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.     

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.     

Phosphatocopina – ostracode-like sister group of Eucrustacea

The Phosphatocopina were long considered as the oldest, Cambrian, record of ostracode Crustacea. However, our detailed analysis of more than 2,500 specimens from the Upper Cambrian ‘Orsten’ of Sweden reveals that Phosphatocopina are neither Ostracoda nor Eucrustacea. The antenna and mandible of the phosphatocopines investigated consist of a prominent limb stem which carries a two-segmented endopod and multi-annulated exopod. This stem portion is now recognised as the fusion product of the coxa and basipod during ontogeny. Phosphatocopina share features, such as the coxa and basipod on antennae and mandibles, as well as ventral body structures such as the prominent pre-oral labrum and a single post-oral cephalic plate, the sternum (with paragnaths on the mandibular sternal portion), exclusively with the Eucrustacea. As a plesiomorphy, the ontogeny of Phosphatocopina starts with a ‘head larva’ with four pairs of limbs, a larva type found in the ground pattern of the Euarthropoda as well as the Crustacea. In contrast, eucrustacean ontogeny begins with a nauplius with three pairs of limbs, a ‘short-head larva’ or orthonauplius. Again, the post-mandibular limbs of phosphatocopines retain the plesiomorphic limb design of a basipod with a setiferous ‘ proximal endite’, whereas Eucrustacea, including the Ostracoda, have their first post-mandibular limb differentiated into a ‘ mouthpart’, the maxillula. Autapomorphies of Phosphatocopina include the small antennula with few terminal setae, a bivalved shield with interdorsum, and the fused coxa and basipod on antenna and mandible. We therefore consider the Phosphatocopina to be the sister group of the Eucrustacea. The respective phosphatocopine species of the Upper Cambrian of southern Sweden are restricted to a particular time zone and may be useful as stratigraphic markers.     

Evaluation of a new character for the phylogenetic analysis of Ostracoda (Crustacea): the podocopan maxillular branchial plate

A data set of the number of setae on the maxillular branchial plate of podocopan ostracods was compiled from published drawings and our own dissections. A total of 168 species in six superfamilies were covered, with the Cytheroidea, Cypridoidea and Bairdioidea being best represented. The number of setae is shown to be independent of the size of the ostracod and the environment in which the species lives, but to have phylogenetic significance at superfamily level. Ontogenetic data suggest that the low number of branchial plate setae in many cytheroidean families is a paedomorphic phenomenon.     

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.     

Myodocopa (Crustacea: Ostracoda) as models for evolutionary studies of light and vision: multiple origins of bioluminescence and extreme sexual dimorphism

Evolutionists often use phylogeny to examine independent evolutionary events in search of generality. Therefore, groups of organisms rich in such independent character transitions are particularly valuable for the study of evolution. With respect to eyes, vision, and light-related characters, one such group is Ostracoda (Crustacea). Phylogenies of ostracods, derived from DNA sequence data and morphological characters, are presented. These inferred relationships largely agree with previous assessments of ostracod phylogeny, with the exception of paraphyletic Philomedidae. Based on methods of character reconstruction using these inferred relationships, different groups of ostracods probably evolved both bioluminescence and extreme sexual dimorphism (females lack eyes, males have large eyes) multiple times. Furthermore, myodocopid ostracods may have evolved compound eyes independently of other arthropods. For these and other reasons, it is proposed that the Ostracoda are an exceptionally important group for studying the evolution of vision- and light-related characters.     

The ontogeny of the platycopid Keijcyoidea infralittoralis (Ostracoda: Podocopa)

Platycopid ostracods such as the genus Keijcyoidea Malz, 1981 (Family Cytherellidae) have a unique body plan and are regarded as a phylogenetically ancient lineage. Their ontogeny, which is important in considering phylogenetical relationships, is very poorly known except for the growth of the carapace; there are nine instars including the adult, as in other podocopan groups. All appendages in all immature instars (A-8 to A-1) of Keijcyoidea infralittoralis Tsukagoshi, Okada & Horne, 2006 are described and illustrated here. The anlagen of the copulatory organs and the sexual dimorphism of carapace size appear in the sixth (A-3) instar, whereas sexual dimorphism in both the fifth and the sixth limbs, a distinctive feature of adults, is not clearly evident until the eighth (A-1) instar. Appendages are added at the moults between the second (A-7) and third (A-6), and between the fifth (A-4) and sixth (A-3) instars. The seventh limb, which platycopid ostracods have lost in the adult stage, is observed as an anlage in the sixth (A-3) and seventh (A-2) instars. During the other moults, there are no significant changes to the body plan. The ontogeny of the Platycopida is compared with that of the Podocopida, and strongly suggests that the phylogenetic position of the Platycopida is as an end-member of the Podocopa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 213–237.     

