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Autohaemorrhaging
Autohaemorrhaging
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Horned lizard showing evidence of autohaemorrhaging

Autohaemorrhaging, or reflex bleeding, is the action of animals deliberately ejecting blood from their bodies. Autohaemorrhaging has been observed as occurring in two variations.[1] In the first form, blood is squirted toward a predator. The blood of these animals usually contains toxic compounds, making the behaviour an effective chemical defense mechanism. In the second form, blood is not squirted, but is slowly emitted from the animal's body. This form appears to serve a deterrent effect, and is used by animals whose blood does not seem to be toxic.[1] Most animals that autohaemorrhage are insects, but some reptiles also display this behaviour.[2]

Some organisms have shown an ability to tailor their autohaemorrhaging response. Armoured crickets will projectile autohaemorrhage over longer distances when attacked from the side, compared to being attacked from an overhead predator.[3]

Insects

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Several orders of insects have been observed to utilize this defence mechanism.

Reptiles

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A West Indian wood snake displaying autohaemorrhaging. The eyes are fully flooded with blood and some is emerging from the mouth.

Lizards

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  • Horned lizards (Phrynosomatidae). At least six species of horned lizards are able to squirt an aimed stream of blood from the corners of their eyes, up to 5 feet (1.5 m).[12]

Snakes

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  • West Indian wood snake (Tropidophis). Thirteen species have been found to expel blood from the mouth and nostrils while also fully flooding both eyes with blood.[13]
  • European grass snake (Natrix natrix), which secretes blood from the lining of the mouth while playing dead.[14]
  • Long-nosed snake (Rhinocheilus lecontei), which exudes blood from the cloaca.[15]
  • Eastern hognose snake (Heterodon platirhinos), which emits blood from the cloacal region.[15]
  • Plain-bellied water snake (Nerodia erythrogaster), which releases blood from the mouth.[15]
  • Western hognose snake (Heterodon nasicus), which releases blood from the mouth.

Consequences of reflexive bleeding

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In some cases, the loss of blood can be substantial. Beetles may lose up to 13% of their net body weight as a consequence of expelling haemolymph.[16] Autohaemorrhaging may result in dehydration. The ejection of blood puts organisms at risk of cannibalism from other members of their species.[17]

An inactive prepupa Asian ladybeetle autohaemorrhaging, resulting from cannibalism by an adult of the same species.

See also

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References

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[edit]
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Autohaemorrhaging

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Autohaemorrhaging, also known as reflex bleeding, is a defensive in which certain animals deliberately eject haemolymph or blood from their bodies to ward off predators, often rendering the fluid noxious or toxic through chemical additives. This phenomenon has been documented across diverse taxa, primarily in and reptiles, where it serves as a last-resort antipredator triggered by stress or physical provocation. In , autohaemorrhaging typically involves the release of haemolymph from specialized body regions, such as joints, antennal sockets, or the , and is prevalent in families like (ladybird beetles) and Meloidae (blister beetles). For instance, in the harlequin ladybird (), repeated reflex bleeding over three weeks significantly reduces haemocyte concentration, protein content, and antimicrobial activity in the haemolymph, thereby weakening the and delaying by approximately two days without affecting overall egg production or body mass. In blister beetles such as Lydus trimaculatus and Mylabris variabilis, the ejected haemolymph contains high levels of (0.03–0.79 mg/g), a potent produced in the , stored in , and transferred during to enhance defense, causing or in predators. Among reptiles, autohaemorrhaging is observed in various snakes and , often involving ocular or oral discharge of during threat responses like death feigning. In dice snakes (Natrix tessellata), about 10.6% of individuals (higher in adults at 12.4% than juveniles at 2.1%) release from the mouth during death feigning, which, when combined with musking and fecal smearing, shortens the feigning duration by roughly two seconds, potentially signaling unpalatability more effectively. A notable example occurs in the Bahamian pygmy boa (Tropidophis curtus barbouri), where gentle head pressure induces rapid exudation from the eyes (within 0.7 seconds) and mouth (0.8 seconds later), clearing in about 1.6 seconds as part of a multi-component defense including and release, though the exact antipredator efficacy remains unconfirmed. Despite its spectacular nature, autohaemorrhaging incurs physiological costs, such as increased susceptibility in , and its prevalence varies by age, sex, and environmental factors across .

