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Adv."],"published-print":{"date-parts":[[2026,10,16]]},"abstract":"<jats:p>Quantifying biomorphological evolution and integrating phenotypic data into large-scale ecological and evolutionary studies are often limited by the scalability, subjectivity, and poor interpretability of existing methods. We introduce HAIKA (Hierarchical Attention-based Interpretable K-dimensional Autoencoder), a self-supervised residual channel-attention autoencoder that extracts continuous morphological representations from unlabeled standardized images and highlights the image regions that contribute to those representations. Applied to insect forewings across broad pterygote lineages, HAIKA outperformed most baseline deep learning models and conventional descriptors in clustering, dimensionality reduction, and morphological embedding. When applied to Hymenoptera (i.e., ants, bees, and wasps), the HAIKA-derived continuous characters improved morphology-based phylogenetic reconstruction and illuminated the evolutionary dynamics of wing shape and venation. Our results reveal that wing morphological evolution in Hymenoptera proceeds at decoupled rates across shape and venation modules and is not consistently aligned with key ecological transitions such as parasitoidism or eusociality. HAIKA provides a scalable, image-based route for phenotypic analyses, broadly applicable across diverse taxa and to both 2D and 3D datasets.<\/jats:p>","DOI":"10.1126\/sciadv.aef0605","type":"journal-article","created":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T17:59:24Z","timestamp":1792000764000},"update-policy":"https:\/\/doi.org\/10.34133\/aaas_crossmark","source":"Crossref","is-referenced-by-count":0,"title":["Deep learning reveals ecological decoupling and rate heterogeneity in Hymenoptera wing evolution"],"prefix":"10.1126","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3813-1802","authenticated-orcid":true,"given":"Ancheng","family":"Peng","sequence":"first","affiliation":[{"name":"Institut de Syst\u00e9matique, \u00c9volution, Biodiversit\u00e9 (UMR 7205), Mus\u00e9um national d\u2019Histoire naturelle, CNRS, SU, EPHE-PSL, UA, Paris F-75005, France."},{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and 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Neff The Solitary Bees: Biology Evolution Conservation (Princeton Univ. Press 2019).","DOI":"10.2307\/j.ctvd1c929"},{"key":"e_1_3_2_78_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ento.45.1.121"},{"key":"e_1_3_2_79_2","doi-asserted-by":"publisher","DOI":"10.1134\/S0013873810060047"},{"key":"e_1_3_2_80_2","doi-asserted-by":"publisher","DOI":"10.1111\/brv.12719"},{"key":"e_1_3_2_81_2","unstructured":"B. Epstein R. Meir Generalization bounds for unsupervised and semi-supervised learning with autoencoders. arXiv:1902.01449 [stat.ML] (2019)."},{"key":"e_1_3_2_82_2","doi-asserted-by":"crossref","unstructured":"Y. Liang X. Li X. Huang Z. Zhang Y. Yao \u201cAn automated data mining framework using autoencoders for feature extraction and dimensionality reduction\u201d in 2024 4th International Conference on Electronic Information Engineering and Computer Communication (EIECC 2024) pp. 710\u2013714.","DOI":"10.1109\/EIECC64539.2024.10929475"},{"key":"e_1_3_2_83_2","first-page":"14357","article-title":"General E(2)-equivariant steerable CNNs","volume":"32","author":"Weiler M.","year":"2019","unstructured":"M. Weiler, G. Cesa, General E(2)-equivariant steerable CNNs. Adv. Neural Inf. Process. Syst. 32, 14357\u201314368 (2019).","journal-title":"Adv. Neural Inf. Process. Syst."},{"key":"e_1_3_2_84_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-45362-4"},{"key":"e_1_3_2_85_2","first-page":"6840","article-title":"Denoising diffusion probabilistic models","volume":"33","author":"Ho J.","year":"2020","unstructured":"J. Ho, A. Jain, P. Abbeel, Denoising diffusion probabilistic models. Adv. Neural Inf. Process. Syst. 33, 6840\u20136851 (2020).","journal-title":"Adv. Neural Inf. Process. Syst."