Daniel W. Franks’s research while affiliated with New York University and other places

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Publications (115)


Analysis of sparse animal social networks
  • Preprint
  • File available

November 2024

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51 Reads

Helen Kate Mylne

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Colin M. Beale

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Daniel W. Franks

Low-density social networks can be common in animal societies, even among species generally considered to be highly social. Social network analysis is commonly used to analyse animal societal structure, but edge weight (strength of association between two individuals) estimation methods designed for dense networks can produce biased measures when applied to low-density networks. Frequentist methods suffer when data availability is low, because they contain an inherent flat prior that will accept any possible edge weight value, and contain no uncertainty in their output. Bayesian methods can accept alternative priors, so can provide more reliable edge weights that include a measure of uncertainty, but they can only reduce bias when sensible prior values are selected. Currently, neither accounts for zero-inflation, so they produce edge weight estimates biased towards stronger associations than the true social network, which can be seen through diagnostic plots of data quality against output estimate. We address this by adding zero-inflation to the model, and demonstrate the process using group-based data from a population of male African savannah elephants. We show that the Bayesian approach performs better than the frequentist to reduce the bias caused by these problems, though the Bayesian requires careful consideration of the priors. We recommend the use of a Bayesian framework, but with a conditional prior that allows the modelling of zero-inflation. This reflects the fact that edge weight derivation is a two-step process: i) probability of ever interacting, and ii) frequency of interaction for those who do. Additional conditional priors could be added where the biology requires it, for example in a society with strong community structure, such as female elephants in which kin structure would create additional levels of social clustering. Although this approach was inspired by reducing bias observed in sparse networks, it could have value for networks of all densities.

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Schematic diagram of the PhytNet convolutional neural network. (A) PhytNet convolutional block design incorporated into PhytNet as sequential blocks in layers 1 and 2. (B) Full PhytNet end‐to‐end design. Layer 1 and layer 2 are composed of a predefined number of sequential convolutional blocks. The number of channels and kernel size of all convolutional layers should be optimised for a given dataset, but these optimised values are constant within layer 1 and layer 2, respectively. The stride values of layer 1 and layer 2 are 1 and 2, respectively. ReLU, rectified linear unit.
Distributions of non‐photochemical quenching (NPQt) (A) and photosynthetic yield (Phi2) (B) of cocoa trees with different disease states. Box plots show the interquartile range with whiskers at 1.5 times the interquartile range from the first and third quartiles. Raw data points are plotted as white circles. Measurements were taken from cocoa trees in five disease states using a MultispeQ v2.0. Black pod rot n = 10, Frosty pod rot n = 5, Healthy n = 9, Witches' broom (adjacent leaf) n = 5, Witches' broom (affected leaf) n = 5.
Mean reflectance spectrum measurements (±1 standard error) (A) and feature importance scores (B) derived from a random forest classifier trained on the same data. Spectroscopy data were gathered from cocoa pods that were either healthy or showing early to middle stage frosty pod rot or black pod rot symptoms. The standard error is shown by grey shading, and the dotted lines at 400 and 720 nm show the bounds of the human‐visible spectrum. Random forest classifier training accuracy = 95.45%; test accuracy = 78.26%.
Infrared images with class activation heatmaps produced using Grad‐CAM and four convolutional neural networks. The models used are PhytNet, ResNet18, EfficientNet‐b0, and EfficientNet‐V2s (left to right). The leftmost column shows raw input images with ground truth labels in white; other white labels are predicted by each model.
Violin and box plots of 10‐fold cross‐validation results for PhytNet and ResNet18 trained (A) and validated (B) on infrared or RGB images of cocoa disease. Shown here is the Gaussian density function, median and interquartile ranges for mean F1, per class F1, precision, and recall. PhytNet was trained only on infrared data, while ResNet18 was trained on infrared or RGB data. The datasets had four classes: Black pod rot (BPR), Frosty pod rot (FPR), Healthy, and Witches' broom disease (WBD). n = 70 images per class of early to midstage diseased or healthy cocoa with a 90%:10% train:validation split.
Tailoring convolutional neural networks for custom botanical data

