An ostracod-like arthropod with appendages preserved from the lower Ordovician of England

A tiny arthropod with a thin, possibly poorly mineralized, bivalved carapace and a pair of annulated, uniramous, probable frontal appendages is described from lower Ordovician marine mudstones in boreholes from central England. It represents only the fifth Ordovician example of a conservation deposit with soft integument preserved. Its systematic position is unresolved, but it may belong to the Ostracoda; if so, it is a rare example of an ostracod with fossilized appendages. Arthropoda, Ostracoda, appendages, Tremadoc Series, Ordovician, England.     

Possible predation damage and repair in a Quaternary marine ostracod

Evidence of damage and repair in a Quaternary fossil brackish-marine ostracod is described. The specimen is a right valve of Bicornucythere bisanensis from the Laizhou Bay, eastern China, showing distortion in the posterior region, interpreted as a possible bite mark resulting from attempted predation on the ostracod. Comparison with undamaged specimens from the same assemblage suggests that the damage occurred just after the ostracod had moulted and was still in a ‘soft-shell’ state, before the new bivalved carapace was calcified, and that subsequent calcification preserved the resulting distortion of the reticulate surface ornament. This implies that the ostracod survived the damage and was able to heal and repair its shell. As far as we are aware, this is the first such example to be documented in Ostracoda although similar occurrences have been recorded in other fossil arthropods.     

Ontogenetic changes in the carapace shape of the non-marine ostracod Eucypris virens (Jurine)

Developmental changes in carapace form (size+shape) during ontogeny have been explored in Eucypris virens (Crustacea, Ostracoda) using elliptic Fourier analysis. Clones from different geographic localities raised under controlled constant conditions (temperature and photoperiod) were used to characterize developmental pathways in the species. A larger data set including field populations and laboratory populations cultured under a range of environmental conditions were used to infer influence of environmental factors on carapace shape changes during ontogeny. Size changes between consecutive juvenile stages support empirical laws describing the doubling of ostracod volume at each moult. Ontogenetic changes point out the remarkable influence of environmental conditions on carapace shape.     

同种介形类壳体的多态性兼及化石介形类的物种鉴定

介形类个体发育过程中壳体形状及各种构造的多态性,以及各种外界环境因素对分形类壳体各种分类特征形态变化的影响是十分明显的。该文较系统地阐述了在介形类个体发育中壳体形状、大小、肌痕、边缘毛细管带、边庭、铰合构造及纹饰的变化,并介绍了成年期雌、雄性壳体及左、右壳瓣在形态上的差异;列举了环境因素(主要是水体温度、含盐度、深度)对介形类壳体各分类特征形态变化的影响。基于以上事实,提出了古生物工作者在介形类化石鉴定中应注意的一些问题并讨论了某些描述术语的具体标准。     

Reproductive strategy of an isopod Onisocryptus ovalis,parasitizing a bioluminescent myodocope ostracod Vargula hilgendorfii

The functional morphology and the reproductive strategy of a parasitic isopod Onisocryptus ovalis in a bioluminescent ostracod Vargula hilgendorfii as its final host were studied based on video and SEM observations. During its lifetime, Onisocryptus ovalis dramatically metamorphoses several times, changing sex from male to female in the final host's carapace. At nearly the last ontogenetical stage, the parasite anchors its body with a pair of thoracopods to the posterodorsal region of the host ostracod's trunk and loses all the other appendages and thus its mobility as well. Thereafter, the parasite reverses bodily orientation during the final moulting so as to locate its mouth in the midst of the host eggs, and finally consumes them, leaving only the egg membrane. Such a mode of feeding of the parasite following the fixation of the body is interpreted in terms of the adaptation to escape elimination from the ostracod carapace by the host's cleaning appendages (the seventh limbs) and to obtain as much space as possible for the parasite's own eggs/embryos at the sacrifice of the mother's mobility. The synchronization between the timing of metamorphosis of the parasite and the reproductive cycle of the host animal can be expected to guarantee the parasite the opportunity to exploit sufficient nutrition from the eggs of the host.     

