the described specimens. In this paper, T. tortuosus is redescribed based on newly collected material from three species of fishes from Argentinean coasts. A detailed description of appendages morphology is given and the validity of T. merluccii Talice, 1936; T. puntaniger Thomé, 1963; T. physiculi Heegaard, 1962 and T. lotellae (Thomson, 1890) is herewith discussed.

Taphonomy of Exceptionally Preserved Crustaceans from the Upper Carboniferous of Southeastern Ireland

The Castlecomer Fauna, a Konservat-Lagerstatte from the Upper Carboniferous of Ireland, is dominated by two species of exceptionally preserved spinicaudatan conchostracan branchiopod crustaceans; over 300 articulated specimens are known from short lengths of core from two boreholes. The spinicaudatans were entombed at various angles to bedding in thin event beds deposited in a paralic environment. Cuticle is the only tissue represented. The fossils are essentially two-dimensional as specimens collapsed following decay-induced loss of the structural strength of the cuticle. Some three-dimensionality was retained, however, as clay-grade material, probably sediment, infiltrated the carapace during or after burial, preventing the valves from being compacted to the same plane as the body of the specimen. Cuticle-lined voids created by decay of the labile tissues (notably within the tubelike antennal rami) were lined or infilled by authigenic clay minerals that were subsequently metamorphosed. The antennae, mandibles, distal parts of the claspers, and the telson are preserved in excellent detail, often retaining all or most of their original relief. Other anatomical features (antennal protopods, head, proximal part of claspers, trunk appendages) are usually preserved in two dimensions, either as a film of carbon or a dark-colored stain. The antennules, maxillules, and maxillae are rarely preserved. Decay experiments confirm that this variation reflects differences in the recalcitrance (thickness and degree of sclerotization) of the cuticle that covered different parts of the body. Complete decay of certain appendages precedes appreciable deterioration in the morphology of others. Some of the disparity exhibited by exceptionally preserved fossil arthropods is likely to be taphonomic rather than biological in origin.

A Silurian ‘marrellomorph’ arthropod

Xylokorys chledophilia, a new arthropod with three-dimensionally preserved soft tissues, is described from the Herefordshire (Silurian) Lagerstätte of England. The head and trunk are covered by a relatively featureless ovoid carapace, which comprises a domed central part and a flange-like border. The head bears five pairs of appendages. The first is uniramous, with dorsal and ventral projections distally. Appendages two to four are biramous and each endopod terminates in two projections. Appendage five is possibly biramous. The hypostome is very long and subrectangular in outline. There are approximately 35 pairs of biramous trunk appendages. Each exopod comprises a long slender shaft bearing numerous fine filaments; each endopod comprises a ribbon-like shaft bearing paddle-like endites. Morphological comparisons and cladistic analyses of X. chledophilia indicate affinity with Vachonisia rogeri from the Lower Devonian Hunsrück Slate, within the marrellomorphs, but assignment to Marrellomorpha is provisional pending revision of other members of this clade. Xylokorys is the first 'marrellomorph' to be reported from the Silurian. It is interpreted as a benthic particle filter feeder, which may also have consumed prey items.