"Modern snakes have lost all hindlimb elements except for the femur. Some alethinophidians (pythons and boids) have a single claw-like element of uncertain homology distal to the femur, used in courtship." — Michael Lee and Michael Caldwell, Philosophical Transactions of the Royal Society B, 19981
"Python embryos initiate formation of hindlimb buds, but leg development is not sustained, resulting in formation of a rudimentary femur and terminal claw." — Francisca Leal and Martin Cohn, Current Biology, 20162
Summary
The parent article, Vestigial Structures, separates two claims: that a structure is useless, and that it is a reduced copy of something larger in an ancestor. This article asks both of the pair of claws beside a python's vent, and finds that the first claim is not made by anyone who has dissected one, and that the second is conceded by nearly every creationist who has written on it.
- The spurs are hind limbs in miniature. Boas and pythons carry on each side of the vent a horny claw over a short femur, hung from a pelvic rod buried in the body wall. Lee and Caldwell's 1998 monograph on the fossil snake Pachyrhachis states the living condition: "Modern snakes have lost all hindlimb elements except for the femur. Some alethinophidians (pythons and boids) have a single claw-like element of uncertain homology distal to the femur, used in courtship."1 Palci and colleagues' 2020 review of every snake pelvis known finds that the ancestor of living snakes "most probably retained all three pelvic elements and rudimentary hindlimbs (femoral spurs)," that the one rod left in blind snakes is the ischium and the one rod left in boas and pythons is the pubis.3 Over 85 percent of living snake species have no limb skeleton at all.4
- They work, and the anatomists have said so all along. The spurs "have musculature, innervation, and vascularisation and can move autonomously," and "are known to be used by males during combat, courtship, and mating."5 Male Bahamian boas carry spurs "nearly twice as long and notably more curved than females."6 Palci and colleagues read the spur, femur and pubis as "related functionally to the widespread use of the hindlimbs in mating and courtship, rather than the usual representation of the bones as non-functional vestiges."3 The useless-organ claim is not in the specialist literature.
- The embryo starts a leg and stops. Python embryos form a hind-limb bud with a real ridge and a real signalling centre; Sonic hedgehog switches on in a few cells and switches off within a day, the ridge decays, and cartilage condensations that "could be anlagen of the femur, tibia and fibula, and digital plate" form and then dissolve.2 The cause is in the limb enhancer of Sonic hedgehog: three deletions shared by pythons and boas,2 and a 17-base deletion, found in all six snakes Kvon and colleagues sequenced,4 that "is the only sequence that is deleted in all snakes but present in all examined limbed vertebrates and fish."4 A mouse given the python enhancer grows truncated limbs; put the 17 bases back and "the resulting knock-in mice with the modified python allele had normal limbs."4
- The fossils give legged snakes, not a lineage. Najash from Patagonia had "a sacrum supporting a pelvic girdle and robust, functional legs outside the ribcage";7 Pachyrhachis, Haasiophis and Eupodophis from the Cretaceous Tethys had femur, tibia, fibula and tarsals but "lack a sacral region."7 Their place on the tree has been disputed since 1997. The "four-legged snake" Tetrapodophis is probably a lizard.8
- The creationist literature concedes the reduction. Answers in Genesis in 2009: "creationists have generally understood this to mean that the original serpentine created kind had legs," and "The leglessness of snakes today represents a corruption, loss, or deactivation of that information."9 This answers the parent article's open question: for the snake pelvis, creationists do treat the structure as a reduced homologue of a leg. What they deny is that the legged ancestor was a lizard, and what they add is Genesis 3:14 as the occasion of the loss. The replies that list functions answer a claim the primary literature does not make.
- The verdict. The spurs pass all three steps of the live vestige test. The creationist position that fits the evidence, a legged snake kind whose legs were switched off, has to carry one enhancer deletion shared by boas, pythons and the advanced snakes, an enhancer that has piled up extra changes only in the snakes with no limb bones left, and legged fossils whose limbs are well formed rather than ruined.
Part 1: What the spurs are
Darwin listed them in 1859 among his rudimentary organs: in some snakes, he wrote, "there are rudiments of the pelvis and hind limbs."10
On either side of the cloaca of a boa or python sits a horny, curved claw. Under the claw is a bone. Lee and Caldwell, describing the modern condition as the baseline for their fossil, put it in one sentence: "Modern snakes have lost all hindlimb elements except for the femur. Some alethinophidians (pythons and boids) have a single claw-like element of uncertain homology distal to the femur, used in courtship."1 Leal and Cohn's 2016 embryology paper gives the adult result of python development as "a rudimentary femur and terminal claw."2 Romer's Osteology of the Reptiles says which snakes keep the thigh bone: the femur "is generally absent in snakes, but a vestige may persist in typical boids, leptotyphlopids, and Anilius."11
Inside, the femur hangs from a pelvis. Which bone of the pelvis is the question Palci, Hutchinson, Caldwell, Smith and Lee set out to settle in 2020, in a review of pelvic and hind-limb elements across living and fossil snakes. Their abstract gives the result. The most recent common ancestor of all living snakes "most probably retained all three pelvic elements and rudimentary hindlimbs (femoral spurs)." Since then "there have been multiple losses of ossified pelvic and hindlimb elements and regaining of ossified pelvic elements." And the two great branches of living snakes have reduced the pelvis by different routes: "The single remaining rod-like element in some scolecophidians is the ischium, whereas the single remaining rod-like element in many basal alethinophidians is the pubis."3 The scolecophidians are the blind snakes and thread snakes; the basal alethinophidians include the pipe snakes, boas and pythons.
What the pelvis has lost is its connection to the spine. Lee and Caldwell list it as a character of modern snakes: "In modern snakes, the sacral contact is lost and the pelvic rudiments, when present, lie within the ribcage."1 A python's pelvic rods are free bones in the muscle of the body wall, behind the last ribs, with a femur and claw on the end. The whale's pelvic rod sits the same way.
How many snakes carry any of this? Kvon and colleagues, choosing six genomes to span the range, describe the two ends: "basal snakes (boa and python) that retained a vestigial pelvic girdle and rudimentary hindlimbs," and "advanced snakes (viper, rattlesnake, king cobra, and corn snake) that completely lost all skeletal limb structures and represent the majority (>85%) of all extant snake species."4 Philip Bell of Creation Ministries International gives the same picture from the creationist side: "some types of snake (e.g. wormsnakes, pipesnakes, boas and pythons) do have a small pelvic girdle and tiny hind limbs. However, the majority of today's snakes (1,600 species of the colubrid family) lack a pelvis."12
The spurs are not the same size in the two sexes. Anzai and colleagues summarise the literature back to Hoge in 1947: "These pelvic spurs are generally larger in males than in females."5 Hoefer, Robinson and Pinou measured it in 32 Bahamian boas in 2021: "mean pelvic spur length in males being 2.6 mm (n = 16; range: 0.5 – 5.0 mm) compared to the 1.5 mm in females (n = 16; range: 0.5 – 3.0 mm)," and against body length "the proportion of spur length relative to SVL was 0.3% in males and 0.1% in females."6
Part 2: What they do
The spurs are moved by the snake. Anzai and colleagues, writing in The Herpetological Bulletin in 2023, summarise sixty years of anatomy: "pelvic spurs have musculature, innervation, and vascularisation and can move autonomously," citing Hoge 1947, Bellairs 1950 and Carpenter and colleagues 1978 and 1979. And they summarise the behaviour: "The pelvic spurs of boids and pythonids are known to be used by males during combat, courtship, and mating."5
Anzai and colleagues watched captive Boa constrictor at two Brazilian breeding facilities, six males and four females in 2019 and five males and three females in 2021. "the males began to rub their spurs gently on the side of the females' bodies, moving in a posterior to anterior direction," and "The spurs repeatedly moved from a resting to an erect state."5 They read the movement as a stimulus that helps the female shift into position for cloacal contact, and add a reason it might matter: "Spur use may be particularly relevant for smaller males because they may not have enough strength to move the body of large females."5 Three of the females later gave birth to 51 young, and the authors set that against a 1983 report of failed births after matings with little spur use: "Our data would appear to reinforce the critical role of pelvic spurs for successful courtship and mating in red-tailed boas."5 That is a suggestion from two facilities, not a controlled test, and the authors' "would appear" is the right strength.