October 2024

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22 Reads

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2 Citations

Premise Automated disease, weed, and crop classification with computer vision will be invaluable in the future of agriculture. However, existing model architectures like ResNet, EfficientNet, and ConvNeXt often underperform on smaller, specialised datasets typical of such projects. Methods We address this gap with informed data collection and the development of a new convolutional neural network architecture, PhytNet. Utilising a novel dataset of infrared cocoa tree images, we demonstrate PhytNet's development and compare its performance with existing architectures. Data collection was informed by spectroscopy data, which provided useful insights into the spectral characteristics of cocoa trees. Cocoa was chosen as a focal species due to the diverse pathology of its diseases, which pose significant challenges for detection. Results ResNet18 showed some signs of overfitting, while EfficientNet variants showed distinct signs of overfitting. By contrast, PhytNet displayed excellent attention to relevant features, almost no overfitting, and an exceptionally low computation cost of 1.19 GFLOPS. Conclusions We show that PhytNet is a promising candidate for rapid disease or plant classification and for precise localisation of disease symptoms for autonomous systems. We also show that the most informative light spectra for detecting cocoa disease are outside the visible spectrum and that efforts to detect disease in cocoa should be focused on local symptoms, rather than the systemic effects of disease.


The evolution of menopause in toothed whales
a, Chronological trimmed phylogenies of Odontocete cetaceans (derived from ref. ³⁵) showing species with lifespan data calculated by and included in this study. The phylogeny shows 1,000 overlapping alternative tree structures. The four independent transitions to menopause are shown with red branches and all branches without menopause are shown in blue. Species names marked with a filled point are those for which we also have estimates of reproductive lifespan derived from ovarian corpora. b, The relationship between body length and ordinary maximum lifespan (age at which 90% of adult years have been lived) in female toothed whales with lifespan data (n = 32). Points (red with menopause; blue without menopause) show the estimated mean value, with error (s.d.) in both the size and lifespan dimensions; this error is carried through the analysis (all points labelled: Supplementary Fig. 1a). The line shows the predicted relationship between body length and lifespan in whales without menopause (posterior mean ± 50% CI (darker ribbon) and ±95% CI (lighter ribbon)). Species with menopause have longer lifespans than expected given their size. c, The relationship between reproductive lifespan and body length in toothed whale species with reproductive lifespan data (n = 18, filled points in a). Points, lines and ribbons as for b. Reproductive lifespan is the age at which ovarian activity is predicted to cease. Species with menopause (red) do not have shorter reproductive lifespans than expected given their size (all points labelled: Supplementary Fig. 1b).
Opportunities for intergenerational help and harm in toothed whales
a,b, Relative grandmother years (a) and mother–daughter reproductive overlap (b) for toothed whale species (n = 18; Fig. 1a ‘filled point’ species) with (red) and without (blue) menopause. Relative grandmother years represent the number of grandoffspring years an adult female can expect to experience, scaled to the species-specific age at maturity. Reproductive overlap is the summed proportion of adult female reproduction that occurs concurrently with the reproduction of a daughter. The left-hand panels show estimates of both grandmother years and reproductive overlap and are derived from 1,000 kinship demography models with parameters derived from the posterior distributions of the lifespan and reproductive lifespan models applied to each species. Points in this panel show the mean estimate of the parameter, whereas the thick and thin error bars show the 50% and 95% CIs, respectively. The right-hand panels show the distribution of posterior estimates from a Bayesian model (Model post.) estimating the mean demographic parameter expected over all species with and without menopause; points show the mean estimate. Species with menopause have more grandmother years but the same reproductive overlap as species without menopause.
Help and harm implications of life history strategies for species with menopause
Comparing the estimated relative grandmother years and reproductive overlap for all five toothed whale species with menopause as: observed in real systems (observed, red areas); in their simulated ancestral non-menopausal state (ancestral, blue areas); and a simulated analogue of the species that keeps reproducing to the end of life (slow life history, yellow areas). Contoured areas show the distribution of the posterior distributions under each state. Kinship demographic models are run separately for each species-state but, for illustration, posterior estimates have been combined and z-scored in species to get comparable between-species estimates. In the menopause state (red) females have more grandmother years but the same reproductive overlap as their ancestral state (blue) and a lower reproductive overlap than in the slow life history state (yellow).
The evolution of menopause in toothed whales

