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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2025.1634037</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Mini Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Centipede predation on vertebrates: a review with the first bat case from Asia</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Kamalakannan</surname>
<given-names>Manokaran</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Mondal</surname>
<given-names>Rahul</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Banerjee</surname>
<given-names>Dhriti</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
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<aff id="aff1">
<sup>1</sup>
<institution>Mammal and Osteology Section, Zoological Survey of India</institution>, <addr-line>Kolkata, West Bengal</addr-line>,&#xa0;<country>India</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Hemiptera Section, Zoological Survey of India</institution>, <addr-line>Kolkata, West Bengal</addr-line>,&#xa0;<country>India</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Zoological Survey of India</institution>, <addr-line>Kolkata, West Bengal</addr-line>,&#xa0;<country>India</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Jing Chen, Institute of Zoology (CAS), China</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Darrell Acu&#xf1;a, University of Santo Tomas, Philippines</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Manokaran Kamalakannan, <email xlink:href="mailto:kamalakannanm1@gmail.com">kamalakannanm1@gmail.com</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>28</day>
<month>07</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>13</volume>
<elocation-id>1634037</elocation-id>
<history>
<date date-type="received">
<day>23</day>
<month>05</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>09</day>
<month>07</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Kamalakannan, Mondal and Banerjee.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Kamalakannan, Mondal and Banerjee</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Centipedes (Chilopoda: Scolopendromorpha), long regarded as generalist arthropod predators, are increasingly recognized for their capacity to subdue and consume small vertebrates. This review synthesizes over a century of published accounts documenting centipede predation on amphibians, reptiles, birds, mammals, and occasionally fish, emphasizing the ecological breadth, behavioral strategies, and taxonomic diversity of both predators and prey. Notable cases include <italic>Scolopendra gigantea</italic> preying on bats in Venezuelan caves, <italic>Scolopendra subspinipes</italic> capturing snakes in urban environments, and <italic>Cormocephalus coynei</italic> exerting top-down control on seabird populations on predator-free islands. We also present the first confirmed case of bat predation by a centipede in Asia, where a <italic>Rhysida</italic> species was observed consuming a <italic>Pipistrellus</italic> bat in a fig tree hollow in West Bengal, India. This observation expands the known biogeography and ecological context of vertebrate predation by centipedes. Our synthesis highlights the underappreciated role of scolopendrid centipedes as mid-level predators capable of influencing small vertebrate populations, particularly in resource-limited or insular ecosystems, and calls for a re-evaluation of their functional position within terrestrial food webs.</p>
</abstract>
<kwd-group>
<kwd>bat</kwd>
<kwd>centipede</kwd>
<kwd>prey</kwd>
<kwd>predation</kwd>
<kwd>
<italic>Pipistrelles</italic>
</kwd>
</kwd-group>
<counts>
<fig-count count="2"/>
<table-count count="1"/>
<equation-count count="0"/>
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<page-count count="8"/>
<word-count count="2854"/>
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<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Natural History Insights</meta-value>
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</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Predation is a central ecological force shaping trophic dynamics, influencing population regulation, species distributions, and evolutionary trajectories (<xref ref-type="bibr" rid="B5">Begon et&#xa0;al., 2006</xref>). While classical food web models often depict vertebrates preying upon invertebrates, an emerging body of literature emphasizes the importance of reverse trophic interactions where invertebrates act as predators of vertebrates (<xref ref-type="bibr" rid="B37">Nyffeler and Kn&#xf6;rnschild, 2013</xref>; <xref ref-type="bibr" rid="B36">Nyffeler and Gibbons, 2022</xref>).</p>