Pythons use the spurs harder. Trivedi and colleagues watched a male Indian rock python with three larger females at a Gujarat zoo on 2 February 2024, and describe the male working the female "not with gentle rubbing but through more forceful poking actions." "This behaviour, reminiscent of finger-walking but executed with the spurs, serves to provoke a reaction from the female, likely due to discomfort or pain caused by the spurs' sharpness."13 Their reading of why is the same as Anzai's: "it is plausible that this behaviour has evolved as a male strategy to navigate the challenges of mating with larger females."13
Senter, Harris and Kent put 33 courtship and combat behaviours onto a snake phylogeny in 2014. Two of the 33 are spur behaviours. The spur rub "is widespread enough in Boidae to infer that it is ancestral for the family," so that "courtship in Boidae included the spur rub by the Paleocene, the date of the earliest known crown-group boid."14 The spur poke in combat "is recorded only in the sister genera Eunectes and Epicrates," the anacondas and the West Indian boas.14 Their Boidae is the broad one that includes the pythons.
The function is real, sex-linked and old, and it is the kind the vestige argument allows. The parent article shows that the modern argument was built on functional vestiges from the start. Here the point is sharper than for the whale, because the people who make the homology claim are the same people who describe the use. Lee and Caldwell's sentence calling the claw a hind-limb element ends "used in courtship."1 Palci and colleagues, having traced the pelvis bone by bone from the oldest fossils to living forms, close their abstract with the function: "many basal alethinophidians share a distinctive configuration of cloacal spur (claw), femur and a sizeable pubis, which is likely to be related functionally to the widespread use of the hindlimbs in mating and courtship, rather than the usual representation of the bones as non-functional vestiges."3 The anatomists never held the popular view.
Part 3: The embryo starts a leg, and the enhancer that stops it
Cohn and Tickle opened this question in Nature in 1999. In python embryos, "Hindlimb buds are initiated, but apical-ridge and polarizing-region signalling pathways that are normally required for limb development are not activated. Leg bud outgrowth and signalling by Sonic hedgehog in pythons can be rescued by application of fibroblast growth factor or by recombination with chick apical ridge."15
Leal and Cohn went back to earlier embryos in 2016 and found that the python bud is closer to a leg than that. Sonic hedgehog does switch on: "SHH mRNA was detected in a small domain of cells at the posterior margin of the hindlimb buds, but this ZPA-like expression pattern is transient, disappearing within 24 hr of oviposition."2 The ridge is there too: "A morphological AER is evident in python hindlimb buds at stage 1," and then "Shortly after termination of SHH, the FGF8 domain begins to degrade."2 Yet the distal genes keep going, and the bud lays down cartilage templates: the SOX9 domains in the python bud "could be anlagen of the femur, tibia and fibula, and digital plate," and "they are transitory structures, as distal hindlimb elements are not found in mature pythons."2 A python embryo sketches a shin and a foot and erases them.
The reason is a stretch of DNA about a million bases from the gene it controls. Leal and Cohn describe the ZRS as "a limb-specific enhancer located ∼1 Mb upstream of SHH, in intron 5 of the LMBR1 gene."2 In pythons it carries three deletions, ΔA, ΔB and ΔC, each of which "occurred in HOXD13 binding sites,"2 and the same three are in boas: "Our identification of the same three mutations in the ZRS of pythons and boas indicates that ΔA, ΔB, and ΔC were present in their last common ancestor."2
Kvon and colleagues, in Cell the same month, tested the enhancer in mice. They compared six snake genomes with limbed vertebrates and fish, and found the boa and python ZRS still about 80 percent identical to a lizard's and changing at the ordinary rate for this enhancer, while "the ZRS of advanced snakes displays a substantially increased number of substitutions compared to other enhancers."4 Then they replaced the mouse enhancer with the snake versions. The cobra sequence gave "a complete loss of Shh expression and a truncated limb phenotype, affecting both the fore- and hindlimbs, that is indistinguishable from the phenotype caused by deletion of the mouse enhancer."4 "The less diverged python ZRS resulted in a similar but a slightly milder phenotype."4 One change stood out: a 17-base deletion in a region conserved from fish to mammals, and "this deletion is the only sequence that is deleted in all snakes but present in all examined limbed vertebrates and fish."4 It removes a binding site for the transcription factor ETS1, and "all examined snakes have lost the E0 and E1 ETS motifs."4 Putting the 17 bases back into the python enhancer was enough: "the resulting knock-in mice with the modified python allele had normal limbs."4
So the boa and python enhancer is a damaged limb enhancer, not a different kind of element, and the damage is graded: light in the snakes that keep a femur, heavy in the snakes that keep nothing.
Not every limb enhancer decayed. Infante and colleagues found in 2015 that "snake genomes retain many sequences orthologous to mammalian limb enhancers,"16 and showed why for one of them: the Tbx4 enhancer HLEB works in both hind limbs and genitals, and in snakes it has "functionally diverged along such a path with snake HLEBs losing hindlimb function but retaining activity within subdomains of the external genitalia."16 That is the finding Fazale Rana cites as shared design. It does not extend to the ZRS, which is limb-specific.
Part 4: The fossil snakes with legs
Five fossils carry the argument, and their describers disagree about what they mean.
Pachyrhachis problematicus, from Cenomanian marine limestone near Ramallah, was reinterpreted by Caldwell and Lee in 1997 as "a primitive snake with a well developed pelvis and hindlimbs," which "retains a well developed sacrum, pelvis and hindlimb (femur, tibia, fibula, tarsals)," and which they placed as "the sister-taxon of all other snakes."17 Their 1998 monograph gives the pelvis: "The three elements of the pelvic girdle are not fused or sutured together. Rather, they appear to have abutted each other, forming a very weak girdle,"1 but the ilium "closely overlies the distal end of the sacral rib, indicating that the ilium was external to the ribcage and suggesting the presence of a sacral contact."1 Below the femur, "Pachyrhachis is more primitive than modern snakes in retaining a normal tibia, fibula, astragalus and calcaneum."1
Haasiophis terrasanctus, from the same beds, was described by Tchernov, Rieppel, Zaher, Polcyn and Jacobs in Science in 2000 as "the most extreme hindlimb development of any known member of that group, as it preserves the tibia, fibula, tarsals, metatarsals, and phalanges." And they drew the opposite tree: their analysis "shows both to be related to macrostomatans, a group that includes relatively advanced snakes such as pythons, boas, and colubroids to the exclusion of more primitive snakes such as blindsnakes and pipesnakes."18 On that reading their legs are either retained or regrown. Leal and Cohn's model takes the second option seriously, proposing that "Re-emergence of the tibia and fibula (zeugopod) and digits (autopod) in Tethyan snakes may have resulted from persistence of embryonic skeletal condensations into adulthood,"2 the same condensations a python embryo forms and erases.