March 2024

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205 Reads

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13 Citations

Nature

Understanding how and why menopause has evolved is a long-standing challenge across disciplines. Females can typically maximize their reproductive success by reproducing for the whole of their adult life. In humans, however, women cease reproduction several decades before the end of their natural lifespan1,2. Although progress has been made in understanding the adaptive value of menopause in humans3,4, the generality of these findings remains unclear. Toothed whales are the only mammal taxon in which menopause has evolved several times⁵, providing a unique opportunity to test the theories of how and why menopause evolves in a comparative context. Here, we assemble and analyse a comparative database to test competing evolutionary hypotheses. We find that menopause evolved in toothed whales by females extending their lifespan without increasing their reproductive lifespan, as predicted by the ‘live-long’ hypotheses. We further show that menopause results in females increasing their opportunity for intergenerational help by increasing their lifespan overlap with their grandoffspring and offspring without increasing their reproductive overlap with their daughters. Our results provide an informative comparison for the evolution of human life history and demonstrate that the same pathway that led to menopause in humans can also explain the evolution of menopause in toothed whales.


Group size modulates kinship dynamics and selection on social traits

December 2023

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137 Reads

The mean relatedness of an individual to others in its social group may change with age. Such kinship dynamics, which are expected to influence sex- and age-specific social behaviours, are sensitive to demography. Existing models of kinship dynamics, however, have focused chiefly on the influences of population-level demographic parameters (e.g., sex-specific dispersal rate and mating pattern). Here, we extend these models to explore the effects of local group size on kinship dynamics and the resulting patterns of selection for sex- and age-specific helping and harming. We show that individuals' average relatedness to others is higher in smaller than larger groups, and consequently, selective pressure for helping or harming is generally stronger in smaller than larger groups. Moreover, relatedness changes faster with age in smaller groups, particularly at younger ages. These patterns favour more extreme social traits in smaller groups and especially so at earlier life history stages. In particular we highlight that, while social systems characterized by bi-sexual philopatry with non-local mating (e.g., whales) are known to favour a shift from more helpful to more harmful behaviour as females age, a trend that has been invoked to explain the evolution of menopause and post-reproductive helping, the timing of this shift is predicted to be sensitive to local group size, potentially favouring earlier female reproductive cessation in smaller groups. Our study generates new insights into the effects of local group properties on kinship dynamics, and suggests that such effects might also help to explain widespread variation in age-linked social traits, such as the timing of menopause in social mammals.


Computer vision for plant pathology: A review with examples from cocoa agriculture

December 2023

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144 Reads

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7 Citations

Plant pathogens can decimate crops and render the local cultivation of a species unprofitable. In extreme cases this has caused famine and economic collapse. Timing is vital in treating crop diseases, and the use of computer vision for precise disease detection and timing of pesticide application is gaining popularity. Computer vision can reduce labour costs, prevent misdiagnosis of disease, and prevent misapplication of pesticides. Pesticide misapplication is both financially costly and can exacerbate pesticide resistance and pollution. Here, we review the application and development of computer vision and machine learning methods for the detection of plant disease. This review goes beyond the scope of previous works to discuss important technical concepts and considerations when applying computer vision to plant pathology. We present new case studies on adapting standard computer vision methods and review techniques for acquiring training data, the use of diagnostic tools from biology, and the inspection of informative features. In addition to an in‐depth discussion of convolutional neural networks (CNNs) and transformers, we also highlight the strengths of methods such as support vector machines and evolved neural networks. We discuss the benefits of carefully curating training data and consider situations where less computationally expensive techniques are advantageous. This includes a comparison of popular model architectures and a guide to their implementation.