Preface: The phylogeny, fossil record and ecological diversity of ostracod crustaceans

After more than two centuries of research, more than 65,000 living and fossil ostracod species have been described and studied, yet much remains to be learned about this ancient, widespread and diverse group of bivalved arthropods. Their higher classification and phylogeny are subjects of vigorous debate, as is their position in the broader picture of crustacean phylogeny. At the same time, major advances in our understanding of ostracod lineages and their relationships are resulting from the application of innovative approaches and techniques. This preface provides a contextual overview of the 15 contributions to this volume, which resulted from the 14th International Symposium on Ostracoda (ISO2001) held in 2001at Shizuoka, Japan. As such it provides a cross-section of topics at the forefront of research on the evolution and diversity of Ostracoda, and indicates directions for future work.     

Larval development of Japanese 'conchostracans': part 1, larval development of Eulimnadia braueriana (Crustacea, Branchiopoda, Spinicaudata, Limnadiidae) compared to that of other limnadiids

As a part of a project to compare phylogenetically the larval or embryonic development of all major taxa of the Branchiopoda (Crustacea), the larval development of the Japanese spinicaudatan clam shrimp Eulimnadia braueriana Ishikawa, 1895, is described. Seven naupliar stages are recognized, based mainly on significant morphological differences between them, but in one case, on size alone. The seven stages range in length from 156 µm to 760 µm. Nauplius 1 is nonfeeding with incompletely developed and nonfunctional feeding structures. Nauplius 2 has apparently functional feeding structures, including a well-developed mandibular gnathobase, setulate protopodal endites of the antennae, and setules on various setae involved in swimming and food manipulation. Nauplius 3 is morphologically identical to Nauplius 2, but more than 50% larger. In nauplius 4, the coxal endite (naupliar process) of the antennae develops a bifid tip. Nauplius 5 has a lateral pair of primordial carapace lobes, and the first 4–5 pairs of trunk limb buds are weakly developed, making the anterior part of the trunk wider than the posterior. In nauplius 6, five pairs of trunk limb buds are visible externally and a small carapace has appeared, reaching approximately to trunk limbs 2; also, the pair of large buds behind the mandibles in previous stages has become divided into a large, anterior, setose bud and two smaller, posterior buds. The identities of these structures as either paragnaths or maxillules/maxillae remain uncertain. In nauplius 7, about six pairs of trunk limb buds are visible externally. The general morphology of the nauplius larvae of E. braueriana is much like those of the well-known Limnadia lenticularis (Linnaeus, 1758) and Eulimnadia texana Packard, 1871, including an elongate, lanceolate labrum; however, because of various heterochronies, the correspondence between the larval sequences of these species is not perfect. There is even less correspondence with the 5-stage larval development reported for Limnadia stanleyana King, 1855, and the spatulate labra of that species and Jmnadia spp. are different from those of other known limnadiid nauplii. The larvae of E. braueriana possess many typical (and synapomorphic) branchiopod features, such as the general morphology of the appendages involved in feeding and the mode of trunk limb development, while the small buds of the first antennae and the exact number and development of the parts of the trunk limbs are typical for the Spinicaudata.     

Thylacocephala (Arthropoda: Crustacea?) from the Cretaceous of Lebanon and implications for thylacocephalan systematics

An intensive study of a collection of arthropods from the Cretaceous of Lebanon, formerly referred to as stomatopod larvae, reveals that these forms belong to the problematical arthropod class Thylacocephala. The species Protozoea hilgendorfi , P. damesi , and Pseuderichthus cretaceus display defining thylacocephalan characters such as a carapace enclosing the entire body bearing a large anterior optic notch; three pairs of large, raptorial appendages; and a posterior battery of small swimming limbs associated with muscle segments. Unique characters of at least the genus Protozoea are the numerous 'pits' covering the entire carapace, elongated anterior rostral and posterior spines, and a dorsal hinge indicative of a truly bivalved carapace. We note several anatomical curiosities that provide some insight into the anatomy, ecology and phylogenetic relationships of these and other thylacocephalan species. Structures on the carapace surface of thylacocephalans may represent sensory organs comparable to the receptors seen on thecostracan crustaceans. The affinities of the three species described are still not entirely clear, but a phylogenetic analysis of the entire group may resolve this.     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian (Series 3, Stage 5) Burgess Shale Formation (Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton‐benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown‐group arthropods), were found to have occurred among basal bivalved arthropods.     

