A New Small-Bodied Species of <Emphasis Type="Italic">Palaeonictis</Emphasis> (Creodonta,Oxyaenidae) from the Paleocene-Eocene Thermal Maximum

Oxyaenid creodonts are extinct carnivorous mammals known from the Paleogene of North America, Europe, and Asia. The genus Palaeonictis is represented by three species that together span the late Paleocene to early Eocene of North America, and at least one species from the early Eocene of Europe. Previously, only a single trigonid of Palaeonictis was known from the interval encompassing the Paleocene-Eocene Thermal Maximum (PETM) in North America. We describe Palaeonictis wingi sp. nov. from the PETM in the Cabin Fork drainage, southeastern Bighorn Basin, Wyoming, based on associated right and left dentaries with P₂-M₂. Palaeonictis wingi sp. nov. is substantially smaller than the other North American congeners, making it similar in size to P. gigantea from the earliest Eocene of Europe and the previously described PETM specimen. We suggest that a form similar to the large-bodied late Paleocene P. peloria from North America gave rise to two smaller species in the earliest Eocene of North America (P. wingi) and Europe (P. gigantea). Palaeonictis wingi may have given rise to P. occidentalis following the PETM in North America. Dispersal of Palaeonictis to Europe coincided with rapid global warming of 5–10°C and related geographic range shifts in plants and other animals during the PETM. It has been suggested that certain mammalian lineages decreased in body size during the PETM, possibly in response to elevated temperature and/or higher CO₂ levels. Results from a dietary analysis of Palaeonictis indicate that it was an omnivore that primarily consumed meat. This suggests that the decreased nutritious quality of vegetation caused by increased CO₂ levels was not the direct contributing factor that caused body size reduction of this lineage during the PETM. Other selective pressures such as temperature, aridity, and prey size may have also contributed to the smaller body size of carnivorous mammals during this interval, although the presence of smaller species could also be explained by latitudinal range shifts of mammals during the PETM.     

New specimens of the mesonychid Dissacus praenuntius from the early Eocene of Wyoming and evaluation of body size through the PETM in North America

The Mesonychia is a group of archaic carnivorous mammals of uncertain phylogenetic affinities with a Holarctic distribution during the Paleogene. Intensive fossil collecting efforts in the Bighorn Basin, Wyoming, have resulted in recovery of the largest sample and most complete specimens yet known of the mesonychid Dissacus praenuntius from the second biozone of the Wasatchian North American Land Mammal Age (Wa-0). The Wa-0 biozone corresponds to the body of the Paleocene-Eocene Thermal Maximum (PETM), a brief but intense global warming event that occurred ~56 myr ago that significantly impacted terrestrial mammal faunas, including dwarfing in many mammal lineages. To evaluate the potential response of this lineage to climate change, we compared the PETM sample of D. praenuntius with those recovered from just before the PETM in the last biozone of the Clarkforkian North American Land Mammal Age (Cf-3) and just after the PETM in the Wa-1 biozone. While the sample size is still too small to say with certainty, tooth size (as a proxy for body weight) of D. praenuntius appears to be smaller during the late PETM than during either the pre-PETM Cf-3, or post-PETM Wa-1 biozones, suggesting the possibility of a muted dwarfing response to the PETM. However, the pattern observed for D. praenuntius differs from that of many other PETM mammals, as the shift to smaller body size is less pronounced and may have only occurred in late Wa-0.     

Invading Europe: did climate or geography trigger early Eocene primate dispersals?

The Palaeocene-Eocene transition is characterized by a significant turnover of mammalian taxa in the fossil record of the northern continents, and primates are among the groups that make their first appearance at this time. One of the many questions that remain to be answered with regard to the earliest evolution of primates is the reason for their sudden and virtually simultaneous appearance in the fossil records of Asia, Europe and North America. The most obvious environmental correlate of the Palaeocene-Eocene transition is a sharp but relatively short-lived warming event leading up to the Palaeocene-Eocene thermal maximum (PETM) and evidenced in the stratigraphic record by a negative delta(13)C excursion. It remains unclear, however, whether or how this warming event may have influenced Palaeocene-Eocene faunal turnovers. This paper explores the hypothesis that environmental changes associated with the PETM facilitated an invasion of Western Europe by primates by comparing the ecological structure of local mammalian fauna immediately before and following the Palaeocene-Eocene transition. The results suggest that changes to the ecological profile of local mammalian fauna were relatively small and did not favour an invasion by primates, although a major uncertainty remains with respect to the availability of arboreal niches. At present it seems more likely that the invasion of western Europe by primates was due to the breakdown of one or more dispersal barriers close to the end of the Palaeocene.     

