Exceptional preservation of reproductive organs and giant sperm in Cretaceous ostracods

The bivalved crustacean ostracods have the richest fossil record of any arthropod group and display complex reproductive strategies contributing to their evolutionary success. Sexual reproduction involving giant sperm, shared by three superfamilies of living ostracod crustaceans, is among the most fascinating behaviours. However, the origin and evolution of this reproductive mechanism has remained largely unexplored because fossil preservation of such features is extremely rare. Here, we report exceptionally preserved ostracods with soft parts (appendages and reproductive organs) in a single piece of mid-Cretaceous Kachin amber (approximately 100 Myr old). The ostracod assemblage is composed of 39 individuals. Thirty-one individuals belong to a new species and genus, Myanmarcypris hui gen. et sp. nov., exhibiting an ontogenetic sequence from juveniles to adults (male and female). Seven individuals are assigned to Thalassocypria sp. (Cypridoidea, Candonidae, Paracypridinae) and one to Sanyuania sp. (Cytheroidea, Loxoconchidae). Our micro-CT reconstruction provides direct evidence of the male clasper, sperm pumps (Zenker organs), hemipenes, eggs and female seminal receptacles with giant sperm. Our results reveal that the reproduction behavioural repertoire, which is associated with considerable morphological adaptations, has remained unchanged over at least 100 million years—a paramount example of evolutionary stasis. These results also double the age of the oldest unequivocal fossil animal sperm. This discovery highlights the capacity of amber to document invertebrate soft parts that are rarely recorded by other depositional environments

Correlation between investment in sexual traits and valve sexual dimorphism in Cyprideis species (Ostracoda)

Assessing the long-term macroevolutionary consequences of sexual selection has been hampered by the difficulty of studying this process in the fossil record. Cytheroid ostracodes offer an excellent system to explore sexual selection in the fossil record because their readily fossilized carapaces are sexually dimorphic. Specifically, males are relatively more elongate than females in this superfamily. This sexual shape difference is thought to arise so that males carapaces can accommodate their very large copulatory apparatus, which can account for up to one-third of body volume. Here we test this widely held explanation for sexual dimorphism in cytheroid ostracodes by correlating investment in male genitalia, a trait in which sexual selection is seen as the main evolutionary driver, with sexual dimorphism of carapace in the genus Cyprideis. We analyzed specimens collected in the field (C. salebrosa, USA; C. torosa, UK) and from collections of the National Museum of Natural History, Washington, DC (C. mexicana). We digitized valve outlines in lateral view to obtain measures of size (valve area) and shape (elongation, measured as length to height ratio), and obtained several dimensions from two components of the hemipenis: the muscular basal capsule, which functions as a sperm pump, and the section that includes the intromittent organ (terminal extension). In addition to the assessment of this primary sexual trait, we also quantified two dimensions of the male secondary sexual trait-where the transformed right walking leg functions as a clasping organ during mating. We also measured linear dimensions from four limbs as indicators of overall (soft-part) body size, and assessed allometry of the soft anatomy. We observed significant correlations in males between valve size, but not elongation, and distinct structural parts of the hemipenis, even after accounting for their shared correlation with overall body size. We also found weak but significant positive correlation between valve elongation and the degree of sexual dimorphism of the walking leg, but only in C. torosa. The correlation between the hemipenis parts, especially basal capsule size and male valve size dimorphism suggests that sexual selection on sperm size, quantity, and/or efficiency of transfer may drive sexual size dimorphism in these species, although we cannot exclude other aspects of sexual and natural selection

A new Late Agenian (MN2a, Early Miocene) fossil assemblage from Wallenried (Molasse Basin, Canton Fribourg, Switzerland)