BISoN: A Bayesian Framework for Inference of Social Networks

July 2023

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48 Reads

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29 Citations

Animal social networks are often constructed from point estimates of edge weights. In many contexts, edge weights are inferred from observational data, and the uncertainty around estimates can be affected by various factors. Though this has been acknowledged in previous work, methods that explicitly quantify uncertainty in edge weights have not yet been widely adopted and remain undeveloped for many common types of data. Furthermore, existing methods are unable to cope with some of the complexities often found in observational data, and do not propagate uncertainty in edge weights to subsequent statistical analyses. We introduce a unified Bayesian framework for modelling social networks based on observational data. This framework, which we call BISoN , can accommodate many common types of observational social data, can capture confounds and model effects at the level of observations and is fully compatible with popular methods used in social network analysis. We show how the framework can be applied to common types of data and how various types of downstream statistical analyses can be performed, including non‐random association tests and regressions on network properties. Our framework opens up the opportunity to test new types of hypotheses, make full use of observational datasets, and increase the reliability of scientific inferences. We have made both an R package and example R scripts available to enable adoption of the framework.


Postreproductive female killer whales reduce socially inflicted injuries in their male offspring

July 2023

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69 Reads

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3 Citations

Current Biology

Understanding the evolution of menopause presents a long-standing scientific challenge1,2,3-why should females cease ovulation prior to the end of their natural lifespan? In human societies, intergenerational resource transfers, for example, food sharing and caregiving, are thought to have played a key role in the evolution of menopause, providing a pathway by which postreproductive females can boost the fitness of their kin.4,5,6 To date however, other late-life contributions that postreproductive females may provide their kin have not been well studied. Here, we test the hypothesis that postreproductive female resident killer whales (Orcinus orca) provide social support to their offspring by reducing the socially inflicted injuries they experience. We found that socially inflicted injuries, as quantified by tooth rake marks, are lower for male offspring in the presence of their postreproductive mother. In contrast, we find no evidence that postreproductive mothers reduce rake marking in their daughters. Similarly, we find no evidence that either reproductive mothers or grandmothers (reproductive or postreproductive) reduce socially inflicted injuries in their offspring and grandoffspring, respectively. Moreover, we find that postreproductive females have no effect on reducing the rake marks for whales in their social unit who are not their offspring. Taken together, our results highlight that directing late-life support may be a key pathway by which postreproductive females transfer social benefits to their male offspring.


Temporal dynamics of mother-offspring relationships in Bigg's killer whales: opportunities for kin-directed help by post-reproductive females

June 2023

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199 Reads

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12 Citations

Age-related changes in the patterns of local relatedness (kinship dynamics) can be a significant selective force shaping the evolution of life history and social behaviour. In humans and some species of toothed whales, average female relatedness increases with age, which can select for a prolonged post-reproductive lifespan in older females due to both costs of reproductive conflict and benefits of late-life helping of kin. Killer whales (Orcinus orca) provide a valuable system for exploring social dynamics related to such costs and benefits in a mammal with an extended post-reproductive female lifespan. We use more than 40 years of demographic and association data on the mammal-eating Bigg's killer whale to quantify how mother-offspring social relationships change with offspring age and identify opportunities for late-life helping and the potential for an intergenerational reproductive conflict. Our results suggest a high degree of male philopatry and female-biased budding dispersal in Bigg's killer whales, with some variability in the dispersal rate for both sexes. These patterns of dispersal provide opportunities for late-life helping particularly between mothers and their adult sons, while partly mitigating the costs of mother-daughter reproductive conflict. Our results provide an important step towards understanding why and how menopause has evolved in Bigg's killer whales.


