Development Mode Research Articles

Development of eodiscinid trilobites.

A comprehensive review of a full developmental sequence of eodiscinid trilobites reported in recent decades from Cambrian Series 2 and 3 strata is presented. These mostly articulated specimens exhibit detailed morphologies with preservation of even delicate structures at different growth stages, such as the bacculae, axial pores, spine pores, pygidial marginal spines and line coaptative structures. Their trunk segmentation schedules displayed a consistent developmental mode in segment generation and liberation, that is tagmosis and somitogenesis occurred heterochronously after each moulting event, providing clues regarding the potential developmental strategy in isopygous and even macropygous trilobites. The fact that the rate of segmentation obviously exceeds that of articulation results in a seemingly prolonged process of the formation of thoracic segments, which might explain why eodiscinid trilobites have a fixed and limited number of thoracic segments. In addition, the relationship between enrollment mechanism and trunk segmentation during eodiscinid ontogeny confirms this highly unusual growth pattern among the Trilobita, revealing why these early-diverging trilobites controlled the rate of segment increase and release during their life cycles, and is thus of interest with regard to the evolution of arthropod body patterning.

Morphological variation in first-formed shells of the Ordovician Paucicrura–Diceromyonia brachiopod lineage of North America

Abstract First-formed shells of several species of Dalmanellidae (Brachiopoda) from the Ordovician (Katian) of North America were measured and compared: Cincinnetina multisecta, Diceromyonia tersa, Diceromyonia storeya, Paucicrura corpulenta, Paucicrura rogata, and Paucicrura sillimani. Sizes and structures of the first-formed shells suggest that members of this family had planktotrophic subadults, with some species showing indications of only an unshelled larval stage and others showing both a larval stage and a shelled juvenile stage. This differs from modern rhynchonelliformean brachiopods, which all possess only a lecithotrophic larval stage. The range of sizes of first-formed shells of most studied species are similar, but P. sillimani of Baffin Island, Canada (middle Katian), has a significantly larger first-formed shell that formed during an extended juvenile stage. This may have enabled the species to colonize newly exploitable habitats during an interval of rapid sea level rise in Laurentia during the Katian. This plasticity of developmental modes in the Dalmanellidae shows not only that using distantly related modern brachiopods as an analog for extinct Paleozoic lineages may be misleading, but also that development can vary within a single lineage and that timing of developmental stages should not be considered a reliable character for use in phylogenetic studies of brachiopods.

Multiple origins of freshwater invasion and parental care reflecting ancient vicariances in the bivalve family Cyrenidae (Mollusca)

Habitat transitions in living organisms are key innovations often coupled with species diversification after their successful adaptation to new environment. The Cyrenidae is among the most well-known heterodont bivalve groups that have successfully invaded freshwater systems from brackish water environments and display diverse lineage-specific developmental modes. Phylogenetic and molecular clock-based divergence time analyses using 12 complete mitochondrial genome sequences suggest that Cyrenidae species independently colonized freshwater habitats during three distinct spatial and geological periods: one from the American continents approximately in the Early Jurassic and the two others from Australasian/East Asian continents in the Early/Middle Cretaceous and the Paleogene-Neogene boundary, respectively. This study provides significant insight into the temporal and spatial patterns of multiple freshwater invasions, aligning with ancient vicariance events inferred from different geological timelines of plate tectonics. Additionally, mitogenome phylogeny confirms the earlier hypothesis of the repeated parallel evolution of parental care system within this bivalve group.

Genome and life-history evolution link bird diversification to the end-Cretaceous mass extinction.

Complex patterns of genome evolution associated with the end-Cretaceous [Cretaceous-Paleogene (K-Pg)] mass extinction limit our understanding of the early evolutionary history of modern birds. Here, we analyzed patterns of avian molecular evolution and identified distinct macroevolutionary regimes across exons, introns, untranslated regions, and mitochondrial genomes. Bird clades originating near the K-Pg boundary exhibited numerous shifts in the mode of molecular evolution, suggesting a burst of genomic heterogeneity at this point in Earth's history. These inferred shifts in substitution patterns were closely related to evolutionary shifts in developmental mode, adult body mass, and patterns of metabolic scaling. Our results suggest that the end-Cretaceous mass extinction triggered integrated patterns of evolution across avian genomes, physiology, and life history near the dawn of the modern bird radiation.