Definition and Purpose

Definition

Autohaemorrhaging, also known as reflex bleeding, is the deliberate release or ejection of haemolymph or from an animal's body as a last-resort antipredator defense mechanism when threatened or attacked. This behavior involves externalizing body fluids, often containing toxins or distasteful compounds, to deter predators by making the prey unpalatable, startling them visually, or signaling unprofitability. It is a costly strategy, as it can lead to significant physiological losses for the animal. In , autohaemorrhaging specifically entails the expulsion of haemolymph—the open circulatory fluid analogous to in vertebrates but lacking and red blood cells—from specialized body sites such as leg joints or integumental ruptures. This contrasts with vertebrates like certain s, where true is ejected, typically carrying oxygen via . The phenomenon is widespread in various insect orders, including Coleoptera and , and occurs in some reptile taxa, though detailed mechanisms vary by group. Autohaemorrhaging manifests in two primary variations: active squirting, where the fluid is forcefully directed toward the predator, often laced with or distasteful chemicals to enhance deterrence; and passive slow emission, involving a gradual oozing that serves as a visual or olfactory warning signal, sometimes without strong toxicity but relying on aposematic cues. In active forms, such as in some blister beetles, haemolymph is deliberately expelled under pressure from joints, while passive release may occur through minor physical damage or reflexive rupture. This defense has fascinated biologists for centuries, with reflex bleeding noted as a response to disturbance in like ladybugs.

Evolutionary Significance

Autohaemorrhaging represents a convergent defensive trait that has evolved independently in and reptiles, distantly related taxa facing similar predation pressures, underscoring its adaptive value as an antipredator strategy. This behavior likely emerged under strong selective forces in environments where rapid, chemical-based deterrence provided survival benefits without necessitating elaborate morphological structures, though it incurs physiological costs such as loss and immune compromise. In , autohaemorrhaging has arisen multiple times across at least four orders, including Coleoptera, , , and , with particular prevalence in beetle families like and Meloidae. The trait's distribution suggests independent in response to diverse predators, including birds, mammals, and , where the release of noxious exploits aversion to bitter or toxic compounds, enhancing escape probabilities despite associated fitness trade-offs like delayed reproduction. Comparative studies indicate it functions as a secondary defense, complementing other tactics in resource-limited settings. Among reptiles, autohaemorrhaging is documented in select lineages, notably the horned lizard genus Phrynosoma, where it is plesiomorphic and retained in six of 13 species, with losses in specific clades such as P. mcallii-modestum-platyrhinos. This ocular blood ejection targets mammalian predators like canids, delivering irritant chemicals that induce learned avoidance, thereby increasing survival rates in encounters. Observations in snake families, including Tropidophiidae, further highlight its sporadic occurrence, often integrated with behaviors like death feigning. Phylogenetic analyses infer multiple independent origins of autohaemorrhaging from extant distributions, with no direct available.

Physiological Mechanisms

In

In , autohaemorrhaging, also known as reflex bleeding, involves the deliberate rupture of thin membranes at exoskeletal joints, such as the femorotibial or intersegmental areas, to release haemolymph under as a defensive response to threats. This process allows haemolymph to exude from sites like leg joints or under the pronotum, where the softer, thinner facilitates controlled leakage without compromising the insect's overall structural integrity. The release is triggered by mechanical stress, such as grasping or attack, leading to the haemolymph emerging as droplets or jets that deter predators through distastefulness or entanglement. Pressure for haemolymph ejection is generated primarily through muscle contractions that elevate hydrostatic pressure within the open circulatory system, often supplemented by body deformation or abdominal compression, enabling expulsion over distances of up to several centimeters. The primary force typically arises from localized activity around the release sites. This mechanism leverages the insect's haemocoel, where haemolymph bathes internal organs, to rapidly mobilize the fluid without requiring specialized vascular rupture. The released haemolymph frequently contains defensive chemicals, such as alkaloids or phenolics, which impart toxicity or repellency; for instance, in ladybugs (), coccinellins like coccinelline are present at concentrations that make the fluid unpalatable to predators. These compounds are biosynthesized via pathways involving precursors, accumulating in the haemolymph to reinforce its role in . In other groups, such as , the haemolymph exhibits an acrid odor due to similar phenolic constituents. Certain exhibit directional control over haemolymph ejection, adjusting the angle and force based on the threat's position through neural reflexes that coordinate muscle responses. For example, in armoured ground crickets (), side-directed attacks prompt greater projection distances (up to 10 cm) from leg seams compared to overhead threats, optimizing coverage toward the predator via proprioceptive feedback. This reflexive modulation enhances the defense's efficacy by targeting the assailant more precisely.

