Developmental Novelties Research Articles

Interlocking of co-opted developmental gene networks in Drosophila and the evolution of pre-adaptive novelty

The re-use of genes in new organs forms the base of many evolutionary novelties. A well-characterised case is the recruitment of the posterior spiracle gene network to the Drosophila male genitalia. Here we find that this network has also been co-opted to the testis mesoderm where is required for sperm liberation, providing an example of sequentially repeated developmental co-options. Associated to this co-option event, an evolutionary expression novelty appeared, the activation of the posterior segment determinant Engrailed to the anterior A8 segment controlled by common testis and spiracle regulatory elements. Enhancer deletion shows that A8 anterior Engrailed activation is not required for spiracle development but only necessary in the testis. Our study presents an example of pre-adaptive developmental novelty: the activation of the Engrailed transcription factor in the anterior compartment of the A8 segment where, despite having no specific function, opens the possibility of this developmental factor acquiring one. We propose that recently co-opted networks become interlocked, so that any change to the network because of its function in one organ, will be mirrored by other organs even if it provides no selective advantage to them.

The Multi-Causal Basis of Developmental Potential Construction

In this article we analyse the issue of what accounts for developmental potential, i.e., the possible phenotypes a developing organism can manifest during ontogeny. We shall argue in favour of two theses. First, although the developing organism is the unit of development, the complete causal basis for its potential to develop does neither lie entirely in itself as a whole nor in any specific part of itself (such as its genome). Thus, the extra-organismal environment must be counted as one of the three necessary, partial and complementary causal bases for development potential. Secondly, we shall defend a constructivist view of the developmental process. If the genome, the developing organism and the extra-organismal environment are to be counted as proper elements of the causal basis for an organism’s developmental potential, the latter is not a given. Rather, it is the result of an interaction-based construction, a process sometimes generating genuine developmental novelty. We will thus argue for an interactionist multi-causal basis view of developmental potential construction. We contend that our view provides a biologically tenable and metaphysically coherent account of developmental dynamics.

Distinct Embryonic Origin and Injury Response of Resident Stem Cells in Craniofacial Muscles.

Craniofacial muscles emerge as a developmental novelty during the evolution from invertebrates to vertebrates, facilitating diversified modes of predation, feeding and communication. In contrast to the well-studied limb muscles, knowledge about craniofacial muscle stem cell biology has only recently starts to be gathered. Craniofacial muscles are distinct from their counterparts in other regions in terms of both their embryonic origin and their injury response. Compared with somite-derived limb muscles, pharyngeal arch-derived craniofacial muscles demonstrate delayed myofiber reconstitution and prolonged fibrosis during repair. The regeneration of muscle is orchestrated by a blended source of stem/progenitor cells, including myogenic muscle satellite cells (MuSCs), mesenchymal fibro-adipogenic progenitors (FAPs) and other interstitial progenitors. Limb muscles host MuSCs of the Pax3 lineage, and FAPs from the mesoderm, while craniofacial muscles have MuSCs of the Mesp1 lineage and FAPs from the ectoderm-derived neural crest. Both in vivo and in vitro data revealed distinct patterns of proliferation and differentiation in these craniofacial muscle stem/progenitor cells. Additionally, the proportion of cells of different embryonic origins changes throughout postnatal development in the craniofacial muscles, creating a more dynamic niche environment than in other muscles. In-depth comparative studies of the stem cell biology of craniofacial and limb muscles might inspire the development of novel therapeutics to improve the management of myopathic diseases. Based on the most up-to-date literature, we delineated the pivotal cell populations regulating craniofacial muscle repair and identified clues that might elucidate the distinct embryonic origin and injury response in craniofacial muscle cells.

Stages of embryonic development in the live-bearing fish, Gambusia holbrooki.

Divergent morphology and placentation of Poeciliids make them suitable model for investigating how evolutionary selection has altered and conserved the developmental mechanisms. However, there is limited description of their embryonic staging, despite representing a key evolutionary node that shares developmental strategy with placental vertebrates. Here, we describe the embryonic developmental stages of Gambusia holbrooki from zygote to parturition using freshly harvested embryos. We defined 40 embryonic stages using a numbered (stages 0-39; zygote to parturition, respectively) and named (grouped into seven periods, ie, zygote, cleavage, blastula, gastrula, segmentation, pharyngula, and parturition) staging system. Two sets of quantitative (ie, egg diameter, embryonic total length, otic vesicle closure index, heart rates, the number of caudal fin rays and elements) and qualitative (ie, three-dimensional analysis of images and key morphological criteria) data were acquired and used in combination to describe each stage. All 40 stages are separated by well-defined morphological traits, revealing developmental novelties that are influenced by narrow perivitelline space, placentation, internal gestation, and sex differentiation. The principal diagnostic features described are quick, reliable, and easy to apply. This system will benefit researchers investigating molecular ontogeny, particularly sexual differentiation mechanisms in G. holbrooki.