Radula formation and development in Limapontia senestra (Quatrefages, 1844) (Heterobranchia, Gastropoda) from the White Sea

Sacoglossa is a monophyletic group of small suctorial herbivorous Нeterobranchia. Due to the biology of feeding, most sacoglossans (except Cylindrobulla) have a distinctive feeding apparatus, which includes a sucking muscular organ as well as a sacoglossan-type radula with one tooth in a transverse row, which pierces algal cells. Although there is obvious interest in the radular morphology and developmental modes of Sacoglossa, as these characters may provide some insight into the evolutionary history and phylogenetic relationships of Sacoglossa, data on radula formation and morphogenesis in this group are sparse. This paper presents original data on the direct development, formation, and morphogenesis of the radula of Limapontia senestra (Quatrefages, 1844) from the White Sea. Based on the data obtained, we describe the development and formation of the radula. They fit into the general pattern of heterochrony in the radula anlage, characteristic of all gastropods. The small number of odontoblasts in the formation of the adult radula of L. senestra is a distinctive feature of the radula of Heterobranchia.

Experimental evolution of multicellularity via cuboidal cell packing in fission yeast

Abstract The evolution of multicellularity represents a major transition in life’s history, enabling the rise of complex organisms. Multicellular groups can evolve through multiple developmental modes, but a common step is the formation of permanent cell–cell attachments after division. The characteristics of the multicellular morphology that emerges have profound consequences for the subsequent evolution of a nascent multicellular lineage, but little prior work has investigated these dynamics directly. Here, we examine a widespread yet understudied emergent multicellular morphology: cuboidal packing. Extinct and extant multicellular organisms across the tree of life have evolved to form groups in which spherical cells divide but remain attached, forming approximately cubic subunits. To experimentally investigate the evolution of cuboidal cell packing, we used settling selection to favor the evolution of simple multicellularity in unicellular, spherical Schizosaccharomyces pombe yeast. Multicellular clusters with cuboidal organization rapidly evolved, displacing the unicellular ancestor. These clusters displayed key hallmarks of an evolutionary transition in individuality: groups possess an emergent life cycle driven by physical fracture, group size is heritable, and they respond to group-level selection via multicellular adaptation. In 2 out of 5 lineages, group formation was driven by mutations in the ace2 gene, preventing daughter cell separation after division. Remarkably, ace2 mutations also underlie the transition to multicellularity in Saccharomyces cerevisiae and Candida glabrata, lineages that last shared a common ancestor >300 million years ago. Our results provide insight into the evolution of cuboidal cell packing, an understudied multicellular morphology, and highlight the deeply convergent potential for a transition to multicellular individuality within fungi.

A hundred-year-old mystery—the reproductive mode and larval morphology of the enigmatic frog genus Allophryne (Amphibia; Anura; Allophrynidae)

Frogs of the Allophrynidae are an enigmatic family from South America. To date, published information is lacking regarding this group’s reproductive biology and larval morphology. Here, we provide the first detailed description of the reproductive mode, developmental mode, and tadpole morphology for Allophryne ruthveni. We developed a captive breeding and rearing protocol for this species and then conducted a series of observations to describe aspects of its reproductive biology. In captivity, this species exhibits aquatic oviposition, where single eggs are laid ungrouped within a simple jelly capsule and are scattered free in the water column before sinking to develop on benthic substrates. We did not observe parental care nor any parental interactions with eggs post-fertilization. Tadpoles are characterized by an oval body, anteroventral oral disc, a labial tooth row formula of 2(2)/3, and a dextral vent tube. The buccopharyngeal cavity is marked by the presence of two pairs of infralabial papilla and four lingual papillae. Cranial morphology is characterized by the presence of the commissura quadratoorbital. This species possesses an additional slip of the m. rectus cervicis and of the m. levator arcuum branchialium III. We discuss our results in comparison with glassfrogs (Centrolenidae).