<p>Across a range of taxa including arachnids, insects, myriapods, and crustaceans, invertebrates have been documented preying on vertebrate animals in both aquatic and terrestrial ecosystems (<xref ref-type="bibr" rid="B27">McCormick and Polis, 1982</xref>; <xref ref-type="bibr" rid="B16">Formanowicz and Bradley, 1987</xref>; <xref ref-type="bibr" rid="B31">Mori and Ohba, 2004</xref>; <xref ref-type="bibr" rid="B37">Nyffeler and Kn&#xf6;rnschild, 2013</xref>; <xref ref-type="bibr" rid="B51">Valdez, 2020</xref>; <xref ref-type="bibr" rid="B36">Nyffeler and Gibbons, 2022</xref>; etc.). Among invertebrate predators, centipedes (class Chilopoda) represent a particularly intriguing and underappreciated group capable of subduing vertebrate prey (<xref ref-type="bibr" rid="B50">Undheim and King, 2011</xref>; <xref ref-type="bibr" rid="B13">Dugon and Arthur, 2012</xref>; <xref ref-type="bibr" rid="B9">Clark, 1979</xref>). Members of the order Scolopendromorpha, especially within the genus <italic>Scolopendra</italic>, are large, nocturnal, and venomous generalists that use modified forelegs (forcipules) to inject venom, immobilize prey, and initiate extra-oral digestion (<xref ref-type="bibr" rid="B50">Undheim and King, 2011</xref>; <xref ref-type="bibr" rid="B13">Dugon and Arthur, 2012</xref>). Although their primary diet consists of arthropods, accumulating evidence reveals that scolopendrid centipedes frequently prey on small vertebrates across diverse environments (<xref ref-type="bibr" rid="B10">Cloudsley-Thompson, 1968</xref>; <xref ref-type="bibr" rid="B17">Forti et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B4">Bauer, 1990</xref>; <xref ref-type="bibr" rid="B30">Molinari et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B48">Srbek-Araujo et&#xa0;al., 2012</xref>, etc.).</p>
<p>These interactions between centipede and vertebrate are often overlooked due to the secretive and nocturnal habits of centipedes, yet they may exert significant top-down pressures on small vertebrate populations, particularly in resource-limited habitats or insular ecosystems (<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al., 2021</xref>). Unlike most terrestrial invertebrates, large scolopendrid centipedes are capable of overpowering prey exceeding their own body mass (<xref ref-type="bibr" rid="B13">Dugon and Arthur, 2012</xref>). This functional capacity positions them uniquely in terrestrial food webs, where they may occupy mid-level predator niches and influence vertebrate community composition. Their role in nutrient transfer and energy flow, especially in tropical and subtropical regions, is only beginning to be understood (<xref ref-type="bibr" rid="B36">Nyffeler and Gibbons, 2022</xref>).</p>
<p>This review synthesizes over a century of records on centipede predation on vertebrates, emphasizing the diversity of prey, attack strategies, ecological contexts, and the evolutionary implications of such interactions. By integrating published literature and novel observations, including the first documented case of bat predation by a centipede in Asia, we aim to expand the ecological narrative surrounding centipedes and highlight their role as key, albeit cryptic, predators of vertebrates.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Literature review</title>
<sec id="s2_1">
<label>2.1</label>
<title>Overview of invertebrate-driven vertebrate mortality</title>
<p>A diverse array of invertebrates have been documented preying on vertebrates, often in ecologically impactful ways. For example, odonate larvae (dragonfly nymphs) are major predators of amphibian larvae and exert strong selective pressures on anuran life history traits (<xref ref-type="bibr" rid="B32">Morin, 1983</xref>; <xref ref-type="bibr" rid="B26">McCollum and Leimberger, 1997</xref>). In tropical ecosystems, army ants (<italic>Eciton burchellii</italic>) are known to kill and consume nestling birds and small reptiles during raids (<xref ref-type="bibr" rid="B43">Rettenmeyer et&#xa0;al., 2011</xref>). Similarly, wasps and spiders have been recorded depredating avian nestlings (<xref ref-type="bibr" rid="B20">Henriques, 1998</xref>; <xref ref-type="bibr" rid="B37">Nyffeler and Kn&#xf6;rnschild, 2013</xref>).</p>
<p>These interactions are not merely incidental; they contribute significantly to mortality regimes and influence prey behavior and habitat use (<xref ref-type="bibr" rid="B46">Sih et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B41">Preisser et&#xa0;al., 2005</xref>). Importantly, they underscore the need to reassess traditional trophic hierarchies and recognize the diverse roles of invertebrates as apex or mesopredators in certain ecological contexts (<xref ref-type="bibr" rid="B35">Nyffeler and Birkhofer, 2017</xref>).</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Centipede predation on amphibians</title>
<p>Predation events of centipedes have been documented across a range of amphibian taxa. In the Caribbean, <italic>Scolopendra alternans</italic> was observed attacking and consuming the toxic cane toad (<italic>Rhinella marina</italic>), a species known for its chemical defenses (<xref ref-type="bibr" rid="B7">Carpenter and Gillingham, 1984</xref>). This highlights the centipede&#x2019;s resistance or indifference to amphibian toxins in some cases. Similarly, in tropical Australia, native centipedes (<italic>Ethmostigmus rubripes</italic>) have been reported killing invasive cane toads, indicating a remarkable reversal of expected predator&#x2013;prey dynamics (<xref ref-type="bibr" rid="B40">Pomeroy et&#xa0;al., 2021</xref>). In Brazil&#x2019;s Atlantic Forest, <italic>Otostigmus tibialis</italic> has been recorded preying on the small arboreal tree frog <italic>Dendropsophus elegans</italic>, showcasing centipede predation within vertical forest strata (<xref ref-type="bibr" rid="B17">Forti et&#xa0;al., 2007</xref>). These observations provide crucial insights into centipedes as opportunistic and generalist predators capable of subduing toxic or agile amphibian prey across diverse ecosystems.</p>