Bayesian inference of toothed whale lifespans

February 2023

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112 Reads

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2 Citations

Accurate measures of lifespan and age-specific mortality are important both for understanding life-history evolution and informing conservation and population management strategies. The most accurate data to estimate lifespan are from longitudinal studies, but for many species - especially those such as toothed whales that are wide-ranging and live in difficult-to-access environments - these longitudinal data are not available. However, other forms of age-structured data - such as from mass-strandings - are available for many toothed species, and using these data to infer patterns of age-specific mortality and lifespan remains an important outstanding challenge. Here we develop and test a Bayesian mortality model to derive parameters of a mortality function from age-structured data while accounting for potential error introduced to these data by mistakes in age estimation, sampling biases and population growth. We then searched the literature to assemble a database of 269 published age-structured toothed whale datasets. We applied our mortality model to derive lifespan estimates for 32 species of female and 33 species of male toothed whale. We also use our model to characterise sex differences in lifespan in toothed whales. Our mortality model allows us to curate the most complete and accurate collection of toothed whale lifespans to date.


The Evolution of Democratic Peace

February 2023

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89 Reads

A major goal in evolutionary biology is to elucidate common principles that drive human and other animal societies towards war or peace. One idea proposed to account for differences between human societies is the "democratic peace" hypothesis, which suggests autocracies are more warlike than democracies because leaders can pursue fights for private gain. Given that animals can make autocratic or democratic collective decisions, we consider whether the logic of democratic peace may apply across taxa. We adapt the classic Hawk-Dove model to consider conflict decisions by groups rather than individuals, finding support for the democratic peace hypothesis without mechanisms involving complex human institutions. We suggest that the degree to which collective decisions are shared may explain variation in the intensity of intergroup conflict in nature.


Citations (69)


... Comparative biology is another approach; for example, Finch and Holmes (2010) argue that postreproductive survival is not limited to organisms with substantial kin networks or especially long lives. Ellis et al. (2018Ellis et al. ( , 2024 have begun to quantify post-reproductive survival across animal species, a promising approach. ...

Reference:

Evolution of the Human Life Cycle, Revisited
The evolution of menopause in toothed whales

Nature

... Computer vision projects in areas like plant pathology and agronomy often have limited data for training and are thus constrained in the choice of model architecture. ResNet (He et al., 2016), EfficientNet (Tan and Le, 2020), and ConvNeXt variants are among the best current neural networks available for such smaller datasets (Sykes et al., 2024). However, these models were developed for general image recognition tasks and are honed to perform well on huge benchmark datasets such as ImageNet, which has 1.4 million images and 1000 classes (Woo et al., 2023). ...

Computer vision for plant pathology: A review with examples from cocoa agriculture

... As long as the benefit to overall reproductive fitness of the lineage outweighs the cost of the female producing no further progeny at late ages, then the menopause can be selectively advantageous. Consistent with the Grandmother hypothesis, female reproductive cessation and postreproductive caregiving has now been observed in multiple species, including humans and killer whales (Hawkes and Coxworth, 2013;Croft et al., 2017;Grimes et al., 2023;Nielsen et al., 2023). It is possible to consider this mechanism in terms of an AB replicator model. ...

Postreproductive female killer whales reduce socially inflicted injuries in their male offspring
  • Citing Article
  • July 2023

Current Biology

... We constructed an influence network for sleep disruption across our dataset, where every individual is represented as a node, and directed edges go from the influencer to the influenced individuals, with an edge value equal to the peak cross correlation. To test whether dominance predicts influence, we performed nodal regression (Hart et al. 2023) by calculating the proportion of interactions in which the focal individual influences others (was a leader, see above). We then fitted a logistic regression with individual dominance and sex as predictors, and the proportion of influenced individuals as response. ...