Both high and low dispersal? Apparently contradictory genetic patterns in the Antarctic littorinid gastropod Laevilacunaria antarctica

How the near-shore marine benthic communities survived Quaternary glaciations in Antarctica is a major question for Southern Ocean biogeographers. Several hypotheses that consider life-history traits, such as bathymetric ranges and developmental modes, have been proposed. Near-shore species with high dispersive potential are expected to show star-like genealogies, with broadly distributed haplotypes surrounded by closely related variants at low frequencies, a consequence of rapid population post-glacial expansion mediated by larvae. By contrast, shallow-water taxa with low dispersal potential are expected to evidence marked genetic structure or even consist of separate cryptic species, due to their survival in small refugial populations subject to repeated bottlenecks. In this study we performed phylogeographic analyses on the high-latitude littorinid Laevilacunaria antarctica across its distribution from maritime Antarctica to South Georgia. This species is a highly abundant near-shore gastropod, commonly found on macroalgae, with low autonomous vagility and exhibits benthic protected development with no free-living dispersive stages. Such characteristics make this species a suitable model to test the above hypotheses. Contrary to low dispersal potential predictions, L. antarctica comprises a single unstructured unit across its distribution in maritime Antarctica, exhibiting a typical high-dispersal star-like, short-branched genealogy centered on a dominant broadly distributed haplotype. The South Georgia population, however, consists of a separate genetic unit, strongly differentiated from those of the maritime Antarctic. We estimate that these two genetic groups separated ~1.2 My, long before the Last Glacial Maximum and evolved independently in these areas. Our results thus provide partial support for the expected pattern for a near-shore marine benthic species with low innate dispersive potential. On a wider geographic scale (maritime Antarctica vs South Georgia), our results in L. antarctica provide support for the expected pattern for a near-shore marine benthic species with low innate dispersive potential. However, on a narrower scale (maritime Antarctica) life-history characters including the close association of the species with macroalgae, would allow dispersal and a species with low dispersal potential exhibits paradoxically an absence of genetic structure across hundreds of kilometers probably mediated by rafting.

Tolerant and Rapid Endochondral Bone Regeneration Using Framework-Enhanced 3D Biomineralized Matrix Hydrogels.

Tissue-engineered bone has emerged as a promising alternative for bone defect repair due to the advantages of regenerative bone healing and physiological functional reconstruction. However, there is very limited breakthrough in achieving favorable bone regeneration due to the harsh osteogenic microenvironment after bone injury, especially the avascular and hypoxic conditions. Inspired by the bone developmental mode of endochondral ossification, a novel strategy is proposed for tolerant and rapid endochondral bone regeneration using framework-enhanced 3D biomineralized matrix hydrogels. First, it is meticulously designed 3D biomimetic hydrogels with both hypoxic and osteoinductive microenvironment, and then integrated 3D-printed polycaprolactone framework to improve their mechanical strength and structural fidelity. The inherent hypoxic 3D matrix microenvironment effectively activates bone marrow mesenchymal stem cells self-regulation for early-stage chondrogenesis via TGFβ/Smad signaling pathway due to the obstacle of aerobic respiration. Meanwhile, the strong biomineralized microenvironment, created by a hybrid formulation of native-constitute osteogenic inorganic salts, can synergistically regulate both bone mineralization and osteoclastic differentiation, and thus accelerate the late-stage bone maturation. Furthermore, both in vivo ectopic osteogenesis and in situ skull defect repair successfully verified the high efficiency and mechanical maintenance of endochondral bone regeneration mode, which offers a promising treatment for craniofacial bone defect repair.

Complex effects of the exo-xis region of the Shiga toxin-converting bacteriophage Φ24B genome on the phage development and the Escherichia coli host physiology

Lambdoid bacteriophages are excellent models in studies on molecular aspects of virus-host interactions. However, some of them carry genes encoding toxins which are responsible for virulence of pathogenic strains of bacteria. Shiga toxin-converting bacteriophages (Stx phages) encode Shiga toxins that cause virulence of enterohemorrhagic Escherichia coli (EHEC), and their effective production depends on Stx prophage induction. The exo-xis region of the lambdoid phage genome consists of genes which are dispensable for the phage multiplication under laboratory conditions; however, they might modulate the virus development. Nevertheless, their exact effects on the phage and host physiology remained unclear. Here, we present results of complex studies on the role of the exo-xis region of bacteriophage Φ24B, one of Stx2b phages. Transcriptomic analyses, together with proteomic and metabolomic studies, provided the basis for understanding the functions of the exo-xis region. Genes from this region promoted lytic development of the phage over lysogenization. Moreover, expression of the host genes coding for DnaK, DnaJ, GrpE, and GroELS chaperones was impaired in the cells infected with the Δexo-xis phage mutant, relative to the wild-type virus, corroborating the conclusion about lytic development promotion by the exo-xis region. Proteomic and metabolomic analyses indicated also modulation of gad and nrf operons, and levels of amino acids and acylcarnitines, respectively. In conclusion, the exo-xis region controls phage propagation and host metabolism by influencing expression of different phage and bacterial genes, directing the virus to the lytic rather than lysogenic developmental mode.