},{"key":"e_1_3_2_86_2","doi-asserted-by":"crossref","unstructured":"T. Karras S. Laine M. Aittala J. Hellsten J. Lehtinen T. Aila \u201cAnalyzing and improving the image quality of StyleGAN\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2020) pp. 8107\u20138116.","DOI":"10.1109\/CVPR42600.2020.00813"},{"key":"e_1_3_2_87_2","doi-asserted-by":"crossref","unstructured":"R. Rombach A. Blattmann D. Lorenz P. Esser B. Ommer \u201cHigh-resolution image synthesis with latent diffusion models\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2022) pp. 10684\u201310695.","DOI":"10.1109\/CVPR52688.2022.01042"},{"key":"e_1_3_2_88_2","doi-asserted-by":"crossref","unstructured":"A. Kirillov E. Mintun N. Ravi H. Mao C. Rolland L. Gustafson T. Xiao S. Whitehead A. C. Berg W.-Y. Lo P. Doll\u00e1r R. Girshick \u201cSegment anything\u201d in Proceeding IEEE\/CVF International Conference on Computer Vision (IEEE 2023) pp. 3992\u20134003.","DOI":"10.1109\/ICCV51070.2023.00371"},{"key":"e_1_3_2_89_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1127647"},{"key":"e_1_3_2_90_2","unstructured":"D. P. Kingma M. Welling \u201cAuto-encoding variational Bayes\u201d in Conference proceedings: papers accepted to the International Conference on Learning Representations (ICLR 2014)."},{"key":"e_1_3_2_91_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-022-01541-z"},{"key":"e_1_3_2_92_2","unstructured":"S. Ioffe C. Szegedy \u201cBatch normalization: Accelerating deep network training by reducing internal covariate shift\u201d in Proceeding 32nd International Conference Machine Learning (2015) pp. 448\u2013456."},{"key":"e_1_3_2_93_2","unstructured":"V. Nair G. E. Hinton \u201cRectified linear units improve restricted Boltzmann machines\u201d in Proceeding 27th International Conference Machine Learning (2010) pp. 807\u2013814."},{"key":"e_1_3_2_94_2","unstructured":"M. Lin Q. Chen S. Yan \u201cNetwork in network\u201d in Conference proceedings: papers accepted to the International Conference on Learning Representations (ICLR 2014)."},{"key":"e_1_3_2_95_2","unstructured":"I. Loshchilov F. Hutter \u201cDecoupled weight decay regularization\u201d in 7th International Conference on Learning Representations (ICLR 2019)."},{"key":"e_1_3_2_96_2","unstructured":"C. M. Bishop Pattern Recognition and Machine Learning (Springer 2006)."},{"key":"e_1_3_2_97_2","doi-asserted-by":"publisher","DOI":"10.1080\/00401706.1970.10488634"},{"key":"e_1_3_2_98_2","doi-asserted-by":"crossref","unstructured":"Z. Wu S. Song A. Khosla F. Yu L. Zhang X. Tang J. Xiao \u201c3D ShapeNets: A deep representation for volumetric shapes\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2015) pp. 1912\u20131920.","DOI":"10.1109\/CVPR.2015.7298801"},{"key":"e_1_3_2_99_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41597-024-04261-5"},{"key":"e_1_3_2_100_2","first-page":"281","article-title":"Some methods for classification and analysis of multivariate observations","volume":"1","author":"MacQueen J.","year":"1967","unstructured":"J. MacQueen, Some methods for classification and analysis of multivariate observations. Berkeley Symp. Math. Stat. Probab. 1, 281\u2013297 (1967).","journal-title":"Berkeley Symp. Math. Stat. Probab."},{"key":"e_1_3_2_101_2","doi-asserted-by":"publisher","DOI":"10.1080\/14786440109462720"},{"key":"e_1_3_2_102_2","first-page":"2579","article-title":"Visualizing data using t-SNE","volume":"9","author":"Van der Maaten L.","year":"2008","unstructured":"L. Van der Maaten, G. Hinton, Visualizing data using t-SNE. J. Mach. Learn. Res. 9, 2579\u20132605 (2008).","journal-title":"J. Mach. Learn. Res."},{"key":"e_1_3_2_103_2","doi-asserted-by":"publisher","DOI":"10.1016\/0377-0427(87)90125-7"},{"key":"e_1_3_2_104_2","doi-asserted-by":"publisher","DOI":"10.1080\/03610927408827101"},{"key":"e_1_3_2_105_2","doi-asserted-by":"publisher","DOI":"10.1109\/TPAMI.1979.4766909"},{"key":"e_1_3_2_106_2","unstructured":"K. Simonyan A. Zisserman \u201cVery deep convolutional networks for large-scale image recognition\u201d in 3rd International Conference on Learning Representations (ICLR 2015)."