Eupodophis descouensi, from Lebanon, was scanned by synchrotron in 2011. Houssaye and colleagues report that the scan "allowed a virtual exhumation of the second, hidden leg of the specimen," that "four tarsals are observed in each limb, but there are no traces of metatarsals or phalanges," and that the limb bones' inner structure "displays a microanatomical organization similar to that of extant terrestrial lizards. This suggests that limb regression in Eupodophis was not due to a qualitative alteration of growth but, more likely, to a local decrease in growth rate or shortening of growth duration."19
Najash rionegrina, from the Upper Cretaceous of Patagonia, is the one with a hip that works. Apesteguía and Zaher in 2006: "Here we report a fossil snake with a sacrum supporting a pelvic girdle and robust, functional legs outside the ribcage." The three marine snakes "lack a sacral region"; Najash "is the most primitive (basal) snake known."7 New skulls and skeletons scanned by Garberoglio and colleagues in 2019 "provide robust support for an extensive basal radiation of early snakes with hindlimbs and pelves, demonstrating that this intermediate morphology was not merely a transient phase between limbed and limbless body plans."20
Tetrapodophis amplectus was announced in Science in 2015 as a snake with four limbs, whose limbs "were adapted for grasping, either to seize prey or as claspers during mating."21 Caldwell and colleagues examined the specimen and its counterpart slab and in 2021 reported "new evidence refuting it as a snake"; four analyses "unambiguously recover Tetrapodophis as a dolichosaur," a marine lizard.8 Creationist writers were cautious about it from the start, and were right to be.
What the record supports, then, is this. There were snakes with a sacrum, a hip outside the ribs and working legs (Najash); snakes with a weak, unfused hip still touching a sacral rib and a full lower leg (Pachyrhachis, Haasiophis); living snakes with free pelvic rods, a femur and a claw (boas, pythons); and living snakes with nothing. Each stage keeps less of the same parts; it is a series, not a lineage. Apesteguía and Zaher themselves warn that the old picture of "progressive loss of their limbs by gradual diminution of their use" has given way to "a more complex scenario of limb reduction, still poorly documented in the fossil record,"7 Palci and colleagues count "multiple losses" and "regaining" of pelvic bones,3 and the Tethyan snakes may be side branches whose legs are secondary. The Jurassic snakes described in 2015 are known from fragments, and as Philip Bell noted for CMI, "No trace of limbs or even limb girdles (pelvis or pectoral) were found in the four 'oldest' fossils."22 Fragments prove nothing either way.
Part 5: Does it pass the vestige test?
The parent article's live definition has three steps: homology with a fuller structure elsewhere, reduction, and a distribution that fits a tree built from other characters.
Step 1 is carried by the bones, the embryo and the enhancer together. The pelvic rods are ischium or pubis by position and by their history in the fossils;3 the bone under the claw is a femur, and in Pachyrhachis a "normal tibia, fibula, astragalus and calcaneum" sit below it, bones that carry lizard names because they are lizard bones.1 The python bud forms an apical ridge, a polarizing region, Sonic hedgehog, FGF8 and HOXD13 in the places a lizard's does, and condenses cartilage for a femur, tibia, fibula and digital plate.2 The limb enhancer is the lizard's at 80 percent, and drives a mouse leg when 17 bases are restored.4 Step 2, reduction, is conceded by every source on both sides; the creationist literature uses the words "tiny," "small" and "reduced" throughout. Step 3 is the tree. The legged snakes are Cretaceous; the sacrum-bearing Najash comes out basal on the analyses that include it;20 the enhancer deletion is in every snake and no limbed animal;4 and the three python deletions are in boas too.2 The parts of step 3 that are contested, the placement of the Tethyan snakes and whether legs re-emerged, do not touch the spurs; they concern which fossil is nearest the root.
A creationist must say one of two things instead. Either the spur is not a limb at all, which no creationist source found says; or it is a limb, reduced within the snake kind from a created snake that had larger ones, which nearly all of them say. Part 6 states that position from its own sources and prices it.
Part 6: The creationist literature, in its own words
The text under everything is Genesis 3:14: "upon thy belly shalt thou go, and dust shalt thou eat all the days of thy life."23 Answers in Genesis reads it with care: "While the Bible does not explicitly state that the serpent had legs, the divine curse implies a transformation from a previous condition, leading some to infer it might have had legs or a different form of movement before the fall," and whether it had legs "remains a point of theological speculation."24 Sarfati and Batten's Creation Answers Book entry says only that "Genesis 3:14 suggests that snakes maybe once had legs."25
The creationist position on the spurs has been consistent for twenty years, and it is a reduction position. Sarfati in 2008, on Eupodophis: "We don't know exactly what God did to the serpent, but one possibility is that He turned off the genetic information to make the legs, hips and other features necessary to walk."26 Answers in Genesis in 2009: "creationists have generally understood this to mean that the original serpentine created kind had legs. Even if God 'muted' the genetic information for legs in snakes (so that none could truly walk), a mutation could cause some of the latent genetic information to be expressed in the form of an errant limb."9 Bell in 2009: "it is possible that some snakes had legs to begin with."12 Elizabeth Mitchell in 2016, on the 17-base deletion: "it is quite possible the 17 absent base pairs apparently responsible for the snake's signature slithery nature departed the snake genome sometime after snakes were created," and the ZRS papers "may have given us a peek into genetic events through which God may have implemented the Curse He spoke upon the serpent in Genesis 3:14, though we cannot be dogmatic about this."27 Cserhati in 2019: "A possible creationist explanation is that God cursed the snake right after the Fall in such a way as to turn off certain genetic elements in its genome, which coded for limbs. As to which species of snake this might have been, we don't know."28 Brian Thomas at ICR: "For all we know, He may have made some snake kinds with no legs and others with the genetic ability to turn leg production on or off."29
This answers the parent article's open question. For the snake pelvis, the creationist literature does not answer the function claim and stop; it grants that snakes are descended from legged snakes, and that the spur and pelvic rods are what is left. What it denies is the step before: "Snakes and lizards are different created kinds of animals, not biological cousins" (Mitchell),27 "God specially created snakes—they did not evolve from other kinds of creatures" (Bell).12
The function replies sit beside this and answer a different claim. Answers in Genesis in 2008: "these so-called 'legs,' which evolutionists admit were too short in this specimen to be used for ambulation, may have been used in copulation. Perhaps no snakes ever 'walked' in any sense."30 Troy Lacey in 2019: creationists "recognize the design feature of an anchoring pelvis and possible stimulators for reproductive structures."31 Rana, for the old-earth Reasons to Believe: "Snakes use their hind limbs—called anal spurs—to grasp during mating. So, in that sense, these structures are functional and could be viewed as an archetypical design deployed by the Creator."32 All of this is true and none of it is in dispute; Lee and Caldwell and Palci and colleagues say the same. "Perhaps no snakes ever 'walked' in any sense" is the one claim here that the fossils contradict, since Najash had a sacrum and "robust, functional legs outside the ribcage."7
The position that fits, a created legged snake kind whose legs were switched off, has three costs.
