# Palaeo-bioinspiration draws on the fossil record to advance innovation

Annabelle Aish, Chris Broeckhoven, Valentin Buffa, Tom Challands, Anton Du Plessis, Tom Fletcher, Eberhard Frey, Romain Garrouste, Alexandra Houssaye, Guillaume Lecointre, et al.

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Annabelle Aish, Chris Broeckhoven, Valentin Buffa, Tom Challands, Anton Du Plessis, et al. Palaeo-bioinspiration draws on the fossil record to advance innovation. Communications Biology, 2025, 8 (1), pp.706. 10.1038/s42003-025-08043-6 . hal-05336802

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# Palaeo-bioinspiration draws on the fossil record to advance innovation

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Annabelle Aish Ⓓ¹, Chris Broeckhoven²,³, Valentin Buffa Ⓓ⁴, Tom Challands Ⓓ⁵, Anton Du Plessis Ⓓ⁶,⁷, Tom Fletcher⁸, Eberhard Frey⁹, Romain Garrouste¹⁰, Alexandra Houssaye¹¹, Guillaume Lecointre Ⓓ¹⁰, Valentina Perricone¹², Luce-Marie Petit¹³, Vikram Shyam¹⁴, Thomas Speck Ⓓ¹⁵,¹⁶,¹⁷,¹⁸ & Michael Habib Ⓓ¹⁹✉

Bioinspiration is an approach to innovation based on the observation of biological systems, of which only 0.1% remain since life began 3.7 billion years ago. Expanding the scope of bioinspiration to the fossil record greatly increases the diversity of potential biological “muses” and provides a means to understand the form, function and origins of current living systems. This extended approach, here termed “palaeo-bioinspiration”, has already been applied to the fields of hydrodynamics, aeromechanics, actuators, protective technology and building construction. To reach its full potential, palaeo-bioinspiration has to overcome the misconception that fossils are “failures” or “primitive”, and temper the widespread belief that extant biodiversity is inherently “optimised”. By encouraging interdisciplinarity, investing in technical infrastructure and collaborating with both Natural Science institutions and industry, palaeo-bioinspiration could become a powerful asset within the bioinspiration domain.

Bioinspiration and biomimetics are approaches that translate observations from nature into ideas for innovative products and processes¹⁻⁴. Inspiration can be gleaned from biological systems across multiple scales, from ecosystems to individual organisms, even down to biomolecules. Bioinspired innovation draws on a vast range of forms and functions, and through “reverse biomimetics”, scientists learn to better understand how living systems operate⁵. While extant biodiversity still holds considerable potential for innovation, it likely represents only 0.1% of all life that has ever existed on earth⁶ (Fig. 1). Fossils, in contrast, can offer a much larger, and hitherto underutilised, source of biological inspiration⁷⁻¹². Furthermore, the fossil record provides valuable context for understanding form-function relationships and the evolutionary origins of intriguing traits observed in living systems. In this article, we emphasise the value of expanding the scope of bioinspiration beyond present-day life and into Deep Time.

## The principles of Palaeo-bioinspiration

Bioinspiration has existed for centuries, but its systematic use as a strategic design tool is relatively recent. The first academic journal dedicated to the field, *Biomimetics* (edited by J. Vincent), only emerged in the last 30 years¹³ amidst growing scientific and technological interest in the form of patents, research grants, publications and products¹⁴. The concept that nature can inspire technical innovation has become a powerful idea⁴,⁵ that has caught the public's attention, especially in terms of its potential to support sustainability¹⁵,¹⁶.

The analysis and comparison of extinct life forms to human technology seems to have been first explored over fifty years ago¹⁷ and perhaps even a hundred years ago¹⁸. Subsequently, an array of different terms have been used to describe the concept of deriving innovation from extinct biological systems (Table 1), including “palaeomimesis”, “palaeobiotechnology” and “palaeomimetics”¹⁹⁻²². We opt for the inclusive term “palaeo-bioinspiration”, which is defined here as “a creative approach based on the observation of palaeobiological systems”.

Regardless of the terminology used, we propose four elements that underpin the methodological advantages of integrating knowledge from palaeontology into modern innovation processes: the “biological library” concept, the evolution of form and function, the process of convergent evolution, and the integration of environmental context (Fig. 2). Together, these elements underscore the vast, yet largely untapped, potential of extinct organisms in driving technological development and design strategies. Accordingly, our article complements and expands on principles set out in Perricone et al.²² (which focuses predominantly on the methodology and applications of a palaeo-bioinspired approach) by promoting a more expansive exploration of palaeontological data in innovation processes across diverse disciplines.

## The vast “biological library” of Deep Time

The fossil record represents a plethora of biological structures, functions and systems that no longer exist today. Bioinspiration based on extant living

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<caption>**Fig. 1 | Hypothetical phylogenetic tree demonstrating the limited species diversity of extant life.**</caption>

The tree represents all life that has existed on Earth; the three branches reaching the outermost edge (highlighted in green) represent species currently alive. Palaeo-bioinspiration takes advantage of the biological diversity of the entire tree.

