A new 'great-appendage' arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages

The uniramous ‘great appendages’ of several arthropods from the Early to Middle Cambrian are a characteristic pair of pre‐oral limbs, which served for prey capture. It has been assumed that the morphological differences between the ‘great‐appendage’ arthropods indicate that raptorial antero‐ventral and anteriorly pointing appendages evolved more than once in arthropod phylogeny. One set of Cambrian ‘great‐appendage’ arthropods has, however, very similar short antero‐ventral appendages with a peduncle of two segments angled against each other (elbowed) and with stout distally or medio‐distally directed spines or long flexible flagellate spines on each of the four distal segments. Moreover, the head appendages of all these forms comprise the ‘great appendages’ and three pairs of biramous limbs. To this set of taxa we can add a new form from the Lower Cambrian Maotianshan Shale of southern China, Haikoucaris ercaiensis n. gen. and n. sp. It is known from three specimens, possibly being little abundant in the faunal community. It can be distinguished from all other taxa by the prominence of the proximal claw segment of its ‘great appendages’ and by only three distal spines (one on each of the distal segments). The similarity of the short, spiky ‘great appendages’ of Haikoucaris with the chelicera of the Chelicerata leads us to hypothesize that this particular type of ‘great appendages’ was the actual precursor of the chelicera. Homeobox gene and developmental data recently demonstrated the homology between the antenna of ateloceratans and the antennula of crustaceans on one side and the chelicera of chelicerates on the other. To this we add palaeontological evidence for the homology between the chelicerae of chelicerates and the ‘short great appendages’ of certain Cambrian arthropods, which leads us to hypothesize that the evolutionary path went from the ‘short great appendages’, by progressive compaction, toward the chelicera with only a two‐spined chela. The new form from China is regarded as the possible latest offshoot, whereas the other ‘great appendages’ arthropods with similar short grasping limbs were derivatives of the stem lineage of the crown‐group Chelicerata. Consequently, the chelicera with a chela with one fixed and one mobile finger is an autapomorphy of the crown group of Chelicerata, whereas a raptorial, but more limb‐like antenna, with more distal spine‐bearing segments, characterized the ground pattern of Chelicerata. Further taxa having ‘great appendages’, including the large Anomalocarididae, are also discussed in the light of their possible affinities to the Chelicerata and possible monophyly of all of these arthropods with raptorial anterior appendages.     

The input and fate of new C in two forest soils under elevated CO2

The aim of this study was to estimate (i) the influence of different soil types on the net input of new C into soils under CO₂ enrichment and (ii) the stability and fate of these new C inputs in soils. We exposed young beech–spruce model ecosystems on an acidic loam and calcareous sand for 4 years to elevated CO₂. The added CO₂ was depleted in ¹³C, allowing to trace new C inputs in the plant–soil system. We measured CO₂‐derived new C in soil C pools fractionated into particle sizes and monitored respiration as well as leaching of this new C during incubation for 1 year. Soil type played a crucial role in the partitioning of C. The net input of new C into soils under elevated CO₂ was about 75% greater in the acidic loam than in the calcareous sand, despite a 100% and a 45% greater above‐ and below‐ground biomass on the calcareous sand. This was most likely caused by a higher turnover of C in the calcareous sand as indicated by 30% higher losses of new C from the calcareous sand than from the acidic loam during incubation. Therefore, soil properties determining stabilization of soil C were apparently more important for the accumulation of C in soils than tree productivity. Soil fractionation revealed that about 60% of the CO₂‐derived new soil C was incorporated into sand fractions. Low natural ¹³C abundance and wide C/N ratios show that sand fractions comprise little decomposed organic matter. Consistently, incubation indicated that new soil C was preferentially respired as CO₂. During the first month, evolved CO₂ consisted to 40–55% of new C, whereas the fraction of new C in bulk soil C was 15–23% only. Leaching of DOC accounted for 8–23% of the total losses of new soil C. The overall effects of CO₂ enrichment on soil C were small in both soils, although tree growth increased significantly on the calcareous sand. Our results suggest that the potential of soils for C sequestration is limited, because only a small fraction of new C inputs into soils will become long‐term soil C.     

Are psyllids affiliated with the threatened plants Acacia ausfeldii,A. dangarensis and A. gordonii at risk of co‐extinction?