First, the switch-off is one event at the base of most living snakes. The 17-base deletion is in all six snakes Kvon and colleagues sequenced, the boa, the python, the king cobra, a rattlesnake, a viper and the corn snake,4 and the three HOXD13-site deletions are shared by boas and pythons. All six are alethinophidians. No blind snake, thread snake or pipe snake enhancer was sequenced in either 2016 paper, so whether the deletion goes back to the root of all snakes is an inference, not a measurement. Leal and Cohn draw it from the bones, and mark it as predicted: "Absence of a tibia, fibula, foot, and toes in scolecophidians (e.g., blind snakes) and alethinophidians suggests ZRS hypofunctionalization in the common ancestor."2 Blind snakes and thread snakes keep pelvic rods and, in the thread snakes, a femur.11 On the sequenced snakes alone, the "serpentine created kind" that had legs has to take in the boas, pythons, vipers, cobras and colubrids, which together make up most of the some 4,000 living species, and the Curse has to fall on their one ancestor. Cserhati's "we don't know" which snake is honest, and Thomas's several kinds with legs on or off would need the same deletion to have happened in each.
Second, the cost sits in the deletion and in the advanced snakes, not in the 80 percent figure. The boa and python ZRS differs from a lizard's at about a fifth of its sites, but Kvon and colleagues found that it "shows a substitution rate similar to other vertebrate ZRS orthologs."4 On the boa and python branch, the enhancer has changed no faster than it has in other vertebrates. A creationist who holds snakes and lizards to be separate created kinds can call that difference created, as for any other gene, and it counts against that view no more than ordinary snake-lizard differences do. What stands out is narrower. The 17-base gap falls in a stretch conserved from fish to mammals and removes an ETS binding site,4 in every snake sequenced. And in the snakes that kept no limb bones at all, the ZRS "displays a substantially increased number of substitutions compared to other enhancers,"4 the pattern of a sequence that use no longer holds in check. For scale only: the functional core Kvon and colleagues swapped into mice is 1.3 kilobases, and 2,000 generations of mutation at a mammalian per-site rate of one in a hundred million would be expected to change about 0.03 of its sites. No model claims that a few thousand years produced the advanced snakes' extra changes, and none should. A young-earth reading must therefore hold that the extra changes were created too, and that the Curse, if it acted here, was at most the 17-base deletion. Mitchell's alternative, that snakes were "simply designed without those 17 base pairs in ZRS,"27 gives up the Curse as a mechanism and leaves the parent article's question: why a created limbless animal carries a limb enhancer that works in a mouse when 17 bases are restored.
Third, the fossils do not look like the re-expression model's prediction. Sarfati's mechanism for legged fossil snakes is a mutation that "turned some of this information back on, but incompletely," ruined by accumulated damage, so that "the only result is tiny deformed legs too small to walk on."26 The legs that have been examined are small but not deformed: Eupodophis's bone microanatomy is that of a terrestrial lizard, grown for a shorter time;19 Pachyrhachis has a normal tibia, fibula, astragalus and calcaneum;1 Haasiophis has a complete foot;18 Najash has a sacrum and legs its describers call functional. Small, well-built legs are what reduced growth gives, not what a damaged program gives.
Objections and responses
Creationist: "The spurs have a function, so they are not vestigial." The parent article answers this in general. Here the sources answer it themselves: the 1998 sentence that calls the claw a hind-limb element ends "used in courtship," and the 2020 review reads the configuration as functional. Function was never the claim.
Creationist: "Losing legs is devolution, not evolution." Agreed, and the parent article grants it. The vestige argument is about ancestry, not about gaining information. The question is what the legged ancestor was.
Creationist: "Snakes and lizards are different kinds; a snake leg is not a lizard leg." Then the created snake carried a lizard's three pelvic bones, a lizard's femur, tibia, fibula, astragalus and calcaneum, a lizard's limb-bud program, and a limb enhancer four-fifths the same as a lizard's. Common design can be said of any of these, but it explains why the enhancer is present, not why it is broken in the same 17 bases in every snake sequenced.
Creationist: "Genesis 3:14 explains the loss." The text says the serpent will go on its belly. Answers in Genesis itself calls legs a "point of theological speculation." Read as a mechanism, the Curse has to act once on the ancestor of every sequenced snake family, has to leave the rest of the boa and python enhancer changing at its ordinary rate, and has to be followed by legged fossils that Sarfati explains by re-expression and Thomas by created variation.
Skeptic: "Tetrapodophis shows a snake with four legs, so the series is complete." It probably shows a dolichosaur. Caldwell and colleagues re-examined the specimen and its counterpart and found "new evidence refuting it as a snake." The creationist writers who doubted it in 2015 were right, and no part of this article rests on it.
Skeptic: "The fossils give a smooth sequence from legs to none." They give stages, not a sequence. The Tethyan snakes with the fullest legs have skulls that Tchernov and colleagues place among advanced snakes, so their legs may be retained or regrown; Palci and colleagues count losses and regainings of pelvic bones; the oldest snakes preserve no limb material. The spurs are the strongest point on that tree because they are in the living animal.
Skeptic: "The enhancer's decay proves deep time on its own." It shows that the divergence is not a few thousand years of mutation, which no young-earth writer claims. A created difference is a design claim, and this article prices it rather than refutes it.
Issues and unanswered questions
- Palci and colleagues' 2020 paper was read only in abstract; its family-by-family table of pelvic elements is paywalled.
- The classic behavioural papers, Carpenter and colleagues 1978, Gillingham and Chambers 1982 and Slip and Shine 1988, are on JSTOR only and were not opened, and Cohn and Tickle 1999 was read in abstract only; their content is here through Anzai 2023, Trivedi 2024, Senter 2014 and Leal and Cohn 2016.
- Senter and colleagues' reconstruction makes the spur rub ancestral for Boidae in the broad sense. Whether pipe snakes or Anilius use their spurs, which would push the behaviour to the base of the alethinophidians, was not found in a primary.
- Kvon and colleagues and Leal and Cohn sequenced the ZRS only in boas, pythons and advanced snakes. Whether blind snakes, thread snakes and pipe snakes carry the 17-base deletion was not found in any primary; a ZRS sequence from a scolecophidian would test whether the deletion reaches the root of all snakes.
- No creationist source prices the shared 17-base deletion, the fast change of the advanced snakes' enhancer, or the well-formed legs of the fossils against the Curse model. Whether a baraminology study places all snakes in one created kind, which the shared deletion requires, was not checked.
- The relationship of the Tethyan legged snakes to the rest is an open mainstream dispute; its two answers are about 95 million years apart from a Flood year either way.
Verdict
The pelvic spurs of boas and pythons are working structures: a mobile claw on a femur, larger in males, used to court and to fight. They are also a hind limb and a hip with the walking parts gone: the femur has lost its shin and foot, the pelvis has lost its joint with the spine, the embryo builds the missing parts and dissolves them, and the enhancer that should sustain the leg is broken in the same 17 bases in every snake sequenced. Fossil snakes with a sacrum and working legs existed. The creationist literature, unusually, agrees that snakes came from legged snakes and lost them. What remains is whether the legged snake was made or descended, and on that the spurs bear through the enhancer they share with lizards and the leg bones their fossils share with lizards. That evidence favours descent; the honest cost of the alternative is a created snake built with a lizard's limb program and a limb enhancer missing the same 17 bases across boas, pythons and advanced snakes.
