Origin of the Ostracoda and their maxillopodan and hexapodan affinities

There are Cambrian fossils attributed to the Ostracoda but the extant subclasses Podocopa and Myodocopa do not appear until the Ordovician. At this time the morphologically similar, free-living ancestors of the now sedentary Thecostraca (Ascothoracida, Acrothoracica and Cirripedia) may have still been extant, and from an ecological point of view it seems likely that, by and large, ostracods replaced them. However, living ostracods have an abbreviated, direct development, and some key aspects of their morphology, such as the nature of the maxillary segment and abdomen, are conjectural. Thus the affinities between these and related taxa remain uncertain; e.g., while some contemporary carcinologists place Ostracoda as a taxon coordinate with the Branchiopoda, Remipedia, Cephalocarida, Maxillopoda, Malacostraca, others tentatively or unequivocally ally them with the Maxillopoda (generally Mystacocarida, Copepoda, Tantulocarida and Thecostraca, and sometimes Branchiura and Pentastomida). Others, largely involved with fossils, have stretched the definition of the Maxillopoda even further, to the point where it seems even less likely a monophyletic taxon. Until recently cladistic analyses utilizing genetic (largely 18S rDNA) as well traditional morphological characteristics have given confusing results regarding the affinities between these taxa, and an important one suggested the Ostracoda might even be diphyletic. Furthermore, a very recent genetic study utilizing protein encoding genes places a podocopine ostracod among the most primitive of the extant crustaceans (Branchiopoda, Cephalocarida Remipedia and Mystacocarida), and then generally at the base of a lineage leading to the Malacostraca, a lineage giving rise to copepods and cirripeds along the way. This indicates these so-called maxillopodan taxa evolved independently from a malacostracan-like ancestor, and if so they are convergent. And finally, from genetic studies it is not only becoming well documented the Crustacea rather than Myriapoda gave rise to the Hexapoda, but it appears the Hexapoda stem from among the lower rather than the higher crustaceans, possibly even from the Ostracoda. Whether there were terrestrial ostracods at the time hexapods appeared in the Lower Ordovician is unknown, but the modest diversity of terrestrial ostracods today are podocopines which also first appeared in the Lower Ordovician. Thus, if current interpretations of living ostracodan and fossil hexapodan body plans are largely correct, it can be hypothesized the Ostracoda are close to the ancestor of the Hexapoda.     

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.     

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.     

New data on the systematics of the Limnocytheridae (Ostracoda,Cytheracea)

A new grouping of genera having fossil as well as recent representatives of the family Limnocytheridae (Ostracoda, Cytheracea) is proposed using morphological characteristics of the appendages, of the shell and of the evolutionary trends observed in this family.     

Phylogenetic implications of the Crustacean nauplius

The plesiomorphic mode of crustacean development is widely accepted to be via a larva called the nauplius. Extant taxa like the Cephalocarida, Branchiopoda, Ostracoda, Mystacocarida, Copepoda, Cirripedia, Ascothoracida, Facetotecta, Euphausiacea and Penaeidea hatch from an egg as a free-living nauplius. Other crustaceans show an embryonic phase of development suggestive of a naupliar organization. Several features of the nauplius larva have been proposed as diagnostic characters for the Crustacea: a median (nauplius) eye; at least three pairs of head appendages (antennules, antennae, mandibles); a posteriorly directed fold (the labrum) extending over the mouth and a cephalic (nauplius) shield. The relationship between trilobite protaspis with at least four appendages and the crustacean nauplius remains unclear, but reports of a copepod orthonauplius with four appendages are rejected. Swimming is suggested to represent the underived mode of locomotion for the crustacean nauplius, and that naupliar swimming directly results in naupliar feeding which also is underived.     

SOME LOWER ORDOVICIAN OSTRACODS FROM WESTERN HUPEH

The specimens of Ostracoda dealt herewith were collected by Messrs.K.C.Yangand A.T.Mu from the Ichang formation of Lower Ordovician during their field tripin western Hupei in the Spring of 1951.Six new species in three different genera ofwhich one is new,are described in the present paper.All of them are found inassociation with graptolites and triobites.The Ichang formation has been divided     

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.     

Appendage morphology of the trilobite Cryptolithus and its implications

The appendages of the Ordovician trilobite Cryptolithus tesselatus are restudied, and particular reference is made to the exite branch. Details formerly borrowed from Triarthrus to complete the reconstruction of the appendages of Cryptolithus are now found in part to be significantly different in the two trilobites. The exite 'filaments' of Cryptolithus evidently are strong spines that pointed ventrally and may have been used to stir up deposited mud. The broadened view of trilobite appendages indicates that some trace fossils (Cruziana semi-plicata type) may be interpreted as being partly made by the exites, while others (C. rugosa type) are perhaps made exclusively by the exites.     

