Beyond venomous fangs: Uloboridae spiders have lost their venom but not their toxicity
Posted on: 13 November 2025
Preprint posted on 8 January 2025
Article now published in BMC Biology at https://bmcbiol.biomedcentral.com/articles/10.1186/s12915-025-02248-1
By blurring the line between venom and digestion, Uloboridae spiders remind us that evolution doesn’t discard complexity — it reshapes it.
Selected by Daniel Fernando Reyes Enríquez, Marcus Oliveira
Categories: biochemistry, evolutionary biology, pharmacology and toxicology
Updated 9 November 2025 with a postLight by Daniel Fernando Reyes Enríquez
postLight: A comparison between the preprint and the published paper
The preprint (Peng et al., 2023; bioRxiv 2023.06.26.546488) proposed that Uloborus plumipes and other Uloboridae spiders lost their venom glands but retained toxin-related genes, which were redeployed within digestive secretions. It primarily presented morphological and transcriptomic evidence, identifying the absence of venom ducts and the expression of venom-like transcripts in the midgut. The study suggested that these toxins were repurposed to support extra-oral digestion and prey immobilization, highlighting an evolutionary case of venom loss coupled with functional compensation through digestive toxicity.
The published paper (BMC Biology, 2025, 23:159) built upon and expanded this foundation with broader experimental validation and comparative analyses. The authors incorporated detailed histology, scanning electron microscopy, and functional assays demonstrating insecticidal activity of midgut extracts. RNA-seq data from multiple tissues, together with comparative genomic analyses of defensin genes, reinforced the notion of toxin repurposing. The discussion was substantially revised to integrate morphological, molecular, and physiological results into a cohesive evolutionary framework, emphasizing the adaptive shift from envenomation to digestive toxicity.
In essence, while the preprint outlined a conceptual and transcriptomic hypothesis of venom system loss, the published article confirmed and contextualized it through enhanced methodology, functional validation, and deeper evolutionary interpretation.
Background
Venom is both a polygenic characteristic, defined by its toxicity and functional role in facilitating both feeding and defense within an organism [7]. It often emerges from the co-option and specialization of pre-existing secretory systems, particularly the salivary glands, as observed across a wide range of venomous animals [6, 4].
In spiders, venom has traditionally been linked to predation through rapid immobilization and initiation of extra-oral digestion [2, 7, 8]. However, the Uloboridae family presents an evolutionary anomaly: they lack a differentiated venom gland, a trait first formally described by Weng et al. (2006). This morphological absence led to the hypothesis that venom genes were not entirely absent but rather rendered redundant by a behavioral shift in prey capture strategy, specifically, the extensive use of silk to immobilize prey and the application of digestive fluids rich in proteolytic enzymes [12, 13, 14].
The study ultimately suggests how venom, as a modular and multifunctional trait, could have been retooled by natural selection [16]. In Uloboridae, this manifests as a shift from a neurochemical weapon to a biochemical digestive strategy, blurring the line between venom and digestive secretion [14].
In this preprint, the authors address a gap in our knowledge by providing a physiological description of analog regions of the venom gland structure in Uloboridae that in most species of spiders corresponds to venom glands. The hypothesis tested by the preprint authors is that uloborids have replaced both a venom gland and fang duct with toxic molecules in their digestive fluids, leaving venom glands redundant and leading to their evolutionary loss over time.
Key findings
Morphological evidence for the lack of venom glands
Histological analysis of the chelicerae and anterior prosoma of Uloborus plumipes confirmed the complete absence of venom glands. Unlike venomous spiders such as Parasteatoda tepidariorum, which exhibit clear venom ducts and glandular tissues, U. plumipes chelicerae are occupied by large flexor muscles with, according to the authors, no trace of venom ducts or secretory cells. Additionally, while electron microscopy did reveal the presence of minute pores at the tips of the fangs, these were too small and structurally dissimilar to the venom canal pores seen in venomous species, suggesting they serve another, currently unknown function. These morphological findings corroborate the hypothesis of a functional and anatomical loss of the venom delivery system in Uloboridae.
RNA-seq reveals the expression of putative toxins in the midgut gland of U. plumipes
Despite the anatomical loss of venom glands, transcriptomic analysis (RNA-seq) revealed high expression levels of toxin-like transcripts within the midgut gland. Among the 124 translated sequences with homology to known venom proteins, a significant portion were neurotoxin-like and enzymatically active proteins such as metalloproteases and serine proteases. These molecules, while typically associated with venom function, were instead upregulated in the midgut, a key site for digestive enzyme secretion. This expression pattern suggests repurposing of ancestral venom genes for use in digestive physiology rather than envenomation.
