Articles published on Reproductive ecology
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- New
- Research Article
- 10.1111/jpy.70193
- Jun 23, 2026
- Journal of phycology
- Aurélien Baud + 5 more
Unlike other red algae, the perennial species Palmaria palmata exhibits pronounced sexual dimorphism with a microscopic female gametophytic phase that does not produce cystocarps. To investigate how these unique features of its life cycle influence its reproductive mode, we analyzed the relative proportion of macroscopic individuals (tetrasporophytes vs. male gametophytes) across multiple geographic scales and between intertidal and subtidal habitats along the western coast of France (892 individuals collected from 17 locations). Because female gametophytes are microscopic and cannot be directly sampled in the field, their frequencies were inferred under two alternative hypotheses: (H1) assuming an equal sex ratio (i.e., the proportion of females equal to males) and (H2) assuming the proportion of females equals tetrasporophytes, given that tetrasporophytes develop directly on a female gametophyte. Under H1, our results predict a demographic advantage of tetrasporophytes over gametophytes. Moreover, tetrasporophytes were significantly more abundant in intertidal than in subtidal zones, likely reflecting their greater resilience to environmental fluctuations. Conversely, under H2, the sex ratio was significantly biased toward female gametophytes and the estimated proportion of tetrasporophytes was lower than that of gametophytes. In addition, the relative contribution of self-fertilization and clonal reproduction were assessed using 11 microsatellite markers. These findings confirm that this species reproduces mainly sexually and suggest that sexual dimorphism may limit selfing and enhance the survival of the tetrasporophyte, which develops directly on the female, thereby compensating for the absence of cystocarps. Overall, our results provide valuable insights into the reproductive strategies and adaptive potential of this commercially important alga.
- New
- Research Article
- 10.1080/03949370.2026.2667305
- Jun 21, 2026
- Ethology Ecology & Evolution
- Matthew P.O Dore + 1 more
Disturbance, hydrology and nest architecture were evaluated as drivers of reproductive outcomes in West African dwarf crocodiles (Osteolaemus tetraspis) over two breeding seasons (2021–2022) in Delta State, Nigeria. Surveys spanned culturally protected Emu (Ndokwa West LG) and more disturbed Okogbe and Esaba (Ehavbaren and Okpahoro) in Ughelli South LG, recording nest fate, architecture, microclimate and proximity to water; egg morphometrics (three Emu sites) and hatchling measurements (Emu vs Okopahoro/USLG) complemented field data. Non-parametric tests (Mann-Whitney; Kruskal-Wallis with Holm adjustment) and Cliff’s δ quantified differences. A disturbance gradient was evident: Emu showed repeated nest reuse and successful hatching, whereas Okogbe and Esaba showed recurrent failure. Architecture tracked hydrology (Emu-Ndokwa West LG: deeper upper-egg placement/chamber bottoms; Ughelli South LG: greater cavity depth/mound height). Mean core temperature was 36.7 °C; clutches ranged between 15 and 27 eggs. Within Emu, eggs differed significantly among sites; between localities, Okogbe/USLG hatchlings were longer and heavier. Findings indicate that human pressure rather than microhabitat alone dominates nesting vulnerability and hatch success; architecture mitigates flood risk; and maternal traits with microenvironment jointly shape offspring phenotypes. Recommended actions include community co-management and targeted enforcement to reduce disturbance, hydrology aware restoration that treats mound elevation and cavity depth as functional design features, and instrumented monitoring (core temperature, water level, maternal metrics) to anticipate failure thresholds and sustain recruitment in a changing swamp forest mosaic.
- Research Article
- 10.1111/jfb.70521
- Jun 4, 2026
- Journal of fish biology
- Malia B Davidson + 4 more
Understanding the timing, duration and effort put into reproduction is vital to successful management of any species. The southeastern United States is home to tremendous levels of aquatic biodiversity and was recently named one of the Earth's biodiversity hotspots. For some species, the fundamentals of age, growth, reproductive timing and ecology remain poorly understood. Percina aurora, the pearl darter, has been recently listed as threatened due to its contracting range, likely due to anthropogenic pressure on these systems. However, management efforts are hindered by the lack of knowledge about the life history of P. aurora. The purpose of this project was to examine the age, growth and reproductive ecology of P. aurora in the Pascagoula River basin of southeastern Mississippi, USA. To do this, we compiled length data from recent field collections and catalogued museum specimens to estimate age and growth, as well as clutch parameters (GSI, egg size) to infer reproductive output. We also used larval fish metabarcoding to confirm successful spawning of P. aurora. These results indicate P. aurora exhibits a protracted, potentially batch-spawning reproductive strategy with consistent spawning timing across rivers, supported by both growth models and larval detections.