},{"key":"e_1_3_2_107_2","doi-asserted-by":"crossref","unstructured":"K. He X. Zhang S. Ren J. Sun \u201cDeep residual learning for image recognition\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2016) pp. 770\u2013778.","DOI":"10.1109\/CVPR.2016.90"},{"key":"e_1_3_2_108_2","doi-asserted-by":"crossref","unstructured":"G. Huang Z. Liu L. Van Der Maaten K. Q. Weinberger \u201cDensely connected convolutional networks\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2017) pp. 4700\u20134708.","DOI":"10.1109\/CVPR.2017.243"},{"key":"e_1_3_2_109_2","doi-asserted-by":"crossref","unstructured":"F. Chollet \u201cXception: Deep learning with depthwise separable convolutions\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2017) pp. 1251\u20131258.","DOI":"10.1109\/CVPR.2017.195"},{"key":"e_1_3_2_110_2","doi-asserted-by":"crossref","unstructured":"C. Szegedy V. Vanhoucke S. Ioffe J. Shlens Z. Wojna \u201cRethinking the inception architecture for computer vision\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2016) pp. 2818\u20132826.","DOI":"10.1109\/CVPR.2016.308"},{"key":"e_1_3_2_111_2","first-page":"4278","article-title":"Inception-v4, inception-resnet and the impact of residual connections on learning","volume":"31","author":"Szegedy C.","year":"2017","unstructured":"C. Szegedy, S. Ioffe, V. Vanhoucke, A. A. Alemi, Inception-v4, inception-resnet and the impact of residual connections on learning. Proc. AAAI Conf. Artif. Intell. 31, 4278\u20134284 (2017).","journal-title":"Proc. AAAI Conf. Artif. Intell."},{"key":"e_1_3_2_112_2","unstructured":"M. Tan Q. V. Le \u201cEfficientnet: Rethinking model scaling for convolutional neural networks\u201d in International Conference Machine Learning (PMLR 2019) pp. 6105\u20136114."},{"key":"e_1_3_2_113_2","doi-asserted-by":"crossref","unstructured":"M. Sandler A. Howard M. Zhu A. Zhmoginov L. C. Chen \u201cMobilenetv2: Inverted residuals and linear bottlenecks\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2018) pp. 4510\u20134520.","DOI":"10.1109\/CVPR.2018.00474"},{"key":"e_1_3_2_114_2","doi-asserted-by":"crossref","unstructured":"B. Zoph V. Vasudevan J. Shlens Q. V. Le \u201cLearning transferable architectures for scalable image recognition\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2018) pp. 8697\u20138706.","DOI":"10.1109\/CVPR.2018.00907"},{"key":"e_1_3_2_115_2","first-page":"21271","article-title":"Bootstrap your own latent: A new approach to self-supervised learning","volume":"33","author":"Grill J.-B.","year":"2020","unstructured":"J.-B. Grill, F. Strub, F. Altch\u00e9, C. Tallec, P. H. Richemond, E. Buchatskaya, C. Doersch, B. A. Pires, Z. D. Guo, M. G. Azar, B. Piot, K. Kavukcuoglu, R. Munos, M. Valko, Bootstrap your own latent: A new approach to self-supervised learning. Adv. Neural Inf. Process. Syst. 33, 21271\u201321284 (2020).","journal-title":"Adv. Neural Inf. Process. Syst."},{"key":"e_1_3_2_116_2","unstructured":"T. Chen S. Kornblith M. Norouzi G. Hinton \u201cA simple framework for contrastive learning of visual representations\u201d in International Conference Machine Learning (2020) pp. 1597\u20131607."},{"key":"e_1_3_2_117_2","doi-asserted-by":"crossref","unstructured":"K. He H. Fan Y. Wu S. Xie R. Girshick \u201cMomentum contrast for unsupervised visual representation learning\u201d in Proceeding IEEE\/CVF Conference Computer Vision Pattern Recognition (IEEE 2020) pp. 9729\u20139738.","DOI":"10.1109\/CVPR42600.2020.00975"},{"key":"e_1_3_2_118_2","doi-asserted-by":"crossref","unstructured":"A. Chattopadhay A. Sarkar P. Howlader V. N. Balasubramanian \u201cGrad-CAM++: Generalized gradient-based visual explanations for deep convolutional networks\u201d in Proceeding 2018 IEEE Winter Conference Application Computer Vision (WACV 2018) pp. 839\u2013847.","DOI":"10.1109\/WACV.2018.00097"},{"key":"e_1_3_2_119_2","doi-asserted-by":"crossref","unstructured":"S. Abnar