A high-precision global biostratigraphy of myodocope ostracods for the Silurian upper Wenlock Series and Ludlow Series

The wide, trans-oceanic geographical distribution of myodocope ostracods during the Silurian (especially during the Ludlow and Pridoli epochs), and their widespread preservation in rocks of that age, permits the establishment of a transcontinental biostratigraphy of comparable resolution to coeval graptolite/chitinozoan/conodont biozones. Seven myodocope biozones, extending from the Homerian Stage, upper Wenlock Series Cyrtograptus lundgreni graptolite biozone to the middle part of the Ludfordian Stage of the Ludlow Series, enable a time-resolution for each biozone of circa 1 million years. These biozones can provide high-resolution correlation across Europe into Arctic Russia and Central Asia. There is also the potential for a myodocope biostratigraphy applicable from the uppermost Silurian (Pridoli) to the Carboniferous.     

The evolutionary pathway of light emission in myodocopid Ostracoda

The evolutionary history of bioluminescence and iridescence in myodocopid ostracods was estimated by phylogenetic analysis of mitochondrial 16S ribosomal RNA sequences. The inferred phylogeny of the myodocopids suggests that the common ancestor of Myodocopida evaluated in this study exhibits iridescence. This type of light emission was once lost and recaptured independently in the descendant lineages. Bioluminescent species also evolved from non-luminous ancestral species. In the suborder Myodocopina, all the bioluminescent species form a monophyletic group, suggesting that bioluminescence evolved only once. Structural differences between two bioluminescent groups in the order Myodocopida suggests independent origins for bioluminescence.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 449–455.     

The Cambrian origin of the circulatory system of crustaceans

The carapace of Recent crustaceans such as myodocope ostracodes and phyllocarids is pervaded with well-developed anastomosing sinuses conveying hemolymph from the metabolizing organs to the dorsal heart. The inner lamella cuticle, which separates the sinuses from seawater, is thin enough to allow gaseous diffusion (e.g., O₂ uptake) over its surface. Comparable radiating and/or anastomosing features, of possible vascular origin, are herein recognized in several possible Crustacea from the Cambrian: cambriid, svealutiid, hipponicharionid and beyrichonid Bradoriida and in Carnarvonia from the middle Cambrian Burgess Shale. The vascular network is basically the same in these groups, consisting of sinuses radiating from supposed adductorial areas or from inferred areas of dorsal attachment of the body. The integumental (carapace sinuses) and branchial (gills) systems of respiration in crustaceans and crustacean-like animals were probably already differentiated by the middle Cambrian. The oldest record of probable integumental circulation is in the bradoriid Petrianna from the early Cambrian of Greenland. Similar circulatory systems may be represented by radiating ridges on the cephalon of other Cambrian arthropod groups such as the arachnomorphs ( Burgessia ) and trilobites ( Naraioa ) and may also be manifest in the carapaces of Ordovician-Devonian leperditicope ostracodes. Organs on the thoracopods of Cambrian supposed crustaceans, such as Canadaspis , resemble the foliaceous thoracic gills of Recent nebaliid phyllocarids and therefore may have served the same (respiratory) function.     

Embryonic development clarifies polyphyly in ostracod crustaceans

The present study describes the embryonic developmental process of the bioluminescent ostracod crustacean Vargula hilgendorfii . Optical microscopy, scanning electron microscopy, DAPI staining and video recording were used for observations. This study is the first detailed report of the embryonic development of a myodocopid ostracod. Contrary to previous studies, cleavage occurred in the yolk sphere and no evident cleavage furrow was found. No nauplius stage was found, and five pairs of appendages developed simultaneously. A bivalved carapace developed from two independent buds of the carapace valves. The buds of the left and right valves are enlarged, and become combined. The combined 'one-piece' carapace was divided by the formation of a hinge, and the usual bivalved carapace was formed. On the 16th day, embryos hatched as juveniles with six pairs of appendages, a pair of immature appendages, a pair of compound eyes, a median eye and a bivalved carapace. An important suggestion for the classification of Ostracoda is derived from the observed development of appendages and carapace, because the subclass Ostracoda is defined mainly by the similarities of appendages and the bivalved carapace. The present observations clearly show that the developmental process of Myodocopa differs from that of Podocopa, and supports polyphyly of the Ostracoda.     