Preliminary Assessment of Sea Star (Echinodermata, Asteroidea) Diversity in the Coastal Magellanic Region (South Chile) and Their Geographical Distribution

Sea stars are a diverse and important component of the Southern Ocean benthos. However, scarce information is available regarding their diversity, distribution, evolution, and taxonomic uncertainties persist for multiple taxa. The Magellanic Region (south of Chile) remains under-sampled despite its pivotal location for species distribution and diversity, being located at the crossroad of three ocean basins. In this study, we assessed the biodiversity of coastal Magellanic sea stars and their affinities with other oceanic bioregions. An integrative approach combining morphological identification with DNA barcoding was implemented to highlight taxonomic discrepancies such as suspected synonymy and unrecognised diversity. Firstly, we identified a total of 15 species from the coastal Magellanic Region and reported the occurrence of Cycethra frigida Koehler, 1917 for the first time in this region. The distribution of these 15 species ranged from only in South America to circumpolar, bipolar, or possibly cosmopolitan. Secondly, we highlighted possible synonymy in two species pairs within the genera Anasterias and Odontaster. This preliminary biodiversity assessment forms an important baseline for monitoring and conservation purposes, especially in the face of distribution shifts as a response to climate change and the increased presence of invasive species. Developmental mode has previously been suggested to be important in shaping biogeographical patterns. However, developmental mode was insufficient to explain the observed patterns, and other factors (e.g., physiological constraints, competition, bathymetrical range, and the possibility of passively rafting on kelp) are suggested to be at least equally important. Finally, an increase in barcoding effort is needed to better capture phylogeographic patterns for each species, both by increasing the number of specimens investigated and by covering a broader geographical range.

Interspecific differences in developmental mode determine early cognitive abilities in teleost fish

Most studies on developmental variation in cognition have suggested that individuals are born with reduced or absent cognitive abilities, and thereafter, cognitive performance increases with age during early development. However, these studies have been mainly performed in altricial species, such as humans, in which offspring are extremely immature at birth. In this work, we tested the hypothesis that species with other developmental modes might show different patterns of cognitive development. To this end, we analysed inhibitory control performance in two teleost species with different developmental modes, the zebrafish Danio rerio and the guppy Poecilia reticulata, exploiting a simple paradigm based on spontaneous behaviour and therefore applicable to subjects of different ages. Zebrafish hatch as larvae 3 days after fertilisation, and have an immature nervous system, a situation that mirrors extreme altriciality. We found that at the early stages of development, zebrafish displayed no evidence of inhibitory control, which only begun to emerge after one month of life. Conversely, guppies, which are born after approximately one month of gestation as fully developed and independent individuals, solved the inhibitory control task since their first days of life, although performance increased with sexual maturation. Our study suggests that the typical progression described during early ontogeny in humans and other species might not be the only developmental trend for animals’ cognition and that a species’ developmental mode might determine variation in cognition across subjects of different age.

Digest: Ecological and life history drivers of avian skull evolution.

What are the factors that drive the patterns and evolutionary rates of morphological characteristics? To answer this question, Hunt et al. (2023) explore shape variation and rates of change in modern avian skulls using morphometric measurements and phylogenetic analyses. They find that habitat density and migration have the strongest influences on avian skull variation, and that denser habitats, longer migratory distances, and precocial developmental modes are all associated with faster rates of morphological evolution.

Fertilization Mode Covaries with Body Size.

AbstractThe evolution of internal fertilization has occurred repeatedly and independently across the tree of life. As it has evolved, internal fertilization has reshaped sexual selection and the covariances among sexual traits, such as testes size, and gamete traits. But it is unclear whether fertilization mode also shows evolutionary associations with traits other than primary sex traits. Theory predicts that fertilization mode and body size should covary, but formal tests with phylogenetic control are lacking. We used a phylogenetically controlled approach to test the covariance between fertilization mode and adult body size (while accounting for latitude, offspring size, and offspring developmental mode) among 1,232 species of marine invertebrates from three phyla. Within all phyla, external fertilizers are consistently larger than internal fertilizers: the consequences of fertilization mode extend to traits that are only indirectly related to reproduction. We suspect that other traits may also coevolve with fertilization mode in ways that remain unexplored.

