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.     

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.     

Morphological design and fossil record of the podocopid ostracod naupliar eye

The naupliar eye of podocopid ostracods is a useful character for considering evolution in photic environments. Based on external morphologies and histological observations, naupliar eyes are here categorized into six types. The fossil record demonstrates that the major evolutionary changes in podocopid naupliar eyes occured after the Ordovician. Eye types 1 and 2 are not joined to the carapace by the hypodermal cells, and these two types are found in Palaeozoic ostracods. Eye type 3, 4, 5 and 6 are extended on eye stalks, tightly joined to the carapace by the hypodermal cells, and use the carapace as the refractive cuticle lens. Eye type 3 appeared in the Permian, and eye types 4, 5 and 6 appeared in the Early Jurassic. The design of the podocopid naupliar eye diversified in the Early Jurassic.     

松辽盆地D80井嫩一段介形类化石壳体特征及生物地层

松辽盆地嫩江组一段富含介形类化石,是系统研究Santonian期陆相介形类的理想层位。论文对中央坳陷区D80井嫩一段的介形类化石进行了系统的分类、生物地层和壳体特征分析。鉴定出介形类化石11属30种,划分出6个介形类化石带,并与前人的研究结果进行了对比;描述了该段介形类化石的壳体结构,共识别出4种壳饰类型,8种壳形结构,基于介形类壳体特征和其他证据,初步探讨了该时期湖盆湖平面的变化特征,显示出嫩一段时期松辽湖盆湖平面经历了由深逐渐变浅的过程。     

甘肃玉门下沟地区早白垩世下沟组介形类

甘肃玉门下沟地区下沟组介形类化石十分丰富,该地区下沟组介形类化石共计9属4亚属,21种,本文描述了其中4新种,即Cypridea(Cyamocypris)xiagouensissp.nov.,Cypridea(Cypridea)subunicostatasp.nov.,Stenestroemiasubpeculiarissp.nov.和Stenestroemiaxiagouensissp.nov.。该介形类化石组合尤以Cypridea最为繁盛,通过分析介形类属种的形态特征和化石组合特征并结合岩性特征,推断下沟组的地质时代为早白垩世巴列姆期;并认为下沟组为水动力较弱的淡水-微咸水河湖相沉积。     

Magnesium and strontium in non-marine ostracod shells as indicators of palaeosalinity and palaeotemperature

Measurements of the Ca, Sr, and Mg contents of individual calcitic shells of non-marine ostracods and their host waters, both in lakes and controlled aquaria, permit the calculation of the distribution coefficients of Sr/Ca and Mg/Ca partitioning in ostracod shells. We report new K_D[Sr] for seven genera of non-marine ostracods and K_D[Mg] for Cyprideis at 25°C.Strontium partitioning is virtually temperature-independent, and is related to the Sr/Ca of the host water, and in Ca²⁺-saturated waters, to the salinity of the water. Magnesium partitioning is dependent on both temperature and Mg/Ca of the host water.For simple closed-basin lakes (crater lakes are ideal), the Sr content of ostracods is a sensitive indicator of salinity and thus evaporation/precipitation changes, which in turn, indicate variations in continental climate. A 10000-year continuous palaeosalinity record established by Sr and Mg contents of fossil ostracods for Lake Keilambete, southeastern Australia, is in close agreement with an independent palaeosalinity estimate based on sediment textures.We suggest rules that allow Sr and Mg analyses of suites of individual fossil ostracod shells from lacustrine sediments to be interpreted in terms of palaeosalinity and palaeotemperature variations.     