Complex controls on ostracod palaeoecology in a shallow coastal brackish‐water lake: implications for palaeosalinity reconstruction

1. The low‐Mg calcite shells of Ostracoda (Crustacea) are often well preserved in the sediments of alkaline lakes. In coastal waters that have undergone large temporal changes in salinity, ostracod assemblages preserved in the sediment record have been used to reconstruct palaeosalinity, often assuming that salinity is the only significant control on the faunas. 2. We evaluate the performance of ostracods as palaeosalinity indicators in Hickling Broad, a shallow brackish coastal lake in Norfolk, U.K., by comparing fossil ostracod assemblages covering two centuries with geochemical inferences and instrumental records of past salinity and water composition along with other palaeolimnological indicators. 3. Despite large changes in the salinity of the lake and the supposed salinity sensitivity of ostracods, the fossil ostracod assemblages do not clearly reflect the salinity trends inferred from the other independent data. Rather, a complex series of changes has occurred in the lake over the past 200 years and factors other than salinity, including eutrophication, toxicity and associated complex alterations in habitat availability have probably influenced ostracod assemblages. In contrast, there is a good broad agreement between inferred or measured salinity and the trace‐element chemistry of ostracod shells. 4. We conclude that ostracod faunas may not always provide unambiguous palaeosalinity records and should therefore not be used to reconstruct salinity changes except as part of a multi‐proxy investigation that includes other palaeoecological and/or geochemical indicators.     

介形类系统分类与起源演化的形态、分子和化石证据

介形类(Ostracoda)因其丰富的化石记录和广布的海陆现生代表类群,而被认为是进化生物学中研究生物多样性产生机制和演变历程的颇具潜力的重要模式生物。介形类在甲壳亚门中的谱系发生位置、起源及其内部各类群间的系统关系还存在诸多争议。基于其体制构造的形态学特征,介形类被归入甲壳亚门下的颚足纲(Maxillopoda),但来自18S rDNA序列数据分析却显示Maxillopoda不是单系群。基于化石记录和壳体形态特征,高肌虫(Bradoriida)长期以来被认为是介形类的一个祖先类群,但保存有软躯体的早寒武世化石的研究表明,Bradoriida不是介形类甚至可能也不属于甲壳类。不同的研究者所强调的壳体和肢体形态特征各异,导致介形类最大的现生类群速足目(Podocopida)的四个超科之间的关系也存在诸多推测。壳体和肢体特征在系统演化意义上的不兼容,需要分子生物学等证据的介入。分子、形态和化石证据的积累及各种信息整合是系统演化研究的必然趋势。     

Phylogenetic relationships of Early–Middle Ordovician ostracods of Baltoscandia

Phylogenetic analysis of the Early and early Middle Ordovician (Tremadoc and Arenig) ostracod species of Baltoscandia suggests a polyphyletic origin for the suborder Beyrichiocopa. Binodicopes, leiocopes and eridostracans are excluded from the beyrichiocopide clade. An independent origin from the basal ostracods is suggested for the binodicopes and eridostracans. The palaeocopes form a strongly supported monophyletic clade. Within this suborder, the ctenonotellid and the tetradellid families together form a monophyletic clade. The tetradellids are paraphyletic, being a stem-group for the ctenonotellids. Nanopsis nanella , the earliest known ostracod from the Tremadoc, is a basal palaeocope. The early eridostracans Conchoprimitia and Incisua , with their uncomplicated carapace morphology, might be the most primitive ostracods.     

Invasive South American floating plants are a successful substrate for native Central African pleuston

Eichhornia crassipes, commonly known as water hyacinth, is a free-floating perennial aquatic plant native to South America, which has been widely introduced on different continents, including Africa. E. crassipes is abundant in both the Congo (Africa) and Amazon (South America) River catchments. We performed a comparative analysis of the ostracod communities (Crustacea, Ostracoda) in the E. crassipes pleuston in the Amazon (South America) and Congo (Africa) River catchments. We also compared the ostracod communities from the invasive E. crassipes with those associated with stands of the native African macrophyte Vossia cuspidata. We recorded 25 species of ostracods associated with E. crassipes in the Amazon and 40 in the Congo River catchments, distributed over 31 ostracod species in E. crassipes and 27 in V. cuspidata. No South American invasive ostracod species were found in the Congolese pleuston. Diversity and richness of Congolese ostracod communities was higher in the invasive (Eichhornia) than in a native plant (Vossia). The highest diversity and abundance of ostracod communities were recorded in the Congo River. The result of principal coordinates analysis, used to evaluate the (dis)similarity between different catchments, showed significant differences in species composition of the communities. However, a dispersion homogeneity test (PERMDISP) showed no significant differences in the variability of the composition of species of ostracods (beta diversity) within Congo and Amazon River catchments. It appears that local ostracod faunas have adapted to exploit the opportunities presented by the floating invasive Eichhornia, which did not act as “Noah’s Ark” by introducing South American ostracods in the Congo River.     

