Search

Abstract: Unresolved taxonomic issues regarding spinicaudatans, clam shrimps that formed the most abundant faunal element in the lacustrine Barremian to Aptian Yixian Formation, have hampered palaeoecological and evolutionary interpretations of this key fossil group. Here, we analyse morphological variability in East Asian clam‐shrimp taxa by quantifying: (1) size and shape; and (2) ornamental features (radial lirae distances). Intergeneric variability was examined using 51 specimens of various East Asian Mesozoic taxa, 16 of which were chosen for a taxonomic revision of the 10 species described from the Yixian Formation of western Liaoning. Geometric morphometrics proved to be effective for the separation of genera, while the quantification of ornamental features was species‐diagnostic. Various authors have previously indicated the possibility that East Asian clam‐shrimp species have been split excessively, a criticism that has been confirmed by our analyses. According to our results, four of the ten species described from the Yixian Formation are valid, of which one might represent a junior synonym of Diestheria jeholensis; five taxon names represent junior synonyms of Eosestheria middendorfii (Jones), and one a nomen dubium. According to lirae distances and analysis of shape, Eosestheria fuxinensis, the original type species of Eosestheria, is synonymous with the type of Diestheria. Therefore, a new type species for Eosestheria is warranted and here fixed as E. middendorfii. This taxonomic revision questions the high level of clam‐shrimp diversity recognized within the Yixian Formation, and indicates that many of the species described in fact represent various ontogenetic stages of E. middendorfii. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Studies of temperature effects on morphology in Spinicaudata have focused on length, with no data on shape. To fill this gap, size and shape variability in response to temperature fluctuations was investigated by rearing the modern spinicaudatan Eulimnadia texana . Two days after hydration, juvenile individuals were separated into four different temperature treatments: 20°C, 23°C, 26°C, and 29°C. Hermaphrodite size and shape were analysed by looking at linear combinations of size variables and using Fourier shape analysis; methods that are also used to describe fossil size and shape for better comparison. Size differences were considerable, with reduced growth at low and high temperatures and accelerated growth at the optimum temperature of 26°C, revealing that the reaction of size to increasing temperature is non-linear. The height of the dorsal margin, which is associated with space for egg production in Eulimnadia texana , accounts for a high amount of size variability in this species and, presumably, in most of the Limnadiidae. Hermaphrodite shapes reared under temperatures of 20°C and 29°C are statistically distinct, while intermediate temperatures yield intermediate shapes. The rate of shape change along temperature is comparatively low between 23°C and 26°C and accelerated at lower and higher temperatures. With increasing temperature, the highest point of the dorsal margin is shifted towards the anterior of the carapace, while it assumes a median position at 20°C. Our result that temperature has strong effects on carapace size and shape implies considerable ecophenotypic variability in Spinicaudata.

Fossil morphological data are time-averaged and generally reflect an overlap of different sources of carapace variability. To examine whether a proposed relationship between size and population density in fossil spinicaudatans is biologically meaningful, we set up rearing experiments involving two extant species: Eulimnadia texana and Eocyzicus argillaquus . Three and five days after hydration, clam shrimp were transferred into cups of various population densities that ranged between 1 and 15 inds/400 ml. Size and shape were measured 14 and 16 days after hydration, respectively. Every second day, we recorded length and sex of E. texana , which matured faster in lower-density cups. According to our growth model, population density and maximal carapace length follow a logarithmic relationship. At maturity, hermaphrodites yielded similar lengths across all population densities (~4.7 mm at 24°C), independent of age. Hence, clam shrimp can put off reproductive maturity as a response to decreased growth under higher density conditions. Growth rate generally decreases at maturity, but that effect is more pronounced in clam shrimp of high population densities, while low-density adults keep growing. For both species, multivariate analyses reveal that carapace size of low-density individuals is significantly larger than carapace size of higher-density individuals, while size values of intermediate densities cannot be distinguished. Shape distinction is strong in hermaphrodites of E. texana : 39.8% of the density-dependent shape variation is associated with relative umbo height, which is generally higher in individuals of smaller population densities. The H/L ratio, which is often used as a simple shape indicator, does not contribute to the main variation in shape, but it forms one of several ratios significant for 18.3% of the shape variability. In turn, the H/L ratio drives 30% of the shape variation in E. argillaquus . In addition, higher densities triggered shifts in ontogenetic growth trajectories in one third of the individuals, which led to aberrant morphologies. The present rearing experiment shows that some of the morphological variability on fossil bedding planes can be explained by population density. Also, it implies a considerable amount of ecophenotypic variability in Spinicaudata that affects our understanding of fossil taxonomy and palaeoecology.