Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.     

Anostraca,Notostraca, Laevicaudata and Spinicaudata of Pannonian region and in its Austrian area

The northwestern area of the Pannonian Lowland extends into Austria. The climatic and hydrologic attributes of this biographic region promote the existence of extremely astatic bodies water lacking any fish and hence the occurrence of Anostraca, Notostraca, Laevicaudata and Spinicaudata. Zoogeographical and ecological features as well as the extinction of species due to anthropogenic influence are described.Dedicated to Prof. Dr F. Berger, Lunz, Austria, on the occasion of his 90th birthdayDedicated to Prof. Dr F. Berger, Lunz, Austria, on the occasion of his 90th birthday     

Sex determination of the Early Cretaceous clam shrimp Eosestheria middendorfii (Yixian Formation,China)

Clam shrimps have been recognized as a key group for the study of reproductive system evolution, owing to the diversity of sexual systems in extant members. However, there are comparatively little data on fossil taxa. In this study, we reveal the sexual system of the Early Cretaceous clam shrimp Eosestheria middendorfii (Yixian Formation, China). This is the first study that restricts the analysis to a single cohort, minimizing the otherwise considerable impact of ecophenotypic variation within this species. In addition, the presence of egg clutches, which identifies some individuals as either female or hermaphroditic, serves as an independent indicator for sex prior to the statistical treatment of the data. Obligate sexuality (‘dioecy’) is the inferred reproductive system for E. middendorfii and sexual dimorphism accounts for about 10% of the adult shape variation. Carapace shape variation resulting from malformation and deformation is more pronounced than the underlying sexual dimorphism. Subtle sexual dimorphism, smaller and slightly more elongate females, lateral egg clutches, and egg diameters of about 140 μm indicate that E. middendorfii is closely allied with the extant family Cyzicidae.     

Horizontal distribution and abundance of cysts of several large branchiopods in temporary pool and ditch sediments

Cysts of three species of anostracans (Branchipusschaefferi, Chirocephalus diaphanus, and Tanymastigites perrieri), two species of Notostracans (Triops numidicus and Lepidurus apus) and one species of Spinicaudata (Leptestheriamayeti) were collected from the upper 2 cm of dry sediments at intervals along transects from a temporary pool in an arid zone of Morocco and from two ditches in France. The horizontal distribution of viable cysts was patchy and is discussed in relation to several ecological parameters prevailing during the aquatic phase. These include vegetation distribution, depth and morphometry of the pond, dominant wind pattern, and granularity of the sediments. The densities reached more than 1700 cysts of C. diaphanus per 100 cm², and 1150 cysts of L. apus per 100 cm² in the two ditches. The mean densities of cysts per 100 cm² calculated for each transect ranged from 250 to 450 for C. diaphanous and 240 to 260 for L. apus (in the ditch populations) and about 1–6 for T. perrieri and L. mayeti, 3–15 for B. schaefferi, and16–58 for T. numidicus in the pool in Morocco. This revised version was published online in August 2006 with corrections to the Cover Date.     

Larval development of Lynceus brachyurus (Crustacea, Branchiopoda, Laevicaudata): redescription of unusual crustacean nauplii, with special attention to the molt between last nauplius and first juvenile

The larval development of "conchostracans" has received only scattered attention. Here I present the results of a study on the larval (naupliar) development and the metamorphosis of Lynceus brachyurus, a member of the bivalved branchiopod order the Laevicaudata. Lynceus brachyurus is the only species of the "Conchostraca" in Denmark. The phylogenetic position of the Laevicaudata has traditionally been a source of controversy, and this study does not solve the question completely. This work focuses on features potentially important for phylogeny. The general appearance of the larvae of L. brachyurus has been known for more than a century and a half, and some of its unique features include a large, larval dorsal shield; a huge, plate-like labrum; and a pair of immovable, horn-like antennules. However, many details relating to limb morphology, potentially important for phylogeny, have not been studied previously. Based on size categories, five or six larval stages can be recognized. The larvae approximately double their length and width during development (length: 230-520 microm). Most morphological features stay largely unchanged during development, but the antennal coxal masticatory spines are significant exceptions: they become bifid after one of the first molts. In all larval stages only the antennae and the mandibles actively move. In late naupliar stages the trunk limbs become visible as rows of laterally placed, undeveloped, and still immovable lobes. Swimming is performed by the antennae, whereas the mandibles appear to be involved mainly in feeding, as in other branchiopod larvae. The last naupliar stage undergoes a small metamorphosis to the first juvenile stage, the details of which in part were studied by following the premolt juvenile condition through the cuticle of the last stage nauplius. Among other changes there is a characteristic change in the shape and morphology of the univalved dorsal naupliar shield to a bivalved juvenile carapace. The general morphologies of the antennae and the mandibles are very similar to those of other branchiopod larvae and fall well within the "branchiopod naupliar feeding apparatus" recognized as a branchiopod synapomorphy by Olesen (2003), but some specific features shared with the larvae of other "conchostracans" are also identified. These special "conchostracan" features include: 1) a similar antennular setation; 2) a similar comb-like setulation of the bifid antennal coxal processes; and 3) mandibular palpsetae with setules condensed. In light of recent suggestions concerning branchiopod phylogeny (Cyclestheria as a sister group to the Cladocera), these similarities probably do not support a monophyletic "Conchostraca" but rather are symplesiomorphies of this taxon. A final decision must await a phylogenetic analysis of a more complete set of characters.     