Multifunctions of the upper lip and a ventral reflecting organ in a bioluminescent ostracod Vargula hilgendorfii (Müller, 1890)

Multifunctions of the upper lip in a bioluminescent myodocopid Vargula hilgendorfii were studied by video observation and histological method. The localization of luciferin and luciferase gland cells within the upper lip was partly successful. Two long protrusions of the upper lip, both of V. hilgendorfii and a non-luminescent species of the same family, immediately anterior to the mouth, were found to show very flexible movement especially while eating, as if smearing on the food surface a secretion from the protrusions (glands), which may support the hypothesized secretion of digestive enzymes from the upper lip. This hypothesis is further supported by the new finding of a pair of ducts which connect the basal part of the upper lip with the posterior digestive duct (stomach). Comparative studies of V. hilgendorfii with several sympatric non-luminescent species of the same family have also revealed that it has a characteristic reflecting organ immediately posterior to the anus. It is a conical small protrusion, as if dangling from the ventral edge of the abdomen at the apex of the cone. It is observable only in live specimens, when the furca, which is located outwardly to the organ, is sufficiently transparent. When illuminated, the reflecting organ reflects the distinct light. The diameter of the mirror (chemical composition provisionally analyzed) is about 6–8% of the carapace length. The organ develops from the very first stage of its ontogeny without reference to sex, which suggests that the function may be related to intraspecific signaling or predatory deterrence.     

A Silurian myodocope with preserved soft-parts: cautioning the interpretation of the shell-based ostracod record

Ostracod crustaceans are the most abundant fossil arthropods. The Silurian Pauline avibella gen. et sp. nov., from the Herefordshire Lagerstätte, UK, is an extremely rare Palaeozoic example with soft-part preservation. Based on its soft-part morphology, especially the exceptionally preserved limbs and presence of lateral eyes, it is assigned to the myodocopid myodocopes. The ostracod is very large, with an epipod on the fifth limb pair, as well as gills implying the presence of a heart and an integrated respiratory–circulatory system as in living cylindroleberidid myodocopids. Features of its shell morphology, however, recall halocyprid myodocopes and palaeocopes, encouraging caution in classifying ostracods based on the carapace alone and querying the interpretation of their shell-based fossil record, especially for the Palaeozoic, where some 500 genera are presently assigned to the Palaeocopida.     

Silurian bolbozoids and cypridinids (Myodocopa) from Europe: pioneer pelagic ostracods

Abstract: Bolbozoid and cypridinid myodocope ostracods from France, the Czech Republic and Sardinia consist of four genera (one new: Silurocypridina) and at least ten species (eight new: Bolbozoe acuta, B. rugosa, B. parvafraga, Parabolbozoe armoricana, Silurocypridina retroreticulata, S. variostriata, S. calva and Calocaria robusta). They have a functional design, palaeogeographical distribution, depositional setting and faunal associates that suggest that they were swimmers in the water column, living in dim light, relatively deep environments, above near bottom dysoxic/anoxic conditions. These myodocope faunas include some possible pelagic species sensu stricto, with widespread palaeogeographical (transoceanic) distribution, and some species that occupied hyperbenthic niches and were capable of making short incursions to the sediment to scavenge on carrion. These data support the model that Silurian myodocopes were pioneer pelagic ostracods.     