Developmental capacity and the early evolution of animals

Disentangling the factors underlying the appearance of macroscopic, often skeletonized, bilaterians during the Ediacaran–Cambrian diversification of animals requires carefully parsing the contributions of ecological opportunity, environmental potential and developmental capacity. The early evolution of animals involved the introduction of genomic, developmental, morphologic and behavioural novelties, identified as the individuation of new characters, which led to the construction of new ecological networks (innovation). Here I employ a recently introduced conceptual framework for novelty and individuation that distinguishes between potentiation, novelty, innovation and adaptive adjustments to the Ediacaran–Cambrian Radiation, and focus on the roles of potentiation and novelty in the expansion of developmental capacity. Comparative developmental studies combined with molecular clock estimates and data from the fossil record suggest that developmental capacity, the potential to generate a range of morphologies, may expand rapidly through developmental novelties without leading directly to morphological novelties, or to innovation. The expected patterns from this framework are markedly different from those in adaptive radiation scenarios. Thematic collection: This article is part of the Advances in the Cambrian Explosion collection available at: https://www.lyellcollection.org/cc/advances-cambrian-explosion

Molecular insights into deuterostome evolution from hemichordate developmental biology.

Hemichordates, along with echinoderms and chordates, belong to the lineage of bilaterians called the deuterostomes. Their phylogenetic position as an outgroup to chordates provides an opportunity to investigate the evolutionary origins of the chordate body plan and reconstruct ancestral deuterostome characters. The body plans of the hemichordates and chordates are organizationally divergent making anatomical comparisons very challenging. The developmental underpinnings of animal body plans are often more conservative than the body plans they regulate, and offer a novel data set for making comparisons between morphologically divergent body architectures. Here I review the hemichordate developmental data generated over the past 20 years that further test hypotheses of proposed morphological affinities between the two taxa, but also compare the conserved anteroposterior, dorsoventral axial patterning programs and germ layer specification programs. These data provide an opportunity to determine which developmental programs are ancestral deuterostome or bilaterian innovations, and which ones occurred in stem chordates or vertebrates representing developmental novelties of the chordate body plan.

Pilot study of bone tissue reparative regeneration of the young Trachemys scripta shell

The turtles skeleton plan is an evolutionary developmental novelty in connection with the formation of a tortoise-specific carapacial ridge, which induces the growth of ribs in the dorsal direction and the subsequent formation of several carapace plates. The bone plates of these reptiles have different histogenesis: the neural and costal plates develop according to the mechanism of indirect osteogenesis by perichondral ossification of the axial skeleton elements (vertebrae and ribs), the peripheral and plastron bones develop according to the mechanism of direct osteogenesis in the dermis of the skin - these are the so-called «skin bones», which are derived from individual osteogenic cells outside the axial skeleton and develop by intramembranous osteogenesis. An experiment was carried out to study the post-traumatic osteogenesis of the carapace bone plates on a freshwater turtles Trachemys scripta at the age of 3 months. Two defects of carapace costal plates with a diameter of 4 mm and a depth to the fascia were performed with further routine histological examination. It was found that already on the 90th day the turtles carapace bones were restored with the formation of a full-fledged bone regenerate by the mechanism of indirect osteogenesis.

MtSUPERMAN plays a key role in compound inflorescence and flower development in Medicago truncatula.

Legumes have unique features, such as compound inflorescences and a complex floral ontogeny. Thus, the study of regulatory genes in these species during inflorescence and floral development is essential to understand their role in the evolutionary origin of developmental novelties. The SUPERMAN (SUP) gene encodes a C2H2 zinc-finger transcriptional repressor that regulates the floral organ number in the third and fourth floral whorls of Arabidopsis thaliana. In this work, we present the functional characterization of the Medicago truncatula SUPERMAN (MtSUP) gene based on gene expression analysis, complementation and overexpression assays, and reverse genetic approaches. Our findings provide evidence that MtSUP is the orthologous gene of SUP in M. truncatula. We have unveiled novel functions for a SUP-like gene in eudicots. MtSUP controls not only the number of floral organs in the inner two whorls, but also in the second whorl of the flower. Furthermore, MtSUP regulates the activity of the secondary inflorescence meristem, thus controlling the number of flowers produced. Our work provides insight into the regulatory network behind the compound inflorescence and flower development in this angiosperm family.

