Non-reproductive Individuals Research Articles

Delayed maturation of the exoskeleton and muscle fibres in the ant Platythyrea punctata

Abstract In many species of eusocial Hymenoptera, workers exhibit a division of labour based on age, wherein young workers remain protected within the nest before transitioning to foraging activities. These changes in tasks and in external stress exposure are accompanied by physiological and anatomical modifications. Although pigmentation changes in the exoskeleton are well documented in ants, limited information is available on muscle morphology and developmental changes in the adult cuticle. Here, we investigated the alterations in the exoskeleton and thoracic muscles across different age groups of the clonal ant Platythyrea punctata, in which reproduction is monopolized by one or a few dominant workers. Our findings revealed a significant thickening of the cuticle and muscle fibres in young workers during the intranidal period, reaching a final stage in forager individuals. We further tested the influence of reproductive status on body development and found variations only in muscle fibres between reproductive and nonreproductive individuals. This suggests that dominant individuals allocate resources in egg-laying by minimizing other metabolic costs. These findings shed light on how eusociality profoundly alters selection pressures and results in temporal shifts in individual development. This strategy probably provides advantages to colonies by minimizing nutritional waste and optimizing resource allocation.

Species- and sex-specific chemical composition from an internal gland-like tissue of an African frog family.

Intraspecific chemical communication in frogs is understudied and the few published cases are limited to externally visible and male-specific breeding glands. Frogs of the family Odontobatrachidae, a West African endemic complex of five morphologically cryptic species, have large, fatty gland-like strands along their lower mandible. We investigated the general anatomy of this gland-like strand and analysed its chemical composition. We found the strand to be present in males and females of all species. The strand varies in markedness, with well-developed strands usually found in reproductively active individuals. The strands are situated under particularly thin skin sections, the vocal sac in male frogs and a respective area in females. Gas-chromatography/mass spectrometry and multivariate analysis revealed that the strands contain sex- and species-specific chemical profiles, which are consistent across geographically distant populations. The profiles varied between reproductive and non-reproductive individuals. These results indicate that the mandibular strands in the Odontobatrachidae comprise a so far overlooked structure (potentially a gland) that most likely plays a role in the mating and/or breeding behaviour of the five Odontobatrachus species. Our results highlight the relevance of multimodal signalling in anurans, and indicate that chemical communication in frogs may not be restricted to sexually dimorphic, apparent skin glands.

Female little brown bats require both building and natural roosts in a mountainous environment with short summers

AbstractThe little brown bat (Myotis lucifugus) is an endangered species that occurs throughout most of North America. The recovery strategy for little brown bats in Canada identifies a need to understand habitat use and protect important bat habitat across the species' Canadian range; however, bat roosting habitat requirements have not been well studied in mountainous environments within the northern part of the species' range. Our objective was to understand the use of building and natural roosts by female little brown bats in the Canadian Rocky Mountains. Over two summers, we captured little brown bats in Banff, Yoho, and Kootenay National Parks, in Alberta and British Columbia, Canada, and tracked 49 females to their day roosts. Buildings were the only roost type used by radio‐tagged bats of all reproductive stages, the dominant roost type used throughout the reproductive period, and the only structures in which we located maternity colonies. Buildings also appeared to shape the distribution of foraging female and reproductive female bats such that few were captured at long distances from towns and building clusters. Compared with bats in more southern and in non‐mountainous areas, bats in our study area had delayed juvenile weaning and volancy, began hibernation early, and had low reproductive rates. We suggest that buildings are a key habitat for female little brown bats to persist at their current levels in our study area, by providing thermal conditions that promote overwinter survival; additional reproductive delays caused by torpor use in cool natural roosts would likely reduce juvenile and adult female overwinter survival below the levels required to sustain populations. Nevertheless, building colonies were unusually small, and bats had relatively low fidelity to particular building roosts, suggesting that building roosts may not consistently meet bats' needs. Some nonreproductive and pregnant females also frequently used natural roosts, particularly after colder nights; these roosts may be important in facilitating torpor and energy conservation for these bats. Because bats are long‐lived with low reproductive output and low juvenile survival, addressing the needs of both reproductive and nonreproductive individuals is important for population persistence.

