Guillaume Dera

Paul Sabatier University - Toulouse III | UPS Toulouse · Laboratoire Géosciences Environnement Toulouse - UM 97 (UMR 5563 / UMRD 234) - GET

The tropical, warm, and equable climate of the Jurassic world is regularly challenged by geoscientists, especially since oxygen isotopes ( 18O) of fossil hardparts have been used to reconstruct the paleotemperature history of seawater. By applying the innovative “SiZer” (significant zero crossings of the derivatives) statistical approach to a newly compiled 18O database for the Jurassic, we demonstrate the occurrence of major and multiscale 18O changes mainly related to climate disturbances. For the first time, two long-term anomalies in 18O are identified during the Toarcian and the Late Jurassic, in conjunction with intensive volcanism in large igneous provinces. These results support a strong influence of repeated volcanic pulses on the modulation of pCO2, temperatures, and polar ice cap volumes over protracted periods. At shorter time scales, 13 relatively rapid (0.5–1 m.y.) and significant warming and cooling events are identified, the causes of which include transient fluctuations in greenhouse gas concentrations related to still-debated mechanisms.

The breakup of Pangea and onset of growth of the Pacific plate led to several paleoenvironmental feedbacks, which radically affected paleoclimate and ocean chemistry during the Jurassic. Overall, this periodwas characterized by intense volcanic degassing from large igneous provinces and circum-Panthalassan arcs, new oceanic circulation patterns, and changes in heat and humidity transports affecting continental weathering. Few studies, however, have attempted to unravel the global interactions linking these processes over the long-term. In this paper, we address this question by documenting the global changes in continental drainage and surface oceanic circulation for thewhole Jurassic period. For this purpose, we present 53 new neodymium isotope values (εNd(t)) measured on well-dated fossil fish teeth, ichthyosaur bones, phosphatized nodules, phosphatized ooids, and clastic sediments from Europe, western Russia, and North America. Combined with an extensive compilation of published εNd(t) data, our results showthat the continental sources of Nd were very heterogeneous across theworld. Volcanic inputs froma Jurassic equivalent of the modern Pacific Ring of Fire contributed to radiogenic εNd(t) values (−4 ε-units) in the Panthalassa Ocean. For the Tethyan Ocean, the average surface seawater signal was less radiogenic in the equatorial region (−6.3), and gradually lower toward the epicontinental peri-Tethyan (−7.4), western Russian (−7.4) and Euro-Boreal seas (−8.6). Different Nd sources contributed to this disparity, with radiogenic Nd influxes from westward Panthalassan currents or juvenile volcanic arcs in open oceanic domains, and substantial unradiogenic inputs fromold Laurasian and Gondwanan shields for the NW Tethyan platforms. Overall, the εNd(t) values of Euro-Boreal, peri-Tethyan, and western Russian waters varied quite similarly through time, in response to regional changes in oceanic circulation, paleoclimate, continental drainage, and volcanism. Three positive shifts in εNd(t) values occurred successively in these epicontinental seas during the Pliensbachian, in the Aalenian–Bathonian interval, and in the mid-Oxfordian. The first and third events are interpreted as regional incursions of warm surface radiogenic currents from low latitudes. The Aalenian–Bathonian shift seems linked to volcanic outbursts in the NW Tethys and/or circulation of deep currents resulting from extensional events in the Hispanic Corridor and reduced influences of boreal currents crossing the Viking Corridor. In contrast, the εNd(t) signals decreased and remained very low(b−8) during the global warming events of the Toarcian and Late Oxfordian–Early Tithonian intervals. In these greenhouse contexts, a latitudinal expansion of humid belts could have extended the drainage pathways toward boreal Nd sources of Precambrian age and increased the supply of very unradiogenic crustal-derived inputs to seawater. Finally, a brief negative εNd(t) excursion recorded in parallel with regional drops in seawater temperature suggests that southward circulation of cold unradiogenic Arctic waters occurred in theNW Tethys in the Callovian–Early Oxfordian. All these results show that changes in surface oceanic circulation resulting from the Pangean breakup could have regionally impacted the evolution of seawater temperatures in the NW Tethys.