Ostracods had colonized estuaries by the late Silurian

The fossil record of terrestrialization documents notable shifts in the environmental and physiological tolerances of many animal and plant groups. However, for certain significant components of modern freshwater and terrestrial environments, the transition out of marine settings remains largely unconstrained. Ostracod crustaceans occupy an exceptional range of modern aquatic environments and are invaluable palaeoenvironmental indicators in the fossil record. However, pre-Carboniferous records of supposed non-marine and marginal marine ostracods are sparse, and the timing of their marine to non-marine transition has proven elusive. Here, we reassess the early environmental history of ostracods in light of new assemblages from the late Silurian of Vietnam. Two, low diversity but distinct ostracod assemblages are associated with estuarine deposits. This occurrence is consistent with previous incidental reports of ostracods occupying marginal and brackish settings through the late Silurian and Devonian. Therefore, ostracods were pioneering the occupation of marginal marine and estuarine settings 60 Myr before the Carboniferous and they were a component of the early phase of transition from marine to non-marine environments.     

Palaeozoic roots of the sigilliid ostracods

Podocopine ostracods rapidly increased in species diversity after the Palaeozoic, and 32 out of the 36 podocopine families are extant (Whatley et al., 1993), but the phyletic origins are known for only a few. The Sigilliacea are unique in having a long and detailed fossil record. The Sigilliidae known from the recent and Mesozoic faunas differ from others ostracods in having large adductor muscle scars with many spots, a character typical rather to Palaeozoic forms. Paleozoic Microcheilinellidae were morphologically close to the sigilliids, but details of their hingement and carapace margin structures were unknown until now.Silicified specimens of Microcheilinella mendelgrammi, new species, from the Early Carboniferous of southern China and Microcheilinella mandelstami, from the Middle Devonian of north-western Poland show merodont hingement and narrow calcified inner lamella. This makes their relationship to the Mesozoic, Tertiary, and recent sigilliids Saipanetta, Cardobairdia, Sigillium, and Kasella likely. The sigilliid lineage was thus established already in the Devonian.Tubulibairdia, with adont hingement and tubulous shell wall, and Microcheilinella s. s., with merodont hingement and no tubules, are distinct genera in different families.     

On the preservation of carapaces of some limnic ostracods: An exercise in actuopalaeontology

The way in which the ostracod carapace decomposes through biological and physico-chemical processes is illustrated with Recent and Neogene non-marine ostracods (candonines, herpetocyprines and cytherids) from lacustrine and groundwater habitats. The degree of carapace preservation for Recent Candoninae, in the Mondsee (Austria) is shown to be dependent on the sedimentary microenvironment. The carapaces of herpetocyprines, because of their special morphological structure, decay very easily. This could explain the rare occurrrence of this ostracod group in Palaeogene deposits in Europe.     

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.     

Erratic rates of molecular evolution and incongruence of fossil and molecular divergence time estimates in Ostracoda (Crustacea)

Dating evolutionary origins of taxa is essential for understanding rates and timing of evolutionary events, often inciting intense debate when molecular estimates differ from first fossil appearances. For numerous reasons, ostracods present a challenging case study of rates of evolution and congruence of fossil and molecular divergence time estimates. On the one hand, ostracods have one of the densest fossil records of any metazoan group. However, taxonomy of fossil ostracods is controversial, owing at least in part to homoplasy of carapaces, the most commonly fossilized part. In addition, rates of evolution are variable in ostracods. Here, we report evidence of extreme variation in the rate of molecular evolution in different ostracod groups. This rate is significantly elevated in Halocyprid ostracods, a widespread planktonic group, consistent with previous observations that planktonic groups show elevated rates of molecular evolution. At the same time, the rate of molecular evolution is slow in the lineage leading to Manawa staceyi, a relict species that we estimate diverged approximately 500 million years ago from its closest known living relative. We also report multiple cases of significant incongruence between fossil and molecular estimates of divergence times in Ostracoda. Although relaxed clock methods improve the congruence of fossil and molecular divergence estimates over strict clock models, incongruence is present regardless of method. We hypothesize that this observed incongruence is driven largely by problems with taxonomy of fossil Ostracoda. Our results illustrate the difficulty in consistently estimating lineage divergence times, even in the presence of a voluminous fossil record.     

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.     

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.     