THE MICRO AND MACRO IN BODY SIZE EVOLUTION

The diversity of body sizes of organisms has traditionally been explained in terms of microevolutionary processes: natural selection owing to differential fitness of individual organisms, or to macroevolutionary processes: species selection owing to the differential proliferation of phylogenetic lineages. Data for terrestrial mammals and birds indicate that even on a logarithmic scale frequency distributions of body mass among species are significantly skewed towards larger sizes. We used simulation models to evaluate the extent to which macro- and microevolutionary processes are sufficient to explain these distributions. Simulations of a purely cladogenetic process with no bias in extinction or speciation rates for different body sizes did not produce skewed log body mass distributions. Simulations that included size-biased extinction rates, especially those that incorporated anagenetic size change within species between speciation and extinction events, regularly produced skewed distributions. We conclude that although cladogenetic processes probably play a significant role in body size evolution, there must also be a significant anagenetic component. The regular variation in the form of mammalian body size distributions among different-sized islands and continents suggests that environmental conditions, operating through both macro- and microevolutionary processes, determine to a large extent the diversification of body sizes within faunas. Macroevolution is not decoupled from microevolution.     

Deep phylogenetic divergence between Scolytoplatypus and Remansus,a new genus of Scolytoplatypodini from Madagascar (Coleoptera,Curculionidae, Scolytinae)

Scolytoplatypodini Blandford is a monotypic tribe of ambrosia beetles found in Asia, Madagascar and Africa. Only three species are currently known from Madagascar and four additional species are here described as new to science. Phylogenetic analyses of morphological and molecular data revealed that four of the seven endemic species are deeply separated from all other species by genetic and distinct morphological characters and therefore placed in a new genus Remansus Jordal. The split between this ancient lineage and Scolytoplatypus Schaufuss was estimated to approximate Palaeocene age (63 Ma), extending the minimum age of ambrosia feeding for this tribe to the beginning of the Palaeocene‒Eocene thermal maximum (PETM). In addition to the ancient origin of Remansus in Madagascar during the Palaeocene, a second origin occurred in Scolytoplatypus no more than 13 Ma. A geographical origin of the latter in South-Eastern Africa was unequivocally inferred from the phylogenies.     

COMPARISONS BETWEEN PALAEOCENE–EOCENE PARATROPICAL SWAMP AND MARGINAL MARINE POLLEN FLORAS FROM ALABAMA AND MISSISSIPPI, USA

Abstract:  Climate warming at the Palaeocene/Eocene boundary ( c . 55.8 Ma) had significant permanent affects on paratropical and warm-adapted vegetation types. Pollen and spore records which document vegetation turnover from the eastern US Gulf Coast have all been taken from sediments of marginal marine depositional environments. Pollen and spores (sporomorphs) are preserved excellently in these marginal marine depositional environments but these assemblages contain grains transported from many different vegetation types and over huge geographic distances. Currently it is unclear whether the turnover from important paratropical areas like the US Gulf Coast is a reflection on actual vegetation change in the local region or from source areas far away in the continental interior. Sporomorph data from 20 former swamps (lignites) from the Nanafalia, Tuscahoma and Hatchetigbee formations in Mississippi and Alabama, USA, are used to test the fidelity of the marine sporomorph record across the Palaeocene–Eocene transition. Data show that extinction is noted in the swamp record (≥7 per cent of Palaeocene taxa) and that swamps were susceptible to immigration in the Early Eocene with the first occurrences of Brosipollis spp. (Burseraceae), Dicolpopollis spp. (Palmae), Nuxpollenites psilatus (Loranthaceae) and Platycarya spp. (Juglandaceae). Swamps have higher within-sample diversity in the Eocene but higher among-sample diversity in latest Palaeocene–earliest Eocene samples, which parallels exactly diversity trends estimated from marine sporomorph assemblages. Palms also increase in abundance in the Eocene. The swamp data demonstrate that the flora growing in these ancient paratropical forests was diverse ( c. 120 taxonomic groups) but incorporated an unusual admixture of plants with modern tropical affinities together with those that now live in modern temperate to subtropical North America.     