Integrating Complex Life Cycles in Comparative Developmental Biology.

The goal of comparative developmental biology is identifying mechanistic differences in embryonic development between different taxa and how these evolutionary changes have led to morphological and organizational differences in adult body plans. Much of this work has focused on direct-developing species in which the adult forms straight from the embryo and embryonic modifications have direct effects on the adult. However, most animal lineages are defined by indirect development, in which the embryo gives rise to a larval body plan and the adult forms by transformation of the larva. Historically, much of our understanding of complex life cycles is viewed through the lenses of ecology and zoology. In this review, we discuss the importance of establishing developmental rather than morphological or ecological criteria for defining developmental mode and explicitly considering the evolutionary implications of incorporating complex life cycles into broad developmental comparisons of embryos across metazoans.

A quantitative assessment of ontogeny and molting in a Cambrian radiodont and the evolution of arthropod development

AbstractRadiodonta is a clade of stem euarthropods of central importance to our understanding of the evolution of this phylum. Radiodonts include some of the largest early Paleozoic animals; however, little is known about their ontogeny. We present an analysis of molting patterns and ontogeny in the radiodont Stanleycaris based on 265 exceptionally preserved specimens from the mid-Cambrian (Wuliuan) Burgess Shale. Ranging in size from 10 to 83 mm, they constitute the most extensive radiodont ontogenetic series known. Using a novel morphospace approach, we show that putative carcasses and exuviae can be quantitatively distinguished by the particular suites of structures preserved and their modes of preservation. We propose that Stanleycaris, and probably other radiodonts, molted via a suture near the anterior of the trunk. Similar anterior molting strategies, with a suture located at the head–trunk boundary, are shared with some Cambrian euarthropods and are potentially ancestral. Allometric analyses suggest that as Stanleycaris body size increased, the head sclerite and neck became relatively broader, while the trunk and flaps became slightly longer. The eyes developed precociously, indicating an important role of visual processing in juveniles. Finally, we find evidence for an initial anamorphic developmental phase, where segment number increased at least from 11 or 12 up to 17, followed by an epimorphic phase, in which growth continued without segment addition. This is consistent with the hypothesis that finite postembryonic segment addition (hemianamorphosis) is ancestral for arthropods and refines the timing of the origin of this important developmental mode.

Genomics of Marine Larval Evolution and Development

Synopsis Functional and ecological genomics in emerging marine models is shaping our views on new and fundamental biological questions. Representing a wealth of morphological, behavioral, and developmental adaptations, marine organisms are a key group for studying ancestral and shared body plan patterns, as well as some of the most intriguing novelties in animals. Here, we outline key questions, resources, and approaches to advance our understanding of marine larva biology and evolution that were covered in the recent symposium “Genomics of marine larval evolution and development.” Over 2 days, researchers from multiple continents, career stages, and research interests, covering a range of taxonomic diversity, contributed to three emerging topics: (1) Understanding the genetic basis of larval morphology and behaviors; (2) Finding the genomic basis for body plan development and novel morphologies; and (3) Constructing genomic tools in development for comparative insights. In this short piece, we highlight some of the most relevant findings on how different developmental modes and larval forms evolve.

Classification and Control Factors of Reservoir Types in Hybrid Sedimentary Rocks: A Case Study on Lucaogou Formation in the Jimusaer Sag, Junggar Basin.