A NEW SPECIES OF BRITTLESTAR (OPHIUROIDEA, ECHINODERMATA) FROM THE HUNSRÜCK SLATE (LOWER EMSIAN, LOWER DEVONIAN) OF GERMANY

Abstract:  Lapworthura lehmanni , a new species of ophiuroid, is described from four specimens from the Lower Devonian Hunsrück Slate of Germany. It is the only known ophiuroid in the Hunsrück Slate with paired but unfused ambulacrals, and it exhibits unique rows of spine-bearing dorsal arm ossicles. The genus Lapworthura Gregory was previously known only from the Ordovician of Scotland and the Silurian of England and Australia.     

The first British record and a new species of the superfamily Terrestricytheroidea (Crustacea, Ostracoda): morphology, ontogeny, lifestyle and phylogeny

Terrestricythere elisabethae sp. nov. is described from a semiterrestrial coastal habitat at two sites in Hampshire, southern England. It is the first record of a living population of the genus outside the Far East (north-west Pacific). Based on extensive collections and from observations of cultures, its morphology is described (including a formal definition of the unique 'visordont' hinge), as well as its ontogeny and lifestyle (encompassing habitat, life cycle, mode of life and locomotion). A further new species from Somerset, south-west England is also recorded on the basis of a single specimen but left in open nomenclature. The affinities of the Terrestricytheroidea are discussed in the context of a tentative phylogeny of podocopan Ostracoda. Both carapace and appendage characters are such as to warrant maintaining it as a separate superfamily, which is more closely related to the Cytheroidea and Darwinuloidea than to the Cypridoidea.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 253–288.     

On the origin of the putative furca of the Ostracoda (Crustacea)

The posterior end of body of the extant ostracods exhibits a pair of variously shaped appendages, commonly designated as furca(e), uropods or caudal rami, used for feeding and/or locomotion. It is here shown that the so-called furca of all extant ostracods has evolved from the (probably epipodal) vibratory plates of a pair of uropods. The transformation comprised the following steps: (a) complete reduction of the uropodal protopodite and endopodite; (b) sclerotisation of the lateral walls of the vibratory plates; (c) transformation of the branchial filaments into spines and/or claws; (d) re-orientation of the plates from posterodorsal to posteroventral. These modifications are suggested to have evolved in parallel with a change in function, from respiratory to locomotory and/or feeding. The most primitive condition, reminiscent of the ancestral state of character, is seen in the Platycopida: the ‘furca’ still appears similar in shape to the vibratory plates of the pair of sixth limbs. In the Podocopida the uropodal plates have been modified into plate-like, more often into rod-shaped rami mainly used for locomotion. In both the Platycopida and Podocopida the anus has remained in its original place, posterior to the ‘furcal’ plates or rami. In the Myodocopida and Halocyprida the uropodal vibratory plates are transformed into heavily developed lamellae bearing sturdy spines. They are activated by a complex apparatus of muscles and sclerites, the development of which necessitated the displacement of the anus from the end of the body towards its present place, anterior to the ‘furca’. The furca of the Ostracoda being not a ‘true’ furca, a change in terminology is proposed: uropodal plates or lamellae in the Platycopida, Palaeocopida and Myodocopida/Halocyprida; uropodal rami in the Podocopida. The so-called furcae of the Ostracoda being homologous structures, it is concluded that all extant ostracods belong to a monophyletic lineage.     

Biology of the Hyolitha

Hyoliths are Paleozoic fossils that have a calcareous exoskeleton consisting of an elongate, usually bilaterally symmetrical cone, a close fitting operculum, and a pair of curved appendages. Their skeletal ultrastructure resembles the crossed-lamellar shell layers of some molluscs. Several specimens from the Ordovician of France and the Cambrian of Antarctica have parts of the gut preserved by infilling matrix, showing that both mouth ad anus were located near the cone aperture. Muscle scars in other hyolith shells indicate that the animal had a series of dorsoventral and longitudinal, or longitudinal and circular muscles, which operated through a hydrostatic skeleton to protract and retract the head, to open and close the operculum, and to move the appendages. Although the shell form and skeletal ultra-structure of hyoliths are of a molluscan type, the muscle insertions suggest that the hyolith cone is not homologous with the dorsal exoskeleton of primitive molluscs. Hyoliths probably constitute a small extinct branch of phylum size, related to the Mollusca and the Sipunculoidea. All three groups may have had common ancestors in the late Precambrian.     

The arthropod Offacolus kingi (Chelicerata) from the Silurian of Herefordshire,England: computer based morphological reconstructions and phylogenetic affinities

The small, non-biomineralized, three-dimensionally preserved arthropod Offacolus kingi Orr et al. from the Wenlock Series (Silurian) of Herefordshire, England, is re-evaluated, and the new family Offacolidae erected. This new study is based on specimens which have been serially ground, reconstructed by computer and rendered in the round as coloured models. Offacolus possesses a prosomal appendage array similar to that of Limulus, but also bears robust and setose exopods on appendages II-V which are unlike those found in any other arthropods. Opisthosomal appendages are similar in number and morphology to the book-gills of Limulus. Cladistic analysis places Offacolus basally within the Chelicerata, as a sister taxon to the eurypterids and extant chelicerates, but more derived than the Devonian Weinbergina.     