Extinction and replacement among predatory mammals in the North American late Eocene and Oligocene: Tracking a paleoguild over twelve million years

Changes in taxonomic and morphologic diversity within the paleoguild of predatory mammals were explored within a mammalian chronofauna spanning a twelve million year interval, from the latest Eocene to the end of the Oligocene of North America (36–24 ma). The timespan encompassed a modest extinction event among terrestrial mammals (circa 34 ma) followed by a period of relative stability. Morphological diversity was assessed with estimates of body mass, relative tooth size, and tooth shape. Principal component, nearest‐neighbor, and minimum‐spanning‐tree analyses were used to compare morphological diversity and species packing within predator paleoguilds in the mid‐Chadronian (37–34 ma), Orellan (34–32), Whitneyan (32–29.5), and early Arikareean (29.5–24) land mammal ages. Species richness of predators throughout the interval was relatively constant, fluctuating between 15 and 18 total taxa. Moreover, despite significant differences in taxonomic composition and a modest extinction event among terrestrial mammals, morphological diversity within the paleoguild was very similar in the Chadronian and Orellan. In the Whitneyan and especially the early Arikareean, the diversity of feeding adaptations among species declined slightly, largely due to the loss of several highly specialized meat‐eaters (creodonts, nimravids) and the addition of small omnivores (canids).     

Fossil evidence of interactions between plants and plant-eating mammals.

We document changes in mammalian dietary and foraging locomotor adaptation, and appearances and developments of angiosperm fruiting strategies and vegetation types since the late Cretaceous in the Euramerican region, and to some extent in low latitude Africa. These changes suggest: (i): an expansion in the exploitation of dry fruits and seeds by mammals on the ground as well as in the trees after the terminal Cretaceous dinosaur extinction; (ii) a relation between large nuts and rodents, which appear in the late Palaeocene and radiate in the late Eocene; (iii) a relation between primates and fleshy fruits established in the early-Middle Eocene when tropical forests reached their maximum latitudinal extent; (iv) a hiatus of several million years in the vertebrate exploitation of leaves after dinosaur extinction and before the first few mammalian herbivores in the Middle Palaeocene, followed by an expansion in the late Eocene when climates cooled and more open vegetation became established.     

Cenozoic climate change and diversification on the continental shelf and slope: evolution of gastropod diversity in the family Solariellidae (Trochoidea)

Recent expeditions have revealed high levels of biodiversity in the tropical deep‐sea, yet little is known about the age or origin of this biodiversity, and large‐scale molecular studies are still few in number. In this study, we had access to the largest number of solariellid gastropods ever collected for molecular studies, including many rare and unusual taxa. We used a Bayesian chronogram of these deep‐sea gastropods (1) to test the hypothesis that deep‐water communities arose onshore, (2) to determine whether Antarctica acted as a source of diversity for deep‐water communities elsewhere and (3) to determine how factors like global climate change have affected evolution on the continental slope. We show that although fossil data suggest that solariellid gastropods likely arose in a shallow, tropical environment, interpretation of the molecular data is equivocal with respect to the origin of the group. On the other hand, the molecular data clearly show that Antarctic species sampled represent a recent invasion, rather than a relictual ancestral lineage. We also show that an abrupt period of global warming during the Palaeocene Eocene Thermal Maximum (PETM) leaves no molecular record of change in diversification rate in solariellids and that the group radiated before the PETM. Conversely, there is a substantial, although not significant increase in the rate of diversification of a major clade approximately 33.7 Mya, coinciding with a period of global cooling at the Eocene–Oligocene transition. Increased nutrients made available by contemporaneous changes to erosion, ocean circulation, tectonic events and upwelling may explain increased diversification, suggesting that food availability may have been a factor limiting exploitation of deep‐sea habitats. Tectonic events that shaped diversification in reef‐associated taxa and deep‐water squat lobsters in central Indo‐West Pacific were also probably important in the evolution of solariellids during the Oligo‐Miocene.     