W. Zuidema \u201cQuantifying attention flow in transformers\u201d in Proceeding 58th Annual Meeting Association Computational Linguistic (ACL 2020) pp. 4190\u20134197.","DOI":"10.18653\/v1\/2020.acl-main.385"},{"key":"e_1_3_2_120_2","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.11036"},{"key":"e_1_3_2_121_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1257570"},{"key":"e_1_3_2_122_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1817794116"},{"key":"e_1_3_2_123_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1541-0420.2005.00440.x"},{"key":"e_1_3_2_124_2","first-page":"1","article-title":"PAST: Paleontological Statistics Software Package for Education and Data Analysis","volume":"4","author":"Hammer \u00d8.","year":"2001","unstructured":"\u00d8. Hammer, D. A. T. Harper, P. D. Ryan, PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontol. Electron. 4, 1\u20139 (2001).","journal-title":"Palaeontol. Electron."},{"key":"e_1_3_2_125_2","doi-asserted-by":"publisher","DOI":"10.1111\/cla.12524"},{"key":"e_1_3_2_126_2","doi-asserted-by":"crossref","unstructured":"A. A. Polilov At the Size Limit-Effects of Miniaturization in Insects (Springer 2016).","DOI":"10.1007\/978-3-319-39499-2"},{"key":"e_1_3_2_127_2","doi-asserted-by":"publisher","DOI":"10.1145\/3236009"},{"key":"e_1_3_2_128_2","unstructured":"H. Goulet J. T. Huber Hymenoptera of the World: An Identification Guide to Families (Research Branch Agriculture Canada Publication 1894\/E 1993)."},{"key":"e_1_3_2_129_2","unstructured":"A. V. Antropov \u201cBody Size Diminution and the Ways of Evolution of Wing Venation in Aculeate Hymenopterous Insects (Insecta: Hymenoptera: Aculeata: Apoidea). An Attempt of Functional Interpretation\u201d in Evolution and Systematics: Lamarck and Darwin in Modern Studies A. V. Sviridov A. I. Shatalkin Eds. (Archiv of Zoology of the Museum of Moscow University 2009) vol. 50 pp. 247\u2013274."},{"key":"e_1_3_2_130_2","doi-asserted-by":"publisher","DOI":"10.1086\/706914"},{"key":"e_1_3_2_131_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.0014-3820.2003.tb00285.x"},{"key":"e_1_3_2_132_2","doi-asserted-by":"publisher","DOI":"10.1080\/106351599260184"},{"key":"e_1_3_2_133_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041-210X.2011.00169.x"},{"key":"e_1_3_2_134_2","doi-asserted-by":"publisher","DOI":"10.1080\/10618600.1996.10474713"},{"key":"e_1_3_2_135_2","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/bty633"},{"key":"e_1_3_2_136_2","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu181"},{"key":"e_1_3_2_137_2","doi-asserted-by":"crossref","unstructured":"H. Wickham ggplot2: Elegant Graphics for Data Analysis (Springer-Verlag 2016).","DOI":"10.1007\/978-3-319-24277-4_9"},{"key":"e_1_3_2_138_2","unstructured":"A. Dosovitskiy L. Beyer A. Kolesnikov D. Weissenborn X. Zhai T. Unterthiner M. Dehghani M. Minderer G. Heigold S. Gelly J. Uszkoreit N. Houlsby \u201cAn image is worth 16\u00d716 words: Transformers for image recognition at scale\u201d in 9th International Conference on Learning Representations (ICLR 2021)."}],"container-title":["Science Advances"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.science.org\/doi\/pdf\/10.1126\/sciadv.aef0605","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T18:02:45Z","timestamp":1792000965000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.science.org\/doi\/10.1126\/sciadv.aef0605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,16]]},"references-count":137,"journal-issue":{"issue":"42","published-print":{"date-parts":[[2026,10,16]]}},"alternative-id":["10.1126\/sciadv.aef0605"],"URL":"https:\/\/doi.org\/10.1126\/sciadv.aef0605","relation":{},"ISSN":["2375-2548"],"issn-type":[{"value":"2375-2548","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,16]]},"assertion":[{"value":"2025-12-27","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-14","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-14","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"eaef0605"}}