Functionally mediated cranial allometry evidenced in a genus of rock-wallabies.

In assessments of skeletal variation, allometry (disproportionate change of shape with size) is often corrected to examine size-independent variation for hypotheses relating to function. However, size-related trade-offs in functional demands may themselves be an underestimated driver of mammalian cranial diversity. Here, we use geometric morphometrics alongside dental measurements to assess craniodental allometry in the rock-wallaby genus Petrogale (all 17 species, 370 individuals). We identified functional aspects of evolutionary allometry that can be both extensions of, and correlated negatively with, static or ontogenetic allometric patterns. Regarding constraints, larger species tended to have relatively smaller braincases and more posterior orbits, the former of which might represent a constraint on jaw muscle anatomy. However, they also tended to have more anterior dentition and smaller posterior zygomatic arches, both of which support the hypothesis of relaxed bite force demands and accommodation of different selective pressures that favour facial elongation. By contrast, two dwarf species had stouter crania with divergent dental adaptations that together suggest increased relative bite force capacity. This likely allows them to feed on forage that is mechanically similar to that consumed by larger relatives. Our results highlight a need for nuanced considerations of allometric patterns in future research of mammalian cranial diversity.

Facing the facts: adaptive trade-offs along body size ranges determine mammalian craniofacial scaling.

The mammalian cranium (skull without lower jaw) is representative of mammalian diversity and is thus of particular interest to mammalian biologists across disciplines. One widely retrieved pattern accompanying mammalian cranial diversification is referred to as 'craniofacial evolutionary allometry' (CREA). This posits that adults of larger species, in a group of closely related mammals, tend to have relatively longer faces and smaller braincases. However, no process has been officially suggested to explain this pattern, there are many apparent exceptions, and its predictions potentially conflict with well-established biomechanical principles. Understanding the mechanisms behind CREA and causes for deviations from the pattern therefore has tremendous potential to explain allometry and diversification of the mammalian cranium. Here, we propose an amended framework to characterise the CREA pattern more clearly, in that 'longer faces' can arise through several kinds of evolutionary change, including elongation of the rostrum, retraction of the jaw muscles, or a more narrow or shallow skull, which all result in a generalised gracilisation of the facial skeleton with increased size. We define a standardised workflow to test for the presence of the pattern, using allometric shape predictions derived from geometric morphometrics analysis, and apply this to 22 mammalian families including marsupials, rabbits, rodents, bats, carnivores, antelopes, and whales. Our results show that increasing facial gracility with size is common, but not necessarily as ubiquitous as previously suggested. To address the mechanistic basis for this variation, we then review cranial adaptations for harder biting. These dictate that a more gracile cranium in larger species must represent a structural sacrifice in the ability to produce or withstand harder bites, relative to size. This leads us to propose that facial gracilisation in larger species is often a product of bite force allometry and phylogenetic niche conservatism, where more closely related species tend to exhibit more similar feeding ecology and biting behaviours and, therefore, absolute (size-independent) bite force requirements. Since larger species can produce the same absolute bite forces as smaller species with less effort, we propose that relaxed bite force demands can permit facial gracility in response to bone optimisation and alternative selection pressures. Thus, mammalian facial scaling represents an adaptive by-product of the shifting importance of selective pressures occurring with increased size. A reverse pattern of facial 'shortening' can accordingly also be found, and is retrieved in several cases here, where larger species incorporate novel feeding behaviours involving greater bite forces. We discuss multiple exceptions to a bite force-mediated influence on facial proportions across mammals which lead us to argue that ecomorphological specialisation of the cranium is likely to be the primary driver of facial scaling patterns, with some developmental constraints as possible secondary factors. A potential for larger species to have a wider range of cranial functions when less constrained by bite force demands might also explain why selection for larger sizes seems to be prevalent in some mammalian clades. The interplay between adaptation and constraint across size ranges thus presents an interesting consideration for a mechanistically grounded investigation of mammalian cranial allometry.

Differential developmental rates and demographics in Red Kangaroo (Osphranter rufus) populations separated by the dingo barrier fence.