Winter filtering, immigrant selection and species composition of insular mammals of Lake Huron

Patterns in species composition of small mammals on islands of northwestern Lake Huron, USA were studied to test the hypothesis that insular community structure was strongly influenced by immigrant selection, l e, selection for the more vagile species Of 17 insectivores and rodents in the mainland pool nine were detected on at least one of the 25 islands studied These species comprised a biased subset of the mainland pool, biased in favor of species with relatively good winter immigration abilities The five true hibernators were absent from all islands studied and the two winter-inactive species occurred on just two islands Frequency of insular occurrence increased with level of winter activity (hibernator vs winter-inactive vs winter-active) and with body size (p < 0.02)
The winter filtering effect remains one of the best documented forms of immigrant selection In general however, insular community structure is influenced by the combined effects of selective extinction as well as selective immigration Common insular species are those which combine relatively high immigration abilities with low resource requirements     

‘On being the right size’ – Do aliens follow the rules?

Aim

To assess whether mammalian species introduced onto islands across the globe have evolved to exhibit body size patterns consistent with the ‘island rule,’, and to test an ecological explanation for body size evolution of insular mammals.

Location

Islands worldwide.

Methods

We assembled data on body mass, geographical characteristics (latitude, maximum elevation) and ecological communities (number of mammalian competitors, predators and prey) for 385 introduced populations across 285 islands, comprising 56 species of extant, non‐volant mammals. We used linear regression, ANCOVA and regression tree analyses to test whether introduced populations of mammals exhibit the island rule pattern, whether the degree of body size change increased with time in isolation and whether residual variation about the general trend can be attributed to the geographical and ecological characteristics of the islands.

Results

Introduced populations follow the predicted island rule trend, with body size shifts more pronounced for populations with greater residence times on the islands. Small mammals evolved to larger body sizes in lower latitudes and on islands with limited topographic relief. Consistent with our hypothesis on the ecology of evolution, body size of insular introduced populations was influenced by co‐occurring species of mammalian competitors, predators and prey.

Conclusion

The island rule is a pervasive pattern, exhibited across a broad span of geographical regions, taxa, time periods and, as evidenced here, for introduced as well as native mammals. Time in isolation impacts body size evolution profoundly. Body size shift of introduced mammals was much more pronounced with increasing residence times, yet far less than that exhibited by native, palaeo‐insular mammals (residence times > 10,000 years). Given the antiquity of many species introductions, it appears that much of what we view as the natural character and ecological dynamics of recent insular communities may have been rendered artefacts of ancient colonizations by humans and commensals.     

SPECIES SELECTION AND THE MACROEVOLUTION OF CORAL COLONIALITY AND PHOTOSYMBIOSIS

Differences in the relative diversification rates of species with variant traits are known as species selection. Species selection can produce a macroevolutionary change in the frequencies of traits by changing the relative number of species possessing each trait over time. But species selection is not the only process that can change the frequencies of traits, phyletic microevolution of traits within species and phylogenetic trait evolution among species, the tempo and mode of microevolution can also change trait frequencies. Species selection, phylogenetic, and phyletic processes can all contribute to large‐scale trends, reinforcing or canceling each other out. Even more complex interactions among macroevolutionary processes are possible when multiple covarying traits are involved. Here I present a multilevel macroevolutionary framework that is useful for understanding how macroevolutionary processes interact. It is useful for empirical studies using fossils, molecular phylogenies, or both. I illustrate the framework with the macroevolution of coloniality and photosymbiosis in scleractinian corals using a time‐calibrated molecular phylogeny. I find that standing phylogenetic variation in coloniality and photosymbiosis deflects the direction of macroevolution from the vector of species selection. Variation in these traits constrains species selection and results in a 200 million year macroevolutionary equilibrium.     

Effects of conspecific and heterospecific residents on patterns of immigration in two species of voles

Studies with birds have shown that presence and density of resident conspecifics and heterospecifics can influence patterns of habitat selection. There have been few studies on the effects of social cues on rates of immigration in mammals. We report results from a long-term live trapping study of immigration in two species of voles, Microtus ochrogaster Wagner, 1842 and Microtus pennsylvanicus Ord, 1815, in bluegrass habitat in east-central Illinois, USA. We compare immigration into control sites from which no individuals of either species were removed with immigration into experimental sites from which either conspecifics or heterospecifics were removed. We focus on characteristics of immigrants and rates of immigration in relation to density in destination habitats. Within each species, immigrants into control and removal sites were similar with respect to body mass and reproductive condition, indicating no major differences in the physical condition of immigrants into sites with established populations and sites without established populations. For both species, density of conspecifics at a site positively influenced rate of immigration at that site. Density of heterospecifics at destination sites did not significantly influence rate of immigration for either species. These results suggest that site selection by dispersing M. ochrogaster and M. pennsylvanicus is characterized by conspecific attraction.     