- Research Article
- 10.1093/reprod/xaag062
- Jun 2, 2026
- Reproduction (Cambridge, England)
- Rhiannon V Eastment + 5 more
In brief: Pharmaceutical pollution can have serious consequences for wildlife reproduction. This paper reveals that exposure to the pervasive oestrogenic pollutant 17ɑ-ethinylestradiol does not impact female reproduction in a nonmodel viviparous fish. Abstract: Pharmaceuticals are fast becoming a major source of environmental pollution. Many pharmaceuticals are endocrine disruptors and can thus interfere with the endocrine system of exposed organisms, leading to a suite of behavioural and physiological alterations. While numerous studies have shown that exposure to pharmaceutical contaminants can disrupt the reproductive ecology of oviparous fishes, far less attention has been given to understanding the impacts on viviparous fishes, despite their ecological and evolutionary importance. Here, we investigated the impacts of the pharmaceutical contaminant 17ɑ-ethinylestradiol (EE2) on the reproductive biology of a marine livebearer, the Southern crested weedfish (Cristiceps australis). Adult weedfish were exposed to one of three nominal concentrations of EE2 (0, 30, and 300 ng/L) for 14 days. We then assessed the impacts of exposure on morphology of both sexes, and female gametogenesis. In doing so, we found no differences in the body length, weight, or condition of fish between treatment groups. We additionally found no differences in gonadosomatic index or gametogenesis between exposed and unexposed females. In contrast to reports in other taxa, our findings suggest that short-term exposure to EE2 does not produce detectable effects on weedfish growth and reproduction. However, the extent to which exposure duration influences these responses is unknown. Ultimately, these results emphasize the importance of assessing the effects of EE2 on a diverse range of organisms if we are to gain a more comprehensive understanding of its ecological impacts.
- Research Article
- 10.1093/jhered/esag042
- May 22, 2026
- The Journal of heredity
- Rebecca Krohman + 4 more
Life-history divergence between Atlantic cod (Gadus morhua) ecotypes in Skagerrak is maintained despite gene flow. Fjord and North Sea ecotypes differ in growth, body size, and reproductive ecology. One environmental heterogeneity in Skagerrak is salinity, as coastal habitats experience substantially lower and more variable salinities than offshore waters. Egg buoyancy is highly sensitive to salinity and represents a critical life-history trait shaping survival and recruitment. Inversions on chromosomes 2, 7, and 12 in cod are linked to salinity adaptation and might be important for egg buoyancy and survival. Here, we conducted a buoyancy test on Atlantic cod eggs, spawned from North Sea and Fjord ecotypes, using Coombs apparatus. Ecotypes and genotypes for the inversions on chromosomes 2, 7, and 12 were determined in all individuals (n = 242). We predicted that inversion genotypes and ecotypes would be associated with egg buoyancy. We found that buoyancy was variable and followed a bimodal distribution sorted by ecotype, with Fjord and North Sea individuals having mean buoyancies at neutral salinity (at 7°C) of 27 and 31ppt, respectively. Therefore, individuals with intermediate buoyancies might be selected against. This finding is consistent with our expectations, as Fjord ecotypes experience lower salinities compared to North Sea ecotypes. Interestingly, there was little evidence that the inversions on chromosomes 2, 7, and 12 impacted buoyancy. Overall, we provide experimental evidence of the relative contributions of ecotypic background and inversions on a key life history trait, which helps to explain the maintained divergence among ecotypes.