Eccentricity, obliquity, and precession are cyclic parameters of the Earth’s orbit whose climatic implications have been widely demonstrated on recent and short time intervals. Amplitude modulations of these parameters on million-year time scales induce ‟grand orbital cycles,” but the behavior and the paleoenvironmental consequences of these cycles remain debated for the Mesozoic owing to the chaotic diffusion of the solar system in the past. Here, we test for these cycles from the Jurassic to the Early Cretaceous by analyzing new stable isotope datasets reflecting fluctuations in the carbon cycle and seawater temperatures. Our results document a prominent cyclicity of ∼9 My in the carbon cycle paced by changes in the seasonal dynamics of hydrological processes and long-term sea level fluctuations. These paleoenvironmental changes are linked to a great eccentricity cycle consistent with astronomical solutions. The orbital forcing signal was mainly amplified by cumulative sequestration of organic matter in the boreal wetlands under greenhouse conditions. Finally, we show that the ∼9-My cycle faded during the Pliensbachian, which could either reflect major paleoenvironmental disturbances or a chaotic transition affecting this cycle.

The reconstruction of past variations in seawater strontium isotope ratios (87Sr/86Sr) from marine carbonate skeletons (MCSs) is based on the assumption that the Sr isotope composition of oceans is homogeneous worldwide. However, fossil materials are mainly from shelf domains whose modern Sr budget is known to be very sensitive to continental Sr fluxes. Their reliability as chemostratigraphic tools or weathering proxies may thus be questioned, especially because no formal appraisal of biological and environmental biases has been done on a global scale. Here, we provide new modern MCS 87Sr/86Sr values from various marine environments that we analyze statistically in regards to extensive compilations of MCS and water data. We show that most MCS 87Sr/86Sr values fall within the range of those of modern oceanic waters (0.709172 ± 23). Others, concerning ~30% of studied localities, may display slight to considerable offsets compared to the 87Sr/86Sr signature of modern seawater (±250 × 10–6). These patterns mainly concern bivalves and gastropods, or more generally epibenthic and eurytopic organisms, from coastal domains with water mass restriction, low salinity, or strong continental supplies (fjords, lagoons, endorheic basins). Beyond classical fluvial freshwater influences, submarine groundwater discharges may lead to a less radiogenic 87Sr/86Sr signature of MCSs even in lagoons with euhaline conditions (e.g., Oualidia lagoon, Morocco). All biases considered, this calls for caution in relative dating, as we calculate that half the values of the Phanerozoic 87Sr/86Sr curve cannot provide time calibration accuracy better than ±3 m.y., which is necessary to discriminate the shorter stages.

Major carbon cycle disturbances reflected by carbon isotope excursions (CIE) have been reported in the Jurassic successions of Euro-Boreal and Mediterranean basins. However, sedimentary and geochemical data from other domains are still required to unravel the global or regional nature of paleoenvironmental disturbances and identify the potential triggers. Here, we present biostratigraphically well-constrained (middle Toarcian to lower Kimmeridgian) bulk-rock δ13C and δ18O data from central Saudi Arabia. Five major negative δ18O excursions are identified in back-reef facies below sequence boundaries and disconformities (Aalenian-Bajocian boundary, upper Bajocian, middle Bathonian, upper Callovian and lower Kimmeridgian), suggesting that these carbonates underwent intense meteoric diagenesis during partial or major subaerial exposures of the Arabian Platform. The δ13C record is less altered and shows correlations with the regional relative sea-level changes influencing the productivity levels and inflows of fluvial (12C-enriched) or oceanic (12C-depleted) waters onto the platform. Overall, most of the δ13C variations recorded at the scale of the Arabian Platform within the lower and middle Jurassic are in agreement with NW Tethyan data. Among these variations, the increase in δ13C values recorded in the middle Toarcian, lower Bajocian, and lower Bathonian zigzag Zone indicate a considerable increase of productivity and 12Corg burial concomitant with global sea-level rises. Conversely, the occurrence of major but diachronous negative CIEs at the Aalenian-Bajocian boundary and around the lower-upper Bajocian transition are interpreted as accelerations of terrestrial 12Corg transfer to the ocean and atmosphere linked to regional but widespread emergences of landmasses. Whether global (eustatic) or regional (tectonic), all these sea-level changes and carbon transfers could be distal consequences of successive episodes of the Pangean breakup.