Secondary extinctions of dependent affiliate species may exceed the numbers of species directly extirpated with habitat loss. We investigated the host specificity, and hence the co‐extinction risk, of multiple psyllid species (Hemiptera: Psylloidea) on the threatened plants Acacia ausfeldii, A. dangarensis and A. gordonii by sampling from these and other sympatric common plants across multiple sites, years and plant phenologies. We found one specialist psyllid species (Psyllidae: Acizzia sp.) on each of the plants A. ausfeldii and A. gordonii but no specialists on A. dangarensis. The A. gordonii specialist only occurred when its host was in flower. Specialist psyllids considered at risk of extinction represented 10% of all psyllid species observed on the three threatened acacias. Potential risks to co‐endangered affiliates need to be considered when implementing threatened plant recovery actions. Conservation of plant populations and plant species diversity is also necessary to maintain high insect diversity.     

Exceptionally well‐preserved isolated eyes from Cambrian ‘Orsten’ fossil assemblages of Sweden

Abstract: Eyes other than those of trilobites are rarely preserved in the fossil record. We describe here a set of six tiny, isolated, three‐dimensionally preserved compound eyes. These secondarily phosphatized eyes were etched from ‘Orsten’ limestone nodules dated to the Agnostus pisiformis Biozone from the Cambrian Alum Shale Formation of Sweden. The ovoid eyes arise from an elongated stalk, their surface being covered by a mosaic of regular and hexagonal‐shaped facets representing the surface of ommatidia. Facet size and pattern change within the same specimen from the posterior to the anterior end. With regard to some morphological criteria, we grouped the material in two different morphotypes, type A and B, the first being represented by specimens of two different developmental stages. From stage to stage, mostly growth in overall size and addition of new ommatidia was noticed. Among the meiobenthic ‘Orsten’ arthropods, only the crustacean Henningsmoenicaris scutula has been described as possessing stalked eyes, but the eyes of the largest specimen with preserved eyes of this species are much smaller than the new eyes and do not display any kind of ommatidia on their visual surface. However, fragments of larger specimens of H. scutula and the co‐occurrence of this species with the new isolated eyes in the sieving residues make it likely that the latter belong to this species but belong to more advanced stages than those described previously of H. scutula. Ontogenetically, the eye stalks of this fossil crustacean elongate progressively, while the regular hexagonal facets, lacking in early stages, appear later on.     

Characterization of Two Sibling Species of the Genus Stylonychia (Ciliata,Hypotricha): S. mytilus Ehrenberg, 1838 and S. lemnae n. sp. I. Morphology and Reproductive Behavior1

All Stylonychia mytilus-like ciliates which were collected and sent to us during the last 20 years belonged either to S. mytilus or to a new species, S. lemnae, which is described here. The only morphological differences are the shape and the size. Stylonychia mytilus that have been starved for one day average 300 μm in length, and S. lemnae starved for a day are 230 μm long. The occurrence of mating types is described.     

DAS THERMO-HYGRO-LABYRINTH

Vorzugstemperatur und Vorzugsluftfeuchtigkeit konnten bisher nur getrennt ermittelt werden, so daß zusammenhängende Präferenzänderungen zwischen Vorzugstemperatur und Vorzugsluftfeuchtigkeit nicht erfaßbar waren. Mit Hilfe einer neuen Apparatur, dem Thermo-Hygro-Labyrinth, wurde es möglich, Vorzugstemperatur und Vorzugsluftfeuchtigkeit gleichzeitig zu bestimmen. Das neue Gerät besteht aus einem “Labyrinth” von Kammern mit unterschiedlichen Temperaturen und Luftfeuchtigkeiten. Sie sind durch Laufröhren miteinander verbunden, so daß die Versuchstiere jede beliebige Kammer aufsuchen können. Die Kammern sind in fünf parallelen Längsreihen mit je sieben Kammern angeordnet. Jede Reihe weist die gleiche Temperatur auf. Es können somit insgesamt fünf Temperaturstufen eingestellt werden. Rechtwinklig zur Längsrichtung gesehen weist jede der sieben Querreihen mit ihren fünf Kammern die gleiche Luftfeuchtigkeit auf. Dabei kann von Querreihe zu Querreihe die Luftfeuchtigkeit stufenweise steigen. Es stehen also maximal sieben Feuchtigkeitsstufen zur Verfügung. Zur Regulierung der Lufttemperatur in den Kammern steht jede Kammerlängsreihe in einem separat temperierbaren Wasserbecken. Die verschiedenen Luftfeuchtigkeiten werden durch gesättigte Salzlösungen bewirkt.     