- Research Article
- 10.1038/s41598-026-53009-1
- May 13, 2026
- Scientific reports
- Mathieu Pruvot + 4 more
Wild pigs (Sus scrofa) have been expanding their range in Canada during recent decades. While high reproductive output is typically reported in wild pigs, reproductive performance at the northern edge of the North American invasive range is not well characterized. In this study, we describe the population structure and reproductive parameters of wild pigs in Alberta, Canada, estimate age of juvenile and sub-adults based on a Gompertz growth model fitted on wild pigs of known age from Texas, and test hypotheses regarding their reproductive ecology in this northern context. With a high proportion of reproductively active females (68%), and litter sizes comparable to other regions (median 5, IQR [4-8]), wild pigs in Alberta maintained a high reproductive potential. However, age estimates suggested low survival within the first year which may limit population growth in this region, potentially working in favor of on-going control program. Continued monitoring of wild pig population structure and reproductive parameters is essential to understanding population dynamics and planning control efforts in relation to changing environmental conditions.
- Research Article
- 10.1002/ajhb.70272
- May 1, 2026
- American journal of human biology : the official journal of the Human Biology Council
- Muhammad Irfan Malik
The maintenance of hygienic conditions and the provision of appropriate medical care during childbirth are essential to reduce the complications and infections that pose health risks to both expectant mothers and their newborn babies. Although skilled birth attendance (SBA) has been very much helpful in reducing the biological risks linked to childbirth, it is pertinent to mention that more than 34% of deliveries in Pakistan continue to take place at home without the proper medical care and supervision. The main objective of this proposed work is to explore the correlation between women's empowerment characterized by resources, agency, and relational autonomy and access to skilled birth attendance within the reproductive and sociocultural context of Pakistan. This study also considers the influence of various socioeconomic and demographic factors on this correlation. The present study utilizes the logistic regression models to estimate average marginal effects, using nationally representative data from currently married women aged 15-49 as mentioned in the Pakistan Demographic and Health Survey 2017-18. This analysis aims to evaluate the relative contributions of different empowerment dimensions, while simultaneously controlling for household and demographic characteristics. Women empowerment is a very important part of skilled birth attendance. The relationship between educational empowerment, digital access, specifically ownership of mobile phones and internet usage, and the rejection of norms related to domestic violence substantially enhances the probability of skilled birth attendance. This phenomenon exhibits effects that are comparable to those of conventional determinants, including household wealth and parity. Significant disparities persist in geographic and socio-economic contexts, adversely affecting mothers in Baluchistan, particularly those with larger families. This study demonstrates that the empowerment of women within the context of Pakistan's reproductive ecology is essential, rather than ancillary, for reducing biological risks associated with childbirth. It provides comparative insights that are pertinent to the fields of human biology and maternal health research.
- Research Article
- 10.1121/10.0043817
- May 1, 2026
- JASA express letters
- Sujay Balebail + 8 more
The long-duration advertisement "hums" produced by type I male plainfin midshipman fish (Porichthys notatus) are tonal signals consisting of a fundamental frequency and multiple harmonics. Using acoustic playback of synthetic, hum-like stimuli under natural conditions, we found that the fundamental frequency is likely required for females' final entry into a type I male's nest, whereas higher harmonics may play a limited role. We found no evidence of phonotaxis in type I and type II males. Additionally, the observed decrease in body size of captured individuals over the breeding season suggests that larger individuals migrate earlier to spawning sites.
- Research Article
- 10.3390/plants15091362
- Apr 29, 2026
- Plants
- Aleksandra V Stogova + 6 more
Phytolacca acinosa Roxb., a perennial herb native to East Asia, is increasingly naturalizing in Europe, yet its reproductive ecology in the secondary range remains poorly understood. This study evaluated seed productivity across central and edge populations in the secondary range, fruit and seed morphometrics, and germination responses to cold storage, acid scarification (simulating bird endozoochory), and light exposure. Fruit production per raceme was influenced by an interaction between insolation and range position: reduced insolation increased fruit set in central populations but decreased it at the range edge. Raceme number per shoot was lower in spontaneous plants compared to cultivated ones. Fresh seeds exhibited strong dormancy with no germination without scarification. Acid scarification significantly enhanced germination, particularly with light exposure, reaching up to 55%. Cold storage did not increase germination percentage but accelerated germination of scarified seeds under light, reducing median germination time from 24 to 21 days. Compared to the congeneric P. americana, P. acinosa shows more stringent dormancy requirements. We conclude that P. acinosa retains deep seed dormancy in its secondary range and relies on bird-mediated endozoochory for both dispersal and dormancy release. At the northern range edge, reduced plant vigor and lower raceme numbers are partially offset by increased flower production per raceme, though fruit set remains constrained. The species does not exhibit the simplified germination requirements often associated with successful invaders; instead, its invasion success appears driven by a bet-hedging strategy combining persistent seed banks with specific dormancy-breaking cues.