<figure><img src="image_5.png" /><figcaption>**Table 1 | List of terminology related to learning from natural systems**</figcaption></figure>

<table><tr><td>Bioinformed</td><td>A design approach that is informed by detailed and accurate information on biological systems or processes (rather than superficial, figurative or one-dimensional analogies)<sup>115</sup></td></tr><tr><td>Bioinspiration</td><td>A creative approach based on the observation of biological systems<sup>116</sup></td></tr><tr><td>Biomimetics</td><td>The interdisciplinary cooperation of biology and technology, or other fields of innovation, with the goal of solving practical problems through the functional analysis of biological systems, their abstraction into models, and the transfer into and application of these models to the solution<sup>116</sup></td></tr><tr><td>Biomimicry</td><td>A philosophy and interdisciplinary design approach taking nature as a model to meet the challenges of sustainable development (social, environmental and economic)<sup>116</sup></td></tr><tr><td>Bionics</td><td>Like life (precursor to biomimicry). Bionics can also refer to using artificial materials and methods to produce activity or movement in a person or animal<sup>117</sup></td></tr><tr><td>Palaeo-bioinspiration</td><td>A creative approach based on the observation of palaeobiological systems</td></tr><tr><td>Palaeobiology</td><td>A discipline that combines biology with palaeontology. Palaeobiological research uses biological field research of current biota and of fossils millions of years old to answer questions about the molecular evolution and the evolutionary history of life</td></tr><tr><td>Palaeobiotechnology</td><td>Fossil-inspired technology<sup>19</sup></td></tr><tr><td>Palaeomimesis</td><td>Seeking inspiration and mimicking extinct lifeforms and ecosystems for modern problem solving<sup>20</sup></td></tr><tr><td>Palaeomimetics</td><td>The study of extinct species and their environment to inspire modern and future solutions<sup>21</sup>. Biomimetic design inspired by extinct species and evolutionary processes<sup>22</sup></td></tr><tr><td>Palaeo-biomimetics</td><td>The transfer of an idea drawn from a fossil organism to a technical application</td></tr><tr><td>Palaeontology</td><td>The scientific study of life of the geologic past (i.e. usually prior to the Holocene) that involves the analysis of plant and animal fossils, including those of microscopic size, preserved in rocks</td></tr><tr><td>Physiomimetics</td><td>Learning from nature<sup>20</sup></td></tr><tr><td>Reverse biomimetics</td><td>The process by which findings that arise during the implementation of biomimetics contribute to a better understanding of biological systems<sup>4</sup></td></tr></table>
**Fig. 2** | Four elements that underpin the advantages of integrating fossils into bioinspiration: the “biological library” concept, the evolution of form and function, the process of convergent evolution, and the integration of environmental context.

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systems can only draw from a snapshot in time. The inclusion of fossils increases the number of potential biological models by several orders of magnitude and provides unique traits for bioinspired innovation⁸⁻¹¹.

Many fossil organisms represent extremes in scale, function, and environmental context that are absent in the modern world, such as the titanosaur *Argentinosaurus* and *Patagotitan*, the largest known terrestrial animals to have ever existed²³. In this way, the fossil record opens our eyes to

the possibility that biological morphologies can scale up (or down) in ways that extant biodiversity cannot reveal. For example, a model that predicts the maximum size of a flying animal to be around 20 kg would appear to be sound when validated against living biodiversity alone, since no living flyers surpass that “limit” and multiple species seem to converge on it. However, even a cursory look at fossil records would quickly indicate that such a model is fatally flawed. The largest flight-adapted animals in the fossil record,
Pterosaurs, have estimated masses around 250 kg, which is an order of magnitude larger. Issues of scaling are not limited to overall body size, either. The scaling of numerous anatomical and performance attributes can be informed by the fossil record. For example, cardiovascular performance models must be able to account for the known existence of very large, very long-necked animals in the fossil record, like sauropod dinosaurs. Gigantism and dwarfism within different lineages also show the scalability of form-function relationships (i.e. modern dragonflies vs the Late Carboniferous giant dragonfly *Meganeura*<sup>24,25</sup>).

Other fossils do not have modern analogues for a specific shape or function, offering insights into unique “solutions” beyond current bio-diversity. Examples include the dermal armour of some placodonts (marine reptiles), the dorsal plates of stegosaurs (dinosaurs), the neural spine sail of *Dimetrodon* (stem mammal), the specialised floating legs of *Chresmoda* (archaeorthopteran insects)<sup>26</sup>, or stem structural adaptations found in tree-like lycopods (*Lycopodiopsida*) and in seed ferns (*Pteridospermatophyta*)<sup>8,12</sup>.

A rough order of magnitude suggests that there are petabytes of information extractable from scientific collections, zoos, botanical gardens, and journals that could provide a vast database for comparative biomechanics.

## Understanding the evolution of forms and functions

Palaeontology is the only bioscience that explores evolutionary systems in their entirety by incorporating the dimension of Deep Time. This provides valuable and unique opportunities to investigate the evolution of forms and functions. Evolutionary context can shed light on how, when, and in which environmental context specific traits evolved. Importantly, palaeontological data can also demonstrate that the evolution of observed traits is not influenced by functional demands alone: species may possess a particular trait simply due to common ancestry, an evolutionary “burden” tied to a particular lineage<sup>27,28</sup>. This tendency of form to persist based on ancestry alone is assessed through the measurement of phylogenetic constraints. This is typically measured as a variable known as “phylogenetic signal”, defined as the covariance in the trait of interest and the total branch length between any two taxa on a phylogeny for which the trait has been measured. Understanding the phylogenetic constraints of a trait can help determine its applicability as a model for innovation.

Robust inferences can be made about fossil taxa' form-function relationships on the basis of modern taxa analyses (whose performance can be estimated). This extension of the understanding and characterisation of form-function relationships to the fossil record (through actualism) considerably increases bioinspiration's potential. The perspectives offered by Deep Time also provide insights into the trade-offs inherent in the evolution of multifunctional living systems. Organisms are the result of compromises amongst essential life functions in relation to a set of environmental pressures<sup>27</sup>. Recognising these evolutionary trade-offs helps avoid possible “naïve” transpositions via the biomimetic method<sup>28</sup>, whilst also providing opportunities in the design of multifunctional technologies.