Decommissioning the dingo barrier fence has been suggested to reduce destructive dingo control and encourage a free transfer of biota between environments in Australia. Yet the potential impacts that over a century of predator exclusion might have had on the population dynamics and developmental biology of prey populations has not been assessed. We here combine demographic data and both linear and geometric morphometrics to assess differences in populations among 166 red kangaroos (Osphranter rufus)-a primary prey species of the dingo-from two isolated populations on either side of the fence. We also quantified the differences in aboveground vegetation biomass for the last 10 years on either side of the fence. We found that the age structure and growth patterns, but not cranial shape, differed between the two kangaroo populations. In the population living with a higher density of dingoes, there were relatively fewer females and juveniles. These individuals were larger for a given age, despite what seems to be lower vegetation biomass. However, how much of this biomass represented kangaroo forage is uncertain and requires further on-site assessments. We also identified unexpected differences in the ontogenetic trajectories in relative pes length between the sexes for the whole sample, possibly associated with male competition or differential weight-bearing mechanics. We discuss potential mechanisms behind our findings and suggest that the impacts of contrasting predation pressures across the fence, for red kangaroos and other species, merit further investigation.

Multiple modes of inference reveal less phylogenetic signal in marsupial basicranial shape compared with the rest of the cranium.

Incorporating morphological data into modern phylogenies allows integration of fossil evidence, facilitating divergence dating and macroevolutionary inferences. Improvements in the phylogenetic utility of morphological data have been sought via Procrustes-based geometric morphometrics (GMM), but with mixed success and little clarity over what anatomical areas are most suitable. Here, we assess GMM-based phylogenetic reconstructions in a heavily sampled source of discrete characters for mammalian phylogenetics-the basicranium-in 57 species of marsupial mammals, compared with the remainder of the cranium. We show less phylogenetic signal in the basicranium compared with a 'Rest of Cranium' partition, using diverse metrics of phylogenetic signal (Kmult, phylogenetically aligned principal components analysis, comparisons of UPGMA/neighbour-joining/parsimony trees and cophenetic distances to a reference phylogeny) for scaled, Procrustes-aligned landmarks and allometry-corrected residuals. Surprisingly, a similar pattern emerged from parsimony-based analyses of discrete cranial characters. The consistent results across methods suggest that easily computed metrics such as Kmult can provide good guidance on phylogenetic information in a landmarking configuration. In addition, GMM data may be less informative for intricate but conservative anatomical regions such as the basicranium, while better-but not necessarily novel-phylogenetic information can be expected for broadly characterized shapes such as entire bones. This article is part of the theme issue 'The mammalian skull: development, structure and function'.

Development of the ethmoid in a wallaby and implications for the homology of turbinal elements in marsupials.

The homologies of the turbinals (scroll bones) of the ethmoid are not well understood, including the potential implication for understanding mammalian phylogeny. Here we examine the postnatal development of this anatomical system in a marsupial mammal because previous work has shown that the adult pattern of five endoturbinals (ethmoturbinals) and two ectoturbinals (frontoturbinals) is conserved. Furthermore, marsupial phylogeny is fairly well resolved and provides a solid evolutionary framework for examining turbinal homologies. In this study, we documented the development of the ethmoid of the tammar wallaby, Notamacropus eugenii, using histology and computed tomography imagery of a growth series of pouch young. The pattern of development of the turbinal elements in the wallaby was compared to that in didelphids, as described in previous work. We found that four ethmoturbinals initially develop, followed later in development by an interturbinal; these five elements then develop into the bony endoturbinals found in adults. These data support the idea that endoturbinal III, derived from an interturbinal, has a distinctive development pattern from the other endoturbinals. This is consistent with what is seen in the didelphid marsupials, Caluromys philander and Monodelphis domestica, suggesting this is a common developmental pattern for marsupials. This article is part of the theme issue 'The mammalian skull: development, structure and function'.

The relative performance of geometric morphometrics and linear-based methods in the taxonomic resolution of a mammalian species complex.

Morphology-based taxonomic research frequently applies linear morphometrics (LMM) in skulls to quantify species distinctions. The choice of which measurements to collect generally relies on the expertise of the investigators or a set of standard measurements, but this practice may ignore less obvious or common discriminatory characteristics. In addition, taxonomic analyses often ignore the potential for subgroups of an otherwise cohesive population to differ in shape purely due to size differences (or allometry). Geometric morphometrics (GMM) is more complicated as an acquisition technique but can offer a more holistic characterization of shape and provides a rigorous toolkit for accounting for allometry. In this study, we used linear discriminant analysis (LDA) to assess the discriminatory performance of four published LMM protocols and a 3D GMM dataset for three clades of antechinus known to differ subtly in shape. We assessed discrimination of raw data (which are frequently used by taxonomists); data with isometry (i.e., overall size) removed; and data after allometric correction (i.e., with nonuniform effects of size removed). When we visualized the principal component analysis (PCA) plots, we found that group discrimination among raw data was high for LMM. However, LMM datasets may inflate PC variance accounted in the first two PCs, relative to GMM. GMM discriminated groups better after isometry and allometry were removed in both PCA and LDA. Although LMM can be a powerful tool to discriminate taxonomic groups, we show that there is substantial risk that this discrimination comes from variation in size, rather than shape. This suggests that taxonomic measurement protocols might benefit from GMM-based pilot studies, because this offers the option of differentiating allometric and nonallometric shape differences between species, which can then inform on the development of the easier-to-apply LMM protocols.