<p>Although quantitative studies on the frequency and ecological impact of centipede-amphibian interactions are lacking, the available evidence suggests that centipedes play a significant, if under-recognized, role in amphibian mortality particularly among juveniles and small-bodied species.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Centipede predation on reptiles</title>
<p>Reptilian prey, especially small lizards and snakes, form a notable component of centipede diets. Species like <italic>Cormocephalus coynei, Scolopendra viridicornis</italic> and <italic>Scolopendra subspinipes</italic> have been recorded attacking geckos, skinks, and colubrids using venom injection and powerful mechanical restraint (<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B54">Vieira et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B12">Deb et&#xa0;al., 2023</xref>). These interactions occurred in microhabitats such as leaf litter, under logs, and within rock crevices, where reptiles are abundant and centipedes can ambush effectively.</p>
<p>Multiple field observations illustrate the breadth of reptilian predation. In India, <italic>Scolopendra hardwickei</italic> has been reported preying on <italic>Oligodon taeniolatus</italic>, a small snake (<xref ref-type="bibr" rid="B47">Smart et&#xa0;al., 2010</xref>), while in Thailand&#x2019;s Sakaerat Biosphere Reserve, <italic>Scolopendra dawydoffi</italic> consumed <italic>Sibynophis triangularis</italic> (<xref ref-type="bibr" rid="B8">Chiacchio et&#xa0;al., 2017</xref>). In the Andaman Islands, <italic>S. dehaani</italic> was documented subduing <italic>Lycodon hypsirhinoides</italic> in forest habitats (<xref ref-type="bibr" rid="B53">Vazifdar et&#xa0;al., 2021</xref>), and <italic>Lycodon zawi</italic> in urban Guwahati (<xref ref-type="bibr" rid="B12">Deb et&#xa0;al., 2023</xref>), suggesting adaptability to varied landscapes.</p>
<p>In urban Singapore, <italic>Scolopendra subspinipes</italic> was recorded preying on multiple fossorial snakes such as <italic>Calamaria schlegeli</italic>, <italic>Calamaria pavimentata</italic>, and <italic>Pseudorabdion longiceps</italic> within green spaces embedded in city infrastructure (<xref ref-type="bibr" rid="B42">Pwa et&#xa0;al., 2023</xref>). Elsewhere, <italic>Scolopendra gigantea</italic> has been observed attacking <italic>Leptodeira bakeri</italic>, a colubrid snake (<xref ref-type="bibr" rid="B18">Goessling et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B52">van Buurt and Dilrosun, 2017</xref>). Additionally, <italic>Scolopendra</italic> sp. captured <italic>Calliophis melanurus</italic>, a venomous elapid, in Mumbai&#x2019;s rocky outcrops (<xref ref-type="bibr" rid="B29">Mirza and Ahmed, 2009</xref>), indicating their ability to handle dangerous prey.</p>
<p>These accounts underscore the ecological versatility of large centipedes and their ability to exploit a wide range of reptilian prey across diverse habitats, from pristine forests and islands to anthropogenic settings such as cities and agricultural margins.</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Centipede predation on birds and fish</title>
<p>Although uncommon, centipede predation on birds is best exemplified by <italic>Cormocephalus coynei</italic> on Phillip Island, Norfolk Island, Australia where it functions as a top terrestrial predator in the absence of mammals. This species preys on thousands of black-winged petrel (<italic>Pterodroma nigripennis</italic>) chicks annually by entering their burrows and using venom to subdue them (<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al., 2021</xref>). <italic>Cormocephalus coynei</italic> also scavenges regurgitated marine fish from seabirds, demonstrating notable dietary flexibility (<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al., 2021</xref>).</p>
<p>Historical reports suggest other <italic>Scolopendra</italic> species may occasionally prey on small birds (<xref ref-type="bibr" rid="B11">Cumming, 1903</xref>; <xref ref-type="bibr" rid="B10">Cloudsley-Thompson, 1968</xref>), though such accounts lack detailed verification. While active predation on fish has not been documented, scavenging behaviors particularly in island or predator-limited ecosystems reveal the trophic adaptability of some centipedes. These interactions challenge conventional views of invertebrate predators and warrant further ecological investigation.</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Centipede predation on mammals</title>