BISoN: A Bayesian Framework for Inference of Social Networks
  • Citing Article
  • July 2023

... En la comunidad de delfines, la orca, a pesar de tener un dimorfismo sexual marcado no hubo diferencias significativas. A pesar de ello, el solapamiento de nicho isotópico entre hembras y machos fue el más bajo (23%) y el tamaño de la elipse corregida (SEAc) fue tres veces más grande de los machos que de las hembras.En el caso de los machos de orcas, algunos tienden a dejar sus grupos y realizar movimientos de forma individual para integrarse a otros grupos, con esto disminuir la competencia sexual intragrupal y aumentar la probabilidad de encontrar pareja(Towers et al., 2019;Nielsen et al., 2023). Además, los machos también abandonan sus grupos natales para buscar alimento de forma particular(Santos y Neto, 2005;Ford, 2009). ...

Temporal dynamics of mother-offspring relationships in Bigg's killer whales: opportunities for kin-directed help by post-reproductive females

... For each species, we also performed opportunistic searches by following references through the literature and identifying potential data sources using the relevant chapters in authoritative edited collections [71][72][73][74] . From our search, we identified and extracted 75 sex-specific age data into a database 76 . In the database, where possible, each dataset represents data from a single population collected at a single time; however, this is not always possible and some datasets represent data collected from several populations or over a longer timescale. ...

Bayesian inference of toothed whale lifespans

... Also, focusing on the group and not individuals prevents investigating other factors potentially influencing observations of some behaviors (Karniski et al., 2015) such as age, sex, kinship, or individual preferences. Sex differences in killer whales has not been studied in details yet, and besides striking differences in maternal investment for resident killer whales (Weiss et al., 2023), other influences of sex on the behaviors of these animals remain unclear. For example, whereas sex differences seem to exist in the hunting techniques displayed by Crozet Island killer whales as females primarily beach in the intentional stranding foraging strategy (Guinet, 1991), both sexes use this stranding method off Argentina (Lopez and Lopez, 1985). ...

Costly lifetime maternal investment in killer whales
  • Citing Article
  • December 2022

Current Biology

... By generating a distribution of possible networks from the observed data rather than a single network, BISoN (Bayesian Inference of Social Networks) allows us to account for this uncertainty, which can drastically affect the performance of statistical models [68]. Specifically, for each group-year, we fitted a Bayesian 'edge model' with a count conjugate prior to our observed network data, which returns a posterior distribution of edge weights for each dyad in our network rather than a point estimate (for more information see [68,69]). We extracted 1000 draws from this posterior distribution and used these draws as the social networks over which we modelled pathogen spread. ...

bisonR - Bayesian Inference of Social Networks with R
  • Citing Preprint
  • November 2022

... Social network data is inherently relational, violating the assumption of independence required for most statistical tests 61 . To control for non-independence in the data, we used parametric regression in lieu of permutation-based tests, given our centrality metrics were regressed against nodal covariates, calves were sampled equally through automated observational methods, and were restricted to their respective group pens 62 . Assumptions (i.e., residual normality distribution plots) were visually checked using the performance package 63 . ...

Common Permutation Methods in Animal Social Network Analysis Do Not Control for Non-independence

Behavioral Ecology and Sociobiology

... However, in societies that feature multiple reproductive females, the ability of males to monopolize reproduction is inherently limited. Moreover, the fitness costs and benefits of suppression behaviour, and attempts to evade suppression, are likely to vary systematically over the lifetime depending on kinship dynamics [96,97] as relatedness to females and males in the group changes. In many social animals, we would expect ARTs to be adopted in a flexible manner over the life course depending on individual condition during development and relative competitive ability as shown here. ...

Patterns and consequences of age-linked change in local relatedness in animal societies

Nature Ecology & Evolution