## Learning from convergent evolution

Fossil biota reveal numerous examples of convergent evolution, where unrelated groups of organisms derive physically similar “solutions” to specific functional challenges. Animal examples include powered flight (observed in mammals, insects, birds, fishes and pterosaurs); thunniform swimming (in bony fishes, sharks, mammals and ichthyosaurs); and camera eyes (in cephalopods, vertebrates, and cnidarians). In plants, convergent evolution is evident in the formation of lignified and/or silicified trunks (serving both the competing functions of mechanical stability and water assimilation/transport), seed dispersal mechanisms, climbing capacities, and motile structures<sup>12,29</sup>.

Studying convergence helps differentiate between *common* adaptive traits—i.e. a general response to the same constraint(s)—and those that are *specific* to particular lineages, often (but not solely) shaped by their phylogenetic heritage. Taking design inspiration from organisms displaying

convergent evolution is particularly robust because many of these “solutions” have appeared in independent evolutionary contexts, overcoming some of the phylogenetic constraints that limit the current approach to bioinspiration<sup>22,30</sup>. Deep Time provides abundant examples of convergent evolution, enabling more robust analyses than purely neontological research can offer.

## Integrating environmental context

The geological and fossil records encompass a broad range of environments determined by atmospheric composition, climate conditions (temperature and precipitation), continental and oceanic configurations, and the composition of oceanic waters (including dissolved oxygen). Many of these past environmental combinations no longer exist today<sup>31-33</sup>. Consequently, fossil organisms exhibit adaptations to a wider range of abiotic conditions when compared to extant species, providing a record of adaptations across a vast range of environmental contexts<sup>21</sup>. Organisms that succeeded under past conditions can thus provide scope for bioinspired innovations adapted to modern-day climatic scenarios of global environmental change (high CO₂ levels, extreme temperatures, anoxic oceans, etc.)<sup>34</sup>, or new industrial/technological contexts (with “novel” conditions).

## Palaeo-bioinspiration: stories of Potential and Success

Here we present a non-exhaustive set of examples demonstrating palaeo-bioinspiration's potential within six different fields.

### Hydrodynamics

In aquatic environments, palaeobiology has supported hydrodynamic drag-reduction breakthroughs. Modern shark scales have inspired riblet textures to reduce drag in swimwear<sup>35</sup> as well as anti-fouling solutions<sup>36,37</sup>. Yet the scales of extinct sharks helped identify the functional background of riblet evolution<sup>38</sup>, improving its technological application. Some extinct fauna were buoyant and mobile in water, despite often being heavily armoured<sup>39</sup>. This combination of strength, buoyancy and hydrodynamic efficiency has inspired self-propelled underwater vehicles<sup>19,40</sup>. In robotics, inspiration from the only taxa with four large paddles, the plesiosaurs (marine reptiles), found that swimming using all four flippers, (rather than just two at a time like the aquatic tetrapods of today, such as penguins, otariids, and sea turtles) was more efficient during transient manoeuvres, but at a rather high cost; using only two flippers could be used, in contrast, for low-cost cruising<sup>40</sup>. Such experiments have inspired autonomous underwater vehicles with different capabilities depending on the context of their use<sup>41</sup>. With regard to invertebrates, giant Mesozoic surface-skating insects (*Chresmoda*, archaeorthopteran insects)<sup>26</sup> could provide a better understanding of effective flotation mechanisms at different scales. Moreover, insects with swimming appendages on their legs might provide inspiration for aquatic biorobotics (for example, beetles of the extinct family Coptoclavidae and water scorpions of the Belostomatidae, around during the Cretaceous period)<sup>42</sup>.

### Aeromechanics

Extinct flyers, both obligate gliders and powered flyers, provide a wealth of original wing morphologies of interest<sup>11</sup>. Flyers in the fossil record include four-winged and delta-winged gliders, as well as forms with hybrid membranous and feathered wings<sup>13-45</sup>. The fossil record also includes the largest flying animals known, giant *azhdarchid pterosaurs*. Pterosaurs possessed compliant, single-spar wings with multi-modal capacity: the wings folded into robust walking limbs that likely provided most of the power for launch<sup>46-48</sup>. A patent filed for a wind-turbine blade retrofit that can increase turbine efficiency by 14% was heavily based upon concepts discovered in pterosaurs<sup>49</sup>. Pterosaur wing morphology also inspired a solid-state aircraft concept for high altitude exploration of Earth, Venus and Mars<sup>50</sup>. A NASA-Carnegie Mellon University collaboration prototyped a folding-wing “quad-launching” Mars explorer, inspired by pterosaur anatomy and reconstructed launch capacity<sup>51</sup>. In terms of insects, dragonflies are the oldest and most diverse flying organisms, providing an abundant source of functional
and environmentally adapted solutions just beginning to be explored<sup>25,52</sup>. Other extinct forms of insects (such as Palaeodictyopteroidea) could present different models of flight, testable through 3D wing reconstruction and mechanical/aerodynamic experimentation (Aracheloff, 2024)<sup>53</sup>.

## Actuators

Fossil plants have inspired novel movement designs, notably through their reticulate pneumatic actuators. Plants are valuable as models for actuator-based movement, in part, because they utilise very different adaptive “solutions” as compared to animal models. Plants have further utility in this regard, as there is substantial variation in the mechanisms of motion. Macroscopic cortical fibre networks are found in some extant (balsa, papaya; small to medium-sized trees) and fossil (e.g. *Lyginopteris oldhamia*, a Palaeozoic semi-self-supporting seed fern) plants. In these plants, uneven secondary wood growth causes an asymmetric deformation of fibre networks in the cortex, enabling inclined stems to remain upright<sup>54</sup>. These have been used as models for 3D reticulated actuator systems, in which an elastic technical hollow tube is surrounded by a net-like structure. Through asymmetric net arrangements, various types of bending movements can be programmed into these plant-inspired pneumatically-actuated soft machines<sup>54,55</sup>.