<p>Mammalian predation by centipedes, particularly involving bats, is well-documented for <italic>Scolopendra gigantea</italic> in Venezuelan caves, where it preys on molossid and mormoopid bats (<xref ref-type="bibr" rid="B30">Molinari et&#xa0;al., 2005</xref>). Similar predation events involving <italic>Scolopendra viridicornis</italic> have been reported in Brazil (<xref ref-type="bibr" rid="B48">Srbek-Araujo et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B34">Noronha et&#xa0;al., 2015</xref>). In the United States, <italic>Scolopendra heros</italic> has been observed preying on <italic>Eptesicus fuscus</italic> in rock crevices in Texas (<xref ref-type="bibr" rid="B24">Lindley et&#xa0;al., 2017</xref>). In addition to bats, rodents have also been recorded as prey. For instance, <italic>Scolopendra galapagoensis</italic> has been observed preying upon the Gal&#xe1;pagos rice rat <italic>Oryzomys bauri</italic> (= <italic>Aegialomys galapagoensis</italic>) (<xref ref-type="bibr" rid="B9">Clark, 1979</xref>), and unidentified rodents have been found in the diet of other large scolopendrid centipedes (<xref ref-type="bibr" rid="B45">Shugg, 1961</xref>). These observations expand the known mammalian prey base of centipedes and highlight their role as opportunistic vertebrate predators. The behavioral and ecological implications of such predation particularly concerning roost site selection, foraging behavior, and predator avoidance remain poorly studied but are likely to be significant.</p>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>Case study: centipede predation on a bat, a new record from Asia</title>
<p>Recently, a team of scientists of Zoological Survey of India carried out a faunal survey in the dry tropical forests of West Bengal, India. The study area, Nakpur a village situated in the Nalhati Block of Birbhum district of West Bengal state in India (24.295684&#xb0;N, 88.011694&#xb0;E) which is primarily occupied by crop agricultural fields. During the insect survey in the study area, a centipede was observed preying on a bat in the hollow of a Sacred Fig (<italic>Ficus religiosa</italic>) tree at 20:30 h on 19 June 2024. This rare interaction was captured through photographs (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>) and video (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary material</bold>
</xref>) to provide significant documentation. The centipede was identified as a species belonging to the genus <italic>Rhysida</italic> (Chilopoda: Scolopendromorpha: Scolopendridae), based on its head, dorsal body, ultimate legs and leg color, body profile (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1a</bold>
</xref>), and spiracle formula one of the diagnostic characters of the genus against other scolopendrids (<xref ref-type="bibr" rid="B22">Joshi et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B49">Sureshan, 2024</xref>; <xref ref-type="bibr" rid="B6">Bonato et&#xa0;al., 2025</xref>), while the bat was identified as a species belonging to genus <italic>Pipistrellus</italic> (Mammalia: Chiroptera: Vespertilionidae) due to its small size (&#x2248; 6 cm), body shape, and internarial groove (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1b</bold>
</xref>) (<xref ref-type="bibr" rid="B3">Bates and Harrison, 1997</xref>; <xref ref-type="bibr" rid="B44">Sharma et&#xa0;al., 2024</xref>). Due to logistical constraints in the field and the non-invasive nature of the observation, molecular confirmation of species identity was not feasible. Both the bat and centipede were observed at the lower part of the tree, approximately 5 feet above the ground, and the tree was home to the Indian flying fox (<italic>Pteropus medius)</italic>, spotted house gecko (<italic>Hemidactylus parvimaculatus</italic>), Asian common toad (<italic>Duttaphrynus melanostictus</italic>), and bee nests. The weather during this observation was warm and humid, reaching a high of around 36&#xb0;C (96.8&#xb0;F)- West Bengal&#x2019;s monsoon season. The centipede was notably longer than the bat (&#x2248; 10 cm). The centipede began to feed on the bat&#x2019;s wing membrane (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1c</bold>
</xref>) and proceeding deeper into the body of the bat (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1d</bold>
</xref>) while it was still alive (<xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1c, e</bold>
</xref>). The authors noted that the predatory actions occurred after the bat had been immobilized, yet its head continued moving while the centipede was feeding. After capturing a one-minute video and taking some photos, the researchers stepped back from the spot to prevent disrupting the centipede&#x2019;s hunting behavior.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Bat predation by a centipede: <bold>(a)</bold> the predator <italic>Rhysida</italic> sp. centipede, <bold>(b)</bold> the prey <italic>Pipistrellus</italic> sp. bat, <bold>(c)</bold> the centipede immobilizes the bat using its forcipules while the bat is still alive; the open eye suggests it remains conscious, <bold>(d)</bold> damage to the bat&#x2019;s wing membrane caused by the centipede, <bold>(e)</bold> the bat&#x2019;s mouth is open as the centipede feeds, indicating a potential struggle or distress response. All images were taken under natural conditions using a flashlight during nighttime observations.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-g001.tif">