Mobility connects: transposable elements wire new transcriptional networks by transferring transcription factor binding motifs.

Transposable elements (TEs) constitute major fractions of plant genomes. Their potential to be mobile provides them with the capacity to cause major genome rearrangements. Those effects are potentially deleterious and enforced the evolution of epigenetic suppressive mechanisms controlling TE activity. However, beyond their deleterious effects, TE insertions can be neutral or even advantageous for the host, leading to long-term retention of TEs in the host genome. Indeed, TEs are increasingly recognized as major drivers of evolutionary novelties by regulating the expression of nearby genes. TEs frequently contain binding motifs for transcription factors and capture binding motifs during transposition, which they spread through the genome by transposition. Thus, TEs drive the evolution and diversification of gene regulatory networks by recruiting lineage-specific targets under the regulatory control of specific transcription factors. This process can explain the rapid and repeated evolution of developmental novelties, such as C4 photosynthesis and a wide spectrum of stress responses in plants. It also underpins the convergent evolution of embryo nourishing tissues, the placenta in mammals and the endosperm in flowering plants. Furthermore, the gene regulatory network underlying flower development has also been largely reshaped by TE-mediated recruitment of regulatory elements; some of them being preserved across long evolutionary timescales. In this review, we highlight the potential role of TEs as evolutionary toolkits in plants by showcasing examples of TE-mediated evolutionary novelties.

Ontogeny of the anuran urostyle and the developmental context of evolutionary novelty

Developmental novelties often underlie the evolutionary origins of key metazoan features. The anuran urostyle, which evolved nearly 200 MYA, is one such structure. It forms as the tail regresses during metamorphosis, when locomotion changes from an axial-driven mode in larvae to a limb-driven one in adult frogs. The urostyle comprises of a coccyx and a hypochord. The coccyx forms by fusion of caudal vertebrae and has evolved repeatedly across vertebrates. However, the contribution of an ossifying hypochord to the coccyx in anurans is unique among vertebrates and remains a developmental enigma. Here, we focus on the developmental changes that lead to the anuran urostyle, with an emphasis on understanding the ossifying hypochord. We find that the coccyx and hypochord have two different developmental histories: First, the development of the coccyx initiates before metamorphic climax whereas the ossifying hypochord undergoes rapid ossification and hypertrophy; second, thyroid hormone directly affects hypochord formation and appears to have a secondary effect on the coccygeal portion of the urostyle. The embryonic hypochord is known to play a significant role in the positioning of the dorsal aorta (DA), but the reason for hypochordal ossification remains obscure. Our results suggest that the ossifying hypochord plays a role in remodeling the DA in the newly forming adult body by partially occluding the DA in the tail. We propose that the ossifying hypochord-induced loss of the tail during metamorphosis has enabled the evolution of the unique anuran bauplan.

CRISPR/Cas9 mutagenesis in Volvox carteri

Volvox carteri and other volvocine green algae comprise an excellent model for investigating developmental complexity and its origins. Here we describe a method for targeted mutagenesis in V.carteri using CRISPR/Cas9 components expressed from transgenes. We used V.carteri nitrate reductase gene (nitA) regulatory sequences to conditionally express Streptococcus pyogenes Cas9, and V.carteri U6 RNA gene regulatory sequences to constitutively express single-guide RNA (sgRNA) transcripts. Volvox carteri was bombarded with both Cas9 vector and one of several sgRNA vectors programmed to target different test genes (glsA, regA and invA), and transformants were selected for expression of a hygromycin-resistance marker present on the sgRNA vector. Hygromycin-resistant transformants grown with nitrate as sole nitrogen source (inducing for nitA) were tested for Cas9 and sgRNA expression, and for the ability to generate progeny with expected mutant phenotypes. Some transformants of a somatic regenerator (Reg) mutant strain receiving sgRNA plasmid with glsA protospacer sequence yielded progeny (at a rate of ~0.01%) with a gonidialess (Gls) phenotype similar to that observed for previously described glsA mutants, and sequencing of the glsA gene in independent mutants revealed short deletions within the targeted region of glsA, indicative of Cas9-directed non-homologous end joining. Similarly, bombardment of a morphologically wild-type strain with the Cas9 plasmid and sgRNA plasmids targeting regA or invA yielded regA and invA mutant transformants/progeny, respectively (at rates of 0.1-100%). The capacity to make precisely directed frameshift mutations should greatly accelerate the molecular genetic analysis of development in V.carteri, and of developmental novelty in the volvocine algae.