In Reptiles

Autohaemorrhaging in reptiles involves the deliberate rupture of specific vascular structures, such as the in or mucosal linings in the oral cavity and nasal passages of snakes, as a reflexive defense mechanism triggered by activation of the . This process is elicited during perceived threats, where neural signals prompt rapid physiological changes leading to blood ejection from weak points in the vascular system. Unlike the open circulatory systems of , reptiles possess a closed that enables precise control over and distribution, allowing for targeted release of true containing and defensive compounds. The buildup of pressure necessary for autohaemorrhaging is achieved through cardiac and , which elevate in the head and ocular regions, forcing through fragile vascular sites. In , specialized muscles encircling the major veins around the eyes contract to restrict outflow, causing the circumocular sinuses—capillary beds near the ocular orbits—to swell and rupture, propelling outward. This can result in ejection distances of up to 1.5 , depending on the and intensity of the stimulus. In snakes, similar pressure increases lead to tears in mucosal linings of the , nostrils, or cloacal vasculature, with emerging without the need for extreme . Reflex triggers for autohaemorrhaging are primarily mediated by , such as adrenaline (epinephrine) and norepinephrine, released in response to handling, predation attempts, or physical restraint. These catecholamines activate the sympathetic branch of the , inducing and to amplify intrathoracic and . The response is highly reflexive, occurring within seconds of stimulation, and serves an evolutionary role in antipredator behavior by startling or repelling assailants through the sudden release of potentially noxious blood. While the exact neural pathways remain under study, the integration of hormonal and muscular components ensures a coordinated ejection that enhances survival without permanent damage in most cases.001[1114:BSVIHL]2.0.CO;2/Blood-Squirting-Variability-in-Horned-Lizards-Phrynosoma/10.1643/0045-8511(2001)001[1114:BSVIHL]2.0.CO;2.full)

Examples in Insects

Beetles and Ladybugs

In the family , commonly known as ladybugs or ladybird beetles, autohaemorrhaging manifests as reflex , where haemolymph is ejected from the femoro-tibial joints of the legs in response to predator contact or disturbance. This defensive response releases droplets of orange or yellow fluid containing alkaloids such as coccinelline and adaline, along with methoxypyrazines like 2-isopropyl-3-methoxypyrazine, which produce a bitter and pungent to repel attackers. These chemicals effectively deter a range of predators, including , birds, and , as demonstrated in observational studies where vertebrates rejected ladybirds after initial contact with the reflex fluid. The behavioral trigger for this ejection is typically mechanical stimulation, such as grasping by a predator, leading to rapid haemolymph discharge that can project several millimeters from the body. In some species, a single event may involve the loss of 10–12% of the beetle's fresh body weight, highlighting the between immediate benefits and potential or nutritional depletion. Seminal research, including early experiments with birds and , has confirmed the efficacy of these alkaloids in reducing predation rates, with ladybirds often surviving encounters that would otherwise be fatal. Autohaemorrhaging also occurs in other families within Coleoptera, often integrated with additional defenses. For instance, in blister s of the family Meloidae, such as Lydus trimaculatus and Mylabris variabilis, haemolymph containing high levels of (0.03–0.79 mg/g) is ejected, acting as a potent irritant or to predators. is biosynthesized in and transferred to females during for offspring defense. In darkling s of the family Tenebrionidae, such as the blue death-feigning Asbolus verrucosus, larvae release haemolymph during when threatened, using the viscous fluid as an adhesive to bind environmental debris like sand particles for and against visual predators. Adults in this family complement reflex bleeding with noxious sprays ejected from pygidial glands, creating a synergistic chemical barrier that enhances overall deterrence.