Martinssonia elongata gen. et sp.n., a crustacean-like euarthropod from the Upper Cambrian 'Orsten' of Sweden

A tiny arthropod, with five growth stages, is described. Three of the instars are metanauplius-like larvae, having unsegmented bodies and four pairs of appendages. The largest stage, with a length of about 1.5 mm, may still be immature. Its body is divided into three tagmata. The cephalon, including five appendiculate segments, h a projecting forehead with a rostral spine and a small shield with a joint between fourth and fifth segments. Eyes are absent. The trunk is composed of seven annular segments, the anterior two with appendages. The caudal end is a long pleotelson-like segment with the anus on its ventral surface. There are seven pairs of appendages: uniramous antennulae, composed of few tubular podomeres; four pairs of biramous postantennular, almost homeomorphic cephalic appendages; two pairs on the trunk, the anterior pair being similar to the cephalic appendages except for the exopodite, the posterior being much smaller, uniramous and apparently rudimentary. Martinssonia was probably benthic, feeding on detritic particles which it stirred up from the bottom. Besides various crustacean-like features, the new form reveals structures different from Crustacea as well as from all other known arthropodan groups. Martinssonia presumably is a descendant of an euarthropodan group, originating from the crustacean branch long before reaching the eucrustacean level of evolution.     

Functional morphology,ontogeny and evolution of mantis shrimp‐like predators in the Cambrian

Abstract: We redescribe the morphology of Yohoia tenuis (Chelicerata sensu lato) from the Cambrian Burgess Shale Lagerstätte. The morphology of the most anterior, prominent, so‐called great appendage changes throughout ontogeny. While its principal morphology remains unaltered, the length ratios of certain parts of the great appendage change significantly. Furthermore, it possesses a special jack‐knifing mechanism, i.e. an elbow joint: the articulation between the distal one of the two peduncle elements and the most proximal of the four spine‐bearing claw elements. This morphology might have enabled the animal to hunt like a modern spearer‐type mantis shrimp, an analogy enhanced by the similarly large and protruding eyes. For comparison, details of specimens of selected other great‐appendage arthropods from the Lower Cambrian Chengjiang Lagerstätte have been investigated using fluorescence microscopy. This revealed that the morphology of the great appendage of Y. tenuis is much like that of the Chengjiang species Fortiforceps foliosa and Jianfengia multisegmentalis. The morphology of the great appendage of the latter is even more similar to the morphology developed in early developmental stages of Y. tenuis, while the morphology of the great appendage of F. foliosa is more similar to that of later developmental stages of Y. tenuis. The arrangement of the elbow joint supports the view that the great appendage evolved into the chelicera of Chelicerata sensu stricto, as similar joints are found in various ingroup taxa such as Xiphosura, Opiliones or Palpigradi. With this, it also supports the interpretation of the great appendage to be homologous with the first appendage of other arthropods.     

The youngest African clymeniids (Ammonoidea, Late Devonian) – failed survivors of the Hangenberg Event

A co-occurrence of the clymeniid Postclymenia evoluta Schmidt, 1924 and the goniatite Acutimitoceras hilarum Korn, 2002 is reported from the Anti-Atlas of Morocco. Both species occur in the same limestone horizon within the Acutimitoceras prorsum Zone, that has yielded an exclusive conodont fauna of the Upper praesulcata Zone (latest Devonian, above the Hangenberg Black Shale). This record is firm evidence that some clymeniids survived the global Hangenberg Event, but soon later became extinct without descendants.     

Evidence of a Cadmium-Thionein and the Glycine Cleavage System in Oxytricha granulifera

ABSTRACT. This work presents the further purification of a Cd-linking protein in Oxytricha granulifera by reverse-phase chromatography. This protein contains 25% cysteine and no aromatic amino acid. it may be considered as a chelatin with some similarity to metallothioneins. During the purification of another Cd-linking compound, we were able to demonstrate that the H protein precursor of glycine cleavage is present in Oxytricha. This is the first finding of the presence of this system in Protozoa.