Developmental push or environmental pull? The causes of macroevolutionary dynamics.

Have the large-scale evolutionary patterns illustrated by the fossil record been driven by fluctuations in environmental opportunity, by biotic factors, or by changes in the types of phenotypic variants available for evolutionary change? Since the Modern Synthesis most evolutionary biologists have maintained that microevolutionary processes carrying on over sufficient time will generate macroevolutionary patterns, with no need for other pattern-generating mechanisms such as punctuated equilibrium or species selection. This view was challenged by paleontologists in the 1970s with proposals that the differential sorting and selection of species and clades, and the effects of biotic crises such as mass extinctions, were important extensions to traditional evolutionary theory. More recently those interested in macroevolution have debated the relative importance of abiotic and biotic factors in driving macroevolutionary patterns and have introduced comparative phylogenetic methods to analyze the rates of change in taxonomic diversity. Applying Peter Godfrey-Smith's distinction between distributional explanations and explanations focusing on the origin of variation, most macroevolutionary studies have provided distributional explanations of macroevolutionary patterns. Comparative studies of developmental evolution, however, have implicated the origin of variants as a driving macroevolution force. In particular, the repatterning of gene regulatory networks provides new insights into the origins of developmental novelties. This raises the question of whether macroevolution has been pulled by the generation of environmental opportunity, or pushed by the introduction of new morphologies. The contrast between distributional and origination scenarios has implications for understanding evolutionary novelty and innovation and how macroevolutionary process may have evolved over time.

Developmental and Evolutionary Perspectives on the Origin and Diversification of Arthropod Appendages.

Jointed, segmented appendages are a key innovation of arthropods. The subsequent diversification of these appendages, both along the body axis and across taxa, has contributed to the evolutionary success of arthropods. Both developmental and fossil data are informative for understanding how these transitions occurred. Comparative analyses help to pinpoint the developmental novelties that distinguish arthropod appendages from the lobopodous appendages of other panarthropods, and that distinguish different appendage types. The fossil record of stem group arthropods is diverse and preserves intermediate steps in these evolutionary transitions, including some that cannot be directly inferred based on extant taxa. These lead to hypotheses that can be tested with comparative developmental data, as well as to reinterpretations of developmental results. One developmental novelty of arthropods is the reiterated deployment of the joint formation network, which divides the appendages into segments. The fossil record raises questions about how this joint formation network was first deployed, given the contrasting morphologies of appendages in stem group versus extant arthropods. The fossil record supports a character tree for appendage diversification showing progressive individuation of appendages in an anterior-to-posterior sequence. However, to date, developmental evidence provides at best limited support for this character tree. Recent interpretations of the fossil record suggest that the labrum of extant arthropods is a greatly reduced protocerebral appendage pair; this hypothesis is consistent with the extensive shared developmental patterning of the labrum and jointed appendages. Reciprocal illumination from fossils and developmental patterning in a phylogenetic context both makes sense of some results and helps motivates questions for future research.

Evolutionary developmental biology (evo-devo) of cestodes

Cestodes (tapeworms) have complex adaptations to their obligatory parasitic life-style. Among these adaptations, they show many evolutionary innovations in their development, including complex life-cycles with multiple hosts and life-stages, several independent origins of asexual reproduction, and the evolution of segmentation as a mean to generate massive reproductive output. Therefore, cestodes offer many opportunities for the investigation of the evolutionary origins of developmental novelties (evo-devo). However, cestodes have not been exploited as major models for evo-devo research due to the considerable technical difficulties involved in their study. In this review, a panoramic view is given of classical aspects, methods and hypothesis of cestode development, together with recent advances in phylogenetics, genomics, culture methods, and comparative analysis of cestode gene expression. Together with the availability of powerful models for related free-living flatworms, these developments should encourage the incorporation of these fascinating parasites into the first-line of evo-devo research.

Making teeth to order: conserved genes reveal an ancient molecular pattern in paddlefish (Actinopterygii).