The royal food of termites shows king and queen specificity.

Society in eusocial insects is based on the reproductive division of labor, with a small number of reproductive individuals supported by a large number of nonreproductive individuals. Because inclusive fitness of all colony members depends on the survival and fertility of reproductive members, sterile members provide royals with special treatment. Here, we show that termite kings and queens each receive special food of a different composition from workers. Sequential analysis of feeding processes demonstrated that workers exhibit discriminative trophallaxis, indicating their decision-making capacity in allocating food to the kings and queens. Liquid chromatography tandem-mass spectrometry analyses of the stomodeal food and midgut contents revealed king- and queen-specific compounds, including diacylglycerols and short-chain peptides. Desorption electrospray ionization mass spectrometry imaging analyses of 13C-labeled termites identified phosphatidylinositol and acetyl-l-carnitine in the royal food. Comparison of the digestive tract structure showed remarkable differences in the volume ratio of the midgut-to-hindgut among castes, indicating that digestive division of labor underlies reproductive division of labor. Our demonstration of king- and queen-specific foods in termites provides insight into the nutritional system that underpins the extraordinary reproduction and longevity of royals in eusocial insects.

Evolution: How sweat bees gained and lost eusociality.

Eusocial insects divide labor between reproductive and non-reproductive individuals. The molecular mechanisms underlying the evolution of these castes have remained mysterious. A comparative genomic study of sweat bees points to a familiar factor as a regulator of behavioral specialization: juvenile hormone.

Constitutive Innate Immunity of Migrant and Resident Long-Nosed Bats (Leptonycteris yerbabuenae) in the Drylands of Mexico

In contrast to birds, the relationship between migration and immunity has been scarcely studied in bats. We examined how the expression of the humoral portion of the constitutive immunity varied in a bat with partial, sex-biased migration: the lesser long-nosed bat (Leptonycteris yerbabuenae (Phyllostomidae)). The lesser long-nosed bat is a nectarivorous species distributed in the arid and semi-arid regions of North and Central America. We evaluated the bacteria-killing abilities (BKAs) of the plasma of male and female lesser long-nosed bats on the Pacific coast in different periods of the year. Because adult males are resident, they were used to explore the effect of reproductive activity on BKA, and we predicted higher values in mating males (i.e., individuals presenting scrotal testicles and a fresh dorsal patch). In contrast to males, most females migrate to cactus deserts in northern Mexico during pregnancy and lactation, and then return to the dry forests of west-central Mexico to mate. We predicted that the combined effect of breeding and migration would have an adverse effect on BKA; therefore, migratory pregnant and lactating females were expected to exhibit a lower BKA than mating females in west-central Mexico. We compared the BKAs of females captured in October and December in central Mexico, and we predicted that migratory females that had recently arrived in October should exhibit a lower BKA than females captured two months later. In addition, we compared the BKAs between lactating females and young in northern Mexico and predicted lower values in recently born individuals. We found that the BKAs of males were higher in reproductive individuals than in non-reproductive individuals. We found a significant difference in the BKAs between females at the two extremes of their migratory range: the values of pregnant females in Sonora and females in December were higher than those of females captured in October. Finally, we found no difference in BKAs between lactating females and young individuals. Our findings indicate that the basal levels of the innate humoral component are heightened in mating males, that this response is reduced in females that recently returned to their mating grounds, and that the constitutive immunity of young individuals matures early, probably in anticipation of the potential to encounter pathogens during their migration to west-central Mexico.

Cascading effects of seed-stem-individual spatial patterns along a grazing gradient.

