BackgroundDespite extensive knowledge on bivalve anatomy and development, the formation and differentiation of the mantle margin and its associated organs remain largely unclear. Bivalves from the family Pectinidae (scallops) are particularly promising to cast some light on these issues, because they exhibit a complex mantle margin and their developmental stages are easily obtained from scallop farms. We investigated the mantle margin of the scallop Nodipecten nodosus (L. 1758) during larval and postmetamorphic development.MethodsA thorough analysis of the mantle margin development in Nodipecten nodosus, from veliger larvae to mature adults, was conducted by means of integrative microscopy techniques, i.e., light, electron, and confocal microscopy.ResultsInitially unfolded, the pallial margin is divided into distal and proximal regions by the periostracum-forming zone. The emergence of the pallial musculature and its neural innervation are crucial steps during bivalve larval development. By the late pediveliger stage, the margin becomes folded, resulting in a bilobed condition (i.e., outer and inner folds), a periostracal groove, and the development of different types of cilia. After metamorphosis, a second outgrowth process is responsible for emergence of the middle mantle fold from the outer surface of the inner fold. Once the three-folded condition is established, the general adult features are rapidly formed.ConclusionsOur data show that the middle mantle fold forms from the outer surface of the inner fold after metamorphosis and that the initial unfolded mantle margin may represent a common condition among bivalves. The first outgrowth process, which gives rise to the outer and inner folds, and the emergence of the pallial musculature and innervation occur during larval stages, highlighting the importance of the larval period for mantle margin morphogenesis in Bivalvia.
Crickets are frequently used as a model in several areas of science, including acoustic communication, behaviour and neurobiology. However, only a few of these studies are placed in an evolutionary framework due to the limited number of phylogenetic hypotheses for true crickets. We present a phylogenetic hypothesis for a newly defined family of crickets, Oecanthidae defin. nov., sister-group of Gryllidae defin. nov. The phylogenetic analyses are based on molecular and morphological data under likelihood and parsimony criteria and molecular data for divergence-times estimation (Bayesian inference). We used 107 terminals from all biogeographic regions and six fossils for the time calibration of the tree. All analyses resulted in Oecanthidae with four subfamilies: Euscyrtinae, Oecanthinae defin. nov., Podoscirtinae defin. nov. and Tafaliscinae defin. nov. Based on our results, we revise the definition and internal classifications of the subfamilies, supertribes and tribes. A new tribe, Phyllogryllini trib. nov. is described. We also update their diagnoses, list the genera of the tribes and list their apomorphies. We provide an identification key for all suprageneric taxa of Oecanthidae, plus all genera of Tafaliscinae. Finally, we discuss the phylogenetic relationships of Oecanthidae, their divergence times, habitat diversity and the importance of ovipositor variation in this clade.
Convergent morphology is a strong indication of an adaptive trait. Marine mussels (Mytilidae) have long been studied for their ecology and economic importance. However, variation in lifestyle and phenotype also make them suitable models for studies focused on ecomorphological correlation and adaptation. The present study investigates mantle margin diversity and ecological transitions in the Mytilidae to identify macroevolutionary patterns and test for convergent evolution. A fossil-calibrated phylogenetic hypothesis of Mytilidae is inferred based on five genes for 33 species (19 genera). Morphological variation in the mantle margin is examined in 43 preserved species (25 genera) and four focal species are examined for detailed anatomy. Trait evolution is investigated by ancestral state estimation and correlation tests. Our phylogeny recovers two main clades derived from an epifaunal ancestor. Subsequently, different lineages convergently shifted to other lifestyles: semi-infaunal or boring into hard substrate. Such transitions are correlated with the development of long siphons in the posterior mantle region. Two independent origins are reconstructed for the posterior lobules on the inner fold, which are associated with intense mucociliary transport, suggesting an important cleansing role in epifaunal mussels. Our results reveal new examples of convergent morphological evolution associated with lifestyle transitions in marine mussels.
Tentacles are remarkable anatomical structures in invertebrates for their diversity of form and function. In bivalves, tentacular organs are commonly associated with protective, secretory, and sensory roles. However, anatomical details are available for only a few species, rendering the diversity and evolution of bivalve tentacles still obscure. In Pteriomorphia, a clade including oysters, scallops, pearl oysters, and relatives, tentacles are abundant and diverse. We investigated tentacle anatomy in the group to understand variation, infer functions, and investigate patterns in tentacle diversity. Six species from four pteriomorphian families (Ostreidae, Pinnidae, Pteriidae, and Spondylidae) were collected and thoroughly investigated with integrative microscopy techniques, including histology, scanning electron microscopy, and confocal microscopy. Tentacles can be classified as middle fold tentacles (MFT) and inner fold tentacles (IFT) according to their position with respect to the folds of the mantle margin. While MFT morphology indicates intense secretion of mucosubstances, no evidence for secretory activity was found for IFT. However, both tentacle types have appropriate ciliary distribution and length to promote mucus transportation for cleaning and lubrication. Protective and sensory functions are discussed based on different lines of evidence, including secretion, cilia distribution, musculature, and innervation. Our results support the homology of MFT and IFT only for Pterioidea and Ostreoidea, considering their morphology, the presence of ciliated receptors at the tips, and branched innervation pattern. This is in accordance with recent phylogenetic hypotheses that support the close relationship between these superfamilies. In contrast, major structural differences indicate that MFT and IFT are probably not homologous across all pteriomorphians. By applying integrative microscopy, we were able to reveal anatomical elements that are essential for the understanding of homology and function when dealing with such superficially similar structures.
Evaluating the association between ecology and the evolution of morphological traits is a central question of macroevolutionary studies. The Arcoidea comprises several lineages of epibyssate (epifaunal) and endobyssate (semi‐infaunal or infaunal) bivalves, which makes them a suitable model for analyzing the implications of similar life habits on phenotype. Also, the families currently assigned to the clade (Arcidae, Cucullaeidae, Glycymerididae, and Noetiidae) exhibit great variation in the mantle margin, a region responsible for interacting with the surrounding environment and provided with sensory structures (e.g., photoreceptor organs). Consequently, this anatomical region includes traits relevant for testing hypotheses of morphological attributes associated with lifestyles. For this purpose, we investigated the detailed anatomy of the mantle margin of three arcoidean species (i.e., Arca imbricata, Barbatia candida, and Arcopsis adamsi) applying integrative microscopy, and complemented the study with basic morphological analysis of 27 species obtained in museum collections. Secretory cells, muscle fibers and sensory organs are described in detail, including sensory ciliated cells of the inner mantle fold, reported for the first time. Our results corroborate a very reduced to absent middle fold in arcoideans, with transfer of sensory functions to the first outer and inner folds. Moreover, our data indicates association of some pallial characters with life habits, i.e., presence of pallial pigmentation and photoreceptor organs in epibyssates, and development of an enlarged pallial curtain in endobyssates, which should stimulate further investigations on these potentially adaptive morphological features.
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