## Protection technology

The diversity of armour morphologies present in the fossil record greatly exceeds that of modern biological systems. The bony plates or osteoderms of *Glyptotherium* (an extinct relative of the modern-day armadillo) are characterised by a specific combination of thick, compacted layers and porosity of the cellular core, which likely provided an effective alliance of strength and high-energy absorption<sup>9</sup>. Some ankylosaurs (a group of armoured dinosaurs) preserved both keratin and bone in their osteoderms, their composite armour showing details of layering and material proportions<sup>56,57</sup>. Mechanical tests of a synthetic version of this armour run by Pacific Light & Hologram in Pasadena, CA, recovered excellent ablative and energy absorption capacity<sup>28</sup>. While the full analysis is still pending publication (test videos were made public through a CBC Broadcast), the company generated multiple forms of useful intellectual property in the development of custom ceramics and polymers used to create the biomimetic armour. Other dinosaur systems have also yielded industry results: the thick skull structure of some dinosaurs (i.e. the pachycephalosaurs) has inspired the development of light and resistant bike helmets<sup>22</sup>.

## Buildings and construction

Palaeontological organisms could inspire more efficacious building materials: principles extracted from fossil bone structure and fibre-reinforced structures (such as those found in trees) can facilitate the creation of efficient and lightweight constructions<sup>4,8,12,59,60</sup>. Recent studies have highlighted that bone is a powerful tool for bioinspiration<sup>61-63</sup> and that material structure inspiration from bones' inner architecture could offer better load resistance than a topologically optimised structure<sup>60</sup>. The same holds true for reticulated ultra-lightweight wood structures, which recently inspired the construction of sustainable fibre-based pavilions<sup>64</sup>.

Some of the largest animals in the fossil record also exemplify extremes in skeletal construction. Both giant dinosaurs and giant pterosaurs evolved exceptionally high stiffness-to-weight ratio skeletons. The inner structure of the limb long bones in sauropods is thinner than predicted from heavy tetrapod models (albeit not “thin-walled” in the engineering sense). The limb elements of the elephant-sized sauropod *Nigersaurus* showed a structure much lighter than that of today’s giant organisms (rhinos and hippos), with proportionally much thinner walls of compact bone<sup>65</sup>. Moreover, linear elastic static and linearised buckling analysis of the elon gate, thin-walled cervical vertebrae of giant pterosaurs recently yielded notable insights<sup>66</sup>. The unique structure, consisting of a “tube within a tube” shape supported by helically distributed trabeculae, resulted in an increase of the buckling load of the whole element by up to 90%, as compared to a simple, non-trabeculated model<sup>66</sup>.

Fossil marine animals may also provide inspiration in terms of structural resistance: the rigidity of stalked crinoids confers a capacity to resist ocean currents, possibly providing ideas for artificial structures requiring these same characteristics (such as wave energy systems)<sup>19</sup>. In addition, fossil-inspired robotic bivalves, capable of burrowing into unstable sediment, have been explored for potential use in anchoring submarine structures<sup>19,67</sup>.

## Global change resilience

The fossil record is arguably the best source of information regarding organismal adaptations to global change, because many palaeo-ecosystems experienced rapid environmental transformation. While these periods of change were often associated with mass extinctions, those lineages with higher-than-average survivorship provide critical insights into which traits confer advantages in times of global transition, informing technological and behavioural responses to modern biodiversity loss and climate change<sup>68,69</sup>. Applications of fossil record analyses need not be limited to the realm of physical technology: simulations based on extinction processes also have the potential to guide management of economic or pandemic developments and stock exchange crises. Moreover, outside these transition periods, fossil taxa have adapted to conditions unlike those of today (their survival over millions of years attests to this success): understanding these adaptations is a great source of potential inspiration.

## Challenges of Palaeo-bioinspiration

Palaeo-bioinspiration faces similar obstacles to its adoption as bioinspiration in general, including variable access to biological material and insufficient interdisciplinary communication. These common hurdles are outlined in more detail in several scientific publications<sup>5,28,70-72</sup>. Over and above these generic barriers, Palaeo-bioinspiration needs to address three significant challenges: working with incomplete information, overcoming the “fossils are failures” misconception, and reconceptualising optimisation.

## Working with incomplete information

One of the most obvious difficulties associated with Palaeo-bioinspiration is the inescapable loss of data that occurs during the death, decay, and fossilisation of extinct organisms (taphonomy) as well as the dispersed nature of most fossils. Added to this is the issue of preservation bias. The quality of fossil preservation depends on environmental factors, namely death milieu, sediment grain size, oxygen content, temperature, humidity, salinity, the presence of scavengers/decomposers, tectonics and rock mechanics, as well as biological tissue material properties. Fossil-bearing rock is not uniformly spread, neither temporarily nor spatially<sup>73-75</sup>. Indeed, most palaeoenvironments were not conducive to preservation, especially those areas prone to erosion (e.g. mountains, fast-moving rivers) as well as areas with high biological turnover (such as rainforests). Furthermore, while the form of an organism may be retained during fossilisation, the original chemical composition and material properties of biological structures are partially lost.