Sources
- 1.
Lee, Michael S. Y., and Michael W. Caldwell. “Anatomy and relationships of Pachyrhachis problematicus, a primitive snake with hindlimbs.” Philosophical Transactions of the Royal Society B 353:1521–1552. 1998. A detailed description of a fossil snake from the Cretaceous of Israel, Pachyrhachis problematicus, which still had hind limbs and a loosely united pelvis. The authors argue that it is the most primitive known snake and the sister group of all other snakes, and they list the features that set living snakes apart. The article uses their description of living snakes as its baseline and quotes their remark that the homology of the claw, meaning which bone it corresponds to, is uncertain.
- 1a.
Page 1550: Character D11 in the authors' list of characters. “Modern snakes have lost all hindlimb elements except for the femur. Some alethinophidians (pythons and boids) have a single claw-like element of uncertain homology distal to the femur, used in courtship. Pachyrhachis is more primitive than modern snakes in retaining a normal tibia, fibula, astragalus and calcaneum.” ↩ ↩ ↩ ↩ ↩ ↩ ↩
- 1b.
Page 1550: Character D8 in the same list. “In modern snakes, the sacral contact is lost and the pelvic rudiments, when present, lie within the ribcage.” ↩
- 1c.
Page 1542: “The three elements of the pelvic girdle are not fused or sutured together. Rather, they appear to have abutted each other, forming a very weak girdle.” ↩
- 1d.
Page 1542: “the expanded dorsal end closely overlies the distal end of the sacral rib, indicating that the ilium was external to the ribcage and suggesting the presence of a sacral contact.” ↩
- 1a.
- 2.
Leal, Francisca, and Martin J. Cohn. “Loss and re-emergence of legs in snakes by modular evolution of Sonic hedgehog and HOXD enhancers.” Current Biology 26(21):2966–2973. 2016. A laboratory study that compared python embryos with anole lizard embryos and followed the genes and cartilage that begin to form a leg. The authors found that python leg development stops early because the switch (enhancer) that drives Sonic hedgehog in the limb has lost activity, and that some leg cartilage forms and then dissolves. The article uses it to show that a python embryo starts a hind leg and then stops, and that three DNA deletions shared by pythons and boas weaken that switch.
- 2a.
Section Results and Discussion, first paragraph: “Python embryos initiate formation of hindlimb buds, but leg development is not sustained, resulting in formation of a rudimentary femur and terminal claw.” ↩ ↩
- 2b.
Section Results and Discussion: “SHH mRNA was detected in a small domain of cells at the posterior margin of the hindlimb buds, but this ZPA-like expression pattern is transient, disappearing within 24 hr of oviposition”. ↩
- 2c.
Section Results and Discussion: “A morphological AER is evident in python hindlimb buds at stage 1”. “Shortly after termination of SHH, the FGF8 domain begins to degrade”. ↩
- 2d.
Under the heading Conservation of the Distal HOXD Regulatory Landscape and Autopodial Development in Pythons. “Comparison of the SOX9 domains in python in anole hindlimbs suggests that the python condensations could be anlagen of the femur, tibia and fibula, and digital plate”. "python in anole" is the published wording. “they are transitory structures, as distal hindlimb elements are not found in mature pythons”. ↩ ↩ ↩
- 2e.
Section Degenerate Evolution of the Python ZRS Underlies Diminished Transcription of SHH: “a limb-specific enhancer located ∼1 Mb upstream of SHH, in intron 5 of the LMBR1 gene”. ↩
- 2f.
Section Degenerate Evolution of the Python ZRS Underlies Diminished Transcription of SHH: “python ZRS mutations A, B, and C each occurred in HOXD13 binding sites”. ↩
- 2h.
Figure 4G: Point 4 of the legend to Figure 4G. “Re-emergence of the tibia and fibula (zeugopod) and digits (autopod) in Tethyan snakes may have resulted from persistence of embryonic skeletal condensations into adulthood.” ↩
- 2i.
Figure 4G: Point 3 of the legend to Figure 4G. The same legend draws known ZRS status with solid outlines and predicted status with broken outlines and gray branches. “Absence of a tibia, fibula, foot, and toes in scolecophidians (e.g., blind snakes) and alethinophidians suggests ZRS hypofunctionalization in the common ancestor.” ↩
- 2a.
- 3.
Palci, Alessandro et al. “The homologies and evolutionary reduction of the pelvis and hindlimbs in snakes, with the first report of ossified pelvic vestiges in an anomalepidid (Liotyphlops beui).” Zoological Journal of the Linnean Society 188(2):630–652. 2020. A review of the pelvis and hind-limb bones of living and fossil snakes, traced from the oldest known snakes to living forms, which also reports ossified pelvic bones in Liotyphlops beui, a snake of the family Anomalepididae. The authors reconstruct the common ancestor of all living snakes as having all three pelvic bones and rudimentary hind limbs with femoral spurs. The article uses it for that ancestor and for the authors' view that the spur, femur and pubis are probably linked to mating and courtship.
- 3a.
Section Abstract: “the most recent common ancestor of all living snakes (Serpentes) most probably retained all three pelvic elements and rudimentary hindlimbs (femoral spurs). Subsequently, there have been multiple losses of ossified pelvic and hindlimb elements and regaining of ossified pelvic elements.” “The single remaining rod-like element in some scolecophidians is the ischium, whereas the single remaining rod-like element in many basal alethinophidians is the pubis. Notably, many basal alethinophidians share a distinctive configuration of cloacal spur (claw), femur and a sizeable pubis, which is likely to be related functionally to the widespread use of the hindlimbs in mating and courtship, rather than the usual representation of the bones as non-functional vestiges.” ↩ ↩ ↩ ↩ ↩ ↩
- 3a.
- 4.
Kvon, Evgeny Z. et al. “Progressive loss of function in a limb enhancer during snake evolution.” Cell 167(3):633–642. 2016. A comparison of the DNA of six snake species with that of limbed animals and fish, followed by mouse experiments on a limb switch (enhancer) called the ZRS. The snake versions of the switch worked less well in mice, the cobra version truncated the mouse limbs, and a modified python version had normal limbs. The tests were in mice, not snakes, and the article uses the study to show that the switch lost its function in stages.
- 4a.
Section Results: “from basal snakes (boa and python) that retained a vestigial pelvic girdle and rudimentary hindlimbs, to advanced snakes (viper, rattlesnake, king cobra, and corn snake) that completely lost all skeletal limb structures and represent the majority (>85%) of all extant snake species”. ↩ ↩
- 4b.
Section Results: “we examined the draft genomes of six snake species including the Burmese python (Python molurus bivittatus) (Castoe et al., 2013), boa constrictor (Boa constrictor constrictor), king cobra (Ophiophagus hannah) (Vonk et al., 2013), speckled rattlesnake (Crotalus mitchellii pyrrhus), viper (Vipera berus berus), and corn snake (Pantherophis guttatus)”. The full snake sample. All six are alethinophidians; no blind snake, thread snake or pipe snake is in it. ↩ ↩
- 4c.