Investigating the provenance of sediments deposited in the Central Atlantic Ocean is of prime importance to decipher the mechanisms leading to the West African Craton drainage reorganization through time. To determine these changes in provenance during the opening of the Atlantic Ocean, we study the major and trace element concentrations and strontium and neodymium isotopic compositions (expressed as εNd) of Cretaceous sedimentary rocks from eight Deep Sea Drilling Project (DSDP) Sites and one exploration well. The absence of significant differences in major and trace element concentrations indicates that the crustal-derived nature of the sediments remained constant over time. The εNd(0) values from the DSDP Sites show a three-step decrease during the Late Cretaceous. The Albian−Middle Cenomanian εNd(0) values are heterogeneous (-5.5 to -14.9) reflecting the existence of at least three subdrainage basins with distinct sedimentary sources (Hercynian/Paleozoic, Precambrian and mixed Precambrian/Paleozoic). During the Late Cenomanian−Turonian interval, εNd(0) values become homogeneous in the deep-water basin (-10.3 to -12.4), showing a negative shift by 2 epsilon units interpreted as an increasing contribution of Precambrian inputs. This generalized εNd(0) decrease continued in the Campanian−Maastrichtian (-15 epsilon units), indicating that Precambrian sources became predominant. These provenance changes seem related to the opening of the South and Equatorial Atlantic Oceans and tectonic uplift triggered by Africa−Europe convergence. Finally, the difference between εNd(0)values of Cretaceous sediments from the Senegal continental shelf (CM1 well) and from the deep-water basins (DSDP Sites 367 and 368) suggests that ocean currents prevented detrital material from the Mauritanides reaching deep-water areas.

The onset of the late Palaeozoic ice age about 340 million years ago has been attributed to a decrease in atmospheric CO2 concentrations associated with expansion of land plants, as plants both enhance silicate rock weathering—which consumes CO2—and increase the storage of organic carbon on land. However, plant expansion and carbon uptake substantially predate glaciation. Here we use climate and carbon cycle simulations to investigate the potential effects of the uplift of the equatorial Hercynian mountains and the assembly of Pangaea on the late Palaeozoic carbon cycle. In our simulations, mountain uplift during the Late Carboniferous caused an increase in physical weathering that removed the thick soil cover that had inhibited silicate weathering. The resulting increase in chemical weathering was sufficient to cause atmospheric CO2 concentrations to fall below the levels required to initiate glaciation. During the Permian, the lowering of the mountains led to a re-establishment of thick soils, whilst the assembly of Pangaea promoted arid conditions in continental interiors that were unfavourable for silicate weathering. These changes allowed CO2 concentrations to rise to levels sufficient to terminate the glacial event. Based on our simulations, we suggest that tectonically influenced carbon cycle changes during the late Palaeozoic were sufficient to initiate and terminate the late Palaeozoic ice age.

The period spanning from 825 to 540 Ma is characterized by major changes in the surficial Earth system. This extraordinary interval starts with the breakup of the Rodinia supercontinent and eruption of a series of large igneous provinces and ends with the assembly of Gondwana, giving rise to the Pan-African orogenies. This paleogeographic reorganization is accompanied by a global climatic cooling, including the paroxysmal Cryogenian “snowball” glacial events. The ⁸⁷Sr/⁸⁶Sr of seawater displays a major long-term rise over this interval that is punctuated by episodic, smaller declines and inflections. We use a coupled deep time climate-carbon numerical model to explore the complex role of tectonics and climate on this distinct evolution in seawater ⁸⁷Sr/⁸⁶Sr. We show that the modulation of the weathering of the erupted large igneous provinces by continental drift explains the changes in seawater ⁸⁷Sr/⁸⁶Sr from 800 to 635 Ma. The subsequent sharp rise in seawater ⁸⁷Sr/⁸⁶Sr from 635 to 580 Ma is the result of erosion of radiogenic crust exposed in the Pan-African orogens. Coeval evolution of atmospheric CO2 displays a decrease from about 80 times the pre-industrial level around 800 Ma to 5 times just before the beginning of the Phanerozoic.