Other Insect Orders

Autohaemorrhaging manifests in diverse forms across several insect orders beyond Coleoptera, often tailored to the species' ecology and predators, though generally involving smaller volumes of haemolymph compared to the copious releases seen in many beetles. In the order , particularly within the family Cercopidae (froghoppers or spittlebugs), adults exhibit reflex bleeding by exuding haemolymph from specialized rupture lines in the pretarsal membranes of their legs when disturbed. This behavior is closely linked to aposematic coloration, signaling unpalatability, and the released haemolymph is typically odoriferous and distasteful, deterring predators such as and birds; for instance, in species like , the haemolymph can inhibit predation without fully preventing attacks but often combined with evasive actions. Within the order , autohaemorrhaging is prominently featured in armoured crickets of the family , such as Acanthoplus discoidalis. These actively squirt haemolymph from pores along their pronotal margins and leg articulations, achieving directional propulsion with projection distances averaging about 4 cm (up to approximately 10 cm) to target approaching threats like spiders or mantises. The ejected fluid is toxic, containing defensive chemicals that cause irritation or toxicity upon contact, enhancing its efficacy as a standoff weapon; this squirting is integrated with other defenses like loud and feigning death, observed in field encounters where it successfully repels predators. In , a related passive defense known as "easy bleeding" occurs in certain sawflies (), where larvae release haemolymph from easily ruptured integumental points across the body upon contact with predators. This slow seepage exposes plant-derived alkaloids sequestered in the haemolymph to deter and other predators, entangling attackers with coagulating fluid and providing time for escape; unlike active ejection, it involves low mechanical resistance of the integument. Studies on species like Athalia rosae demonstrate its role in reducing ant , though it incurs physiological costs similar to more vigorous forms. Overall, these instances across , , and highlight autohaemorrhaging's , with less voluminous discharges than in beetles but frequently augmented by pheromones or toxins.

Examples in Reptiles

Lizards

Horned lizards in the genus Phrynosoma employ autohaemorrhaging as a striking antipredator , rupturing specialized blood vessels in their circumorbital sinuses to eject streams of blood from the eyes toward threats. This behavior is particularly effective against canine predators, such as coyotes and foxes, which are common in their arid habitats. The defensive display often follows initial attempts at or flight, escalating when the lizard is grasped or closely approached. At least eight species of Phrynosoma have been confirmed to perform blood-squirting, including P. cornutum (Texas horned lizard) and P. solare (regal horned lizard), out of the 17 recognized in the genus. The ejection can reach distances of up to 1.5 meters and is remarkably accurate, often directed at the predator's face or mouth. In experimental trials simulating predation, larger individuals squirted more frequently and expelled greater blood volumes, suggesting a correlation with body size that enhances defensive efficacy. Vascular rupture in the ocular region allows for rapid release under increased intraocular pressure, a physiological adaptation unique among vertebrates to this form of autohaemorrhaging. This autohaemorrhagic response integrates with other behaviors, such as to appear larger, of cranial spines for passive defense, and of a rigid, immobile posture to mimic a non-prey item. The squirted blood deters attackers due to its foul taste and irritating properties, primarily attributed to elevated concentrations derived from the lizards' diet of harvester ants (Pogonomyrmex spp.). These chemical defenses exploit the predators' aversion to the ants' , turning the lizard's own blood into an unpalatable deterrent. Field studies in North American desert ecosystems, including the Chihuahuan and Sonoran Deserts, document this behavior in natural encounters, where deploy it against native predators. Observations indicate that individuals can repeat the blood ejection up to five times during a single predatory event before depleting reserves, allowing temporary escape while the threat is repelled. Such repeatability underscores the mechanism's reliability, though it incurs costs like blood loss and potential in harsh environments.

Snakes

Autohaemorrhaging in snakes primarily occurs among colubrid species, where it serves as a defensive response to predation threats. The European grass snake (Natrix natrix) secretes blood from the lining of the mouth during thanatosis, or death feigning, to mimic a diseased or unpalatable carcass. Similarly, the dice snake (Natrix tessellata), another colubrid, releases blood around the mouth while exhibiting tonic immobility, often inverting its body and gaping to enhance the display. Other snake genera demonstrate varied ejection sites for autohaemorrhaging. Hognose snakes (Heterodon spp.) emit blood from the mouth and nostrils as part of an elaborate bluff, combining it with hissing, flattening the head, and rolling onto the back. The West Indian wood snake (Tropidophis spp.), a dwarf boa, expels blood from the eyes, mouth, and nostrils when stressed, with observations showing rapid filling of the eyes followed by drops from the mouth after gentle head pressure. In contrast to the specialized ocular squirting in certain , snakes exhibit more flexible multi-site bleeding to broadly deter predators. This behavior in snakes is frequently paired with other defenses, such as musking—releasing foul cloacal secretions—and , which coat the body and amplify the illusion of decay. The ejected blood may contribute to deterrence through visual cues or chemical properties, with some studies indicating it can be distasteful or irritating to predators, including mammals, potentially due to bitter compounds. Recent research highlights the synergistic effects of autohaemorrhaging with these behaviors; in dice snakes, individuals combining bleeding during death feigning with prior musking and fecal smearing spent significantly less time immobilized—approximately 2 seconds shorter—allowing quicker escape from simulated predator handling.