Homology of Holocene ostracode biramous appendages with those of other crustaceans: the protopod, epipod, exopod and endopod

Unambiguously biramous appendages with a proximal precoxa, well-defined coxa and basis, setose plate-like epipod originating on the precoxa, and both an endopod and exopod attached to the terminal end of the basis are described from several living Ostracoda of the order Halo-cyprida (Myodocopa). These limbs are proposed as the best choice for comparison of ostracode limbs with those of other crustaceans and fossil arthropods with preserved limbs, such as the Cambrian superficially ostracode-like Kunmingella and Hesslandona. The 2nd maxilla of Metapolycope (Cladocopina) and 1st trunk limb of Spelaeoecia, Deeveya and Thaumatoconcha (all Halocypridina) are illustrated, and clear homologies are shown between the parts of these limbs and those of some general crustacean models as well as some of the remarkable crustacean s.s. Orsten fossils. No living ostracodes exhibit only primitive morphology; all have at least some (usually many) derived characters. Few have the probably primitive attribute of trunk segmentation (two genera of halocyprid Myodocopa, one order plus one genus of Podocopa, and the problematic Manawa); unambiguously biramous limbs are limited to a few halo-cyprids. Homologies between podocopid limbs and those of the illustrated primitive myodocopid limbs are tentatively suggested. A setose plate-like extension, often attached basally to a podocopid protopod, is probably homologous to the myodocopid epipod, which was present at least as early as the Triassic. Somewhat more distal, less setose, and plate-like extensions, present on some podocopid limbs (e.g., mandible), may be homologous instead to the exopod (clearly present on myodocopid mandibles). The coxa (or precoxa) is by definition the most basal part of the limb. A molar-like tooth is present proximally on the mandibular protopod of many ostracodes; it is the coxal endite and projects medially from the coxa (or proximal protopod). The Ostracoda is probably a monophyletic crustacean group composed of Myodocopa and Podocopa. All have a unique juvenile (not a larva) initially with three or more limbs. Except that juveniles lack some setae and limbs, they are morphologially similar to the adult. Thus the following suite of characters in all instars may be considered a synapomorphy uniting all Ostracoda: (1) Each pair of limbs is uniquely different from the others. (2) The whole body is completely enclosed within a bivalved carapace that lacks growth lines. (3) No more than nine pairs of limbs are present in any instar. (4) The body shows little or no segmentation, with no more than ten dorsally defined trunk segments. No other crustaceans have this suite of characters. A probable synapomorphy uniting the Podocopa is a 2nd antenna with exopod reduced relative to the endopod.     

Homology and homoeomorphy in ostracod limbs

The functional modifications of myodocopan and podocopan ostracod limbs constitute a rich data set with which to carry out phylogenetic analyses, but efforts are hindered by lack of consensus on homologies. Homoeomorphy presents particular difficulties; for example, the furca is post-anal in Myodocopa but pre-anal in Podocopa, suggesting homoeomorphy, not homology. Homoeomorphies also exist between ostracod appendages and those of other Crustacea, for example the oral cone and styliform mandibulae of Paradoxostomatidae (Ostracoda) and Siphonostomatoida (Copepoda), both adaptations to commensal or parasitic lifestyles. Such clear manifestations of homoeomorphy, arising independently in different lineages as a result of similar functional requirements imposed on plesiomorphic appendage structures, warn of the possibility of more subtle examples which, if unrecognized, would lead to misinterpretations of character states used in phylogenetic analysis. For instance, the branchial plates found on third, fourth and fifth limbs of podocopans may not be homologous with the branchial plates on the fifth and sixth limbs of myodocopans. Limb homologies of podocopan ostracods (primarily as represented by various podocopid taxa) are investigated. Evidence is presented, based on studies of morphology and musculature, that podocopid branchial plates are exopodites (arising from the basis), while those of myodocopans are epipodites (arising from the coxa or precoxa). In Podocopida, moreover, the protopodites of post-mandibular limbs appear to be undifferentiated, comprising only a basis, while those of Myodocopa clearly exhibit a basis, coxa and often a precoxa. These differences argue against monophyly of the Ostracoda. The absence of epipodites, combined with the lack of a coxa in post-mandibular limbs, is potentially indicative of closer affinities between podocopans and Cambrian stem-group crustaceans (including Phosphatocopida) than between podocopans and myodocopans. The possible derivation of podocopid third, fourth and fifth limbs from a stem-group crustacean limb is demonstrated. The hypothesis is advanced that podocopan ostracods (represented today by Podocopida, Platycopida and Palaeocopida) are derived from much nearer the base of the crown-group Crustacea than myodocopans.