Evolutionary radiations have been extensively studied especially in the fossil record and in the context of postcrisis recoveries. The concept of adaptive radiation that emerges from this very broad topic explicitly involves the effect of adaptation driven by ecological opportunity and is considered to be of the foremost importance. It is essential to be able to detect adaptive radiation because it points up factors that predispose a clade to radiate. Adaptive radiation has received much attention in recent decades based mostly on studies dealing with recent clades, but data from the fossil record are still scarce. This study begins to fill this gap with the example of Lower Jurassic ammonoids (through c. 8 Myr of history). A survey of several clades, using both taxonomic and disparity-based approaches, shows that they diversified successively through time, but not systematically, in terms of species numbers and morphological variety. Some clades seem to have exhibited adaptive radiation and to have become rapidly extinct. One clade (which engendered nearly all post–Lower Jurassic ammonoids) has a fossil record that begins with low diversity and disparity but is superseded by a sustained radiation pattern. The results are discussed in the light of the Modern Synthesis and its continuation into an Extended Evolutionary Synthesis.

The Fe–Mn oxide fraction leached from deep-sea sediments has been increasingly used to reconstruct the Nd isotope composition of deep water masses, that can be used to track changes in oceanic circulation with a high temporal resolution. Application of this archive to reconstruct the Nd isotope composition of bottom seawater in shallow shelf environments remained however to be tested. Yet as the Nd isotope composition of seawater on continental margins is particularly sensitive to changes in erosional inputs, establishment of neritic seawater Nd isotope evolution around areas of deep water formation would be useful to discriminate the influence of changes in oceanic circulation and in isotopic composition of erosional inputs on the Nd isotope record of deep waters. The purpose of this study is to test the potential of Fe–Mn coatings leached from foraminifera tests to reconstruct the Nd isotope composition of seawater in shelf environments for deep-time intervals. Albian to Turonian samples from two different outcrops have been recovered, from the Paris Basin (Wissant section, northern France) and from the Western Interior Seaway (Hot Spring, South Dakota, USA), that were deposited in epicontinental seas. Rare Earth Element (REE) spectra enriched in middle REEs in the foraminifera leach at Wissant highlight the presence of Fe–Mn oxides. The similarity of the Nd isotopic signal of the Fe–Mn oxide fraction leached from foraminifera tests with that of fish teeth suggests that Fe–Mn oxides coating foraminifera can be good archives of shelf bottom seawater Nd isotopic composition. Inferred bottom shelf water Nd isotope compositions at Wissant range from −8.5 to −9.7 ε-units, about 1.5–2 ε-units higher than that of the contemporaneous local detrital fraction. At Hot Spring, linear REE spectra characterizing foraminifera leach may point to an absence of authigenic marine Fe–Mn oxide formation in this area during the Late Cenomanian–Early Turonian, consistent with dysoxic to anoxic conditions at Hot Spring, contemporaneous to an Oceanic Anoxic Event. The similarity of the Nd isotopic signal of the carbonate matrix of foraminifera with that of fish teeth suggests that it records the Nd isotope composition of bottom shelf seawater as well. Inferred bottom shelf water Nd isotope compositions at Hot Spring are quite radiogenic, between −7 and −6 ε-units, about 2.5–4 ε-units higher than that of the contemporaneous local detrital fraction. In contrast, in both sections Fe–Mn oxides leached directly from the decarbonated sediment tend to yield a less radiogenic Nd isotopic composition, typically between 0.2 and 0.8 ε-units lower, that is intermediate between that of fish teeth and of the detrital fraction. This suggests the contribution of pre-formed continental Fe–Mn oxides to the Nd isotopic signal, along with authigenic marine oxides, or a detrital contamination during leaching

The paleoecological disturbances recorded during the Early Toarcian warming event (183 Myr ago), including marine anoxia, sea level rise, seawater acidification, carbonate production crisis, and species extinctions, are often regarded as past examples of Earth’s possible responses to the rapid emergence of super greenhouse conditions. However, physical mechanisms explaining both the global and local expressions of paleoenvironmental events are still highly debated. Here we analyze the paleoclimatic and paleoceanographic consequences of increases in atmospheric pCO2 levels at a multiscale resolution using a fully coupled ocean–atmosphere model (FOAM). We show that, in association with stronger high-latitude precipitation rates and enhanced continental runoff, the demise of polar sea ice due to the global warming event involved a regional freshening of Arctic surface seawaters. These disturbances lead to progressive slowdowns of the global oceanic circulation accountable for widespread ocean stratification and bottom anoxia processes in deep oceanic settings and epicontinental basins. In agreement with very negative oxygen isotope values measured on fossil shells from the NW Tethys, our simulations also show that recurrent discharges of brackish and nutrient-rich Arctic surface waters through the Viking Corridor could have led to both vertical and geographical gradients in salinity and seawater d18O in the NW Tethyan seas. Locally contrasted conditions in water mass density and rises in productivity rates due to strong nutrient supplies could partly explain the regional severity of the anoxic event in the restricted Euro-boreal domains, as it has been previously suggested and modeled regionally.