- Research Article
- 10.17129/botsci.3742
- Apr 21, 2026
- Botanical Sciences
- Guadalupe Amancio + 2 more
Annonaceae is a family of the order Magnoliales with a high species richness and a characteristic floral structure and composition. This lineage inhabits tropical forests around the world, and its evolutionary history has been addressed through the study of its reproductive mechanisms, mainly in Asian and South American species. In Mexico there are 76 reported species of which about 60 % are endemic, however, there are few studies of the reproductive ecology of these plants. In this review we examine the reproductive ecology of the Mexican Annonaceae, floral morphology and biology, phenology, pollination systems, and seed dispersal mechanisms. In general, Mexican Annonaceae have protogynous flowers, they are pollinated by insects, and few species of the genus Desmopsis have reproductive systems by cloning. Herbarium data and publications indicate that they have seasonal flowering patterns linked to rainfall. Beetles, flies, and ants can be the main pollinators, and seed dispersal depends on vertebrates. This review highlights the urgent need for in-depth study of reproductive ecology and specialized reproductive interactions in Mexico's threatened tropical systems.
- Research Article
- 10.1071/zo25064
- Apr 21, 2026
- Australian Journal of Zoology
- Joshua R Keen + 11 more
The Pilbara Olive Python (Liasis olivaceus barroni) is a large-bodied python endemic to the Pilbara and northern Gascoyne regions of Western Australia. Despite its national listing as Vulnerable, data on its morphology, sexual size dimorphism, and reproductive ecology remain limited. We present data from a 4-year monitoring and radio-tracking program comparing measurements from 100 individuals sampled across 12 locations. Allometrically scaled morphometric analyses found minimal evidence of sexual size dimorphism, with males and females showing largely overlapping body proportions. Breakpoint analyses of body mass relative to snout–vent length indicated that males reach sexual maturity at smaller sizes (~1320 mm SVL) than females (~1805 mm SVL). Observations of reproductive activity showed a defined seasonal cycle, with females ovulating in September, gravid in October, and brooding in refuges between October and February. Juveniles were most frequently detected from November to February, with neonate-sized individuals detected in January. Both natural (caves, rocky outcrops, burrows) and artificial shelters (engineered rock wall) were used for brooding, highlighting flexibility in reproductive behaviour. These findings provide the first population-level assessment of sexual size dimorphism and reproductive timing in this subspecies and offer critical baseline data to inform conservation management.
- Research Article
- 10.1007/s00338-026-02871-6
- Apr 20, 2026
- Coral Reefs
- Brett M Taylor + 4 more
punctuated by annual recruitment pulses of widely varying magnitude. Interannual variability in recruitment is likely driven by oceanographic conditions, fluctuations in resource availability, and associated changes in fecundity. The species is genetically distinct in the Mariana Islands, suggesting a self-recruiting population where recruitment failure could have significant local impacts. These findings underscore the importance of integrating place-based ecological knowledge with empirical life-history data to inform sustainable, culturally-grounded management of reef fisheries.
- Research Article
- 10.1002/njz2.70027
- Apr 14, 2026
- New Zealand Journal of Zoology
- L Grey + 3 more
Insects use sensilla to detect chemical and physical stimuli, mediating behaviours such as mate finding, foraging and mechanosensory responses. The distribution and density of sensilla can be examined using scanning electron microscopy. Investigating these structures can help elucidate rarely observed behaviours (e.g. mate finding and foraging). The mating behaviour of the Geodorcus stag beetles of New Zealand is not well understood, as most species (eight of ten) are listed as ‘threatened’, ‘at risk’ or ‘data deficient’ under the New Zealand Threat Classification System. We examined the most common species of Geodorcus , G. helmsi , and compared sensilla on the maxillary palps, antennae and legs of males and females. Male antennae were more densely packed with sensilla and had sensilla campaniformia, absent in females. Additionally, male antennae were larger than those of females, scaling proportionally with body size, suggesting that males have greater sensory capacity and are likely attracted to a pheromone the females emit. Our study sheds new insight into the sensory biology, mating behaviour and life history of G. helmsi . More broadly, it contributes to understanding the reproductive ecology of Geodorcus . Understanding how these beetles interact and locate mates can inform conservation management, for example, by guiding captive breeding programmes and pheromone‐based monitoring strategies for threatened Geodorcus species.