Section Results: “while the ZRS enhancer of basal snakes shares ~80% nucleotide identity with the orthologous region from limbed lizards and shows a substitution rate similar to other vertebrate ZRS orthologs, the ZRS of advanced snakes displays a substantially increased number of substitutions compared to other enhancers”. ↩ ↩ ↩
- 4d.
Section Snake Enhancer Knock-In Causes Severe Limb Truncation in Mice: “replacing the mouse ZRS with the orthologous cobra sequence resulted in a complete loss of Shh expression and a truncated limb phenotype, affecting both the fore- and hindlimbs, that is indistinguishable from the phenotype caused by deletion of the mouse enhancer”. ↩
- 4e.
Section Snake Enhancer Knock-In Causes Severe Limb Truncation in Mice: “The less diverged python ZRS resulted in a similar but a slightly milder phenotype.” ↩
- 4f.
The passage on the snake-specific deletion. “this deletion is the only sequence that is deleted in all snakes but present in all examined limbed vertebrates and fish”. ↩ ↩ ↩
- 4g.
“the resulting knock-in mice with the modified python allele had normal limbs”. ↩ ↩ ↩
- 4h.
The passage on ETS binding sites. “all examined snakes have lost the E0 and E1 ETS motifs”. ↩ ↩
- 4a.
- 5.
Anzai, Rafael Kenji et al. “Pelvic spur use during courtship and mating in the red-tailed boa Boa constrictor.” The Herpetological Bulletin 163:35–36. 2023. A short report of observations on captive red-tailed boas at two Brazilian breeding facilities, made in 2019 and 2021. The 2019 observations covered six males and four females, the 2021 observations five males and three females, and the report describes males rubbing their spurs along the females' sides before mating. The article uses it as the study that shows boas using their spurs in courtship, and every snake in it was captive, so it describes captive behaviour only.
- 5a.
Page 35: “pelvic spurs have musculature, innervation, and vascularisation and can move autonomously”. “The pelvic spurs of boids and pythonids are known to be used by males during combat, courtship, and mating”. ↩ ↩
- 5b.
Page 35: “These pelvic spurs are generally larger in males than in females”. ↩
- 5c.
Page 35: “the males began to rub their spurs gently on the side of the females' bodies, moving in a posterior to anterior direction”. “The spurs repeatedly moved from a resting to an erect state.” ↩
- 5d.
Page 35: “Spur use may be particularly relevant for smaller males because they may not have enough strength to move the body of large females.” ↩
- 5e.
Page 36: “Our data would appear to reinforce the critical role of pelvic spurs for successful courtship and mating in red-tailed boas.” ↩
- 5a.
- 6.
Hoefer, Sebastian et al. “Size matters: Sexual dimorphism in the pelvic spurs of the Bahamian Boa (Chilabothrus strigilatus strigilatus).” Herpetology Notes 14:201–203. 2021. A short field study of 32 Bahamian boas from Eleuthera, measuring pelvic spur length and body length and checking sex by probing or dissection. It found that males' spurs were about twice as long as females' (2.6 mm against 1.5 mm on average), more curved, and that spur length grew faster with body size in males. The article uses it as a simple, minimally invasive way to tell male and female boas apart, and the sample came from one island and was collected opportunistically, with 28 of the 32 boas found dead on the road.
- 6a.
Page 201, section Abstract: “We found that male Bahamian Boas possessed spurs nearly twice as long and notably more curved than females and that the proportion of the spurs to body size differed significantly between the sexes.” ↩
- 6b.
Page 202: “mean pelvic spur length in males being 2.6 mm (n = 16; range: 0.5 – 5.0 mm) compared to the 1.5 mm in females (n = 16; range: 0.5 – 3.0 mm)”. “the proportion of spur length relative to SVL was 0.3% in males and 0.1% in females”. ↩
- 6a.
- 7.
Apesteguía, Sebastián, and Hussam Zaher. “A Cretaceous terrestrial snake with robust hindlimbs and a sacrum.” Nature 440:1037–1040. 2006. A research paper in Nature describing a fossil snake from the Upper Cretaceous of Patagonia with a sacrum (the part of the spine that joins the hip), a pelvic girdle, and legs outside the ribcage. The authors' family-tree analysis makes it the most primitive snake known, and they argue that snakes came from land animals rather than sea animals. The article uses it as the one legged snake whose hip works.
- 7a.
Section Abstract: “Here we report a fossil snake with a sacrum supporting a pelvic girdle and robust, functional legs outside the ribcage.” “the new fossil is the most primitive (basal) snake known”. ↩ ↩ ↩
- 7b.
Section Abstract: “other known fossil snakes with developed hindlimbs, the marine Haasiophis, Pachyrhachis and Eupodophis, lack a sacral region”. ↩
- 7c.
Section Abstract: “It has commonly been thought that snakes underwent progressive loss of their limbs by gradual diminution of their use. However, recent developmental and palaeontological discoveries suggest a more complex scenario of limb reduction, still poorly documented in the fossil record.” ↩
- 7a.
- 8.
Caldwell, Michael W. et al. “Tetrapodophis amplectus is not a snake: re-assessment of the osteology, phylogeny and functional morphology of an Early Cretaceous dolichosaurid lizard.” Journal of Systematic Palaeontology 19(13):893–952. 2021. A research paper by Michael W. Caldwell and colleagues that re-examined the fossil first-hand, including its undescribed counterpart slab, and ran family-tree analyses on it. The authors conclude that it is a dolichosaur, a group of lizards, and that it shows adaptations to water rather than to snake life. The article uses it to show that the four-legged snake is probably a lizard.
- 9.
Answers in Genesis. “The Snake Can Walk?” News to Note. Sep 19, 2009. A short news item from Answers in Genesis, a creationist ministry, on a snake found in China with a leg, complete with claws. It says creationists have generally understood Genesis 3:14 to mean that the original serpent kind had legs, and that today's leglessness is a loss of that information. The article cites it as the creationist concession that snake leglessness is a reduction from a legged original.
- 9a.
“creationists have generally understood this to mean that the original serpentine created kind had legs. Even if God 'muted' the genetic information for legs in snakes (so that none could truly walk), a mutation could cause some of the latent genetic information to be expressed in the form of an errant limb.” “The leglessness of snakes today represents a corruption, loss, or deactivation of that information”. ↩ ↩
- 9a.
- 10.
Darwin, Charles. On the Origin of Species. John Murray. First edition. 1859. The first edition of Darwin's 1859 book. The article uses it because Darwin listed the snake's pelvis and hind limbs among his rudimentary organs in that edition.
- 10a.
Page 450: “there are rudiments of the pelvis and hind limbs.” ↩
- 10a.
- 11.
Romer, Alfred Sherwood. Osteology of the Reptiles. University of Chicago Press. 1956 (edition of 1976). A book on the bones of reptiles, first published in 1956. The article uses it to say which snakes still keep a thigh bone.
- 12.
Bell, Philip. “Of snakes, lizards and mosasaurs—evolutionists puzzle over snake origins.” Creation 31(3):15–17. Jun 2009. A magazine article by Philip Bell for Creation Ministries International, a creationist ministry. It reviews the debate over whether snakes came from lizards or from mosasaurs, and argues that God created snakes as their own kind. The article cites it as the creationist version of the pelvis facts reported by Kvon and colleagues: some living snakes keep a small pelvis and tiny hind limbs.