Studying the seed trait-stem trait-individual spatial pattern system is helpful for understanding the developmental direction of plant dynamics and populations under grazing disturbance as well as the antagonistic relationship between animals and plants, but few systematic analyses of this spatial pattern system have been carried out. Kobresia humilis is the dominant species in alpine grasslands. We studied K. humilis seed traits and their relationship with K. humilis reproductive individuals, the relationship between reproductive and vegetative stems, and the weights and spatial patterns of reproductive and nonreproductive individuals under four grazing treatments: no grazing (control), light grazing, moderate grazing and heavy grazing. We explored the relationship among seed size and seed number with reproductive stems and vegetative stems along the grazing gradient and assessed the spatial pattern changes between reproductive and nonreproductive individuals. The results showed the following: (1) Seed size increased with increasing grazing intensity, and the coefficient of variation for seed size and seed number in the heavy grazing treatment was greater than 0.6. (2) The structural equation model showed that grazing treatment had a positive effect on seed number, seed size and reproductive stem number and a negative effect on reproductive stem weight. (3) Grazing treatment did not affect the resource allocation to reproductive stems and vegetative stems per unit length of reproductive K. humilis individuals. (4) Compared with the number of reproductive individuals in the no grazing treatment, the number in the heavy grazing treatment decreased significantly, and the negative correlation between reproductive individuals and nonreproductive individuals changed from a full-scale negative correlation to a small-scale negative correlation and a large-scale positive correlation. Our study showed that grazing could activate and change the resource allocation pattern of dominant species in a grassland and have significant positive effects on reproductive stem number, reproductive stem weight, seed number and seed size. Along a grazing intensity gradient, with the increase in distance between reproductive and nonreproductive individuals, the transformation of intraspecific relationships from a negative correlation to a positive correlation is an ecological strategy conducive to population survival.

Co-expression Gene Networks and Machine-learning Algorithms Unveil a Core Genetic Toolkit for Reproductive Division of Labour in Rudimentary Insect Societies.

The evolution of eusociality requires that individuals forgo some or all their own reproduction to assist the reproduction of others in their group, such as a primary egg-laying queen. A major open question is how genes and genetic pathways sculpt the evolution of eusociality, especially in rudimentary forms of sociality-those with smaller cooperative nests when compared with species such as honeybees that possess large societies. We lack comprehensive comparative studies examining shared patterns and processes across multiple social lineages. Here we examine the mechanisms of molecular convergence across two lineages of bees and wasps exhibiting such rudimentary societies. These societies consist of few individuals and their life histories range from facultative to obligately social. Using six species across four independent origins of sociality, we conduct a comparative meta-analysis of publicly available transcriptomes. Standard methods detected little similarity in patterns of differential gene expression in brain transcriptomes among reproductive and non-reproductive individuals across species. By contrast, both supervised machine learning and consensus co-expression network approaches uncovered sets of genes with conserved expression patterns among reproductive and non-reproductive phenotypes across species. These sets overlap substantially, and may comprise a shared genetic "toolkit" for sociality across the distantly related taxa of bees and wasps and independently evolved lineages of sociality. We also found many lineage-specific genes and co-expression modules associated with social phenotypes and possible signatures of shared life-history traits. These results reveal how taxon-specific molecular mechanisms complement a core toolkit of molecular processes in sculpting traits related to the evolution of eusociality.

Home-Range Size and Space Use of Territorial Bonelli’s Eagles (Aquila fasciata) Tracked by High-Resolution GPS/GSM Telemetry

High-resolution GPS/GSM dataloggers provide spatial information of the highest quality, which outperform previous tracking methods, such as Argos telemetry or conventional VHF ground-tracking. As a result, this has improved our knowledge of home-range behavior and spatial ecology of many species, including large raptors. In this paper, we use high-resolution GPS/GSM dataloggers to assess the home-range size and the role of sex, season (breeding or non-breeding season), and breeding status (reproductive or non-reproductive individuals) on the space use of Bonelli’s eagle (Aquila fasciata). To this end, 51 territorial individuals (25 females and 26 males) were equipped with GPS/GSM transmitters and were tracked over 7 years (2015–2021) in eastern Spain. Overall, we recorded 4,791,080 fixes that were analyzed through kernel density methods (50%, 75%, and 95% fixed kernels). The average individual home-range size according to the 95%, 75%, and 50% kernels was 54.84 ± 20.78 km2, 24.30 ± 10.18 km2, and 11.17 ± 4.90 km2, respectively. Overall, the home-range size of individuals occupying the same territory was similar, mainly due to the cooperative hunting behavior exhibited by the species. We did not find interannual differences in the home-range size (95% fixed kernel) of the majority of individuals, showing a strong territorial fidelity of the breeding pairs. In general, females’ home-range size was slightly smaller than males’ size due to the decrease in activity in the breeding season as a result of laying, incubation, and chick attendance at nests. No seasonal variation in the 95% kernel was found, but it was found in the 75% and 50% kernels. In regard to the breeding status, higher home-range size was recorded in the non-reproductive individuals. Moreover, we found a low neighbor overlap among the territories (4.18% ± 3.06%), which evidences a high level of intraspecific competition in the Bonelli’s eagle. Finally, this study highlights the advantages of the use of accurate telemetry information to improve our understanding of the spatial ecology of the endangered Bonelli’s eagle, which ultimately will serve to better inform management actions for its conservation.