However, palaeontologists are used to working with taphonomy and assessing its significance. They can select the least affected specimens (appropriate for form-function analyses), resort to the building of a “global” specimen based on various incomplete ones, or use retrodeformation, for example, to overcome these challenges<sup>76,77</sup>. Moreover, although the material properties of fossil organisms are usually altered substantially by fossilisation, working estimates of the original material properties can often be made. Some biomaterials are fairly consistent in their construction and properties, and data from extant species can be used as a guide in these cases. In rare cases, details of preservation can yield indications of the original material properties. For example, histology of the elongate neck “ribs” in many giant, long-neck dinosaurs (called sauropods) shows that these structures were actually ossified tendons<sup>78</sup>. Ossified tendons in living relatives (birds) have material properties different from those of cortical bone, so the identification of these bony structures as tendons shifts (and narrows) the range of plausible material properties.
The locomotion, feeding, reproductive habitats, physiology and behaviour of fossilised organisms all require reconstruction based on available evidence, which can vary greatly amongst specimens. Furthermore, mechanical properties, physiology, environmental constraints, etc., can only be reconstructed based on the natural and physical laws observed *today*, assuming these worked alike in the past (constancy of cause and effect throughout space and time<sup>29,80</sup>). This untestable assumption is called actualism, or in geology, better known as uniformitarianism, a discipline of analytic philosophy.

The principle of actualism was coined by Charles Lyell in the early 1830s<sup>81</sup>; ‘Lyell’s Law’ was subsequently commented on and updated<sup>82–86</sup>. Actualism or uniformitarianism is the only methodological concept that allows the reconstruction of fossils’ material properties, physical constraints, metabolic demands, intrinsic mechanics, etc., and, most importantly, their evolutionary pathways. In other words, this approach uses the study of extant life forms as an essential “gateway” for the interpretation of the fossil record. It does not seek to compare fossil life forms with closely related extant ones; rather, it focuses on their comparison with structurally and mechanically similar extant living constructions. This method vastly expands the scope for discovery of extinct life forms relevant for palaeo-bioinspired solutions because, by comparing structural differences between past and extant life forms, unexpected, innovative and surprising solutions come to light.

Modern palaeontologists and evolutionary biologists have also developed powerful techniques to overcome some limitations of the fossil record, particularly in relation to form-function relationships. These include both physical and experimental methods of analysis borrowed from engineering, such as computed tomography, 3D modelling, finite element analysis, laser doppler anemometry, computational fluid dynamics, machine learning, and robotics. Information on extant organisms and their biomaterials (including non-mineralised hard tissue like cartilage, keratin, chitin, wood, and sclerenchyme fibres) can be used as input data for numerical and analogue simulations. Phylogenetically close living relatives can help to identify unpreserved tissue correlates in fossils, which can then be interpreted based

on structural analogues. Tissues, surface characteristics, interspecific relationships (parasitism/predator-prey), growth habits and heights<sup>87,88</sup>, colours<sup>89</sup>, physiology<sup>90</sup>, movement<sup>91</sup> and even sounds<sup>92</sup> may be reconstructed with moderate to high confidence in many cases. Modern technology also permits the extraction of biomolecules from fossil tissues, unleashing the possibility of developing antimicrobial pharmaceuticals inspired by long-extinct organisms<sup>19</sup>.

Nonetheless, species known only from the fossil record are invariably data-density poor compared to their living counterparts, and there are significant differences between information gleaned from vertebrate and invertebrate fossils (see Box 1). Palaeo-bioinspiration will always be most effective when combined with the use of living model systems: fossil model systems provide a vast array of forms and long spans of time, while living model systems provide precise information and a higher density of data per species.

## Overcoming historical assumptions

A notable barrier to the broader adoption of Palaeo-bioinspiration as a design methodology is the idea that extinct organisms are somehow the “losers of evolution”<sup>93</sup>. This is, of course, incorrect: many extinct species or genera existed for millions of years and were very successful both in terms of their evolutionary lifetime and geographical distribution<sup>21</sup>. Asteroids, volcanic activity, abrupt climate change, changes in resource availability or competition (as well as more chronic changes, such as orogenesis, sea level rise or glaciation) may have given rise to their extinction rather than any evolutionary “inadequacy”<sup>20</sup>. The popular notion of a “primitive” species or group, either extant (e.g. the platypus as a “primitive” mammal) or extinct (e.g. placoderms as “primitive fishes”), encourages the notion that “fossils are failures”. It implies that the destiny of something “primitive” is to be replaced by something “more evolved” or “better adapted”, which implies the former is unlikely to provide data useful for technical solutions. This view is erroneous. Character states or traits can be qualified as “primitive” or “derived”, but not taxa themselves. Indeed, to avoid such misconceptions, in 1950, Hennig devised the adjectives “plesiomorph” for “primitive” and

# Box 1 | Vertebrate vs. invertebrate fossils as sources of palaeo-bioinspiration

Many mechanical properties of fossil vertebrates can be reconstructed based on the application of uniformitarianism or actualism (assisted by graphical software, for example<sup>77</sup>), even if they look very different to extant vertebrate life. One reason is that their soft tissues can be reconstructed from traces of muscle insertions (muscle scars, inner bone structural features) or from structurally similar or evolutionarily close modern organisms (phylogenetic bracketing)<sup>118,119</sup>. The functional aspects of bone articulations in vertebrates can also be reconstructed to a reasonably high fidelity<sup>120,121</sup>. Overall, constructional and mechanical approaches are underpinned by abundant structural data in fossil vertebrates.

In contrast, fossil invertebrates are much more difficult to interpret with respect to constructional morphology because their support systems are based on hydroskeletons that are never preserved in the fossil record. Invertebrates that lack any hard parts (e.g. non-mineralising Cnidaria, “naked” Mollusca, Tunicata, jawless Annelida, Nematoda, Enteropneusta or other hitherto unknown taxonomic groups) at best leave faint traces (“soft fossils”<sup>122</sup>), which barely yield sufficient details for reconstruction—although there are a few biomechanical studies on such life forms that could feed into palaeo-bioinspiration<sup>123–125</sup>.