The Mesozoic, perhaps the longest period of warmth during the Phanerozoic Earth history, has been repeatedly affected by short‐lived cold interludes lasting about one million years. While the origin of these cold snaps has been classically attributed to a temporary atmospheric CO 2 drawdown, quantified mechanisms explaining these instabilities of the carbon cycle are still lacking. Based on a climate carbon cycle model, we show that the general demise of carbonate platforms accompanying these short‐lived cold interludes is a powerful mechanism capable of generating a fast atmospheric CO 2 decrease and a moderate sea level drop associated with ice sheet buildup. The temporary nature of the carbonate production decline explains the relative short time of these cold events but makes it possible to account for ice sheet waxing and waning.

The Pliensbachian–Toarcian crisis (Early Jurassic) is one of the major Mesozoic paleoecological disturbances when ca. 20% of marine and continental families went extinct. Contemporaneously, profound paleobiogeo-graphical changes occurred in most oceanic domains including a disruption of ammonite provincialism during the Early Toarcian. Here, we quantitatively reappraise the structure and evolution of paleobiogeographical patterns displayed by ammonite faunas before, during, and after the biological crisis, over a time-interval including 13 biochronozones. The high-resolution study presented here involves the use of hierarchical Cluster Analyses, non-metric Multi-Dimensional Scaling methods, and Bootstrap Spanning Network approaches that we apply to a large database including 772 ammonite species from 16 northwestern Tethyan and Arctic basins. Our results confirm a robust faunal dichotomy between Euro-Boreal and Mediterranean areas throughout the Pliensbachian, with the first emergence of an Arctic biome during the cooling regressive event of the Spinatum Zone. Whatever its complexity, Pliensbachian provincialism could be directly linked to paleogeographical barriers and to latitudinal paleoclimatic and paleoecological contrasts. During the Early Toarcian, this pattern was progressively lost, with northward expansions of Mediterranean ammonites during the Tenuicostatum Zone, followed by a strong interprovincial mixing during the Falciferum Zone. This faunal homogenization results from the combination of several parameters including a major sea-level rise facilitating basinal connections, a global warming event stretching the spatial range limits of southern taxa, and a mass extinction preferentially removing endemic species. Ammonite provincialism, although slightly different, was progressively re-established during the cooling regressive trend of the Middle Toarcian. These results therefore suggest a paramount influence of paleoclimatic, eustatic, and extinction constraints on the paleobiogeography of Early Jurassic ammonites, even if some threshold effects or independent biological factors may sporadically complicate the patterns.

The Pliensbachian–Toarcian interval was marked by major environmental disturbances and by a second-order mass extinction. Here, we reappraise the taxonomic, spatiotemporal and selective dynamics of extinctions over the whole interval, by analysing a high-resolution dataset of 772 ammonite species from NW Tethyan and Arctic domains. On average, 40–65% of ammonite species disappeared during each subchronozone, but higher extinction pulses (reaching 70–90%) prevailed from the Margaritatus to the Dispansum Chronozone. The main extinctions, corresponding to the Gibbosus, Pliensbachian–Toarcian boundary, Semicelatum, Bifrons–Variabilis, and Dispansum events, differed in their dynamics, suggesting episodes of ecological stress related to climate change, regression, disturbance in the carbon cycle or anoxia. The multi-pulsed volcanic activity in the Karoo–Ferrar province could well have triggered these ecological changes. In addition, ammonites experienced a morphological bottleneck during the Gibbosus event, 1 Ma before the Early Toarcian diversity collapse. Typically, drops in richness were related both to high extinctions and to declines in origination rates. This feature could result from strengthened ecological stresses related to the temporal overlap of environmental disturbances. After the Early Toarcian crisis, the recovery of ammonites was rapid (2 Ma) and probably influenced by a coeval marine transgression.