Cranial anatomy of the mekosuchine crocodylian Trilophosuchus rackhami Willis, 1993.

One of the best-preserved crocodylian fossil specimens from the Cenozoic of Australia is the holotype of the mekosuchine Trilophosuchus rackhami, from the middle Miocene (13.56 ± 0.67 Ma) Ringtail Site at Riversleigh, northwestern Queensland. Although lacking most of the snout, the holotype skull of T. rackhami (QMF16856) has an exceptionally well-preserved cranium. Micro-CT scanning of the holotype has allowed for all the preserved cranial bones to be digitally disarticulated, facilitating an unprecedented insight into the cranial anatomy of not just T. rackhami, but any mekosuchine. Trilophosuchus rackhami was a small-bodied crocodylian and one of the most morphologically distinct mekosuchines, characterized by a unique combination of cranial characteristics several of which are exclusive to the species. Fossil material that is definitively referrable to the species T. rackhami is currently known solely from the middle Miocene Ringtail Site. However, an isolated parietal from Hiatus Site at Riversleigh demonstrates that Trilophosuchus also occurred during the late Oligocene (~25 Ma), extending the range of the genus by more than 10 million years. The new description of T. rackhami also allowed for a reevaluation of its phylogenetic relationships. Our results reaffirm the placement of T. rackhami as a member of Mekosuchinae within the subclade Mekosuchini. In all analyses, Mekosuchinae was consistently found to be monophyletic and part of the larger crocodylian clade Longirostres. However, the assignment of Mekosuchinae as a subset of Crocodylidae is brought into question, suggesting that the status of Mekosuchinae as a subfamily should be reconsidered.

Morphometric classification of kangaroo bones reveals paleoecological change in northwest Australia during the terminal Pleistocene.

Specimen identification is the backbone of archeozoological research. The challenge of differentiating postcranial skeletal elements of closely related wild animals in biodiverse regions can prove a barrier to understanding past human foraging behaviours. Morphometrics are increasingly being employed to classify paleozoological animal remains, however, the potential of these methods to discriminate between wild animal groups has yet to be fully realised. Here we demonstrate the applicability of a traditional morphometric approach to taxonomically classify foot and ankle bones of kangaroos, a large and highly diverse marsupial family. Using multiple discriminant analysis, we classify archaeological specimens from Boodie Cave, in northwest Australia and identify the presence of two locally extinct macropod species during the terminal Pleistocene. The appearance of the banded hare-wallaby and northern nail-tail wallaby in the Pilbara region at this time provides independent evidence of the ecological and human responses to a changing climate at the end of the last Ice Age. Traditional morphometrics provides an accessible, inexpensive, and non-destructive tool for paleozoological specimen classification and has substantial potential for applications to other diverse wild faunas.

Not like night and day: the nocturnal letter-winged kite does not differ from diurnal congeners in orbit or endocast morphology.

Nocturnal birds display diverse adaptations of the visual system to low-light conditions. The skulls of birds reflect many of these and are used increasingly to infer nocturnality in extinct species. However, it is unclear how reliable such assessments are, particularly in cases of recent evolutionary transitions to nocturnality. Here, we investigate a case of recently evolved nocturnality in the world's only nocturnal hawk, the letter-winged kite Elanus scriptus. We employed phylogenetically informed analyses of orbit, optic foramen and endocast measurements from three-dimensional reconstructions of micro-computed tomography scanned skulls of the letter-winged kite, two congeners, and 13 other accipitrid and falconid raptors. Contrary to earlier suggestions, the letter-winged kite was not unique in any of our metrics. However, all species of Elanus have significantly higher ratios of orbit versus optic foramen diameter, suggesting high visual sensitivity at the expense of acuity. In addition, visual system morphology varies greatly across accipitrid species, likely reflecting hunting styles. Overall, our results suggest that the transition to nocturnality can occur rapidly and without changes to key hard-tissue indicators of vision, but also that hard-tissue anatomy of the visual system may provide a means of inferring a range of raptor behaviours, well beyond nocturnality.