Canalized gene expression during development mediates caste differentiation in ants

Ant colonies are higher-level organisms consisting of specialized reproductive and non-reproductive individuals that differentiate early in development, similar to germ–soma segregation in bilateral Metazoa. Analogous to diverging cell lines, developmental differentiation of individual ants has often been considered in epigenetic terms but the sets of genes that determine caste phenotypes throughout larval and pupal development remain unknown. Here, we reconstruct the individual developmental trajectories of two ant species, Monomorium pharaonis and Acromyrmex echinatior, after obtaining >1,400 whole-genome transcriptomes. Using a new backward prediction algorithm, we show that caste phenotypes can be accurately predicted by genome-wide transcriptome profiling. We find that caste differentiation is increasingly canalized from early development onwards, particularly in germline individuals (gynes/queens) and that the juvenile hormone signalling pathway plays a key role in this process by regulating body mass divergence between castes. We quantified gene-specific canalization levels and found that canalized genes with gyne/queen-biased expression were enriched for ovary and wing functions while canalized genes with worker-biased expression were enriched in brain and behavioural functions. Suppression in gyne larvae of Freja, a highly canalized gyne-biased ovary gene, disturbed pupal development by inducing non-adaptive intermediate phenotypes between gynes and workers. Our results are consistent with natural selection actively maintaining canalized caste phenotypes while securing robustness in the life cycle ontogeny of ant colonies.

Seasonal Changes in Locomotor Activity Patterns of Wild Social Natal Mole-Rats (Cryptomys hottentotus natalensis)

Differences in individual locomotor activity patterns may be linked to a number of ecological factors, such as changes in ambient temperature or photoperiod. Observations on subterranean mammals suggest that they exhibit diel rhythms despite the lack of visual cues in their underground burrows, but it is unknown how seasonality and individual characteristics affect their activity. In this study we use RFID technology to monitor daily activity patterns of wild, social Natal mole-rats (Cryptomys hottentotus natalensis) during the summer and winter to investigate how their activity varies with season and whether their activity depends on individual characteristics such as body mass, sex and reproductive status. We found that in winter, individuals were more active during the time with the highest soil temperatures, whereas in summer, they showed a bimodal activity pattern during early morning and late afternoon coinciding with cooler soil temperatures. Individual characteristics, including reproductive status, did not affect general activity indicating that reproductive and non-reproductive individuals contribute equally to cooperative behaviors. We suggest that the activity patterns may be a behavioral adaptation to avoid extreme burrow temperatures and a mechanism to maintain a stable core body temperature. We highlight the advantages of RFID technology to study wild small mammal movements.

Altruism and Phenoptosis as Programs Supported by Evolution.

Phenoptosis is a programmed death that has emerged in the process of evolution, sometimes taking the form of an altruistic program. In particular, it is believed to be a weapon against the spread of pandemics in the past and an obstacle in fighting pandemics in the present (COVID). However, on the evolutionary scale, deterministic death is not associated with random relationships (for example, bacteria with a particular mutation), but is a product of higher nervous activity or a consequence of established hierarchy that reaches its maximal expression in eusocial communities of Hymenoptera and highly social communities of mammals. Unlike a simple association of individuals, eusociality is characterized by the appearance of non-reproductive individuals as the highest form of altruism. In contrast to primitive programs for unicellular organisms, higher multicellular organisms are characterized by the development of behavior-based phenoptotic programs, especially in the case of reproduction-associated limitation of lifespan. Therefore, we can say that the development of altruism in the course of evolution of sociality leads in its extreme manifestation to phenoptosis. Development of mathematical models for the emergence of altruism and programmed death contributes to our understanding of mechanisms underlying these paradoxical counterproductive (harmful) programs. In theory, this model can be applied not only to insects, but also to other social animals and even to the human society. Adaptive death is an extreme form of altruism. We consider altruism and programmed death as programmed processes in the mechanistic and adaptive sense, respectively. Mechanistically, this is a program existing as a predetermined chain of certain responses, regardless of its adaptive value. As to its adaptive value (regardless of the degree of “phenoptoticity”), this is a characteristic of organisms that demonstrate high levels of kinship, social organization, and physical association typical for higher-order individuals, e.g., unicellular organisms forming colonies with some characteristics of multicellular animals or colonies of multicellular animals displaying features of supraorganisms.