<alt-text content-type="machine-generated">A sequence of five images showing a centipede interacting with a bat inside a rocky crevice. Each image provides a different perspective of the centipede, which is approximately three centimeters long, engaging with the bat. The setting is a dark, confined space surrounded by rock. The images illustrate the centipede's movement around the bat, with the bat appearing passive throughout the interaction.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>Previous studies (<xref ref-type="bibr" rid="B30">Molinari et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B48">Srbek-Araujo et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B34">Noronha et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B24">Lindley et&#xa0;al., 2017</xref>) have reported bat predation by <italic>Scolopendra</italic> species in the Americas, mainly in caves and artificial structures. This new record suggests that arboreal predation on bats by centipedes may be more widespread than previously thought.</p>
<p>Our analysis on centipede predation on vertebrate reveals a clear taxonomic bias in centipede predation towards reptiles, particularly snakes (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>; <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Among the 16 centipede species with documented vertebrate predation, reptiles constitute the most frequently reported prey class. This trend is especially evident in large-bodied <italic>Scolopendra</italic> species such as <italic>Scolopendra subspinipes</italic>, <italic>Scolopendra gigantea</italic>, and <italic>Scolopendra heros</italic>, which have been recorded subduing snakes often comparable to their own size (<xref ref-type="bibr" rid="B51">Valdez, 2020</xref>; <xref ref-type="bibr" rid="B14">Easterla, 1975</xref>; <xref ref-type="bibr" rid="B52">van Buurt and Dilrosun, 2017</xref>; <xref ref-type="bibr" rid="B42">Pwa et&#xa0;al., 2023</xref>). The elongated, limbless morphology of snakes may facilitate subjugation and consumption by centipedes using their powerful forcipules and envenomation tactics (<xref ref-type="bibr" rid="B23">Lewis, 1981</xref>). Moreover, snakes often inhabit microhabitats also frequented by centipedes such as forest floors, under logs, or leaf litter thereby increasing encounter rates. This dietary inclination suggests that scolopendrid centipedes, though generalist predators, may exert predatory pressure on certain reptile populations, especially in insular or disturbed ecosystems where snakes are among the few available vertebrate prey.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Documented instances of centipede predators and their vertebrate prey species (amphibians, reptiles, birds, mammals and fishes).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="center">Centipede predator species</th>
<th valign="top" align="center">Vertebrate prey species</th>
<th valign="top" align="center">Common name</th>
<th valign="top" align="center">Habitat</th>
<th valign="top" align="center">Location</th>
<th valign="top" align="center">Setting</th>
<th valign="top" align="center">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="6" align="left">&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;<inline-graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-i001.tif"/> Amphibians</th>
</tr>
<tr>
<td valign="top" align="left">
<italic>Ethmostigmus rubripes</italic>
</td>
<td valign="top" align="left">
<italic>Rhinella marina</italic>
</td>
<td valign="top" align="left">Cane toad</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Australia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B40">Pomeroy et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Otostigmus tibialis</italic>
</td>
<td valign="top" align="left">
<italic>Dendropsophus elegans</italic>
</td>
<td valign="top" align="left">Elegant forest tree frog</td>
<td valign="top" align="left">Forest</td>
<td valign="top" align="left">S&#xe3;o Miguel Arcanjo, Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B17">Forti et&#xa0;al. (2007)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra alternans</italic>
</td>
<td valign="top" align="left">
<italic>Rhinella marina</italic>
</td>
<td valign="top" align="left">Cane toad</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Caribbean (not specified)</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B7">Carpenter and Gillingham (1984)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="2" align="left">
<italic>Scolopendra</italic> sp.</td>
<td valign="top" align="left">
<italic>Kaloula pulchra</italic>
</td>
<td valign="top" align="left">Banded bullfrog</td>
<td valign="top" align="left">Forest</td>
<td valign="top" align="left">Thailand</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B21">Hodges and Goodyear (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scinax fuscovarius</italic>
</td>
<td valign="top" align="left">Fuscous-blotched treefrog</td>
<td valign="top" align="left">Forest</td>
<td valign="top" align="left">Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B15">Folly et&#xa0;al. (2019)</xref>
</td>
</tr>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="6" align="left">&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;<inline-graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-i002.tif"/> Reptiles</th>
</tr>
<tr>