A fairer way to compare researchers at any career stage and in any discipline using open-access citation data.

The pursuit of simple, yet fair, unbiased, and objective measures of researcher performance has occupied bibliometricians and the research community as a whole for decades. However, despite the diversity of available metrics, most are either complex to calculate or not readily applied in the most common assessment exercises (e.g., grant assessment, job applications). The ubiquity of metrics like the h-index (h papers with at least h citations) and its time-corrected variant, the m-quotient (h-index ÷ number of years publishing) therefore reflect the ease of use rather than their capacity to differentiate researchers fairly among disciplines, career stage, or gender. We address this problem here by defining an easily calculated index based on publicly available citation data (Google Scholar) that corrects for most biases and allows assessors to compare researchers at any stage of their career and from any discipline on the same scale. Our ε'-index violates fewer statistical assumptions relative to other metrics when comparing groups of researchers, and can be easily modified to remove inherent gender biases in citation data. We demonstrate the utility of the ε'-index using a sample of 480 researchers with Google Scholar profiles, stratified evenly into eight disciplines (archaeology, chemistry, ecology, evolution and development, geology, microbiology, ophthalmology, palaeontology), three career stages (early, mid-, late-career), and two genders. We advocate the use of the ε'-index whenever assessors must compare research performance among researchers of different backgrounds, but emphasize that no single index should be used exclusively to rank researcher capability.

First record of a tomistomine crocodylian from Australia.

Based on the known fossil record, the majority of crocodylians from the Cenozoic Era of Australia are referred to the extinct clade Mekosuchinae. The only extant crocodylians in Australia are two species of Crocodylus. Hence, the viewpoint that Crocodylus and mekosuchines have been the only crocodylians inhabiting Australia during the Cenozoic has remained largely undisputed. Herein we describe Australia's first tomistomine crocodylian, Gunggamarandu maunala gen. et sp. nov., thus challenging the notion of mekosuchine dominance during most of the Cenozoic. The holotype specimen of Gunggamarandu maunala derives from the Pliocene or Pleistocene of south-eastern Queensland, marking the southern-most global record for Tomistominae. Gunggamarandu maunala is known from a large, incomplete cranium that possesses a unique combination of features that distinguishes it from other crocodylians. Phylogenetic analyses place Gunggamarandu in a basal position within Tomistominae, specifically as a sister taxon to Dollosuchoides from the Eocene of Europe. These results hint at a potential ghost lineage between European and Australian tomistomines going back more than 50 million years. The cranial proportions of the Gunggamarandu maunala holotype specimen indicate it is the largest crocodyliform yet discovered from Australia.

Evolution: Bend it like basal synapsids.

Mammals can amplify their strides through unique up-and-down spinal movements. This ability was long considered to have evolved from lizard-like ancestors with spines moving sideways. A new study now suggests that, instead, it derived from an extinct, previously unknown spinal form.

Relative demographic susceptibility does not explain the extinction chronology of Sahul's megafauna.

The causes of Sahul's megafauna extinctions remain uncertain, although several interacting factors were likely responsible. To examine the relative support for hypotheses regarding plausible ecological mechanisms underlying these extinctions, we constructed the first stochastic, age-structured models for 13 extinct megafauna species from five functional/taxonomic groups, as well as 8 extant species within these groups for comparison. Perturbing specific demographic rates individually, we tested which species were more demographically susceptible to extinction, and then compared these relative sensitivities to the fossil-derived extinction chronology. Our models show that the macropodiformes were the least demographically susceptible to extinction, followed by carnivores, monotremes, vombatiform herbivores, and large birds. Five of the eight extant species were as or more susceptible than the extinct species. There was no clear relationship between extinction susceptibility and the extinction chronology for any perturbation scenario, while body mass and generation length explained much of the variation in relative risk. Our results reveal that the actual mechanisms leading to the observed extinction chronology were unlikely related to variation in demographic susceptibility per se, but were possibly driven instead by finer-scale variation in climate change and/or human prey choice and relative hunting success.

Testing hypotheses of marsupial brain size variation using phylogenetic multiple imputations and a Bayesian comparative framework.