Telomerase-Positive Somatic Tissues of Honeybee Queens (Apis mellifera) Display No DNA Replication

Telomere biology is closely linked to the process of aging. The restoration of telomere length by maintaining telome­rase activity in certain cell types of human adults allows for the proliferative capacity of the cells and preserves the regeneration potential of the tissue. The absence of telome­rase, that leads to telomere attrition and irreversible cell cycle arrest in most somatic cells, acts as a protective mechanism against uncontrolled cancer growth. Nevertheless, there have been numerous studies indicating noncanonical functions of telomerase besides those involved in telomere lengthening. Eusocial insects serve as a great system for aging research. This is because eusocial reproductives, such as queens and kings, have a significantly extended lifespan compared to nonreproductive individuals of the same species. We report that the somatic tissues of honeybee queens (Apis mellifera) are associated with upregulated telomerase activity; however, this upregulation does not fully correlate with the rate of DNA replication in the tissues. This indicates a noncanonical role of telomerase in the somatic tissues of honeybee queens.

Non-kin Cooperation in Ants

Eusociality represents an extreme form of social behavior characterized by a reproductive division of labor. Eusociality necessarily evolved through kin selection, which requires interactions among related individuals. However, many eusocial taxa also show cooperation between non-kin groups, challenging the idea that cooperative actions should only occur among relatives. This review explores the causes and consequences of non-kin cooperation in ants. Ants display a diversity of behaviors that lead to non-kin cooperation within and between species. These interactions occur among both reproductive and non-reproductive individuals. The proximate and ultimate mechanisms leading to non-kin cooperative interactions differ substantially depending on the biotic and abiotic environment. We end this review with directions for future research and suggest that the investigation of non-kin cooperative actions provides insight into processes leading to social evolution.

Social Evolution With Decoupling of Multiple Roles of Biogenic Amines Into Different Phenotypes in Hymenoptera

Convergent evolution of eusociality with the division of reproduction and its plastic transition in Hymenoptera has long attracted the attention of researchers. To explain the evolutionary scenario of the reproductive division of labor, several hypotheses had been proposed. Among these, we focus on the most basic concepts, i.e., the ovarian ground plan hypothesis (OGPH) and the split-function hypothesis (SFH). The OGPH assumes the physiological decoupling of ovarian cycles and behavior into reproductive and non-reproductive individuals, whereas the SFH assumes that the ancestral reproductive function of juvenile hormone (JH) became split into a dual function. Here, we review recent progress in the understanding of the neurohormonal regulation of reproduction and social behavior in eusocial hymenopterans, with an emphasis on biogenic amines. Biogenic amines are key substances involved in the switching of reproductive physiology and modulation of social behaviors. Dopamine has a pivotal role in the formation of reproductive skew irrespective of the social system, whereas octopamine and serotonin contribute largely to non-reproductive social behaviors. These decoupling roles of biogenic amines are seen in the life cycle of a single female in a solitary species, supporting OGPH. JH promotes reproduction with dopamine function in primitively eusocial species, whereas it regulates non-reproductive social behaviors with octopamine function in advanced eusocial species. The signal transduction networks between JH and the biogenic amines have been rewired in advanced eusocial species, which could regulate reproduction in response to various social stimuli independently of JH action.

Small-scale genetic structure and mating patterns in an extensive sessile oak forest (Quercus petraea (Matt.) Liebl.).