The evolution of mammal body sizes: responses to Cenozoic climate change in North American mammals

Explanations for the evolution of body size in mammals have remained surprisingly elusive despite the central importance of body size in evolutionary biology. Here, we present a model which argues that the body sizes of Nearctic mammals were moulded by Cenozoic climate and vegetation changes. Following the early Eocene Climate Optimum, forests retreated and gave way to open woodland and savannah landscapes, followed later by grasslands. Many herbivores that radiated in these new landscapes underwent a switch from browsing to grazing associated with increased unguligrade cursoriality and body size, the latter driven by the energetics and constraints of cellulose digestion (fermentation). Carnivores also increased in size and digitigrade, cursorial capacity to occupy a size distribution allowing the capture of prey of the widest range of body sizes. With the emergence of larger, faster carnivores, plantigrade mammals were constrained from evolving to large body sizes and most remained smaller than 1 kg throughout the middle Cenozoic. We find no consistent support for either Cope's Rule or Bergmann's Rule in plantigrade mammals, the largest locomotor guild (n = 1186, 59% of species in the database). Some cold‐specialist plantigrade mammals, such as beavers and marmots, showed dramatic increases in body mass following the Miocene Climate Optimum which may, however, be partially explained by Bergmann's rule. This study reemphasizes the necessity of considering the evolutionary history and resultant form and function of mammalian morphotypes when attempting to understand contemporary mammalian body size distributions.     

Middle Eocene habitat shifts in the North American western interior: A case study

There has been great interest in the global warming events that heralded the onset of the Eocene and particularly the response of mammalian faunas to these events. However, little information is available on the subsequent deterioration of tropical habitats in the interior of North America after these major warming episodes. The decline of tropical habitats is thought to have begun during the middle Eocene in the interior of North America, but until now, no studies have been able to document the details of this event. Recent fossil collection and stratigraphic studies from sites in southwestern Wyoming and northeastern Utah that span the middle Eocene offer a unique opportunity to evaluate changes in habitat in the western interior. Using a discriminant function analysis, habitats were reconstructed for a sequence of eight stratigraphically controlled middle Eocene assemblages. Adaptive profiles (diets, substrate use, and body masses) of fossil mammal communities were statistically compared to those of extant faunas from a variety of Neotropical habitats. Previously published magnetostratigraphic data from Utah provided a means to correlate our stratigraphic sections to the geomagnetic polarity time scale and the oxygen isotope record. The discriminant model shows that there was a significant change in the mammalian community ecology near the end of the late middle Eocene that is likely reflective of a habitat shift. When correlated to the time scale and oxygen isotope record, this key transition from forested habitats typical of the tropical early Eocene to more open woodlands began about 42 million years ago in this region of the Rocky Mountains.     

Ecological context and the importance of body and gnathopod size for pairing success in two amphipod ecomorphs

Ecological context generates interspecific variation in mating behavior by imposing differential constraints on the action of sexual selection, but operation of these constraints in nature is not well understood. We used field and laboratory studies to examine the importance of body size and size of sexually dimorphic appendages, the gnathopods, for pairing success in two freshwater amphipod species within the Hyalella azteca species complex. We focused on a large-bodied species found in habitats where ecological factors tend to favor large body size, and a small-bodied species in habitats where small body size is favored by size-selective predation. A field study indicated that although male pairing success was greater for larger males in both species, pairing success increased throughout the range of variation in male size in the large species, whereas, in the small species, pairing success was low for smaller individuals, but roughly constant across intermediate to larger sizes. A laboratory mate choice experiment was consistent with the field study, finding that females of the large species exhibited a preference for larger males that was independent of absolute male size, but females of the small species were indifferent when choosing between males of intermediate to larger size. Species also differed in the direction of sexual size dimorphism in the field, with males being the larger sex in the large species but the smaller sex in the small species, a pattern consistent with the species differences in mate preference. Large gnathopod size relative to body size was associated with enhanced pairing success across all body sizes for the large species, but, in the small species, large gnathopod size enhanced pairing success only in smaller males. Species differences in mating behavior appear consistent with change driven by differing forms of the interaction between sexual and natural selection.     