Toxins in spider digestive fluids: an evolutionary adaptation?
The presence of venom-like toxins in digestive fluids is not unique to Uloboridae. Other spider species, including the venomous Nephila cruentata and Acanthoscurria geniculata, have been shown to secrete similar proteins via their digestive systems. In Uloboridae, however, this adaptation is more extreme according to the authors, representing a full shift in toxin deployment from venom injection to external application during prey digestion [9]. This suggests that the incorporation of venom components into digestive fluids may be a widespread and evolutionarily advantageous trait among spiders, perhaps originally serving as a supplementary prey immobilization system and, in Uloboridae, evolving into the primary mechanism.
Why I think this preprint is important
Given how convergent evolution has produced a clade of organisms in which all but two families have venom glands and some sort of duct to inoculate venom, the family Uloboridae showcases a singular path through its evolutionary history. This study continues the route to understand the type of selective pressures that could drive an archaic venom system into an auxiliary secretion in the digestive system.
Questions and comments
Comment 1: Hypothesis of venom loss: Regarding this point, it seems unlikely that venom evolved in the family as a defined product of the gland capable of producing toxins and was lost. However, what could be comparable to examples in nature is that at one point in time, a common ancestor of the clade had selective positive pressures that started allocating genes of proteases and putative archaic forms of now toxin families into a secretion in the midway of being salivary secretion and venom [1, 4, 9]. But success in a niche by developing hunting patterns that do not depend on venom to be effective could have been a major driver to evolve venom gland and duct in the initial stages. Therefore, the homology of proteins that show some relation to toxins found in other organisms might be an example of convergent evolution at a far earlier stage [12, 14]. Subsequently, it bends the definition of a toxin, since compared to a digestive enzyme, the majority of venom protein families act similarly. Except for neurotoxic proteins that are more related to prey capture, but without a defined activity, perhaps there are more related to digestive enzymes. Since the definition of venom is intrinsic to a venom gland, perhaps the authors would strengthen their hypothesis by suggesting a redefinition of venom that includes cases such as the Uloboridae family.
Q1. Functional specificity: This study identifies numerous toxin-like transcripts upregulated in the midgut gland and predicts some to be neurotoxic. However, the electrophysiological assays only tested effects on sodium and potassium channels [8]. Given the negative results, why were alternative neural targets (e.g., calcium channels, acid-sensing ion channels, or glutamate receptors) not explored, especially since these are common targets in other arachnid and insect venoms? Further functional assays or broader electrophysiological screening would strengthen the claim that the midgut toxins play a neurotoxic or prey-immobilizing role.
Q2. Behavioral or ecological observations: The manuscript posits that digestive fluid toxicity substitutes for venom in prey immobilization, yet there is a lack of direct in vivo observations or behavioral assays showing prey death post-application of digestive fluids in natural hunting scenarios [5, 11]. Can the authors provide experimental video evidence or more detailed ethological observations to support this behavioral hypothesis? Controlled behavioral trials or field observations showing the timeline of prey immobilization post-silk wrapping and digestive fluid application would provide compelling support.
Q3. Taxonomic and genomic sampling within Uloboridae: The conclusions are heavily based on Uloborus plumipes, with only limited comparative references to other Uloboridae species. Given that evolutionary loss and repurposing can be lineage-specific, how generalizable are these findings across the entire Uloboridae family? A broader phylogenetic sampling, including genomic or transcriptomic data from other Uloboridae genera, would help validate the evolutionary trends proposed [4, 8, 14].
Q4. Limitations of transcriptome-based toxin identification: The pipeline relies on homology to known venom proteins and differential expressions to define toxin-like transcripts. However, these methods may misclassify or miss lineage-specific toxins, especially small peptides like defensins. How did the authors control false positives/negatives, and could the lack of proteomic validation (e.g., LC-MS of the digestive fluid) affect the conclusions? Integration with proteomic analyses, particularly of midgut secretions, would validate transcriptome predictions and confirm that these toxin-like genes translate into active proteins [7].
References
[1] Bond, J. E., Garrison, N. L., Hamilton, C. A., Godwin, R. L., Hedin, M. & Agnarsson, I. (2014). Phylogenomics resolves a spider backbone phylogeny and rejects a prevailing paradigm for orb web evolution. Current Biology 24, 1765–1771.