Considerable controversy exists about which hypotheses and variables best explain mammalian brain size variation. We use a new, high-coverage dataset of marsupial brain and body sizes, and the first phylogenetically imputed full datasets of 16 predictor variables, to model the prevalent hypotheses explaining brain size evolution using phylogenetically corrected Bayesian generalized linear mixed-effects modelling. Despite this comprehensive analysis, litter size emerges as the only significant predictor. Marsupials differ from the more frequently studied placentals in displaying a much lower diversity of reproductive traits, which are known to interact extensively with many behavioural and ecological predictors of brain size. Our results therefore suggest that studies of relative brain size evolution in placental mammals may require targeted co-analysis or adjustment of reproductive parameters like litter size, weaning age or gestation length. This supports suggestions that significant associations between behavioural or ecological variables with relative brain size may be due to a confounding influence of the extensive reproductive diversity of placental mammals.

Ontogenetic origins of cranial convergence between the extinct marsupial thylacine and placental gray wolf.

Phenotypic convergence, describing the independent evolution of similar characteristics, offers unique insights into how natural selection influences developmental and molecular processes to generate shared adaptations. The extinct marsupial thylacine and placental gray wolf represent one of the most extraordinary cases of convergent evolution in mammals, sharing striking cranial similarities despite 160 million years of independent evolution. We digitally reconstructed their cranial ontogeny from birth to adulthood to examine how and when convergence arises through patterns of allometry, mosaicism, modularity, and integration. We find the thylacine and wolf crania develop along nearly parallel growth trajectories, despite lineage-specific constraints and heterochrony in timing of ossification. These constraints were found to enforce distinct cranial modularity and integration patterns during development, which were unable to explain their adult convergence. Instead, we identify a developmental origin for their convergent cranial morphologies through patterns of mosaic evolution, occurring within bone groups sharing conserved embryonic tissue origins. Interestingly, these patterns are accompanied by homoplasy in gene regulatory networks associated with neural crest cells, critical for skull patterning. Together, our findings establish empirical links between adaptive phenotypic and genotypic convergence and provides a digital resource for further investigations into the developmental basis of mammalian evolution.

Global elongation and high shape flexibility as an evolutionary hypothesis of accommodating mammalian brains into skulls.

Little is known about how the large brains of mammals are accommodated into the dazzling diversity of their skulls. It has been suggested that brain shape is influenced by relative brain size, that it evolves or develops according to extrinsic or intrinsic mechanical constraints, and that its shape can provide insights into its proportions and function. Here, we characterize the shape variation among 84 marsupial cranial endocasts of 57 species including fossils, using three-dimensional geometric morphometrics and virtual dissections. Statistical shape analysis revealed four main patterns: over half of endocast shape variation ranges from elongate and straight to globular and inclined; little allometric variation with respect to centroid size, and none for relative volume; no association between locomotion and endocast shape; limited association between endocast shape and previously published histological cortex volumes. Fossil species tend to have smaller cerebral hemispheres. We find divergent endocast shapes in closely related species and within species, and diverse morphologies superimposed over the main variation. An evolutionarily and individually malleable brain with a fundamental tendency to arrange into a spectrum of elongate-to-globular shapes-possibly mostly independent of brain function-may explain the accommodation of brains within the enormous diversity of mammalian skull form.

Ovotestes suggest cryptic genetic influence in a reptile model for temperature-dependent sex determination.

Sex determination and differentiation in reptiles is complex. Temperature-dependent sex determination (TSD), genetic sex determination (GSD) and the interaction of both environmental and genetic cues (sex reversal) can drive the development of sexual phenotypes. The jacky dragon (Amphibolurus muricatus) is an attractive model species for the study of gene-environment interactions because it displays a form of Type II TSD, where female-biased sex ratios are observed at extreme incubation temperatures and approximately 50 : 50 sex ratios occur at intermediate temperatures. This response to temperature has been proposed to occur due to underlying sex determining loci, the influence of which is overridden at extreme temperatures. Thus, sex reversal at extreme temperatures is predicted to produce the female-biased sex ratios observed in A. muricatus. The occurrence of ovotestes during development is a cellular marker of temperature sex reversal in a closely related species Pogona vitticeps. Here, we present the first developmental data for A. muricatus, and show that ovotestes occur at frequencies consistent with a mode of sex determination that is intermediate between GSD and TSD. This is the first evidence suggestive of underlying unidentified sex determining loci in a species that has long been used as a model for TSD.

Australian Rodents Reveal Conserved Cranial Evolutionary Allometry across 10 Million Years of Murid Evolution.