The Early Cambrian colonization of pelagic niches exemplified by Isoxys (Arthropoda)

The anatomy of the bivalved arthropod Isoxys (Early and Middle Cambrian) is reconstructed, based on new evidence from soft parts and exoskeletal design and on a critical review of previous work. Isoxys had a long segmented body flanked with a pair of short antennules, followed by a series of 14 biramous appendages provided with long paddle-like exopods concealed under a widely open bivalved carapace folded dorsally and bearing long cardinal spines. The close resemblance between Isoxys and Recent pelagic crustaceans (halocyprid ostracods, larval stages of malacostracans) indicates that Isoxys was probably an active epipelagic swimmer (evidence from soft parts, carapace design and distributional pattern). Some species (e.g. I. auritus and I. paradoxus from the Maotianshan Shale biota; Early Cambrian) may have lived in the vicinity of the bottom either permanently or temporarily, whereas others may have had ecological preferences for more open-marine settings. The spinosity of Isoxys had a possible role in predatorial deterrence rather than in buoyancy control or in retarding sinking within the water column. The presence of Isoxys in the Maotianshan Shale of S. China indicates that arthropods had already colonized midwater niches by the Early Cambrian. The midwater communities of the Maotianshan Shale comprised numerous other invertebrates, such as abundant medusiform eldonids, vetulicolids, chordates and possibly early vertebrates. This contradicts the opinion that pelagic communities remained poorly developed until late Cambrian/Ordovician times and that the occupation of the midwater niches largely post-dates the initial diversification of the benthic faunas.     

The soft anatomy of Isoxys minor from the Guanshan fauna,lower Cambrian of Southwest China

Isoxys is a very common Cambrian bivalved arthropod, specimens of which are normally preserved only as valves. The discovery of the soft anatomy of Isoxys may greatly assist understanding affinities and functional morphology. Isoxys minor Luo and Hu in Luo et al., 2008 is the most common representative of all animal species known from the lower Cambrian Guanshan fauna (Cambrian Series 2, Stage 4) at the Shitangshan Section, near Kunming, Yunnan Province, Southwest China. Here we describe and reconstruct the morphology of I. minor on the basis of newly illustrated fossils and a few new specimens that bear soft-parts including new discovery of frontal grasping appendages. Like the soft anatomy of other known Isoxys, it bears a pair of large stalked eyes, a pair of specialized frontal grasping appendages, approximately 12–14 paired biramous limbs, and a helm-like tail exposed outside the valves.     

Functional morphology of Palaeozoic ostracods: phylogenetic implications

Recent discussions of ostracod systematics have focused on soft anatomy, both as seen in extant groups and as recorded by rare examples of special fossil preservation. The position of the fossil Palaeocopina and Leperditicopida, for which no substantial soft part evidence has yet been found, remains in the view of post-Palaeozoic workers uncertain, with some doubt as to whether they should be retained within the Ostracoda. The evolution of carapace bauplans (e.g. the development of brood pouches and lobal structures in palaeocopids as well as the development of adductor muscle scar patterns, calcified inner lamellae and carapace incisures in podocopines) is discussed in relation to presumed soft anatomy. It seems possible to distinguish between plesiomorphic (ancestral, simple) and apomorphic (derived, advanced) characters and consider their significance in ostracod systematics. Although the presumed ‘protostracod’ is not known, the combination of soft anatomy, carapace architecture and behaviour (feeding techniques, brood care) provide evidence of a general body plan which appeared (at the latest) during the Ordovician and continuously evolved towards the anatomy of modern ostracods. In parallel lineages, plesiomorphic forms have died out (leperditicopids and most palaeocopines as well as metacopines), while apomorphic lineages (‘drepanellid archetype’ of palaeocopines; resistant platycopines, podocopines and myodocopines) have survived all extinction events. The evidence supports the retention of the Palaeocopina (and probably the Leperditicopida) in the Ostracoda.     