[2] Casewell, N. R. (2017). Evolution: gene co-option underpins venom protein evolution. Current Biology 27, R647–R649.
[3] Casewell, N. R., Wagstaff, S. C., Harrison, R. A., Renjifo, C. & Wu¨ster, W. (2011). Domain loss facilitates accelerated evolution and neofunctionalization of duplicate snake venom metalloproteinase toxin genes. Molecular Biology and Evolution 28, 2637–2649.
[4] Cordes, M. H. J. & Binford, G. J. (2018). Evolutionary dynamics of origin and loss in the deep history of phospholipase D toxin genes. BMC Evolutionary Biology 18, 194.
[5] Dimitrov, D. & Hormiga, G. (2021). Spider diversification through space and time. Annual Review of Entomology 66, 225–241.
[6] Drukewitz, S. H., Bokelmann, L., Undheim, E. A. B. & von Reumont, B. M. (2019). Toxins from scratch? Diverse, multimodal gene origins in the predatory robber fly Dasypogon diadema indicate a dynamic venom evolution in dipteran insects. GigaScience 8, giz081.
[7] Fry, B. G., Koludarov, I., Jackson, T. N. W., Holford, M., Terrat, Y., Casewell, N. R., Undheim, E. A. B., Vetterb, I., Alia, S. A., Low, D. H. W. & Sunagar, K. (2015). Seeing the woods for the trees: understanding venom evolution as a guide for biodiscovery. In Venoms to Drugs: Venom as a Source for the Development of Human Therapeutics (ed. G. F. KING), pp. 1–36.
[8] Lüddecke, T., Herzig, V., Reumont, B. M., & Vilcinskas, A. (2021). The biology and evolution of spider venoms. Biological Reviews. doi:10.1111/brv.12793
[9] Misof, B., Liu, S., Meusemann, K., Peters, R. S., Donath, A., Mayer, C., Frandsen, P. B., Ware, J., Flouri, T., Beutel, R. G., Niehuis, O., Petersen, M., Izquierdo-Carrasco, F., Wappler, T., Rust, J., et al. (2014). Phylogenomics resolves the timing and pattern of insect evolution. Science 346, 763–767.
[10] Morgenstern, D. & King, G. F. (2013). The venom optimization hypothesis revisited. Toxicon 63, 120–128.
[11] Nisani, Z., Dunbar, S. G. & Hayes, W. K. (2007). Cost of venom regeneration in Parabuthus transvaalicus (Arachnida: Buthidae). Comparative Biochemistry and Physiology – A Molecular and Integrative Physiology 147, 509–513.
[12] Pekar, S. , Bocanek, O. , Michalek, O. , Petrakov a, L. , Haddad, C. R., Sˇedo, O. & Zdrahal, Z. (2018a). Venom gland size and venom complexity— essential trophic adaptations of venomous predators: a case study using spiders. Molecular Ecology 27, 4257–4269.
[13] Pekar, S. , Líznarova, E. , Bocˇanek, O. & Zdrahal, Z. (2018b). Venom of preyspecialized spiders is more toxic to their preferred prey: a result of prey-specific toxins. Journal of Animal Ecology 87, 1639–1652.
[14] Schöneberg, Y., Audisio, T. L., Hamadou, A. B., Forman, M., Král, J., Kořínková, T., Líznerová, E., Mayer, C., Prokopcová, L., Krehenwinkel, H., Prost, S., & Kennedy, S. (2024). Three novel spider genomes unveil spidroin diversification and Hox cluster architecture: Ryuthela nishihirai (Liphistiidae), Uloborus plumipes (Uloboridae) and Cheiracanthium punctorium (Cheiracanthiidae). Molecular Ecology Resources. https://doi.org/10.1111/1755-0998.14038
[15] Silva, L. M., Carvalho Botelho, A. C., Nacif-Pimenta, R., Martins, G. F., Alves, L. C., Brayner, F. A., Fortes-Dias, C. L. & Paolucci Pimenta, P. F. (2008). Structural analysis of the venom glands of the armed spider Phoneutria nigriventer (Keyserling, 1891): microanatomy, fine structure and confocal observations. Toxicon 51, 693–706.
[16] Weng, J. L., Barrantes, G. & Eberhard, W. G. (2006). Feeding by Philoponella vicina (Araneae, Uloboridae) and how uloborid spiders lost their venom glands. Canadian Journal of Zoology 84, 1752–1762
doi: https://doi.org/10.1242/prelights.42070
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