Ray-finned fishes (Actinopterygii) are the dominant vertebrate group today (+30 000 species, predominantly teleosts), with great morphological diversity, including their dentitions. How dental morphological variation evolved is best addressed by considering a range of taxa across actinopterygian phylogeny; here we examine the dentition of Polyodon spathula (American paddlefish), assigned to the basal group Acipenseriformes. Although teeth are present and functional in young individuals of Polyodon, they are completely absent in adults. Our current understanding of developmental genes operating in the dentition is primarily restricted to teleosts; we show that shh and bmp4, as highly conserved epithelial and mesenchymal genes for gnathostome tooth development, are similarly expressed at Polyodon tooth loci, thus extending this conserved developmental pattern within the Actinopterygii. These genes map spatio-temporal tooth initiation in Polyodon larvae and provide new data in both oral and pharyngeal tooth sites. Variation in cellular intensity of shh maps timing of tooth morphogenesis, revealing a second odontogenic wave as alternate sites within tooth rows, a dental pattern also present in more derived actinopterygians. Developmental timing for each tooth field in Polyodon follows a gradient, from rostral to caudal and ventral to dorsal, repeated during subsequent loss of teeth. The transitory Polyodon dentition is modified by cessation of tooth addition and loss. As such, Polyodon represents a basal actinopterygian model for the evolution of developmental novelty: initial conservation, followed by tooth loss, accommodating the adult trophic modification to filter-feeding.

A new embryonic pattern in parasitic wasps: divergence in early development may not be associated with lifestyle

Comparative embryogenesis of Encarsia formosa and Encarsia pergandiella (Hymenoptera Aphelinidae), two endoparasitoids of whiteflies (Hemiptera Aleyrodidae), revealed two strongly diverging developmental patterns. Indeed, the centrolecithal anhydropic egg of E. formosa developed through a superficial cleavage, as it occurs in Nasonia vitripennis, Apis mellifera, and Drosophila melanogaster. In contrast, the alecithal hydropic egg of E. pergandiella developed through holoblastic cleavage within a specialized extra-embryonic membrane (EEM). Since this developmental pattern evolved independently in several lineages of hymenopteran endoparasitoids, departures from the superficial cleavage mode have been argued to be strongly canalized in response to a shift from ecto- to endoparasitic lifestyle. Coexistence of both developmental patterns in two congeneric species suggests that alterations of early embryonic development may not be correlated with lifestyle. In addition, embryogenesis of E. pergandiella exhibited the following developmental novelties compared to other species possessing a hydropic egg: (i) polar body derivatives early acquired a cytoskeletal boundary prior to any other cellularization event; (ii) cellularization was asynchronous, starting with an early differentiation of a single apical blastomere at the end of the third cleavage; (iii) appearance of cytoskeletal boundaries of embryo blastomeres occurred between the third and fourth cleavages; (iv) the EEM originated through asynchronous participation of three separate lineages of cleavage nuclei, one of which associated with the polar body derivatives in a syncytium. Our results confirm a scenario of high plasticity in the early developmental strategies of hymenopteran endoparasitoids.

Developmental residue and developmental novelty - different modes of adipose-fin formation during ontogeny

The adipose fin is a structure with questionable function, for a long time even regarded as a rudimentary element without any function. A comparative study of adipose fin development revealed two different modes: a salmoniform-type where the adipose fin develops directly from the larval fin fold simultaneously with the other median fins. And a characiform-type where the adipose fin develops anew after the reduction of the larval fin fold. Compared to the first mode the latter starts late in ontogeny, when all other median fins already have their shape, fin rays and colouration. The characiform-type of adipose fin development contradicts the view that this structure is just a larval fin fold remainder. It also disputes the hypothesis of the adipose fin as a structure without any function. Such a hypothesis cannot explain a systematically widely distributed structure which even exhibits different modes of ontogenetic development in different taxa. (© 2013 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Development of an embryonic skeletogenic mesenchyme lineage in a sea cucumber reveals the trajectory of change for the evolution of novel structures in echinoderms.