A redescription of Cythere japonica Hanai, 1959 (Podocopida: Ostracoda)

Cythere japonica was proposed by Hanai (1959) as a new species of the genus Cythere , from the Pleistocene Sawane Formation in Sado Island, Niigata Prefecture, Japan. The characters of carapace were already known, but the appendages have not been previously described. The authors formerly considered that this species should be separate from the genus Cythere sensu stricto because it has twice as many sieve-type pore systems as typical Cythere species, and a markedly higher carapace. The existence of living Cythere japonica in the tidal zone of north-west Japan is confirmed, and its taxonomic position re-examined on the basis of its appendages and the ontogeny of pore systems. The appendages, except for the copulatory organ, are almost identical with those of other Cythere species, and their pore systems share the same pattern in and before the A-4 moult stage. On the basis of these features this species should be retained in the genus Cythere. Phylogenetic relationships are considered on the basis of the ontogeny of pore systems.
The abdominal segments of podocopid Ostracoda, which have always been regarded as difficult to observe because of their fusion, are shown clearly by the SEM.     

Generation of bioengineered feather buds on a reconstructed chick skin from dissociated epithelial and mesenchymal cells

Various kinds of in vitro culture systems of tissues and organs have been developed, and applied to understand multicellular systems during embryonic organogenesis. In the research field of feather bud development, tissue recombination assays using an intact epithelial tissue and mesenchymal tissue/cells have contributed to our understanding the mechanisms of feather bud formation and development. However, there are few methods to generate a skin and its appendages from single cells of both epithelium and mesenchyme. In this study, we have developed a bioengineering method to reconstruct an embryonic dorsal skin after completely dissociating single epithelial and mesenchymal cells from chick skin. Multiple feather buds can form on the reconstructed skin in a single row in vitro. The bioengineered feather buds develop into long feather buds by transplantation onto a chorioallantoic membrane. The bioengineered bud sizes were similar to those of native embryo. The number of bioengineered buds was increased linearly with the initial contact length of epithelial and mesenchymal cell layers where the epithelial‐mesenchymal interactions occur. In addition, the bioengineered bud formation was also disturbed by the inhibition of major signaling pathways including FGF (fibroblast growth factor), Wnt/β‐catenin, Notch and BMP (bone morphogenetic protein). We expect that our bioengineering technique will motivate further extensive research on multicellular developmental systems, such as the formation and sizing of cutaneous appendages, and their regulatory mechanisms.     

Appendage Homologies and the First Record of Eyes in Platycopid Ostracods, with the Description of a New Species of Keijcyoidea (Crustacea: Ostracoda) from Japan

A median nauplius eye is reported for the first time in a platycopid ostracod, a group hitherto considered to be blind. A new species of the platycopid ostracod genus Keijcyoidea is described from coastal rocky marine habitats on the Pacific coast of Japan. Observations of living specimens in the laboratory show that it is capable of burrowing to a depth of several millimeters in sandy sediment, using the first two head appendages (antennulae and antennae) and the furca. Females brooded a maximum of five eggs in the posterior brood space of the carapace. The homologies and phylogenetic implications of the trunk segmentation and limbs are discussed, paying particular attention to the sexually dimorphic fifth and sixth limbs; the copulatory appendages of both sexes are interpreted as being attached to trunk segments T6–T7 (counting from the posterior; T1 = posteriormost segment).     