- Research Article
1
- 10.1098/rspb.2025.2728
- Apr 1, 2026
- Proceedings. Biological sciences
- Juliette Champenois + 5 more
Artificial light at night (ALAN) disrupts biological rhythms across taxa, yet behavioural mechanisms linking exposure to population consequences remain poorly understood. Long-distance migrants encounter ALAN across their annual cycle but remain understudied despite experiencing stronger performance impacts than sedentary species. We experimentally introduced ALAN into nest-boxes of a migratory passerine, the collared flycatcher (Ficedula albicollis), breeding in Gotland, Sweden. Nestlings were exposed to ALAN from 2 days post-hatching until fledging, and we video-recorded parental and nestling activity over 24 h on day 8 post-hatching. Across a high-resolution behavioural dataset (32 100 nestling and 3709 parental events), ALAN-exposed nestlings begged more frequently and for longer at night compared with dark controls, revealing disrupted circadian activity. These effects cascaded to parents: both females and males began feeding earlier and ceased later, while reducing hourly feeding rates relative to controls. Consequently, ALAN nestlings fledged at older ages, consistent with delayed development, though reproductive success (number of fledged offspring) was unaffected. Our study provides clear empirical support for socially mediated behavioural mechanisms whereby both parents respond to ALAN, disrupting family level coordination, developmental trajectories and parental investment. This temporal destabilization of social synchrony uncovers a novel ecological pathway through which anthropogenic light alters behavioural timekeeping and life-history trajectories in the wild.
- Research Article
- 10.4257/oeco.2026.3001.02
- Mar 13, 2026
- Oecologia Australis
- Amadeu Dos Santos-Neto + 2 more
Restinga ecosystems in Brazil are characterized by harsh environmental conditions and high biodiversity, yet many of their endemic plant species remain poorly studied. Understanding the reproductive ecology of these species is essential for developing effective conservation strategies. This study investigated the ecological and reproductive strategies of Myrcia ovina, an endemic and critically endangered shrub in Restinga. We monitored phenological events over a 12-month period and analyzed vegetative and reproductive patterns with circular statistics and correlations with climatic variables (temperature and precipitation). Observations also included vegetative behavior and potential ecological drivers of reproductive timing. Our findings indicate that Myrcia ovina is an evergreen species with continuous leaf exchange, which is an important trait for maintaining photosynthesis in nutrient-poor soils. The species exhibits a marked annual flowering peak during the dry season, likely aligning with pollinator availability and reducing herbivory pressure. Fruit development peaks at the beginning of the rainy season, when increased water availability may support seed maturation. However, seed dispersal may be less effective during this period due to reduced frugivore activity in the presence of more abundant food sources. The positive correlation between bud production and preceding precipitation highlights the role of historical moisture availability in reproductive success. These results demonstrate how Myrcia ovina is finely tuned to the seasonal dynamics of the Restinga and offer important insights into the adaptive strategies of Myrtaceae species in tropical coastal systems. This study contributes valuable knowledge for biodiversity conservation in these vulnerable ecosystems.
- Research Article
- 10.55041/ijsmt.v2i3.036
- Mar 10, 2026
- International Journal of Science, Strategic Management and Technology
- Dr.Hema Makne
Climate change is increasingly recognized as one of the most important drivers of biodiversity change worldwide. Birds are particularly sensitive to environmental variations because their life cycles, including breeding and feeding behaviour, are closely associated with climatic conditions. The Indian peafowl (Pavo cristatus), the national bird of India, is widely distributed across forests, agricultural landscapes, and rural ecosystems throughout the country. Although the species is currently categorized as “Least Concern,” recent studies indicate that environmental changes may influence its behaviour, habitat use, and reproductive ecology. Reproductive activities in birds are strongly governed by climatic factors such as rainfall patterns, temperature fluctuations, vegetation growth, and food availability. In India, the breeding season of the Indian peafowl generally coincides with the monsoon period when ecological conditions are favourable for reproduction and chick survival (Yogeshwari & Varunprasath, 2020). However, increasing climatic variability, irregular rainfall patterns, and habitat modification may disrupt these ecological relationships. This review synthesizes recent research (2020–2026) focusing on the reproductive ecology of Pavo cristatus in India and discusses how climate change may influence breeding behaviour, nesting success, and population dynamics. The review also highlights important research gaps and emphasizes the need for long-term ecological monitoring and conservation strategies to ensure the sustainability of peafowl populations in changing environmental conditions.
