Maria João Fernandes Martins Created 2026-10-20 Updated 2026-11-05
A researcher who coauthored an analysis of sexual dimorphism and extinction patterns in fossil ostracods. The interpretation used body-form differences as a proxy for reproductive investment, making the proxy assumption part of the argument.[ref]
High male sexual investment as a driver of extinction in fossil ostracods by Maria João Fernandes Martins and others. 2018, dimorphism and extinction estimates
Four documented examples, each tied to its actual context, are:
- Geographic range: widespread marine bivalve genera survived the end-Cretaceous event better than geographically restricted genera. Spatial spread can reduce exposure to some geographically heterogeneous stresses, although this does not guarantee survival of a global perturbation.[ref]Selectivity of end-Cretaceous marine bivalve extinctions by David Jablonski and others. Abstract: lower extinction intensity in geographically widespread genera
- Body size: large-bodied mammals were disproportionately lost during the late Quaternary. The global analysis documents size selectivity across continents and trophic groups; this is not evidence that large body size always raises extinction risk in every geological interval.[ref]Body size downgrading of mammals over the late Quaternary by Felisa Smith and others. Abstract: size-selective losses over the studied late-Quaternary intervals
- Physiological buffering and respiratory capacity: poor capacity to tolerate elevated carbon dioxide and associated oxygen stress has been linked to higher vulnerability among end-Permian marine groups. This is a paleophysiological interpretation informed by living relatives and fossil traits, rather than a direct measurement of a fossil animal’s tolerance.[ref]Paleophysiology and end-Permian mass extinction by Andrew H. Knoll and others. Abstract and section Extinction in the oceans: hypercapnia vulnerability and physiological selectivity
- Sexual dimorphism as a proxy for male reproductive investment: greater male–female carapace differences in fossil cytheroid ostracods were associated with higher estimated extinction rates. The measured fossil trait is dimorphism; interpreting it as investment requires the biological correspondence observed in living relatives.[ref]High male sexual investment as a driver of extinction in fossil ostracods by Maria João Fernandes Martins and others. Abstract: dimorphic fossil carapaces and association with estimated extinction rates
These examples demonstrate event- and group-dependent selectivity. They do not define one universal vulnerability ranking: the bivalve study, for instance, did not find body-size selectivity at the same end-Cretaceous genus-level event.
Extinction selectivity means that the probability of extinction depends systematically on biological or ecological traits, rather than being equal across all species. An empirical association needs a specified event, group and taxonomic level; a trait protective in ordinary background turnover may not protect during every mass extinction.
Geographic range: at the end-Cretaceous crisis, geographically widespread marine bivalve genera had lower extinction intensity than narrowly distributed genera. This makes range breadth a demonstrated selective trait for that dataset. The same analysis found no clear selectivity by body size, so that result should not be generalized to every trait or event.[ref]
Selectivity of end-Cretaceous marine bivalve extinctions by David Jablonski and others. Abstract: geographic-range selectivity and absence of body-size selectivity
Body size: large-bodied mammals suffered disproportionately during late-Pleistocene and later Quaternary losses, including the loss of many giant herbivores and carnivores. A global analysis over the last years found strong size-selective losses across continents and trophic modes. Large size can correlate with slow reproduction, high resource demand and exposure to human hunting, but the observed size pattern and its possible mechanisms should be distinguished.[ref]
Body size downgrading of mammals over the late Quaternary by Felisa Smith and others. Abstract: size-selective extinction across continents, trophic modes and intervals
Sexual dimorphism and reproductive investment: a study of fossil cytheroid ostracods found much higher estimated extinction rates in species with stronger male–female carapace dimorphism, interpreted as greater male investment in reproduction. This supplies a third empirical trait association; the observed relationship is not a universal claim that sexual selection is always harmful.[ref]
High male sexual investment as a driver of extinction in fossil ostracods by Maria João Fernandes Martins and others. Abstract and main result: approximately tenfold rate contrast between high- and low-investment species
There is no general basis for assigning behaviour a smaller causal role in macroevolution. Its immediate effects are usually easier to measure within populations, but those effects can alter lineage formation and persistence. The comparison needs a definition of “contribution”: the proportion of evolutionary changes initiated by behaviour, their magnitude, their necessity, or the strength of available evidence would give different answers. Microevolution concerns change within biological populations; macroevolution concerns patterns at and above the level of species, including speciation, adaptive radiation and extinction. They are connected processes with different objects of measurement.