Abundant assemblage of Ostracoda (Crustacea) in Mexican Miocene amber sheds light on the evolution of the brackish-water tribe Thalassocypridini

Amber inclusions of fully aquatic animals are rare, most of them isolated specimens of water-bound insect larvae. In contrast, we here describe an ample, speciose fauna of 262 fully aquatic individuals of Ostracoda (Crustacea) from Mexican amber of Early Miocene age. This fauna was trapped whilst under water swimming around trunks of resin-producing trees in a brackish lagoon. Ostracod crustaceans are typically around a millimeter in length, and are known for their mostly well-calcified bivalved carapaces that account for their unparalleled fossil record in arthropods. However, in the Chiapas amber we found representatives of the tribe Thalassocypridini, which are characterised by lightly calcified carapaces and therefore lack a substantial fossil record. Embedded in amber, this ‘drawback’ becomes a clear benefit because the unobstructed view onto the appendages otherwise hidden in the carapace allowed us to identify nine ostracod species, six of them new to science. The exceptional number of individuals permitted insights into population composition, reproduction, taphonomy and into micro-environmental parameters of the amber’s place of formation. Based on ecological data available for Recent Thalassocypridini species, we posit that this tribe reached a peak radiation in open lagoonal settings during the Miocene, and shifted towards anchialine environments by the Recent.

http://zoobank.org/lsid:zoobank.org:pub:3C93A06B-F86B-404F-985D-139897F97EF3     

Origin of the novel chemoreceptor Aesthetasc "Y" in Ostracoda: morphogenetical thresholds and evolutionary innovation

SUMMARY The morphology and developmental processes of the two types of ostracod chemoreceptors, the Aesthetasc "Y" and the "Grouped setae," were compared. Cypridoidea and Pontocypridoidea, belonging to Cypridocopina, have a large baseball bat-like seta as an autapomorphic character on the second antenna, whereas most ostracod taxa with plesiomorphic characters bear "Grouped setae" consisting of multiple setae on the second antenna. Their budding positions, morphology, and ontogenetic changes were compared, and our deduction is that the Aesthetasc "Y" originated from "Grouped setae-like" organ in the Paleozoic. The morphogenetic processes in the molting period of these chemoreceptors were compared at the cellular level. The observations suggest that the "Grouped setae" are formed by hypodermal cells and share sheath cells corresponding to those of the Aesthetasc "Y" as a common constraint in the molting process of setae. We conclude that modification of the morphogenetic processes in the molting period of the "Grouped setae" gave rise to the Aesthetasc "Y" as a novel organ in the evolutionary pathway of the Ostracoda.     

Morphology and affinities of Eridostracina: Palaeozoic ostracods with moult retention

Ostracods are by far the most abundant living group of arthropods in the fossil record. Traditionally, eridostracines were classified as members of the Class Ostracoda. They have also been considered to represent extinct marine spinicaudatan (conchostracan) branchiopods. The ostracod affinity of the Eridostracina is evident in a number of features such as the muscle scars pattern, the hinge structure, the presence of an adductorial sulcus reflected as a ridge on the internal surface and the separation at the dorsal margin of successive valves. The eridostracines might be a polyphyletic group, containing aberrant representatives of ostracods, with ancestors probably among the conchoprimitid, leperditellid and beyrichioidean ostracod species. The Eridostracina represent an extinct group of small marine crustaceans with a multilayer structure of the calcified carapace, formed through the retention of unshed moults during the growth process. Details of the morphology of the eridostracine Cryptophyllus socialis from the Upper Devonian of Russia are reconstructed using the process of exfoliation of successive exuviae. ‘Double-sided’ hingement structures were found in the accumulated exuviae. It is suggested that the main function of these structures was the strengthening of the connection between the accumulated valves. The hingement of Cryptophyllus represents a vestigial structure, which has lost its original function as a pivot, a role documented in the ancestors of that genus. Tubular structures were found attached to the internal side of the calcite layer. It is suggested that they also represent vestigial pore canals, having lost their original function as sensory receptors. External surfaces of the attached exuviae bear imprints of the tubular structures of the overlying exuviae. These imprints originated probably due to the strong pressure of the new cuticle against the old one, during the very short moulting time. During this process, the freshly formed cuticle was at its final size, but still soft and non-calcified. A number of three-dimensionally preserved cell-like structures were recovered inside the interlayer chambers.