- Research Article
- 10.1038/s41597-026-06554-3
- Mar 2, 2026
- Scientific data
- Guiliang Xin + 6 more
Bischofia polycarpa (2n = 68), belonging to Phyllanthaceae family, is a native deciduous tree with naturally distribution ranging from southern Qinling Mountains and Huaihe River basin to the northern regions of Fujian and Guangdong, China. It holds significant horticultural, ornamental, and medicinal value and serves as a crucial winter food resource for wild birds. Herein, we report a de novo genome assembly for B. polycarpa, utilizing a combination of PacBio HiFi Reads and Hi-C data. In total, the genome size reaches 585.68 Mb with a contig N50 of 12.62 Mb, and 99.06% (580.18 Mb) of the assembly successfully anchored on 34 chromosomes. The genome comprises approximately 62.77% repetitive sequences and 32,554 protein-coding genes, of which 96.15% could be functionally annotated. The BUSCO analysis reveals a genome completeness of 95.42% (n = 1,540), including 1,499 (92.87%) single-copy BUSCOs and 41 (2.54%) duplicated BUSCOs. This high-quality genome of the Phyllanthaceae enriches our understanding of the genetic underpinnings of plant reproductive ecology.
- Research Article
1
- 10.1002/ajhb.70225
- Mar 1, 2026
- American journal of human biology : the official journal of the Human Biology Council
- R G Bribiescas + 1 more
The research field of reproductive ecology continues to be a major contributor to the scientific advancement of evolutionary anthropology and human biology in general. Primary contributions to human evolutionary biology include a greater understanding of the physiological mechanisms that manage lifetime reproductive effort, resource allocation, life history trade-offs, demographic variation in fertility, the adaptive traits that define humans (Homo sapiens), non-human primates, and our hominid ancestors as well as novel insights into reproductive health challenges such as cancer. Here we present a brief overview of the foundation on which this research path is based, including a summary of current research advances in human reproductive ecology, particularly within the scope of human variation. Future research directions, unanswered questions, and engagement with reproductive health challenges are discussed.
- Research Article
- 10.1016/j.pld.2026.03.010
- Mar 1, 2026
- Plant Diversity
- Yingqun Feng + 4 more
Climate modulation of phylogenetic and functional constraints on fruit volume: A case study of Chinese angiosperms
- Research Article
- 10.1002/ece3.73131
- Mar 1, 2026
- Ecology and evolution
- Benjamin Marcy-Quay + 1 more
Pedigree-based estimation methods leverage the fact that each offspring in a cohort is genotypically "marked" by its parents and represent a recent and promising toolset for estimating population dynamics. This includes pedigree accumulation estimators that model the "accumulation" of inferred unique parents within a given cohort to estimate parental abundance. Unlike close-kin mark-recapture approaches, which rely on intercohort comparisons, pedigree accumulation modeling can be completed solely using intracohort samples. This is particularly advantageous for semelparous species, where intercohort pairs are impossible and adult life stages can be difficult to sample without affecting their likelihood of successfully reproducing. Previous work has evaluated a range of estimators for such datasets, concluding that the non-parametric Chao estimator provides the most accurate and precise estimates for feasible levels of sampling effort. We used simulated data to evaluate an alternative estimator based on hierarchical modeling and data augmentation in a Bayesian framework. Results indicate that estimates from the hierarchical Bayesian estimator had comparable accuracy and better precision than both the previously tested Chao1 estimator and the improved iChao formulation across a range of sample sizes and sex ratios. Furthermore, the Bayesian estimator was far more robust to simulated errors in pedigree reconstruction, especially the presence of false negatives. Hierarchical Bayesian pedigree accumulation models can also provide additional insight into underlying reproductive ecology through their use of an explicit observation process, allowing for the incorporation or estimation of species- and population-specific reproductive dynamics. More broadly, the parametric nature of these models offers opportunities to efficiently pool information among datasets as well as to propagate uncertainty within more complex models.