The palaeopsychrosphere in the Devonian

The interpretation of Palaeozoic marine benthonic ostracods of the Thuringian ‘Ecotype’ or ‘Mega‐assemblage’ as indicative of a palaeopsychrosphere has been controversial. We review the evidence and conclude that the characteristics and distribution of these ostracods are consistent with the existence of deep, cold, well‐oxygenated water masses, formed by high‐latitude sinking of surface waters, in the Devonian oceans, comparable with those of the modern ocean that constitute the psychrosphere (waters below the thermocline with temperature <10°C). We present a new palaeoceanographic model for the Frasnian–Famennian (Late Devonian) Kellwasser events that resulted in the extinction of 75% of marine ostracod taxa, mostly neritic or pelagic forms, while the deep water Thuringian Mega‐assemblage was relatively unaffected. We offer an explanation for the unlikely preservation of examples of such a deep water (bathyal to abyssal) ostracod fauna that involves upwelling of deep cold waters on continental margins.     

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.     

Size and environmental salinity in the modern euryhaline ostracod Cyprideis torosa (Jones, 1850), a biometrical study

Samples from living populations of the ostracod Cyprideis torosa, taken at various inland brackish-water localities in The Netherlands, are biometrically analyzed. A negative correlation is found between the mean size of the carapace (adult females) and the salinity level of the environment. The results are discussed and compared with published data on the size-salinity relationship in ostracods.     

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.     

Extinction pattern of marine Ostracoda across the Pliensbachian-Toarcian boundary in the Cordillera Ibérica, NE Spain: Causes and consequences

This paper discusses the extinction pattern of the Pliensbachian-Toarcian boundary (PTB) ostracod assemblages at the Almonacid de la Cuba section (Cordillera Ibérica, NE Spain), which has been recently proposed as auxiliary boundary stratotype for the PTB. The ostracod record shows that the main Early Jurassic ostracod extinction event occurred not at the end of the Pliensbachian, but near the top of the Mirabile ammonite Subzone, Tenuicostatum ammonite Zone (Early Toarcian). On the basis of the evaluation of PTB ostracod record, a new causal explanation for the Early Toarcian ostracod turnover is proposed. This paper suggests that a reorganization of surface and deep-water circulations caused by the opening of the Hispanic Corridor could have generated a mild cooling episode, finally affecting the survival of healdioid ostracods.     

Butterflies of the Cambrian benthos? Shield position in bradoriid arthropods

Mode of preservation and method of recovery strongly influences our understanding of the life habits of extinct organisms. Bradoriid arthropods were abundant, and diverse members of early Cambrian ecosystems and most life reconstructions display these animals with the two shields of the carapace open in a ‘butterfly’ configuration. This favoured reconstruction is largely based on the abundance of ‘crack‐out’ specimens preserved in this position (e.g. Kunmingella from the early Cambrian of China). In contrast, large collections of acid processed bradoriids from the Arrowie Basin of South Australia (Cambrian Stage 3) are preserved with a narrow gape at the ventral margin or completely closed with the carapace folded along the dorsal midline. The relative abundance of conjoined, closed (or partially closed) specimens from the lower Cambrian Hawker Group succession suggests that at least some bradoriid taxa were capable of withdrawing appendages and tightly closing the shields, challenging the common view that the majority of bradoriids usually held their carapaces open in a ‘butterfly’ configuration during life. New data show that layers of the bradoriid carapace are continuous through the dorsal fold with no evidence for complex articulating structures as in ostracod hinges. The relatively pliable, sclerotized or lightly mineralized calcium phosphate composition of the carapace and the simple, flexible dorsal fold facilitated opening and closing of the shields. Despite not being closely related, ostracods share close biomechanical and ecological similarities with bradoriids. The evolution of more complex articulating hinge structures – together with well‐developed musculature – in ostracods during the Early Ordovician, may have provided more efficient means for shield articulation and movement, thus promoting the ecological success of ostracods throughout the Phanerozoic.