Oxygen isotopes of biogenic apatite have been widely used to reassess anomalous temperatures inferred from oxygen isotope ratios of ancient biogenic calcite, more prone to diagenetic alteration. However, recent studies have highlighted that oxygen isotope ratios of biogenic apatite differ dependent on used analytical techniques. This questions the applicability of the phosphate–water fractionation equations established over 25 years ago using earlier analytical techniques to more recently acquired data. In this work we present a new phosphate–water oxygen isotope fractionation equation based on oxygen isotopes determined on fish raised in aquariums at controlled temperature and with monitored water oxygen isotope composition. The new equation reveals a similar slope, but an offset of about + 2‰ to the earlier published equations. This work has major implications for paleoclimatic reconstructions using oxygen isotopes of biogenic apatite since calculated temperatures have been underestimated by about 4 to 8 °C depending on applied techniques and standardization of the analyses.

This paper describes a tectonostratigraphic model of the synrift evolution of the Early Jurassic High Atlas rift of Morocco. The model is constrained by mapping of a set of inverted extensional blocks, by facies analysis of carbonate platform and turbiditic to hemipelagic synrift deposits, and by high-resolution (n 3 100 ka) biostratigraphy of the Early Jurassic succession. The chronostratigraphic packages of the High Atlas of Rich vary significantly in thickness, facies and architecture from one tectonic block to another. Our study shows how synrift strain varied in space and time over a long time interval (14 Ma) around the High Atlas rift. Initially, in Sinemurian time, the High Atlas rift was affected by low-strain normal faulting that controlled the growth of an extensive, low-gradient carbonate platform, except in the northern domain (towards the rift axis), where hemipelagic deposition related to high-rate faulting prevailed. Subsequently, in Carixian–Domerian time, a rapid increase in accommodation space and block subsidence caused by highstrain normal faulting brought about localized drowning of the carbonate platform and the development of calciturbidites and of starved deposits towards the rift axis. During this interval, high-strain, upper-crustal normal faulting migrated rapidly (over a period of 5 Ma) towards the rift periphery.

A set of published, unpublished, and new clay mineral data from 60 European and Mediterranean localities allows us to test the reliability of clay minerals as palaeoclimatic proxies for the Pliensbachian–Toarcian period (Early Jurassic) by reconstructing spatial and temporal variations of detrital fluxes at the ammonite biochronozone resolution. In order to discuss their palaeoclimatic meaning, a compilation of low-latitude belemnite δ18O, δ13C, Mg/Ca, and 87Sr/86Sr values is presented for the first time for the whole Pliensbachian–Toarcian period. Once diagenetic and authigenic biases have been identified and ruled out, kaolinite content variation is considered as a reliable palaeoclimatic proxy for the Early Jurassic. Major kaolinite enrichments occur during times of low δ18O, high Mg/Ca, and increasing 87Sr/86Sr, implying warm climates and efficient runoffs during the Davoei, Falciferum and Bifrons Zones. Conversely, cooler and drier times such as the Late Pliensbachian or the Late Toarcian are characterized by low hydrolysis of landmasses, and correspond to kaolinite depleted intervals. Secondary factors as modifications of sources or hydrothermalism may sporadically disturb the palaeoclimatic signal (e.g., in the Bakony area during the Late Pliensbachian). In addition, a spatial comparison of clay assemblages displays significant kaolinite enrichments towards northern parts of the Peritethyan Realm, probably related to the latitudinal zonation of hydrolyzing conditions. This implies enhanced runoffs on northern continental landmasses that reworked kaolinite-rich sediments from subtropical soils and/or Palaeozoic substrata.

For decades, theoretical morphological studies of different groups of organisms have been successfully pursued in biological, paleontological, and computational contexts, often with distinct modeling approaches and research questions. A regular influx of new perspectives and varied expertise has contributed to the emergence of a veritable multidisciplinary outlook for the-oretical morphology. The broadening of this discipline is reflected in a substantial increase in the number of models, leading to a bewildering diversity that has yet to be scrutinized. In this work, we tackle this issue in a synthetic fashion, with a quantitative meta-analysis that allows an objective comparison of theoretical morphological models treated as entities. By analogy with empirical morphospace analyses of actual organisms, we performed a multivariate ordination of a represen-tative sample of models, producing a metaspace of models in which patterns of similarity and dif-ference are visualized. A phenetic tree was used to characterize the relationships between models. Four major groups have been identified, and their disparity analyzed. We suggest this typology as a useful starting point to identify a core set of fundamental principles and protocols for better interpretation of the plethora of current models and for more efficient construction of models in the future. This in turn can help in diversifying the scope of macroevolutionary, developmental, and bioenvironmental questions in theoretical morphology.