- 12a.
“some types of snake (e.g. wormsnakes, pipesnakes, boas and pythons) do have a small pelvic girdle and tiny hind limbs. However, the majority of today's snakes (1,600 species of the colubrid family) lack a pelvis.” ↩
- 12b.
“it is possible that some snakes had legs to begin with”. ↩
- 12c.
“God specially created snakes—they did not evolve from other kinds of creatures.” ↩
- 12a.
- 13.
Trivedi, Krunal et al. “Insights into the use of pelvic spur in mating behavior of Indian Rock Python.” Zoo's Print 39(8):11–13. Aug 21, 2024. A short report, number 258 in the Reptile Rap series of Zoo's Print, on one observed mating. The authors watched one male Indian rock python, with three larger females, at a zoo in Gujarat on 2 February 2024, and saw him use his spurs to position a female. The article uses it because the male pressed harder with his spurs than the gently rubbing boas described by Anzai and colleagues.
- 13a.
Pages 11–13: “not with gentle rubbing but through more forceful poking actions”. Page 13: “This behaviour, reminiscent of finger-walking but executed with the spurs, serves to provoke a reaction from the female, likely due to discomfort or pain caused by the spurs' sharpness.” ↩
- 13b.
Page 13: “it is plausible that this behaviour has evolved as a male strategy to navigate the challenges of mating with larger females”. ↩
- 13a.
- 14.
Senter, Phil et al. “Phylogeny of courtship and male-male combat behavior in snakes.” PLoS ONE 9(9):e107528. 2014. A study that mapped 33 courtship and combat behaviours from 76 species in five families of snakes onto a family tree, and used the fossil record to date when each behaviour appeared. The sample is under 4 percent of the species in its clade, and the authors say many gaps remain. The article uses it for its two spur behaviours: spur rubbing in courtship, which is old in Boidae, the family of boas and pythons, and the spur poke in combat, which appears in only a few genera.
- 14a.
Page 7: BC is the paper's short name for a behavioural component. “This BC is widespread enough in Boidae to infer that it is ancestral for the family.” “Ancestrally, courtship in Boidae included the spur rub by the Paleocene, the date of the earliest known crown-group boid”. ↩
- 14b.
Page 7: Under the heading Spur poke. “This BC is recorded only in the sister genera Eunectes and Epicrates”. ↩
- 14a.
- 15.
Cohn, Martin J., and Cheryll Tickle. “Developmental basis of limblessness and axial patterning in snakes.” Nature 399:474–479. 1999. A laboratory study of python embryos that examined the genes that lay out the body and the signals that grow the limbs. The authors found that python hindlimb buds start to form but the signals that would grow them are not switched on, and that adding a growth factor or chick tissue restores that signalling. The article cites it as the study that opened the question of why python legs stop.
- 15a.
Section Abstract: “Hindlimb buds are initiated, but apical-ridge and polarizing-region signalling pathways that are normally required for limb development are not activated. Leg bud outgrowth and signalling by Sonic hedgehog in pythons can be rescued by application of fibroblast growth factor or by recombination with chick apical ridge.” ↩
- 15a.
- 16.
Infante, Carlos R. et al. “Shared enhancer activity in the limbs and phallus and functional divergence of a limb-genital cis-regulatory element in snakes.” Developmental Cell 35(1):107–119. 2015. A comparison of DNA switches (enhancers) from mice, lizards and snakes, asking which switches that drive limb genes also work in the genitals. The authors tested one switch, HLEB, from the Tbx4 gene, which works in lizard limbs and genitals, and found that in snakes it has lost its limb function but keeps activity in parts of the genitals. The article uses it to show that shared switches do not reach the ZRS, which works only in limbs.
- 16a.
Section Results: “Despite the ancient limb reduction that occurred in the snake lineage, we found that snake genomes retain many sequences orthologous to mammalian limb enhancers.” ↩
- 16b.
Section Discussion: “Our analyses of HLEB orthologs suggest that this element has functionally diverged along such a path with snake HLEBs losing hindlimb function but retaining activity within subdomains of the external genitalia.” ↩
- 16a.
- 17.
Caldwell, Michael W., and Michael S. Y. Lee. “A snake with legs from the marine Cretaceous of the Middle East.” Nature 386:705–709. 1997. A research paper in Nature on Pachyrhachis problematicus, a marine squamate from the Cretaceous of the Middle East that had been read as a lizard. The authors argue from its skeleton that it is a snake with a well developed pelvis and hind limbs, and that it is the sister group of all other snakes, the branch that split off before the rest. The article uses it for the reclassification of this fossil as a legged snake.
- 17a.
Section Abstract: “a primitive snake with a well developed pelvis and hindlimbs”. “Pachyrhachis is the sister-taxon of all other snakes.” “unlike other snakes, Pachyrhachis retains a well developed sacrum, pelvis and hindlimb (femur, tibia, fibula, tarsals)”. ↩
- 17a.
- 18.
Tchernov, Eitan et al. “A fossil snake with limbs.” Science 287(5460):2010–2012. 2000. A short research report in Science on a 95-million-year-old snake from the Middle East that has the most extreme hind limb development known in its group. The authors' family-tree analysis places it, and the older Pachyrhachis, among the macrostomatans, a group of relatively advanced snakes such as pythons, boas and colubroids. The article cites it for the leg bones and for the tree the authors draw, which puts legged snakes among advanced rather than primitive snakes.
- 18a.
Section Abstract: “A 95-million-year-old fossil snake from the Middle East documents the most extreme hindlimb development of any known member of that group, as it preserves the tibia, fibula, tarsals, metatarsals, and phalanges.” “shows both to be related to macrostomatans, a group that includes relatively advanced snakes such as pythons, boas, and colubroids to the exclusion of more primitive snakes such as blindsnakes and pipesnakes.” ↩ ↩
- 18a.
- 19.
Houssaye, Alexandra et al. “Three-dimensional pelvis and limb anatomy of the Cenomanian hind-limbed snake Eupodophis descouensi (Squamata, Ophidia) revealed by synchrotron-radiation computed laminography.” Journal of Vertebrate Paleontology 31(1):2–7. 2011. A research paper that studied the type specimen of Eupodophis descouensi, the specimen the species is named from, with a non-destructive X-ray scan that builds three-dimensional images from thin slices. The scan could see a second, hidden leg, and the authors compare the pelvis and legs with those of other legged snakes and of living lizards. The article uses it for the second leg and for the finding that the leg bones are built like those of lizards, and the study rests on this one specimen.
- 19a.
Section Abstract: “This experiment allowed a virtual exhumation of the second, hidden leg of the specimen.” “As in Haasiophis, four tarsals are observed in each limb, but there are no traces of metatarsals or phalanges.” “displays a microanatomical organization similar to that of extant terrestrial lizards. This suggests that limb regression in Eupodophis was not due to a qualitative alteration of growth but, more likely, to a local decrease in growth rate or shortening of growth duration.” ↩ ↩
- 19a.
- 20.