<td valign="top" rowspan="2" align="left">
<italic>Cormocephalus coynei</italic>
</td>
<td valign="top" align="left">
<italic>Oligosoma lichenigera</italic>
</td>
<td valign="top" align="left">Lord Howe Island skink</td>
<td valign="top" rowspan="2" align="left">Island</td>
<td valign="top" rowspan="2" align="left">Phillip Island, Norfolk Island, Australia</td>
<td valign="top" rowspan="2" align="left">Field</td>
<td valign="top" rowspan="2" align="left">
<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Christinus guentheri</italic>
</td>
<td valign="top" align="left">G&#xfc;nther&#x2019;s island gecko</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Cormocephalus</italic> sp.</td>
<td valign="top" align="left">
<italic>Oedura lesueurii</italic>
</td>
<td valign="top" align="left">Lesueur&#x2019;s gecko</td>
<td valign="top" align="left">Rock crevices</td>
<td valign="top" align="left">Australia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B39">Pike et&#xa0;al. (2010)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra cingulata</italic>
</td>
<td valign="top" align="left">
<italic>Dalmatolacerta oxycephala</italic>
</td>
<td valign="top" align="left">Sharp-snouted rock lizard</td>
<td valign="top" align="left">Rocky area</td>
<td valign="top" align="left">Kor&#x10d;ula Island, Croatia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B55">Zimi&#x107; and Jeli&#x107; (2014)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra dawydoffi</italic>
</td>
<td valign="top" align="left">
<italic>Sibynophis triangularis</italic>
</td>
<td valign="top" align="left">Triangulate collared snake</td>
<td valign="top" align="left">Biosphere reserve</td>
<td valign="top" align="left">Sakaerat, Thailand</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B8">Chiacchio et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="2" align="left">
<italic>Scolopendra dehaani</italic>
</td>
<td valign="top" align="left">
<italic>Lycodon hypsirhinoides</italic>
</td>
<td valign="top" align="left">Andaman wolf snake</td>
<td valign="top" align="left">Forest</td>
<td valign="top" align="left">Andaman Islands</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B53">Vazifdar et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Lycodon zawi</italic>
</td>
<td valign="top" align="left">Zaw&#x2019;s wolf snake</td>
<td valign="top" align="left">Urban</td>
<td valign="top" align="left">Guwahati, India</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B12">Deb et&#xa0;al. (2023)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra galapagoensis</italic>
</td>
<td valign="top" align="left">
<italic>Pseudalsophis biserialis</italic>
</td>
<td valign="top" align="left">Gal&#xe1;pagos racer</td>
<td valign="top" align="left">Island</td>
<td valign="top" align="left">Gal&#xe1;pagos Islands</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B38">Ortiz-Catedral et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra gigantea</italic>
</td>
<td valign="top" align="left">
<italic>Leptodeira bakeri</italic>
</td>
<td valign="top" align="left">Baker&#x2019;s cat-eyed snake</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Aruba</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B18">Goessling et&#xa0;al. (2012)</xref>; <xref ref-type="bibr" rid="B52">van Buurt and Dilrosun (2017)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra hardwickei</italic>
</td>
<td valign="top" align="left">
<italic>Oligodon taeniolatus</italic>
</td>
<td valign="top" align="left">Streaked Kukri snake</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">India</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B47">Smart et&#xa0;al. (2010)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra heros</italic>
</td>
<td valign="top" align="left">
<italic>Rhinocheilus lecontei</italic>
</td>
<td valign="top" align="left">Long-nosed snake</td>
<td valign="top" align="left">Desert</td>
<td valign="top" align="left">Texas, USA</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B14">Easterla (1975)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="2" align="left">
<italic>Scolopendra</italic> sp.</td>
<td valign="top" align="left">
<italic>Ameivula ocellifera</italic>
</td>
<td valign="top" align="left">Spix&#x2019;s whiptail lizard</td>
<td valign="top" align="left">Natural monument area</td>
<td valign="top" align="left">Grota do Angico, Sergipe, Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B33">Moura et&#xa0;al. (2015)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Calliophis melanurus</italic>
</td>
<td valign="top" align="left">Slender coral snake</td>
<td valign="top" align="left">Rock crevices</td>
<td valign="top" align="left">Mumbai, India</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B29">Mirza and Ahmed (2009)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra</italic> sp<italic>inosissima</italic>
</td>
<td valign="top" align="left">
<italic>Hemibungarus mcclungi</italic>
</td>
<td valign="top" align="left">McClung&#x2019;s Philippine coral snake</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Luzon Island, Philippines</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B1">Acu&#xf1;a et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="3" align="left">