Oaks (Quercus) are major components of temperate forest ecosystems in the Northern Hemisphere where they form intermediate or climax communities. Sessile oak (Quercus petraea) forests represent the climax vegetation in eastern Germany and western Poland. Here, sessile oak forms pure stands or occurs intermixed with Scots Pine (Pinus sylvestris). A large body of research is available on gene flow, reproduction dynamics, and genetic structure in fragmented landscapes and mixed populations. At the same time, our knowledge regarding large, contiguous, and monospecific populations is considerably less well developed. Our study is an attempt to further develop our understanding of the reproduction ecology of sessile oak as an ecologically and economically important forest tree by analyzing mating patterns and genetic structure within adult trees and seedlings originating from one or two reproduction events in an extensive, naturally regenerating sessile oak forest. We detected positive spatial genetic structure up to 30 meters between adult trees and up to 40 meters between seedlings. Seed dispersal distances averaged 8.4 meters. Pollen dispersal distances averaged 22.6 meters. In both cases, the largest proportion of the dispersal occurred over short distances. Dispersal over longer distances was more common for pollen but also appeared regularly for seeds. The reproductive success of individual trees was highly skewed. Only 41 percent of all adult trees produced any offspring while the majority did not participate in reproduction. Among those trees that contributed to the analyzed seedling sample, 80 percent contributed 1–3 gametes. Only 20 percent of all parent trees contributed four or more gametes. However, these relatively few most fertile trees contributed 51 percent of all gametes within the seedling sample. Vitality and growth differed significantly between reproducing and nonreproducing adult trees with reproducing trees being more vital and vigorous than nonreproducing individuals. Our study demonstrates that extensive, apparently homogenous oak forests are far from uniform on the genetic level. On the contrary, they form highly complex mosaics of remarkably small local neighborhoods. This counterbalances the levelling effect of long‐distance dispersal and may increase the species’ adaptive potential. Incorporating these dynamics in the management, conservation, and restoration of oak forests can support the conservation of forest genetic diversity and assist those forests in coping with environmental change.

Queen loss increases worker survival in leaf-cutting ants under paraquat-induced oxidative stress.

Longevity is traded off with fecundity in most solitary species, but the two traits are positively linked in social insects. In ants, the most fecund individuals (queens and kings) live longer than the non-reproductive individuals, the workers. In many species, workers may become fertile following queen loss, and recent evidence suggests that worker fecundity extends worker lifespan. We postulated that this effect is in part owing to improved resilience to oxidative stress, and tested this hypothesis in three Myrmicine ants: Temnothorax rugatulus, and the leaf-cutting ants Atta colombica and Acromyrmex echinatior. We removed the queen from colonies to induce worker reproduction and subjected workers to oxidative stress. Oxidative stress drastically reduced survival, but this effect was less pronounced in leaf-cutting ant workers from queenless nests. We also found that, irrespective of oxidative stress, outside workers died earlier than inside workers did, likely because they were older. Since At. colombica workers cannot produce fertile offspring, our results indicate that direct reproduction is not necessary to extend the lives of queenless workers. Our findings suggest that workers are less resilient to oxidative stress in the presence of the queen, and raise questions on the proximate and ultimate mechanisms underlying socially mediated variation in worker lifespan. This article is part of the theme issue 'Ageing and sociality: why, when and how does sociality change ageing patterns?'

Bone remodeling in the longest living rodent, the naked mole-rat: Interelement variation and the effects of reproduction.