Garberoglio, Fernando F. et al. “New skulls and skeletons of the Cretaceous legged snake Najash, and the evolution of the modern snake body plan.” Science Advances 5(11):eaax5833. 2019. A research paper in Science Advances that used micro-CT, a detailed X-ray scan, on the first three-dimensionally preserved skulls of Najash, the legged snake from Patagonia. The team reports eight new skulls and three articulated skeletons, and a new family-tree analysis that places the legged snakes at the base of the snake tree. The article uses it for the claim that early snakes with hind limbs were a varied group, not a passing stage.
- 21.
Martill, David M. et al. “A four-legged snake from the Early Cretaceous of Gondwana.” Science 349(6246):416–419. 2015. A research report in Science announcing a four-limbed fossil from the Early Cretaceous Crato Formation of Brazil. The authors describe a serpent-like body, hooked teeth and a skull they read as snake-like, and argue that snakes came from burrowing ancestors rather than marine ones. The article cites it as the original announcement of a snake with four limbs.
- 21a.
Section Abstract: “The structure of the limbs suggests that they were adapted for grasping, either to seize prey or as claspers during mating.” ↩
- 21a.
- 22.
Bell, Philip. “A four-legged fossil snake.” Creation Ministries International. 2015. A web article from Creation Ministries International, a creationist ministry. The page was updated in 2016 and 2021. The article cites it for the point that the snakes described in 2015 are known only from fragments.
- 22a.
“No trace of limbs or even limb girdles (pelvis or pectoral) were found in the four 'oldest' fossils.” ↩
- 22a.
- 23.
Genesis 3:14, King James Version. n.d. “upon thy belly shalt thou go, and dust shalt thou eat all the days of thy life”. The verse in which God curses the serpent. The article calls it the text under the whole question of snake legs. ↩
- 24.
Answers in Genesis. Garden of Eden. n.d. Retrieved Sep 26, 2026. An undated topic page from Answers in Genesis, a creationist ministry, on whether the serpent of Genesis 3 had legs. It says the Bible does not state that the serpent had legs, and that the question is one of theological speculation. The article cites it as the ministry's careful reading of the curse on the serpent.
- 24a.
Section What Did the Serpent Look Like?: “While the Bible does not explicitly state that the serpent had legs, the divine curse implies a transformation from a previous condition, leading some to infer it might have had legs or a different form of movement before the fall.” “whether or not the serpent had legs remains a point of theological speculation”. ↩
- 24a.
- 25.
Sarfati, Jonathan, and Don Batten. “Vestigial organs: What do they prove?” The Creation Answers Book. Creation Ministries International. 2006. An entry in a creationist reference book on vestigial organs, the structures that look like leftovers of earlier use. The article cites it for the creationist reading that Genesis 3:14 suggests snakes may once have had legs.
- 25a.
Section Legless lizards: “Genesis 3:14 suggests that snakes maybe once had legs.” ↩
- 25a.
- 26.
Sarfati, Jonathan. “Another leggy snake? What should creationists think?” Creation Ministries International. Apr 16, 2008. A creationist article by Jonathan Sarfati on Eupodophis, a fossil snake with two short hind legs that was discovered in 2000. Sarfati calls the researchers' claim to have found a femur, tibia and fibula reasonable, but argues the legs are too small to walk on. The article cites it for the creationist position that God may have switched off the genetic information for legs. The page was updated on 29 April 2008.
- 26a.
“We don't know exactly what God did to the serpent, but one possibility is that He turned off the genetic information to make the legs, hips and other features necessary to walk.” ↩
- 26b.
“If this is so, a mutation might have turned some of this information back on, but incompletely.” “So the only result is tiny deformed legs too small to walk on.” ↩
- 26a.
- 27.
Mitchell, Elizabeth. “Four-Legged Snake Fails as a Transitional Form and Perhaps Even as a Snake.” Answers in Genesis, News to Know. Nov 22, 2016. A news commentary by Elizabeth Mitchell of Answers in Genesis, a creationist ministry. It argues that the four-legged fossil Tetrapodophis is not a transitional form, and that snake leglessness may trace to a regulatory gene that lacks a 17-base-pair piece of DNA, a loss of information rather than new information. The article cites it for the claim that snakes and lizards are different created kinds, and for the 17 missing base pairs.
- 27a.
“it is quite possible the 17 absent base pairs apparently responsible for the snake's signature slithery nature departed the snake genome sometime after snakes were created”. “may have given us a peek into genetic events through which God may have implemented the Curse He spoke upon the serpent in Genesis 3:14, though we cannot be dogmatic about this.” ↩
- 27b.
“Snakes and lizards are different created kinds of animals, not biological cousins.” ↩
- 27c.
“Or were snakes simply designed without those 17 base pairs in ZRS, leaving their Sonic hedgehog genes free to perform their other functions while leaving snakes forever legless by design.” ↩
- 27a.
- 28.
Cserhati, Matthew. “Snakes losing legs is not evolution!” Creation Ministries International. Dec 11, 2019. A creationist article by Matthew Cserhati on creation.com, arguing that snakes losing legs is not evolution. It proposes that God turned off genes for limbs at the Curse, and says the species that this happened to is not known. The article cites it with other creationist writers for the view that the leg loss was a reduction.
- 28a.
“A possible creationist explanation is that God cursed the snake right after the Fall in such a way as to turn off certain genetic elements in its genome, which coded for limbs. As to which species of snake this might have been, we don't know.” ↩
- 28a.
- 29.
Thomas, Brian. “Snakes with Legs?” Creation Science Update. Institute for Creation Research. Aug 10, 2015. A short update by Brian Thomas for the Institute for Creation Research, a creationist ministry. The page was updated on 2 November 2016. The article cites it for the view that God may have made some snake kinds legless and others able to switch leg production on or off.
- 29a.
“For all we know, He may have made some snake kinds with no legs and others with the genetic ability to turn leg production on or off.” ↩
- 29a.
- 30.
Answers in Genesis. “Fossilized Snake with Legs Found In Lebanon.” News to Note. Apr 12, 2008. A news item from Answers in Genesis, a creationist ministry, on a fossil snake from Lebanon whose two hind limbs an X-ray study confirmed. The item says the legs are about 2 cm long and have no toes. The article cites it for the creationist suggestion that the legs were used in mating, not for walking.
- 30a.
“these so-called 'legs,' which evolutionists admit were too short in this specimen to be used for ambulation, may have been used in copulation. Perhaps no snakes ever 'walked' in any sense.” ↩
- 30a.
- 31.
Lacey, Troy. “Snakes Appear to Live a Cursed Life.” Answers in Genesis. Apr 6, 2019. An Answers in Genesis article by Troy Lacey. The article cites it for the creationist reading that the snake's pelvis is a design feature, an anchor for reproductive structures.
- 31a.
“recognize the design feature of an anchoring pelvis and possible stimulators for reproductive structures”. ↩
- 31a.
- 32.
Rana, Fazale. “Genetics Discovery in Snakes Adds Legs to the Case for Creation.” Reasons to Believe. Nov 9, 2015. An article by Fazale Rana for Reasons to Believe, an old-earth creationist ministry. It reports a University of Georgia study in which enhancers that drive limb growth in lizards are also active in the snake genome, where they help form the phallus. Rana reads the shared sequences as common design rather than common descent. The article cites it for the claim that snake spurs are functional, used to grasp during mating.
- 32a.
From endnote 2. “Snakes use their hind limbs—called anal spurs—to grasp during mating. So, in that sense, these structures are functional and could be viewed as an archetypical design deployed by the Creator.” ↩
- 32a.