Furthermore, any hard parts, if present, do not reflect the morphology of the respective soft tissues. The latter is especially true when these hard parts are suspended in soft tissue, which disintegrate during decay (e.g. jawed Annelida and Chaetognatha). The presence of any type of exoskeleton supports fossilisation and therefore enhances the diversity of the

fossil record dramatically. Both exo- and endo-skeletons showed up in invertebrates during the so-called (and disputed) ‘Cambrian Explosion’<sup>126,127</sup> allowing their mechanical properties to be studied (for ammonites<sup>128–130</sup>; for trilobites<sup>131</sup>; for arthropods in general<sup>132,133</sup>).

With the discovery that the internal pressure of the haemolymph in invertebrates represented an antagonist to their flexor muscles (hydraulic leg extension<sup>134</sup>) the uniformitarianism gateway was open for the application of this discovery to the fossil record<sup>135</sup>. The hydromechanical properties of extant spider legs have resulted in biomimetic experiments based on the 3D reconstruction of the limb segment articulations<sup>136,137</sup>. Meanwhile, 3D preserved fossil arthropods have been discovered<sup>133,138</sup>. Applying graphical tools like Blender, the retro-deformation of entire invertebrates is possible, including the mechanical specifications of the intersegmental joints of limbs, mouthparts and body<sup>133,138</sup>. The study of flight in basal dragonflies<sup>24</sup> presents a better understanding of the role of key innovations such as the presence of biological materials likely to enter into resonance<sup>139</sup>. This has led to at least one artificial wing prototype Aracheloff<sup>53</sup>. Large (wingspan 0.3 m) fossil Chresmoda capable of moving on top of the water surface are known from Mesozoic records. Modelling of surface tension dynamics in these taxa could expand on knowledge from extant water striders<sup>140</sup>.

Given that the fossil record is full of invertebrates without any extant allies, the functional investigation of their morphology will certainly result in innovative palaeo-bioinspired applications in hitherto unknown fields.
"apomorph" for "derived" for character states⁹⁴. These notions are relative to the taxonomic group in question. For example, the presence of feathers is a "primitive" (or "ancestral") trait among birds but a "derived" (or "advanced") trait among archosaurs (the common ancestor of crocodiles, pterosaurs, birds and their descendants). No taxon is simply "primitive" in and of itself. Refuting the existence of "primitive" taxa was a fundamental part of the late 20th-century paradigm shift in phylogenetic systematics. The concept of "grades" was abandoned as a valid taxon because it could not be defined independently, only by its "primitiveness" in relation to "more evolved" taxa (an approach criticised by "cladists" like Gould). And even the palaeontologists of the era were convinced that so-called "primitive fossils" were still adapted to the environmental conditions of their time: these species were never considered "failures" of evolution, even by those who believed them to be "primitive". Unfortunately, for the layperson, the term "primitive" can erroneously be understood as "obsolete" or "inferior". Palaeo-bioinspiration should raise awareness of (and capitalise upon) the fact that extinct organisms were no less viable in their respective environments than organisms alive today.

## Reconceptualising optimisation

The notion that "fossils are failures" often goes hand in hand with the misguided belief that today's species are somehow "optimised" or even "optimal" and thus can provide superior bioinspired solutions¹,⁵,²². This misrepresents the processes of evolution, environmental change, phenotypic plasticity and extinction events²⁰,²¹,²⁷,²⁸. Adaptations, and natural selection that principally shapes them, produce compromises, not perfections⁹⁵. Inaccurate public perceptions about palaeoecology are also prevalent: past habitats are often presented as "static ancient worlds" as opposed to dynamic environments offering new ecological niches throughout Earth's history.

While evolutionary processes do change the physiological and/or mechanical performance of organisms, they result in multiple factor "optimisations" within the context of phylogenetic heritage. These trade-offs generate functional requirements in a specific tissue or organ that are (partially) contradictory, i.e. it becomes impossible to improve one function without making the other worse, known as Pareto efficiency²⁸. Evolutionary processes often "resolve" this problem by combining "good enough" solutions for each functional requirement. One example of this is the opposition between mechanical stabilisation and water conduction in tree stems. In conifers, this issue is temporarily resolved by the formation of early wood in spring (mainly for water conduction) and late wood in autumn (mainly for stabilisation), whereas in broad-leaved trees, a spatial separation in water conduction vessels and stabilising wood fibres has evolved⁹⁶,⁹⁷.

For palaeo-bioinspiration to be effective, innovators must first adequately identify their challenges and then use phylogenetically-informed methodologies to resolve them. Both neontological and palaeontological model systems are at their most useful when applied to complex optimisation problems across multiple variables; conversely, "traditional" methods of innovation and design may perform better than biologically-inspired approaches where single-variable optimisation is desired. Regardless of the variables of interest, phylogenetic constraints must be considered (and preferably quantified) for the proper application of palaeo-bioinspiration as a methodology. Phylogenetic context is the only effective way to differentiate between functional constraints (that can inform bioinspired design) and phylogenetic constraints.

## Future directions

### Building on the foundations of "extant" bioinspiration

An important first step in the development of palaeo-bioinspiration will be joining forces with colleagues involved in neontological bioinspiration, which has already established itself as a thriving domain of interdisciplinary research, design and innovation⁴,⁵. Indeed, the research that supports palaeo-bioinspiration and (extant) bioinspiration is already inexorably connected. When analysing and modelling the functions of extinct organisms, researchers have no choice but to refer to extant analogues. But the

reverse can also be true: the inclusion of the fossil record in bioinspired design processes can be critical to properly modelling and predicting the performance of innovations based on extant systems¹⁶,⁴⁷. Moreover, as so much of Earth's biodiversity is extinct, many (if not most) of the extreme morphologies are also exclusive to the fossil record. The inclusion of even a modest sample of fossil forms might immediately demonstrate a foundational error in models validated only on extant species. As already outlined, this is particularly clear with regard to issues of scaling.