Hybrid sedimentary rocks (HSR) represent a significant reservoir type in fine-grained sediments. However, the classification and understanding of HSR reservoirs, including their storage mechanisms and identification of optimal "sweet spots," have been limited due to the lack of clarity regarding the multiple sources of components and their mixing processes. This study focuses on the Lucaogou formation of Jimusaer Sag and aims to highlight the reservoir classification principles, controlling factors, and evolutionary patterns associated with the components of HSR, beginning with examining the microscopic pore structure. The analysis of the microscopic pore structure characteristics reveals the presence of five distinct reservoir types within the HSR. The quality of these reservoirs is governed by various factors, including the composition and support mode of particles, diagenesis, provenance, and sedimentary microfacies. In regions near a provenance with strong hydrodynamic conditions, the HSR predominantly exhibits type I and type II reservoirs, characterized by numerous coarse-grained components and a granular-support mode. As the distance from the provenance increases, transitioning into medium hydrodynamic conditions, the HSR shifts to an interbedded-support mode, primarily developing type III reservoirs. In areas far from the provenance with weak hydrodynamic conditions, HSR reservoir types primarily consist of type IV and type V. Additionally, diagenetic effects such as compaction and calcite cementation further deteriorate intergranular and dissolution pores, consequently diminishing reservoir quality. Notably, during the mixing deposition processes of sand and dolomite, the developmental mode of HSR shifts from type I to type II and type III. Likewise, in the mixing deposition of mud and sand, the HSR transitions from type II to type III and type IV. Similarly, the mixing deposition of dolomite and mud leads to a change in the developmental mode of HSR from type III to type IV and type V. Moreover, this study effectively predicts the occurrence of "sweet spots" using reservoir classification, which reveals their continuous distribution. These findings provide a geological foundation for evaluating "sweet spots" and testing the oil production in HSR reservoirs.

Atypical tracheid organization in proximal wood of late Palaeozoic Sigillaria approximata Fontaine et White (Lycopsida)

Abstract It is thought that arborescent lycopsid cambial expansion kept pace with radial wood growth via tangential broadening of the fusiform initials, resulting in progressive outward expansion in tracheid diameters and an absence of de novo tracheid files. This pattern appears in distal lycopsid axes but has not been investigated in proximal-most trunk vasculature where wood was thickest and primary xylem thinnest. Here, a ground-level trunk vasculature fossil of Sigillaria approximata in transverse section is described. This proximal vasculature diverges from the expectations of the current lycopsid wood production model in two main ways: first, inner and outer wood tracheids have approximately the same lumen diameters, such that there is no trend towards centrifugally increasing lumen diameters; and second, de novo cell files commonly appear within the wood. The outwards rate of cambial circumference expansion closely tracks the outwards rate of new wood tracheid file appearance, suggesting that the addition of new files is the primary way the cambium expanded proximally. Because this vasculature can only be studied in transverse section, the developmental mode producing new files cannot be determined. However, evidence from this specimen is sufficient to demonstrate that arborescent lycopsid wood production was more complex than previously understood.

Cephalopod ontogeny and life cycle patterns

Life cycle definitions provide the background for conceptualizing meaningful questions to address the mechanisms that generate different life cycle patterns. This review provides explicit definitions and explanations of the steps in a cephalopod life cycle, from fertilization to death. Each large step, or phase, is characterized by a particular developmental process and morphology. Each phase is composed of smaller developmentally distinct steps, or stages. The cephalopod life cycle is comprised of all or some of the following phases: Embryonic, Paralarval, Juvenile, Subadult, Adult and Senescent, and each life cycle is taxon-specific. All cephalopods have direct development and maintain a consistent body plan throughout ontogeny (i.e., no true larval phase and no metamorphosis). Most cephalopods have a life cycle marked by a long early life and a short adult life followed by senescence. Cephalopods have two developmental modes: they produce either small planktonic hatchlings as paralarvae, or large hatchlings as juveniles. All cephalopods go through a Hatchling stage soon after eclosion during which they rely on two modes of nutrition: endogenous (yolk) and exogenous (prey). Many cephalopods with planktonic paralarvae will become benthic early in their life cycle during their Settlement stage or remain pelagic during their Metapelagic stage. Juvenile growth is fast and ontogenetic changes (outside of gonadal maturation) generally cease at the end of the Juvenile phase. The Subadult phase begins when the definitive adult morphology (except for size and body proportions) is acquired (e.g., full complement of photophores). Sexual organs undergo most of their development during the Subadult phase. The Adult phase starts with spawning competency and concludes when gonads are spent. The Senescent phase begins with spent gonads and ends with death. Using this new terminology, we examine the patterns of cephalopod life cycles and find that there are four main patterns based on the presence of a Paralarval phase and the habitat occupied by each phase: Holopelagic (all phases are pelagic), Holobenthic (all phases are benthic), Merobenthic and Meropelagic (phases alternate between benthic and pelagic environments). In these two last patterns, the main difference is the presence of a Paralarval phase in Merobenthic species. The definitions and terminology proposed here provide a unifying framework for future ecological, evolutionary and life cycles research on cephalopods.