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.     

Seasonal morphometric variation in sexual and asexual, North American and Australian, populations of the aphid, Acyrthosiphon pisum

The lengths of the body and appendages of the aphid Acyrthosiphon pisum (Harris) (Homoptera: Aphididae) vary seasonally in sexual North American and asexual Australian populations. The first generation of spring aphids in North America and winter aphids in Australia have short appendages in relation to body length. Excluding this phenotype, North American and Australian aphids cannot be discriminated morphometrically. The short appendages in North America are associated with a specialized morph called a fundatrix; the short appendages of Australian aphids are caused by exposure to low temperatures during prenatal development. The same temperature-sensitive mechanism operates in sexual and asexual North American aphids, but does not explain the short appendages of the fundatrix, which appear to arise through a separate mechanism. The short appendages are caused neither by a maternal effect from winged mothers, although such an effect exists, nor by seasonal changes in body length and allometry, nor by microevolutionary changes. The temperature-induced shortening of appendages is a seasonal polymorphism, which mimics the short appendages seen in fundatrices. The two types of phenotypic plasticity have the same consequence in sexual and asexual populations of the same species and may be an example of convergent evolution.     

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.     

A WESTWARD EXTENSION OF THE UPPER ASHGILLIAN HIRNANTIA FAUNA

The very distinctive Hirnantia fauna of uppermost Ordovician age is here recorded from Ireland, where it is found in association with the upper parts of the Chair of Kildare reef limestone. The most important hrachiopod members of the Hirnantia assemblage are Cliftoma, Cryptothyrella, Plectothyrella, Hirnantia, Eostropheodonta , and Dalmanella which are illustrated and their occurrences at other European localities indicated and discussed, particularly with respect to their differing abundances. The nature of the fauna, in conjunction with some lithological pointers, indicates a shallow water lagoonal environment of deposition. With respect to the Kildare succession, the presence of the Hirnantia fauna indicates that the upper part of the Kildare limestone at least belongs to Zone 8 of the Ashgillian; and it is believed that the North of England Keisley Limestone is also of this age, rather than of Lower or Middle Ashgillian age.     

松辽盆地白垩纪介形类生物地层学特征

本文详细论述了松辽盆地白垩系介形类属、种数量及壳饰变化在纵向上的分布规律和介形类组合特征的生物地层学意义。认为介形类属的分布是岩石地层组划分的重要依据,种的分布控制了岩石地层段的划分,并可将萨尔图油层、葡萄花油层和高台子油层进一步细分。同时指出松辽盆地介形类的盛衰与湖盆的发育关系密切,介形类的演化周期与湖盆的发育、收缩和沉积旋回相一致,在湖盆最发育的时期,也是介形类繁盛的高峰,湖盆发育的初期和末期,往往是介形类的发生和衰退期。     

Aral Sea Ostracoda as environmental indicators

Fluctuations in the level and chemistry during its history have played a major part in shaping the floral and faunal communities of the Aral Sea. Of the eleven species of Ostracoda (Crustacea) known to have been living in the Aral Sea in 1960, only one survives today due to the anthropogenically induced salinity increase of the past three decades. The origins of a mixed fresh- and brackish-water ostracod fauna are discussed, and it is concluded that some of the major faunal elements must have reached the Aral Sea Basin during a past high water level phase when connection existed with the Caspian Sea. The taxonomic position of key taxa is clarified, and the major elements of the pre-1960 Aral Sea ostracod fauna are illustrated from Holocene sequences. Aral Sea, Ostracoda, Holocene, palaeolimnology.     

Evolutionary history of regeneration in crinoids (Echinodermata)

The fossil record indicates that crinoids have exhibited remarkable regenerative abilities since their origin in the Ordovician, abilities that they likely inherited from stem-group echinoderms. Regeneration in extant and fossil crinoids is recognized by abrupt differences in the size of abutting plates, aberrant branching patterns, and discontinuities in carbon isotopes. While recovery is common, not all lost body parts can be regenerated; filling plates and overgrowths are evidence of non-regenerative healing. Considering them as a whole, Paleozoic crinoids exhibit the same range of regenerative and non-regenerative healing as Recent crinoids. For example, Paleozoic and extant crinoids show evidence of crown regeneration and stalk regrowth, which can occur only if the entoneural nerve center (chambered organ) remains intact. One group of Paleozoic crinoids, the camerates, may be an exception in that they probably could not regenerate their complex calyx-plating arrangements, including arm facets, but their calyxes could be healed with reparative plates. With that exception, and despite evidence for increases in predation pressure, there is no compelling evidence that crinoids have changed though time in their ability to recover from wounds. Finally, although crinoid appendages may be lost as a consequence of severe abiotic stress and through ontogenetic development, spatiotemporal changes in the intensity and frequency of biotic interactions, especially direct attacks, are the most likely explanation for observed patterns of regeneration and autotomy in crinoids.