Ultimately, biology is the guiding thread that connects different types of bioinspiration⁹⁵. And, in referring to "biology", we include the theoretical framework that goes along with it, i.e. the entire theory of evolution. Bioinspiration should not be limited to exploring how organisms function in the here and now. A critical part of explaining organismal structure is historical (taking Seilacher's triangle into account⁹⁸), and promoting a more efficient bioinspiration (based on either extant or extinct organisms) should start with a shared evolutionary understanding informed by palaeontology.

Methods and tools already developed in the bioinspiration field can be adapted and applied effectively to palaeo-bioinspiration, as described in Perricone et al. in relation to the *Pachycephalosaurus*-inspired bike helmet²². In the future, the evolution of palaeo-bioinspiration will require additional research and case studies to transform it into a recognised part of the bioinspiration field. The realisation of a palaeo-bioinspiration database could be a useful support, including a collection of morpho-functional "solutions" as well as evolutionary frameworks, processes, and "traps" to avoid²².

## Supporting interdisciplinarity

One of palaeo-bioinspiration's key strengths is its scope for resolving engineering challenges. The potential of palaeo-bioinspiration will be best realised through interdisciplinary teams of palaeontologists, geologists, biologists, material scientists, designers and engineers. Currently, much research occurs in silos, impeding effective and goal-oriented "bioinspired" collaboration⁹⁹. Funding for interdisciplinary and international palaeo-bioinspired research would enable initiatives that encourage dialogue and networking. Building interest in palaeo-bioinspiration among applied scientists will be important for the development of strong teams and enhanced funding opportunities that bridge disciplines. Focusing on transposing morphological traits to technology by way of high-profile examples can demonstrate palaeo-bioinspiration's advantages and help it gain recognition. Moreover, the fact that many fossil organisms inspire awe could help generate the positive connotations needed to popularise palaeo-bioinspiration⁴.

Just as the bioinspiration field strives to train "horizontal biologists"⁹⁹,¹⁰⁰, palaeo-bioinspiration will benefit from training palaeontological generalists who can support biomimetic projects from start to finish, seeking specialist advice when needed. The promise of scientific insights could encourage the involvement of a broader palaeontological community: this intellectual "bi-directionality" already occurs within extant bioinspiration. For example, palaeontologists can benefit from reverse engineering studies where "solutions" are applied iteratively to uncover the functions of organisms and their constructional subunits. In addition, reverse biomimetics (a heuristic loop comprising engineering biology and biomimetics) can improve our understanding of palaeontological organisms¹⁰¹, their evolution and even interpolate "missing" morphologies in the fossil record by using information and methods from biomimetics⁵,¹⁰²⁻¹⁰⁴.

## Investing in essential technical infrastructure and "virtual" palaeontology

Bringing the past to life requires not only classical descriptive and comparative methods but also modern research technology¹⁵. Advanced 3D imaging technologies, including micro-computed tomography (µCT),aminography and synchrotron tomography, are essential to discovering and describing new fossil structures, as part of the field of "virtual palaeontology"²². Laboratory micro-CT is the most widely available of these and has been shown to be useful for a wide range of biomimetic studies¹⁰⁶,¹⁰⁷.
Finite element analyses model how load is distributed and can, for example, enable the estimation of degrees of freedom within joints (and other physical capacities) of fossil organisms¹⁰⁵, sometimes revealing previously undetected differences between similar structures. 3D/4D printing and additive manufacturing present opportunities for recreating complex palaeontological forms, allowing them to subsequently be tested mechanically, without putting the original fossils at risk¹⁰⁸. 3D printing has advanced past the original “prototyping” use and today assembles high-quality final products. One key advantage is its complex manufacturing capability, which in turn supports greater biomimetic design capacity¹⁰⁶. From a computational perspective, the hardware and software needed for 3D image analysis and simulations can be costly (particularly for a ‘traditional’ palaeontological lab), a factor that needs to be considered when initiating palaeo-bioinspired projects.

## Collaborating with Natural History Museums and other institutions

Museums possess powerful means to promote palaeo-bioinspiration as a concept, through public outreach activities, exhibitions, training and seminars. Natural History museums and other related institutions (universities, geological heritage institutions, etc.) are also rich sources of palaeontological specimens, literature and scientific expertise. The palaeo-biological library is ever expanding, and therefore, museums, which house these fossil specimens, are integral to the future of innovation. Facilitating access to these resources will be fundamental for palaeo-bioinspiration research. Given the unique value and fragility of many (palaeontological) specimens, a growing number of institutions have invested in 3D scanning and digitisation of specimens. In Europe, these include the Consortium of European Taxonomic Facilities (CETAF)’s “DiSSCo” initiative¹⁰⁹ (Distributed System of Scientific Collections) co-funded by the European Commission, which seeks to digitally unify all European natural science resources. In the US, the Advancing Digitisation of Biodiversity Collections project (ADBC¹¹⁰) has been funded by the National Science Foundation (NSF), which has also co-funded the creation of MorphoSource¹¹¹ and the Paleobiology Database¹¹². These digitisation projects allow excellent remote access to specimen data for those interested in palaeo-bioinspiration. Palaeo-bioinspiration projects also require direct input from palaeontologists to succeed (as with “extant” bioinspiration projects¹¹³), yet finding the right expertise can be challenging. Efforts are underway, for example, as part of Europe’s TETTRIS project¹¹⁴, to improve online access to taxonomists via a dynamic catalogue, thereby facilitating connections between palaeontological experts and bioinspiration practitioners.