AbstractAmong vertebrates, placental mammals are particularly variable in the covariance between cranial shape and body size (allometry), with rodents being a major exception. Australian murid rodents allow an assessment of the cause of this anomaly because they radiated on an ecologically diverse continent notably lacking other terrestrial placentals. Here, we use 3D geometric morphometrics to quantify species-level and evolutionary allometries in 38 species (317 crania) from all Australian murid genera. We ask whether ecological opportunity resulted in greater allometric diversity compared with other rodents or whether conserved allometry suggests intrinsic constraints and/or stabilizing selection. We also assess whether cranial shape variation follows the proposed rule of craniofacial evolutionary allometry (CREA), whereby larger species have relatively longer snouts and smaller braincases. To ensure we could differentiate parallel versus nonparallel species-level allometric slopes, we compared the slopes of rarefied samples across all clades. We found exceedingly conserved allometry and CREA-like patterns across the 10-million-year split between Mus and Australian murids. This could support both intrinsic-constraint and stabilizing-selection hypotheses for conserved allometry. Large-bodied frugivores evolved faster than other species along the allometric trajectory, which could suggest stabilizing selection on the shape of the masticatory apparatus as body size changes.

Skull shape of a widely distributed, endangered marsupial reveals little evidence of local adaptation between fragmented populations.

The biogeographic distribution of diversity among populations of threatened mammalian species is generally investigated using population genetics. However, intraspecific phenotypic diversity is rarely assessed beyond taxonomy-focused linear measurements or qualitative descriptions. Here, we use a technique widely used in the evolutionary sciences-geometric morphometrics-to characterize shape diversity in the skull of an endangered marsupial, the northern quoll, across its 5,000 km distribution range along Northern Australia. Skull shape is a proxy for feeding, behavior, and phenotypic differentiation, allowing us to ask whether populations can be distinguished and whether patterns of variation indicate adaptability to changing environmental conditions. We analyzed skull shape in 101 individuals across four mainland populations and several islands. We assessed the contribution of population, size, sex, rainfall, temperature, and geography to skull shape variation using principal component analysis, Procrustes ANOVA, and variation partitioning analyses. The populations harbor similar amounts of broadly overlapping skull shape variation, with relatively low geographic effects. Size predicted skull shape best, coinciding with braincase size variation and differences in zygomatic arches. Size-adjusted differences in populations explained less variation with far smaller effect sizes, relating to changes in the insertion areas of masticatory muscles, as well as the upper muzzle and incisor region. Climatic and geographic variables contributed little. Strikingly, the vast majority of shape variation-76%-remained unexplained. Our results suggest a uniform intraspecific scope for shape variation, possibly due to allometric constraints or phenotypic plasticity beyond the relatively strong allometric effect. The lack of local adaptation indicates that cross-breeding between populations will not reduce local morphological skull (and probably general musculoskeletal) adaptation because none exists. However, the potential for heritable morphological variation (e.g., specialization to local diets) seems exceedingly limited. We conclude that 3D geometric morphometrics can provide a comprehensive, statistically rigorous phenomic contribution to genetic-based conservation studies.

Shifting spaces: Which disparity or dissimilarity measurement best summarize occupancy in multidimensional spaces?

Multidimensional analysis of traits are now common in ecology and evolution and are based on trait spaces in which each dimension summarizes the observed trait combination (a morphospace or an ecospace). Observations of interest will typically occupy a subset of this space, and researchers will calculate one or more measures to quantify how organisms inhabit that space. In macroevolution and ecology, these measures called disparity or dissimilarity metrics are generalized as space occupancy measures. Researchers use these measures to investigate how space occupancy changes through time, in relation to other groups of organisms, or in response to global environmental changes. However, the mathematical and biological meaning of most space occupancy measures is vague with the majority of widely used measures lacking formal description. Here, we propose a broad classification of space occupancy measures into three categories that capture changes in size, density, or position. We study the behavior of 25 measures to changes in trait space size, density, and position on simulated and empirical datasets. We find that no measure describes all of trait space aspects but that some are better at capturing certain aspects. Our results confirm the three broad categories (size, density, and position) and allow us to relate changes in any of these categories to biological phenomena. Because the choice of space occupancy measures is specific to the data and question, we introduced https://tguillerme.shinyapps.io/moms/moms, a tool to both visualize and capture changes in space occupancy for any measurement. https://tguillerme.shinyapps.io/moms/moms is designed to help workers choose the right space occupancy measures, given the properties of their trait space and their biological question. By providing guidelines and common vocabulary for space occupancy analysis, we hope to help bridging the gap in multidimensional research between ecology and evolution.