The geographic distribution of the European Branchiopods (Anostraca,Notostraca, Spinicaudata,Laevicaudata)

The Large Branchiopod fauna of Europe consists of 72 species and sub-species: 50 anostracans, 8 notostracans, 12 spinicaudatans, and 2 laevicaudatans. Their geographical distribution is mapped, and remarks on ecological requirements of species are given.     

Epizootic algae and Protozoa on fresh water branchiopods (Anostraca,Notostraca and Spinicaudata) in Moroccan temporary ponds

Epizootic algae and Protozoa occur on the exoskeleton of fresh water branchiopods in temporary ponds in Morocco. Algal colonization seemed linked with turbid waters and access by the epibionts to the nutritive and oxygenated flows created by the rhythmical beat of the thoracic legs of the branchiopods. It also seemed linked to the mode of life (planktonic or benthic) of the host, and to their exposition to light. Appendages of anostracans, which swim ventral side up, bear a rich community of Chlorophytes, while in conchostracans, where body and appendages are protected by a bivalve carapace, and which swim ventral side down, epibionts colonize the valves, particularly near the hinge and umbo. The lack of algae on notostracans may result from their habit of living in the deeper part of ponds. Protozoans fixed on notostracan shields and on conchostracan valves or appendages, seem to benefit principally from the flow of nutrient- and oxygen-rich water created by the host.     

Ecological stasis in Spinicaudata (Crustacea,Branchiopoda)? Early Cretaceous clam shrimp of the Yixian Formation of north‐east China occupied a broader realized ecological niche than extant members of the group

The palaeoecology of Spinicaudata, the dominant group of benthic invertebrates in many pre‐Cenozoic freshwater environments, remains poorly understood. In analogy with extant taxa, it has been oversimplified and often reduced to shallow, temporary environments characterized by few trophic levels, implying ecological stasis from the Devonian to the Recent. We excavated 43 horizons of the Lower Cretaceous Yixian Formation (Anjiagou and Hengdaozi beds) to evaluate whether spinicaudatan ecology can be simplified to such an extent. Sedimentological evidence suggests general perennial conditions during the excavated lake interval. Based on 33 226 specimen counts, we identified three arthropod‐dominated macrobenthic associations and two assemblages. Response curves indicate that the spinicaudatan Eosestheria middendorfii was exceptionally tolerant to environmental gradients, followed, in decreasing order, by mayfly larvae, water boatmen and oligochaetes. Many spinicaudatan‐yielding layers represent death after reproductive senescence rather than mass‐mortality events. Spinicaudatan size varies significantly according to faunal association. A forward stepwise regression model suggests that growth responded to population density and diversity: lower densities and higher diversities triggered fast growth, and vice versa. The proposed strong density effect on carapace size has been corroborated by rearing experiments: as for E. middendorfii, natural log regression provided the best fit for the two extant species Eulimnadia texana and Eocyzicus argillaquus. Hence, E. middendorfii was a tolerant and morphologically variable species adapted to perennial waters and the frequent presence of higher trophic levels. Although there are rare records of extant taxa co‐occurring with fish, the described Early Cretaceous environment is commonly not inhabited by extant Spinicaudata.     

Multispecies coexistence of branchiopods (Anostraca,Notostraca & Spinicaudata) in temporary ponds of Chaouia plain (western Morocco): sympatry or syntopy between usually allopatric species

The study of the structure of communities of Crustacean branchiopods Anostraca, Notostraca and Spinicaudata of temporary ponds in Morocco revealed the existence of a geographical area with a particularly diversified number of species. Over a few hectares, 11 species, among the 18 known in Morocco, are present. While most of them are usually allopatric and belong to distinct associations, indicative of particular ecological and climatic conditions, they can coexist in Chaouia plain, either in adjoining ponds (sympatric species) or in the same pond (syntopic species). In one pond we could find up to 10 phyllopods: 6 Anostraca, 2 Notostraca and 2 Spinicaudata.This faunistic diversity may result from abiotic features (the location of the ponds at the boundary of two climatic areas) and from a heterogeneous geological substratum, creating a patchwork of temporary ponds with various mesologic conditions.Secondarily, this diversity may result from different life history characteristics of each coexisting species. As all species hatch synchronically, coexistence is only possible if their growth rates and life span are different. This implies that horizontal and vertical distribution, and exploitation of food supplies in the habitat differ for each ecophase of the syntopic species (spatial and trophic exclusion).     

Rediscovery of Leptestheria dahalacensis and Eoleptestheria ticinensis (Crustacea: Branchiopoda: Spinicaudata): an overview on presence and conservation of clam shrimps in Austria

According to recent literature, five of the six known Austrian conchostracan species are extinct. However, interim results of a current study on large freshwater branchiopods in Austria show that five species still occur at a restricted number of sites in the Pannonian region of Lower Austria. The clam shrimps Leptestheria dahalacensis and Eoleptestheria ticinensis were rediscovered in May 1994 in the flood plains of the river Morava near Marchegg. Imnadia yeyetta and Cyzicus tetracerus have been known to the authors in the same region since 1981, and 1992, respectively. Limnadia lenticularis occurs in the flood plains of the rivers Morava and Danube. Lynceus brachyurus, the only Austrian representative of the Laevicaudata, was not found and most probably got extinct. All Austrian clam shrimp species are considered to be endangered. Main threats are agricultural activities and artificial changes of the hydrologic conditions. Conservational measures are discussed for their effectivity.