At the population scale, animal behaviour changes who encounters which resources, survives, mates and reproduces. Habitat preference changes exposure to ecological natural selection; mate choice changes reproductive success; dispersal changes the spatial distribution of alleles. Behaviour can therefore affect the evolution of other traits even if the behaviour itself does not evolve. Conversely, an environmentally induced change in behaviour is not automatically evolutionary change. Learning and phenotypic plasticity can occur within one lifetime, whereas an evolutionary claim requires persistence through inherited differences or an explicitly specified process of cultural inheritance.
A simple population genetics calculation makes the distinction concrete. Let be an individual's number of copies of a focal allele, its contribution to reproduction, and . With unbiased Mendelian segregation and no other transmission change, the expected allele frequency after selection satisfies:Behaviour can change and thereby change the covariance, whether selection acts directly on a behavioural difference or on another trait whose consequences depend on behaviour. The equation does not imply that every correlation between a behaviour and reproductive success is genetic: common environment, age and condition can generate such correlations. A common garden experiment, a pedigree and measurements across environments help identify the inherited component.
Behaviour may expose a previously uncommon phenotype to selection. Foraging in a new habitat can make a different resource profitable, after which selection can refine feeding structures or behavioural response thresholds. Laboratory competitions between spadefoot tadpoles from populations differing in expression of a carnivore ecomorph found differences in resource capture and morphology consistent with adaptive refinement of a formerly plastic response. Expression without the inducing cue also supported genetic assimilation. [ref]
Plasticity-led evolution: evaluating the key prediction of frequency-dependent adaptation by Nicholas A. Levis and David W. Pfennig. laboratory competition and expression of the carnivore ecomorph in spadefoot tadpoles
This is a combination of behaviour and development, not proof that behaviour universally comes first. Genetic accommodation requires evolution of the response, rather than assuming that an acquired habit is copied directly into DNA.The opposite effect is also possible. Through phenotypic buffering, movement or flexible foraging may keep individuals in suitable conditions and reduce selection on physiological or morphological differences. Thus flexibility can promote occupation of a new environment, delay genetic divergence, or prevent local extinction. The sign depends on whether behaviour changes exposure to selection or allows animals to avoid it. A short-term adaptive response is consequently not evidence for a universal acceleration of evolutionary change.
At the boundary between populations and species, the behavioural effects are particularly direct. Assortative mating, host choice and learned signals can reduce gene flow. In the Big Bird lineage of Darwin's finches, field observations and genomic ancestry documented an immigrant Geospiza conirostris breeding with a resident Geospiza fortis and the formation of an endogamous descendant lineage. Song and distinctive morphology contributed to mating within that lineage. [ref]
Rapid hybrid speciation in Darwin's finches by Sangeet Lamichhaney and others. Science 359, pp. 224–228; immigrant ancestry, endogamy and the Big Bird lineage
The evidence links immigration, hybridization and mate recognition to rapid reproductive separation; it does not show that song alone caused isolation or establish the lineage's indefinite future persistence. Nevertheless, behaviour participates in an event conventionally classified as macroevolution.Over longer periods, these mechanisms can influence the rates of lineage origination and disappearance. Behaviour that permits colonization can expose populations to ecological opportunity and eventual adaptive radiation. Specialization in host use or mating signals can strengthen reproductive barriers. Conversely, costly reproductive competition or inflexible resource use can increase vulnerability. In fossil ostracods, greater sexual dimorphism was associated with greater estimated extinction risk, using carapace differences as a proxy for male reproductive investment. [ref]
High male sexual investment as a driver of extinction in fossil ostracods by Maria João Fernandes Martins and others. Nature 556, pp. 366–369; dimorphism as a proxy and fossil extinction estimates
This is unusually direct evidence about actual lineage loss, but an anatomical proxy does not preserve courtship or experimentally isolate sexual selection from every correlated attribute. It supports a possible connection rather than a general law that strong sexual selection causes extinction.Evaluating a claim about relative importance also requires correcting unequal evidence. Contemporary behavioural experiments can manipulate cues and follow reproduction, whereas most ancient behaviour must be inferred from traces, anatomy or living relatives. Living-only comparative studies omit extinct lineages and can confuse a trait associated with diversification with a trait that evolved after diversification. Comparisons should allow for shared ancestry, environmental opportunity and incomplete sampling, and distinguish an increased speciation rate from a reduced extinction rate. Counting observed behavioural changes over years against anatomical changes over millions of years is not a meaningful common denominator.
Behaviour has demonstrable population-level effects and plausible, sometimes well-documented effects on speciation and extinction. The stronger conclusion is that its evolutionary influence is context-dependent and transmitted across scales, while the evidence becomes harder to identify over deep time. A qualified statement about evidential accessibility is defensible; a general ranking of causal importance is not.