Excavations of two fossiliferous layers in the Wallenried sand- and marl pit produced a very diversified vertebrate fauna. New material allows the reassessment of the taxonomic position of the ruminant taxa Andegameryx andegaviensis and endemic Friburgomeryx wallenriedensis. An emended diagnosis for the second species is provided and additional material of large and small mammals, as well as ectothermic vertebrates, is described. The recorded Lagomorpha show interesting morphological deviations from other Central European material, and probably represent a unique transitional assemblage with a co-occurrence of Titanomys, Lagopsis and Prolagus. Rodentia and Eulipotyphla belong to typical and well-known species of the Agenian of the Swiss Molasse Basin. Abundant small mammal teeth have allowed us to pinpoint the biostratigraphic age of Wallenried to late MN2a. The biostratigraphic age conforms to data derived from the charophyte assemblages and confirms the oldest occurrence of venomous snake fangs. The palaeoenvironmental context is quite complex. Sedimentary structures and fauna (fishes, frogs, salamanders, ostracods) are characteristic for a humid, lacustrine environment within a flood plain system

On <i>Afrocypris barnardi</i> G. O. Sars, 1924 (Ostracoda), a second giant ostracode with additional appendages

Afrocypris barnardi G. O. Sars, 1924, a giant temporary pool ostracode from South Africa, is here redescribed, based on new material. It is only the second time that this species has been found. The present morphological analyses show that the copulatory appendages of both sexes have unusual characteristics. In the female, at least three paired appendage-like structures (R-appendages) are associated with the Female Reproductive Organ (FRO). This is the second time that such structures are described from non-marine ostracodes, Liocypris grandis (G. O. Sars, 1924) having six pairs of such structures. The male copulatory organs have a Penile Attachment uniting the proximal sides of both hemipenes, and in Afrocypris barnardi an asymmetrical, finger-like structure is associated with this PA. The discovery of female R-appendages in a second species strengthen the interpretation of these structures as (part of) true appendages, and reinforces the hypothesis that the FRO is homologous to more than one ancestral somite. However, it remains unclear how many body segments are involved in the origin of the FRO, and three, five, or even six somites are all possible. In addition, these R-appendages can be ancestral, atavistic, or even apomorphic. The position of the two taxa with such appendages in a phylogeny of the Cyprididae as a whole will constitute a test to discriminate between these three possibilities

Experimental assessment of the fecundity of Eucypris virens (Ostracoda, Crustacea) under natural sex ratios

1. The adaptive significance of the observed variations in sex ratios in non-marine ostracods is unclear. This study quantified the fecundity of females taken from a presumed fully sexual Eucypris virens population that were experimentally combined with different proportions of males (male : female sex ratios: 1 : 1, 1 : 2, 1 : 4, 1 : 8 and 0 : 1). 2. The results yielded no indications that female fecundity is altered by short-term variations in the proportion of males, at least not within the range of sex ratios that are common in natural ostracod populations. Complete removal of males, however, did strongly reduce hatching success of dried eggs. This suggests the need for multiple mating events during the reproductive lifetime of the female. It also emphasizes the need for a minimum number of males, although this minimum number evidently may be rather low, as we found a high number of spermatozoa in the seminal receptacles after a single mating event. 3. The sex ratio in the source population was strongly female biased (1 : 3.4; n = 514), whereas in the hatchling assemblages reared in the laboratory, males and females were found in equal proportions (1 : 1.0; n = 1516), irrespective of the prevailing sex ratio. This clear discrepancy is intriguing, and points to the importance of epigenetic factors for the determination of field sex ratios

Morphology of dry-resistant eggs in parthenogenetic Heterocypris incongruens

It has been known that many organisms evolved to survive in temporary or ephemeral inland waters. Many of them have dry-resistant eggs against desiccation. The structural feature of egg shell is important because only this will ensure to survive the dry period. Structural features of egg shell in the parthenogenetic Heterocypris incongruens (Ramdohr, 1808) was investigated by scanning electron microscope. Results showed that egg shell structure consists of two distinct layers; an outer layer with holes or alveoli and an inner layer consisting of two dense sublayers. Also, structural similarities in egg-shell of H. incongruens and some other crustaceans which combat desiccation problem will be discussed