Distribution of Recent ostracod species in the Lake Qinghai area in northwestern China and its ecological significance

In order to interpret the climate-change data using the ecological characteristics of the ostracods from the drill cores in Lake Qinghai, the distribution of Recent ostracods was studied in Lake Qinghai area. A total of 34 species belonging to the Ostracoda class of Crustacea were collected from different bodies of water in the Lake Qinghai area, and the ecological information for Recent ostracod species was studied. Among these 34 species, Cypris pubera, Eucypris dulcifons, Ilyocypris sp. 1, Ilyocypris sp. 2, Fabaeformiscandona caudata, Fabaeformiscandona hyalina, Herpetocypris reptans, Prionocypris gansenensis, Potamocypris villosa, Potamocypris smaragdina, Paralimnocythere compressa and Subulacypris sp. were first reported by us in the Lake Qinghai area. Some of the species identified exhibited cosmopolitan distributions, at least in the Holarctic region, but P. gansenensis and Ilyocypris echinata appeared to be restricted to the cold regions in northwestern China.The ecological significance of the primary ostracod species in the Lake Qinghai area was described according to the observations made during our time in the field and according to data from a number of reports. Our results indicate that the species diversity and abundance of ostracods may be related to water salinity in the Lake Qinghai area. The ecological information for ostracods can be used to distinguish different water environments and types based on the characteristics of one species or of an assemblage of several species.     

Ultrastructure and cuticle formation of the carapace in the myodocopan ostracod exemplified by Euphilomedes japonica (Crustacea: Ostracoda)

The ultrastructure and formation of the cuticle of a myodocopan ostracod, Euphilomedes japonica, are investigated utilizing scanning and transmission electron microscopy. The outer lamella cuticle consists of four layers; epicuticle, exocuticle, endocuticle, and membranous layer like in the cuticle of other arthropods. The exocuticle and endocuticle are well-calcified and the organic matrix develops within the both cuticles. The outermost layer of new cuticle (epicuticle) is secreted first and the inner layers (exocuticle, endocuticle and membranous layer) are added proximally in the pre-, and postmoult stages. The calcification takes place in the whole area of carapace at the same time together with the synthesis of organic matrix within the endocuticle. This study demonstrates that the ultrastructure and formation of the cuticle in myodocopans are different from those in podocopans, and that the myodocopan carapaces have achieved a structural diversity for adaptation to different lifestyles.     

Thermopsis thermophila n. gen. n. sp. from hot springs in Nevada, U.S.A. (Crustacea, Ostracoda)

Thermopsis thermophila n. gen. n. sp., a new freshwater ostracod species is described from hot springs in Nevada, U.S.A. The animals were collected within a temperature range of 40–55°C. The new genus belongs to the Ostracoda Podocopida Cypridoidea Cyprididae Cypridopsinae.     

Recent benthonic ostracod assemblages as indicators of the Tsushima warm current in the southwestern sea of Japan

One hundred and sixty-seven species of podocopid ostracods are reported from 26 recent surface sediment samples collected on the shelf of the Tsushima Strait in the southwestern Sea of Japan. Four biofacies (A–D) are recognised based on Q-mode cluster analysis. Among these, biofacies B is distributed along the Tsushima Warm Current and includes typical East China Sea species. This fact suggests that the migration of the shelf-living ostracods between the open sea and the adjacent semi-enclosed marginal sea mainly occurs in association with the inflow of this sea current. The distribution of the recent ostracod assemblages from the Tsushima Strait are expected to be a sensitive indicator of past and future changes in the Tsushima Warm Current. One new ostracod species of the family Trachyleberididae, Trachyleberis ikeyai, is described and illustrated herein. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: K. Martens     

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.     

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.     

The ontogeny of appendages and carapace of Neonesidea schulzi (Ostracoda: Bairdiidae) from the Red Sea coast,Egypt

The ostracod genus Neonesidea is broadly distributed in shallow marine waters. The ontogeny of the N. schulzi (Bairdiidae) is described in detail by studying the development of the appendages and variations in carapace form, size and structure. Neonesidea schulzi has eight post-embryonic instars, and a gap in its ontogenetic development during instar A-6, where no new Anlage is added. The Anlagen of the copulatory organs and the forked terminal claw of second antenna appear in the seventh (A-1) instar, and the first thoracic legs of podocopid ostracods are shown to descend from the thoracic region. For the first time in ostracods, observations of moulting from sixth and seventh instars are presented.     

Differential Expression of Duplicated Opsin Genes in Two EyeTypes of Ostracod Crustaceans

In the first molecular study of ostracod (Crustacea) vision, we present partial cDNA sequences of ostracod visual pigment genes (opsins). We found strong support for differential expression of opsins in ostracod median and compound eyes and suggest that photoreceptor specific expression may be a general phenomenon in organisms with multiple receptors. We infer that eye-specific expression predates the divergence of the two species examined, Skogsbergia lerneri and Vargula hilgendorfii, because eye-specific opsin orthologs are present in both species. We found multiple opsin loci in ostracods, estimating that at least eight are present in Skogsbergia lerneri. All opsins from both ostracod species examined are more closely related to each other than to any other known opsin sequences. Because we find no evidence for gene conversion or alternative splicing, we suggest the occurrence of many recent gene duplications. Why ostracods may have retained multiple recent opsin gene duplicates is unknown, but we discuss several possible hypotheses.[Reviewing Editor: Martin Kreitman]