BackgroundThe mechanisms by which the conserved genetic “toolkit” for development generates phenotypic disparity across metazoans is poorly understood. Echinoderm larvae provide a great resource for understanding how developmental novelty arises. The sea urchin pluteus larva is dramatically different from basal echinoderm larval types, which include the auricularia-type larva of its sister taxon, the sea cucumbers, and the sea star bipinnaria larva. In particular, the pluteus has a mesodermally-derived larval skeleton that is not present in sea star larvae or any outgroup taxa. To understand the evolutionary origin of this structure, we examined the molecular development of mesoderm in the sea cucumber, Parastichopus parvimensis.ResultsBy comparing gene expression in sea urchins, sea cucumbers and sea stars, we partially reconstructed the mesodermal regulatory state of the echinoderm ancestor. Surprisingly, we also identified expression of the transcription factor alx1 in a cryptic skeletogenic mesenchyme lineage in P. parvimensis. Orthologs of alx1 are expressed exclusively within the sea urchin skeletogenic mesenchyme, but are not expressed in the mesenchyme of the sea star, which suggests that alx1+ mesenchyme is a synapomorphy of at least sea urchins and sea cucumbers. Perturbation of Alx1 demonstrates that this protein is necessary for the formation of the sea cucumber spicule. Overexpression of the sea star alx1 ortholog in sea urchins is sufficient to induce additional skeleton, indicating that the Alx1 protein has not evolved a new function during the evolution of the larval skeleton.ConclusionsThe proposed echinoderm ancestral mesoderm state is highly conserved between the morphologically similar, but evolutionarily distant, auricularia and bipinnaria larvae. However, the auricularia, but not bipinnaria, also develops a simple skelotogenic cell lineage. Our data indicate that the first step in acquiring these novel cell fates was to re-specify the ancestral mesoderm into molecularly distinct territories. These new territories likely consisted of only a few cells with few regulatory differences from the ancestral state, thereby leaving the remaining mesoderm to retain its original function. The new territories were then free to take on a new fate. Partitioning of existing gene networks was a necessary pre-requisite to establish novelty in this system.

Bipartite life cycle of coral reef fishes promotes increasing shape disparity of the head skeleton during ontogeny: an example from damselfishes (Pomacentridae)

BackgroundQuantitative studies of the variation of disparity during ontogeny exhibited by the radiation of coral reef fishes are lacking. Such studies dealing with the variation of disparity, i.e. the diversity of organic form, over ontogeny could be a first step in detecting evolutionary mechanisms in these fishes. The damselfishes (Pomacentridae) have a bipartite life-cycle, as do the majority of demersal coral reef fishes. During their pelagic dispersion phase, all larvae feed on planktonic prey. On the other hand, juveniles and adults associated with the coral reef environment show a higher diversity of diets. Using geometric morphometrics, we study the ontogenetic dynamic of shape disparity of different head skeletal units (neurocranium, suspensorium and opercle, mandible and premaxilla) in this fish family. We expected that larvae of different species might be relatively similar in shapes. Alternatively, specialization may become notable even in the juvenile and adult phase.ResultsThe disparity levels increase significantly throughout ontogeny for each skeletal unit. At settlement, all larval shapes are already species-specific. Damselfishes show high levels of ontogenetic allometry during their post-settlement growth. The divergence of allometric patterns largely explains the changes in patterns and levels of shape disparity over ontogeny. The rate of shape change and the length of ontogenetic trajectories seem to be less variable among species. We also show that the high levels of shape disparity at the adult stage are correlated to a higher level of ecological and functional diversity in this stage.ConclusionDiversification throughout ontogeny of damselfishes results from the interaction among several developmental novelties enhancing disparity. The bipartite life-cycle of damselfishes exemplifies a case where the variation of environmental factors, i.e. the transition from the more homogeneous oceanic environment to the coral reef offering a wide range of feeding habits, promotes increasing shape disparity of the head skeleton over the ontogeny of fishes.

Evolution of morphogenesis in 360‐million‐year‐old conodont chordates calibrated in days

Highly rhythmic increments of crown tissue are identifiable in conodont oral apparatus elements from the Late Devonian of the Holy Cross Mountains, Poland; individual laminae being of thickness comparable with daily increments of vertebrate tooth enamel and fish otoliths. Abundant occurrence of such specimens enables bed-by-bed (stratophenetic) studies of the process of evolution at the population level and quantitative presentation of the evolution of ontogeny in the sampled geological section covering several million years. The morphologic transformation is expressed as expansion of a juvenile asymmetry to later stages of the ontogeny and in decrease of the mature element width, which was due to a change of the mineral tissue secretion rate. It was not just a simple extension of a juvenile character into the later stage of the ontogeny (heterochrony) but rather a true developmental novelty. The evolution was gradual and very slow. The proposed quantitative approach to growth increments in the mineral skeleton of ancient chordates introduces real-time units to evolutionary developmental studies connected with direct paleontological evidence on the course of evolution.