<italic>Scolopendra subspinipes</italic>
</td>
<td valign="top" align="left">
<italic>Calamaria pavimentata</italic>
</td>
<td valign="top" align="left">Collared reed snake</td>
<td valign="top" rowspan="3" align="left">Urban greenery</td>
<td valign="top" rowspan="3" align="left">Singapore</td>
<td valign="top" rowspan="3" align="left">Field</td>
<td valign="top" rowspan="3" align="left">
<xref ref-type="bibr" rid="B42">Pwa et&#xa0;al. (2023)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Calamaria schlegeli</italic>
</td>
<td valign="top" align="left">Red-headed reed snake</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Pseudorabdion longiceps</italic>
</td>
<td valign="top" align="left">Dwarf reed snake</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra viridicornis</italic>
</td>
<td valign="top" align="left">
<italic>Gymnodactylus geckoides</italic>
</td>
<td valign="top" align="left">Naked-toed gecko</td>
<td valign="top" align="left">Semiarid environment</td>
<td valign="top" align="left">Northeastern Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B54">Vieira et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra morsitans</italic>
</td>
<td valign="top" align="left">
<italic>Tympanocryptis tetraporophora</italic>
</td>
<td valign="top" align="left">Eyrean earless dragon</td>
<td valign="top" align="left">Road</td>
<td valign="top" align="left">Australia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B28">McFadden and McFadden (2025)</xref>
</td>
</tr>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="6" align="left">&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;<inline-graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-i003.tif"/> Birds</th>
</tr>
<tr>
<td valign="top" align="left">
<italic>Cormocephalus coynei</italic>
</td>
<td valign="top" align="left">
<italic>Pterodroma nigripennis</italic>
</td>
<td valign="top" align="left">Black-winged petrel (Nestlings)</td>
<td valign="top" align="left">Island</td>
<td valign="top" align="left">Phillip Island, Norfolk Island, Australia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al. (2021)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="3" align="left">
<italic>Scolopendra</italic> sp.</td>
<td valign="top" align="left">Bird</td>
<td valign="top" align="left">Unidentified bird</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Not specified</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B11">Cumming (1903)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">Bird</td>
<td valign="top" align="left">Unidentified bird</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Not specified</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B10">Cloudsley-Thompson (1968)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">Bird</td>
<td valign="top" align="left">Unidentified bird</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Not specified</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B2">Anonymous (1985)</xref>
</td>
</tr>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="6" align="left">&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;<inline-graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-i004.tif"/> Mammals</th>
</tr>
<tr>
<td valign="top" align="left">
<italic>Rhysida</italic> sp.</td>
<td valign="top" align="left">
<italic>Pipistrellus</italic> sp.</td>
<td valign="top" align="left">Bat</td>
<td valign="top" align="left">Tree hollow</td>
<td valign="top" align="left">India</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">This study</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra galapagoensis</italic>
</td>
<td valign="top" align="left">
<italic>Oryzomys bauri</italic> (= <italic>Aegialomys galapagoensis</italic>)</td>
<td valign="top" align="left">Gal&#xe1;pagos rice rat</td>
<td valign="top" align="left">Island</td>
<td valign="top" align="left">Gal&#xe1;pagos Islands, Ecuador</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B9">Clark (1979)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="3" align="left">
<italic>Scolopendra gigantea</italic>
</td>
<td valign="top" align="left">
<italic>Mormoops megalophylla</italic>
</td>
<td valign="top" align="left">Ghost-faced bat</td>
<td valign="top" rowspan="3" align="left">Caves</td>
<td valign="top" rowspan="3" align="left">Venezuela</td>
<td valign="top" rowspan="3" align="left">Field</td>
<td valign="top" rowspan="3" align="left">
<xref ref-type="bibr" rid="B30">Molinari et&#xa0;al. (2005)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Lionycteris curasoae</italic>
</td>
<td valign="top" align="left">Southern long-nosed bat</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Pteronotus davyi</italic>
</td>
<td valign="top" align="left">Davy&#x2019;s naked-backed bat</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra heros</italic>
</td>
<td valign="top" align="left">
<italic>Eptesicus fuscus</italic>
</td>
<td valign="top" align="left">Big brown bat</td>
<td valign="top" align="left">Rock crevices</td>
<td valign="top" align="left">Texas</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B24">Lindley et&#xa0;al. (2017)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra</italic> sp.</td>