Defensive Functions and Consequences

Defensive Benefits

Autohaemorrhaging serves as a potent antipredator strategy by releasing haemolymph or blood laden with noxious compounds, thereby deterring assailants through chemical means. In , particularly ladybird beetles (), the haemolymph expelled via reflex bleeding is rich in alkaloids such as coccinellins, including coccinelline, which impart toxicity and bitterness that cause aversion or illness in predators ranging from to birds and small mammals. These chemicals trigger immediate rejection behaviors, such as grooming or retreat, effectively halting predation attempts. In reptiles, such as horned lizards (Phrynosoma spp.), the blood contains small peptides derived from metabolized , which elicit profound revulsion and distaste in mammalian predators like coyotes and dogs. The abrupt ejection of fluids during autohaemorrhaging also exploits visual and startle responses, surprising predators and creating a brief for escape, particularly against visually hunting such as birds or diurnal mammals. This sudden display amplifies the chemical signal by drawing attention to the distasteful release, enhancing its deterrent impact. Laboratory and field studies provide empirical support for these benefits, demonstrating substantial reductions in attack persistence. For instance, in interactions between and ladybird larvae, contact with reflex blood causes predators to invariably abort attacks and engage in self-cleaning, achieving near-complete deterrence. Similarly, in horned lizards, blood-squirting elicits aversion in approximately 85% of encounters with dogs, significantly lowering the likelihood of continued assault. These outcomes underscore substantial reductions in predation success across tested scenarios involving insects and reptiles. Autohaemorrhaging often integrates synergistically with complementary defenses, amplifying survival probabilities. In horned lizards, it bolsters cryptic and spiny cranial projections, forming a multi-layered strategy that confounds predators before chemical ejection becomes necessary. In insects like , reflex bleeding complements aposematic coloration, where the sudden bleed reinforces warning signals and facilitates evasion. For example, the horned lizard's targeted squirting deters approaching dogs by combining startle with chemical repulsion in a single act.

Physiological Costs

Autohaemorrhaging entails substantial physiological burdens for the organisms that utilize it, stemming from the expulsion of in or in reptiles, which depletes vital fluids containing nutrients, proteins, and immune cells. In ladybird beetles (Harmonia axyridis), a typical reflex bleeding episode results in the loss of 0.44–0.52 µl of , while extreme cases can reach up to 2.2 µl, approximating 5% of the adult's live body mass. This volume reduction induces , impairing circulation and nutrient distribution, which can lead to decreased mobility and functional equivalents of through diminished oxygen-carrying capacity. In reptiles, such as the (Phrynosoma cornutum), a single blood-squirting event may expel 0.5–53% of total , causing acute hemodynamic instability, reduced cardiovascular performance, and heightened risk of circulatory collapse. Frequent autohaemorrhaging episodes compound these effects by compromising immune competence. In a 2020 experimental study on ladybird beetles, individuals subjected to reflex bleeding twice weekly exhibited significantly lower hemocyte concentrations (P < 0.05), reduced protein levels (P < 0.05), and diminished antimicrobial activity against the bacterium (P < 0.05), with a near-significant decline against (P = 0.073). Such suppression weakens overall hemocyte function, elevating susceptibility to bacterial infections and parasitoids, thereby increasing disease vulnerability in the post-bleeding period. Reproductive fitness also suffers due to the energetic demands of replenishment and fluid . The same study found that repeated bleeding delayed the age at first in female ladybirds by approximately 2 days (P = 0.037), with greater loss correlating to later onset of oviposition (P = 0.047), though cumulative output over 30 days was unchanged. Concurrently, the loss of water and electrolytes disrupts , promoting and osmotic stress that diverts metabolic resources from production. The resultant physiological debilitation post-autohaemorrhaging amplifies predation risks by rendering the animal more susceptible to secondary attacks or conspecific owing to impaired locomotion and vitality.

References

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