Ostracodes bathyaux du Crétacé terminal - Éocène moyen en Atlantique tropical (Plateau de Demerara, Leg 207)

Ostracode faunal assemblages which lived in bathyal environments are analysed from uppermost Maastrichtian - middle Eocene sediments of sites 1260 and 1261, drilled on the distal margin of Demerara, off Surinam (Leg 207, western tropical Atlantic). As for numerous other groups, the Cretaceous/Tertiary mass extinction event led to the disappearance of numerous bathyal ostracode species and it was followed by a very slow recovery during the Palaeocene. It appears that this extinction event was more devastating for detritus-feeder ostracode species than for filter or silt-eater groups, which crossed the K/T boundary without any or little morphological change. This extinction selectivity may be explained by the drop in productivity of surface waters which took place at the K/T boundary. Finally, psychrospheric ostracode species were encountered in Middle Eocene sediments, confirming thus the general cooling of deep oceanic realm recognised in general for this time interval.     

Of mice and mammoths: evaluations of causal explanations for body size evolution in insular mammals

Aim We investigated the hypothesis that the insular body size of mammals results from selective forces whose influence varies with characteristics of the focal islands and the focal species, and with interactions among species (ecological displacement and release). Location Islands world‐wide. Methods We assembled data on the geographic characteristics (area, isolation, maximum elevation, latitude) and climate (annual averages and seasonality of temperature and precipitation) of islands, and on the ecological and morphological characteristics of focal species (number of mammalian competitors and predators, diet, body size of mainland reference populations) that were most relevant to our hypothesis (385 insular populations from 98 species of extant, non‐volant mammals across 248 islands). We used regression tree analyses to examine the hypothesized contextual importance of these factors in explaining variation in the insular body size of mammals. Results The results of regression tree analyses were consistent with predictions based on hypotheses of ecological release (more pronounced changes in body size on islands lacking mammalian competitors or predators), immigrant selection (more pronounced gigantism in small species inhabiting more isolated islands), thermoregulation and endurance during periods of climatic or environmental stress (more pronounced gigantism of small mammals on islands of higher latitudes or on those with colder and more seasonal climates), and resource subsidies (larger body size for mammals that utilize aquatic prey). The results, however, were not consistent with a prediction based on resource limitation and island area; that is, the insular body size of large mammals was not positively correlated with island area. Main conclusions These results support the hypothesis that the body size evolution of insular mammals is influenced by a combination of selective forces whose relative importance and nature of influence are contextual. While there may exist a theoretical optimal body size for mammals in general, the optimum for a particular insular population varies in a predictable manner with characteristics of the islands and the species, and with interactions among species. This study did, however, produce some unanticipated results that merit further study – patterns associated with Bergmann’s rule are amplified on islands, and the body size of small mammals appears to peak at intermediate and not maximum values of latitude and island isolation.     

Relationship between exercise capacity and brain size in mammals

Background

A great deal of experimental research supports strong associations between exercise, cognition, neurogenesis and neuroprotection in mammals. Much of this work has focused on neurogenesis in individual subjects in a limited number of species. However, no study to date has examined the relationship between exercise and neurobiology across a wide range of mammalian taxa. It is possible that exercise and neurobiology are related across evolutionary time. To test this hypothesis, this study examines the association between exercise and brain size across a wide range of mammals.

Methodology/Principal Findings

Controlling for associations with body size, we examined the correlation between brain size and a proxy for exercise frequency and capacity, maximum metabolic rate (MMR; ml O₂ min⁻¹). We collected brain sizes and MMRs from the literature and calculated residuals from the least-squares regression line describing the relationship between body mass and each variable of interest. We then analyzed the correlation between residual brain size and residual MMR both before and after controlling for phylogeny using phylogenetic independent contrasts. We found a significant positive correlation between maximum metabolic rate and brain size across a wide range of taxa.

Conclusions

These results suggest a novel hypothesis that links brain size to the evolution of locomotor behaviors in a wide variety of mammalian species. In the end, we suggest that some portion of brain size in nonhuman mammals may have evolved in conjunction with increases in exercise capacity rather than solely in response to selection related to cognitive abilities.