<td valign="top" align="left">Rodent</td>
<td valign="top" align="left">Unidentified rodent</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">Not specified</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B45">Shugg (1961)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Scolopendra subspinipes</italic>
</td>
<td valign="top" align="left">
<italic>Mus musculus</italic>
</td>
<td valign="top" align="left">House mouse</td>
<td valign="top" align="left">&#x2014;</td>
<td valign="top" align="left">China</td>
<td valign="top" align="left">Experiment</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B25">Luo et&#xa0;al. (2018)</xref>
</td>
</tr>
<tr>
<td valign="top" rowspan="2" align="left">
<italic>Scolopendra viridicornis</italic>
</td>
<td valign="top" align="left">
<italic>Eptesicus furinalis</italic>
</td>
<td valign="top" align="left">Argentine brown bat</td>
<td valign="top" align="left">Mosaic floor</td>
<td valign="top" align="left">Southeastern Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B48">Srbek-Araujo et&#xa0;al. (2012)</xref>
</td>
</tr>
<tr>
<td valign="top" align="left">
<italic>Molossus molossus</italic>
</td>
<td valign="top" align="left">Velvety free-tailed bat</td>
<td valign="top" align="left">Roof</td>
<td valign="top" align="left">Brazil</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B34">Noronha et&#xa0;al. (2015)</xref>
</td>
</tr>
<tr>
<th valign="top" align="center"/>
<th valign="top" colspan="6" align="left">&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;&#x2003;<inline-graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-i005.tif"/> Fishes</th>
</tr>
<tr>
<td valign="top" align="left">
<italic>Cormocephalus coynei</italic>
</td>
<td valign="top" align="left">Marine fishes from seabird regurgitates</td>
<td valign="top" align="left">Fish</td>
<td valign="top" align="left">Coastal</td>
<td valign="top" align="left">Phillip Island, Norfolk Island, Australia</td>
<td valign="top" align="left">Field</td>
<td valign="top" align="left">
<xref ref-type="bibr" rid="B19">Halpin et&#xa0;al. (2021)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Stacked bar chart showing the number of prey instances consumed by different centipede predator species and also indicate different group of vertebrate. This visualization highlights variation in feeding breadth and prey specialization across predator species.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1634037-g002.tif">
<alt-text content-type="machine-generated">Bar chart showing the number of prey instances across various centipede predator species, categorized by prey type: amphibians (blue), reptiles (orange), birds (yellow), mammals (gray), and fish (navy). Each predator species, labeled along the x-axis, displays a varying combination of these prey types, with Scolopendra gigantea showing the highest instances, particularly in birds.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>Vertebrate predation by centipedes, though often overlooked, is neither rare nor ecologically insignificant. Our review reveals that large scolopendrid centipedes occupy a unique trophic niche, with documented ability to subdue vertebrate prey ranging from amphibians and reptiles to birds and mammals across diverse habitats worldwide. The new record from India involving a <italic>Rhysida</italic> centipede feeding on a live bat in an arboreal setting represents the first such case from Asia and challenges prior assumptions that such interactions are confined to cave or ground environments. Collectively, these findings underscore the ecological versatility, behavioral sophistication, and predatory impact of scolopendrid centipedes. Future research should prioritize quantitative assessments of their predation rates, prey selection, and ecological consequences, especially in tropical and subtropical ecosystems where they may serve as cryptic yet influential components of vertebrate mortality regimes.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="author-contributions">
<title>Author contributions</title>
<p>MK: Conceptualization, Supervision, Writing &#x2013; review &amp; editing, Methodology, Investigation, Writing &#x2013; original draft, Data curation, Software, Project administration, Validation, Formal Analysis. RM: Writing &#x2013; review &amp; editing, Data curation. DB: Visualization, Resources, Funding acquisition, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s7" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. This study was supported by the core funding from the Zoological Survey of India, Ministry of Environment, Forest and Climate Change, Government of India.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We thank Dr. P. M. Sureshan, former senior scientist at the Zoological Survey of India, for his valuable assistance in identifying the centipede genus.</p>
</ack>
<sec id="s8" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s9" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fevo.2025.1634037/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fevo.2025.1634037/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Video1.mp4" id="SM1" mimetype="video/mp4"/>
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