The pattern of bone remodeling of one of the most peculiar mammals in the world, the naked mole-rat (NMR), was assessed. NMRs are known for their long lifespans among rodents and for having low metabolic rates. We assessed long-term in vivo bone labeling of subordinate individuals, as well as the patterns of bone resorption and bone remodeling in a large sample including reproductive and non-reproductive individuals (n=70). Over 268 undecalcified thin cross-sections from the midshaft of humerus, ulna, femur and tibia were analyzed with confocal fluorescence and polarized light microscopy. Fluorochrome analysis revealed low osteogenesis, scarce bone resorption and infrequent formation of secondary osteons (Haversian systems) (i.e., slow bone turnover), thus most likely reflecting the low metabolic rates of this species. Secondary osteons occurred regardless of reproductive status. However, considerable differences in the degree of bone remodeling were found between breeders and non-breeders. Pre-reproductive stages (subordinates) exhibited quite stable skeletal homeostasis and bone structure, although the attainment of sexual maturity and beginning of reproductive cycles in female breeders triggered a series of anabolic and catabolic processes that up-regulate bone turnover, most likely associated with the increased metabolic rates of reproduction. Furthermore, bone remodeling was more frequently found in stylopodial elements compared to zeugopodial elements. Despite the limited bone remodeling observed in NMRs, the variation in the pattern of skeletal homeostasis (interelement variation) reported here represents an important aspect to understand the skeletal dynamics of a small mammal with low metabolic rates. Given the relevance of the remodeling process among mammals, this study also permitted the comparison of such process with the well-documented histomorphology of extinct therapsids (i.e., mammalian precursors), thus evidencing that bone remodeling and its endocortical compartmentalization represent ancestral features among the lineage that gave rise to mammals. It is concluded that other factors associated with development (and not uniquely related to biomechanical loading) can also have an important role in the development of bone remodeling.

Rewinding Extinction in the Northern White Rhinoceros: Genetically Diverse Induced Pluripotent Stem Cell Bank for Genetic Rescue.

Extinction rates are rising, and current conservation technologies may not be adequate for reducing species losses. Future conservation efforts may be aided by the generation of induced pluripotent stem cells (iPSCs) from highly endangered species. Generation of a set of iPSCs from multiple members of a species can capture some of the dwindling genetic diversity of a disappearing species. We generated iPSCs from fibroblasts cryopreserved in the Frozen Zoo®: nine genetically diverse individuals of the functionally extinct northern white rhinoceros (Ceratotherium simum cottoni) and two from the closely related southern white rhinoceros (Ceratotherium simum simum). We used a nonintegrating Sendai virus reprogramming method and developed analyses to confirm the cells' pluripotency and differentiation potential. This work is the first step of a long-term interdisciplinary plan to apply assisted reproduction techniques to the conservation of this highly endangered species. Advances in iPSC differentiation may enable generation of gametes in vitro from deceased and nonreproductive individuals that could be used to repopulate the species.

Variation of social strategies within and between individual black-tailed prairie dogs (Cynomys ludovicianus) during the reproductive season

Sociality describes the tendency for members of a species to associate in a group based on social attraction. To balance the trade-offs of sociality, the fitness consequences (and therefore proclivity towards being social) should be expected to vary across individuals and within the same individual, across time. We used social network analysis to evaluate the causes and consequences of within- and between-individual variation in sociality in a wild population of black-tailed prairie dogs (Cynomys ludovicianus) during the reproductive season. We hypothesised that individuals would adjust their behaviours based on a key life-history event: reproduction. We assessed this variation by comparing differences in social behaviour between individuals in different life history stages and within individuals across the reproductive season. We predicted breeding females would increase their agonistic interactions to protect their offspring from infanticide during reproduction, while non-reproductive individuals would increase their affiliative interactions to maintain stability in their social network and increase their indirect fitness. We found that the number of social interactions decreased after reproduction for all individuals. Breeding females did not increase their agonistic interactions compared to other groups, but instead had overall decreased social interactions during reproduction, which corresponded with increased fitness. For non-breeding females, an increase of affiliative social interactions correlated with increased indirect fitness as we predicted. Our multiple-level, temporally structured analyses revealed under-realised complexities regarding within- and between-individual variation in social interactions, suggesting reproductive status dictates different social strategies to optimise fitness. Social living with conspecifics confers a range of costs and benefits, the trade-offs of which are expected to change based on life history. Therefore, individuals should modulate their sociality across life history transitions to optimise fitness. Reproductive status is known to affect socio-behavioural decisions across taxa. We used a highly social species, black-tailed prairie dogs, to investigate variation in social structure across the reproductive season, and understand both the causes and consequences of individual variation in social structure. Prairie dogs offered easily observable, dynamic social behaviours and quantifiable fitness consequences to truly demonstrate acute social change. Our results suggest that there is variation of social interactions not only within individuals over time, but also between reproductive groups in the same time period. Our analyses suggest that social variation has important consequences for fitness and should further stimulate study of temporally based analyses of social behaviour.