## Building links with industry

While interdisciplinary research initiatives usually precede bioinspired innovations, industry partnerships ultimately support their commercialisation. As with bioinspiration based on neontology, it is essential to understand the needs and challenges of industrial sectors (“technology pull”) as well as effectively present the benefits of palaeobiologically-inspired innovation (“biology push”). Both these aspects should ideally be represented in collaborations with industry²². Client-oriented approaches that incorporate expertise from the social sciences, art, and communication sectors will, in turn, provide the most effective product storytelling.

## Conclusions

Bioinspiration based on extant biological systems is now a dynamic domain, with the scope to address a myriad of technical and environmental challenges that humanity currently faces. Exploring the fossil record could expand bioinspired innovation's capacity considerably, drawing on 99.9% of all life forms and the vast pool of biological phenotypes buried in Deep Time. This approach, termed palaeo-bioinspiration, is garnering attention, yet still faces several barriers to adoption. Beyond the inherent challenges associated with palaeontology itself, fossil organisms still tend to be misaligned as “dead ends” or “evolutionary failures” in the public consciousness and, as such,

inappropriate drivers of innovation. However, extinction is an erroneous measure of success¹⁹. It does not diminish the value of traits that allowed organisms to cope with complex problems, including those of interest to modern engineers and designers. These prejudices need to be overcome within engineering sciences, the industrial sector, and the public (as end users) for palaeo-bioinspiration to reach its full potential. Interdisciplinary research and design require greater support, and palaeontologists need to be integrated into the process of palaeo-bioinspired innovation from the outset. Such collaborations have the potential to yield substantial benefits to both innovating engineers and to palaeontologists. This “win-win” aspect can be leveraged to help build positive collaborative relationships between the fields of work. Finally, bioinspiration has made tremendous progress in the last twenty years and encompasses many of the same opportunities and challenges as the sub-field of palaeo-bioinspiration. Palaeo-bioinspiration can build on these strong foundations whilst offering a supplementary array of ideas and expertise, creating a new niche within the bioinspiration landscape.

Received: 29 July 2024; Accepted: 4 April 2025;

Published online: 02 June 2025

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## Acknowledgements

The authors would like to thank the following colleagues for their participation in discussions that informed the writing of this paper: Christine Böhmer, David Hone, Stephen Howe, Rick Lind, Elisabeth Martin-Silverstone, Tom Masselter, James Nebelsick, Andrew Parker, Anita Roth-Nebelsick, Olga Speck, Jean-Sébastien Steyer, Igor Yadroitsev and Ina Yadroitsava.

## Author contributions

A.A. created the framework for the manuscript, had a lead role in the writing, and was responsible for the core concept of the perspective (lead author role). M.H. had a lead role in editing and editorial correspondence, as well as significant writing (senior author role). T.C. led the preparation of figures and significant writing and editorial contributions. All other authors (C.B., V.B., A.D.P., T.F., E.F., R.G., A.H., G.L., V.P., L.M.P., V.S. and T.S.) contributed equally to writing.

## Competing interests

The authors declare no competing interests
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<footnote n="1">¹Bioinspire-Museum, Direction Générale Déléguée à la Recherche, l'Expertise, la Valorisation et l'Enseignement (DGD REVE), Muséum National d'Histoire Naturelle, 57 Rue Cuvier, CP 17, 75005 Paris, France. ²Advanced Concepts Team, European Space Agency, Keplerlaan 1, 2201 AZ Noordwijk, The Netherlands. ³Laboratory of Functional Morphology, Department of Biology, University of Antwerp, Universiteitsplein 1, 2610 Antwerp, Belgium. ⁴Evolutionary Studies Institute, University of the Witwatersrand, Private Bag 3, WITS, 2050 Johannesburg, South Africa. ⁵School of Geosciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh EH9 3FE Scotland, UK. ⁶Research Group 3D Innovation, Stellenbosch University, Stellenbosch 7602, South Africa. ⁷Comet Technologies Canada Inc, Montreal H3B 1A7, Canada. ⁸School of Geography, Geology and the Environment, University of Leicester, Leicester, United Kingdom. ⁹Sonnenbergstraße 27, 75180 Pforzheim, Germany. ¹⁰Institut de Systématique, Évolution et Biodiversité, UMR ISYEB 7205 CNRS MNHN SU EPHE UA, Muséum National d'Histoire Naturelle, CP 50, 45 Rue Buffon, 75005 Paris, France. ¹¹UMR 7179 CNRS/MNHN, Département Adaptations du vivant, Muséum National d'Histoire Naturelle, 57 rue Cuvier CP-55, 75005 Paris, France. ¹²Stazione Zoologica Anthon Dhorn of Naples, Villa Comunale, 80121 Naples, NA, Italy. ¹³Centre d'études et d'expertise en biomimétisme, 62 rue du Faubourg Saint-Martin, 60300 Senlis, France. ¹⁴NASA Glenn Research Center, Cleveland, OH, USA. ¹⁵Plant Biomechanics Group @ Botanic Garden, University of Freiburg, Schänzlestraße 1, D-79104 Freiburg, Germany. ¹⁶Freiburg Center for Interactive Materials and Bioinspired Technologies (FIT), Georges-Köhler-Allee 105, D-79110 Freiburg, Germany. ¹⁷Freiburg Materials Research Center (FMF), Stefan-Meier-Straße 21, D-79104 Freiburg, Germany. ¹⁸Cluster of Excellence livMatS @ FIT - Freiburg Center for Interactive Materials and Bioinspired Technologies, University of Freiburg, Freiburg, Germany. ¹⁹Dinosaur Institute, Natural History Museum of Los Angeles County, Los Angeles, CA, USA. e-mail: biologyinmotion@gmail.com</footnote>