Pelvic spurs (also known as vestigial legs) are external protrusions found around the cloaca in certain superfamilies of snakes belonging to the greater infraorder Alethinophidia . [1] These spurs are made up of the remnants of the femur bone, which is then covered by a corneal spur, or claw-like structure. [1] This femur derives from ancestral hind limbs found in the most recent common ancestor of modern snakes and the other reptiles of the clade Toxicofera , many of which have fully functional front and hind limbs. [2] Due to the fact that the spurs derive from the ancestral state of functional legs, but are no longer functional for locomotion specifically, these structures meet the criteria for being considered vestigial. [3] Nonetheless, uses for the structures have been thoroughly documented. Species that have external spurs have corresponding muscles, neurological structures, and vascularization to allow for independent movement. [3] The spurs are more pronounced and visible in male specimens and have been observed in use during courtship behavior. [4] The spurs are specifically used in the clasping and stimulation of females by males during courtship and mating. In certain species, males will also use their spurs to engage in combat with one another. [3]
The fossil record of snakes is not expansive. [5] Nonetheless, multiple fossilized specimens document the progression of the development of leglessness within the suborder Serpentes. The species of extinct snake Najash rionegrina was first described in 2006, and has been proposed as the earliest branching taxa of the suborder Serpentes. [6] The fossils were found in the Patagonia region of Argentina, and were dated to the Upper Cretaceous period. Najash rionegrina exhibited a sacrum, pelvic girdle, and robust hind limb structures outside of the ribcage, all of which led researchers to the conclusion that these hind limbs were functional for locomotion. [6] The significance of this finding is great, as there were three other known species of legged snakes from this time period, Pachyrhachis problematicus, Haasiophis terrasanctus and Eupodophis descouensi, but all were predicted to have been marine species, and all of them lacked the sacrum region found in N. rionegrina. [7] [6] The paleontologists thus concluded via phylogenetic analysis that N. rionegrina is the most ancient taxa within Serpentes, and the three extinct species previously described were more closely related to modern-day snakes belonging to Alethinophidia . [6]
Recent analysis of numerous fossil records supported these findings and further demonstrated the reduction of pelvic and hind limb structures within these lineages. [5] [2] Further evidence for these structures being plesiomorphic of can be found in the pelvis of some living taxa such as Candoia carinata or Eunectes murinus. These taxa possess a triradiate pelvis, which can also be observed in the skeletons of modern lizards. [2]
The presence of pelvic spurs in extant species of Serpentes is limited. Most members of Scolecophidia do not possess spurs, nor do members of the most populous group within Alethinophidia , the Caenophidia . However, spurs are present among the basal clades of Alethinophidia, including in Booidea and Pythonoidea , [8] among Amerophidia , and among one member of Uropeltoidea , the Cylindrophiidae . These basal clades are sometimes referred to as primitive snakes, as they are considered to be the earliest diverging taxa of Alethinophidia, the clade which includes the majority of described living snake species. [9] The presence and use of spurs across Booidea and Pythonoidea is well documented - in these superfamilies, spurs can be observed as tools for courtship and competition between males, and are sexually dimorphic. [3]
While these are the most well-known taxa to possess spurs, evidence does exist for the presence of ossified vestigial structures in other taxa. A 2019 publication provided evidence for similarly ossified structures in the species Liotyphlops beui of the infraorder Scolecophidia. [2] Members of this sister group of Scolecophidia are poorly understood due to their cryptic nature and are typically small in size, fossorial, and worm-like. [8] This 2019 study is the first described occurrence of these structures within the family Anomalepididae , which is one of three families within Scolecophidia. [2]
Numerous studies have been conducted on the use of pelvic spurs by males in the superfamilies Booidea and Pythonoidea. The sexually dimorphic nature of the spurs was formally described by William H. Stickel and Luccille F. Stickel in 1946 in the genus Enygrus (more commonly known today as Candoia ). [10] The authors of this study noted that others in the field had made similar observations before, but had not formally researched the topic specifically. Spurs were found to be significantly larger in males, while in females spurs were much shorter, and were sometimes externally absent. [10]
Research has since found the use of pelvic spurs in dominance displays in numerous species. The use of spurs alongside biting was observed in displays between males in a captive group of Indian pythons (Python molurus), which subsequently formed a linear dominance hierarchy. The position in this hierarchy was directly correlated with a male’s number of successful instances of mating. [11] Another study found the use of spurs in madagascan boas ( Sanzinia madagascariensis ), an arboreal species. In this species, researchers did not observe biting. [12] In fact, the authors observed that the heads of combating males were frequently out of sight of one another. In its place, observed males would tightly grip on to each other using the posterior regions of their bodies, orienting their spurs to be perpendicular to their body while doing so. [12] The authors postulate that this form of spur-based combat is adaptive to the species’ arboreal habitat. [12] It is also worth noting that combat between males in species that lack spurs, such as species of Caenophidia, relies on a distinct repertoire of behaviors that differs from species possessing spurs. [13]
Similar to the described observations of sexual dimorphism prior to their publication, William H. Stickel and Luccille F. Stickel also noted that observations had been made by others of pelvic spur use by males on females during mating. [10] These observations have since been formally investigated and published in multiple species.
One such study found dynamic use of spurs by males during different phases of courtship and mating in Burmese pythons (Python molurus bivittatus), with anterior to posterior spur movements varying in speed of undulation by phase. [14] Furthermore, the males exhibited use of the spurs to better position or adjust the female’s cloaca for mating. The authors note that spur use for combat can also be observed in this species between males. [14] A separate study found similar results in diamond pythons (Morelia spilota), a species that does not exhibit any bouts of combat between males. [15] In this species, multiple males were observed mating with the same female simultaneously, and males did not engage in combat. Instead, it was observed that spurs were used by males to manipulate the tail of the female. [15] Corroborating these results in a New World species, a study in 2023 documented the use of spurs in mating by red-tailed boas, Boa constrictor . [3] It was observed that males repeatedly alternated the orientation of the spurs between horizontal and vertical positions, which the authors suggest stimulated muscle contractions in the female, which would allow for better alignment between the individuals for mating. [3]
Anacondas or water boas are a group of large boas of the genus Eunectes. They are a semiaquatic group of snakes found in tropical South America. Three to five extant and one extinct species are currently recognized, including one of the largest snakes in the world, E. murinus, the green anaconda.
Snakes are elongated, limbless reptiles of the suborder Serpentes. Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joints than their lizard ancestors, enabling them to swallow prey much larger than their heads. To accommodate their narrow bodies, snakes' paired organs appear one in front of the other instead of side by side, and most have only one functional lung. Some species retain a pelvic girdle with a pair of vestigial claws on either side of the cloaca. Lizards have independently evolved elongate bodies without limbs or with greatly reduced limbs at least twenty-five times via convergent evolution, leading to many lineages of legless lizards. These resemble snakes, but several common groups of legless lizards have eyelids and external ears, which snakes lack, although this rule is not universal.
The Boidae, commonly known as boas or boids, are a family of nonvenomous snakes primarily found in the Americas, as well as Africa, Europe, Asia, and some Pacific islands. Boas include some of the world's largest snakes, with the green anaconda of South America being the heaviest and second-longest snake known; in general, adults are medium to large in size, with females usually larger than the males. Six subfamilies comprising 15 genera and 54 species are currently recognized.
The ball python, also called the royal python, is a python species native to West and Central Africa, where it lives in grasslands, shrublands and open forests. This nonvenomous constrictor is the smallest of the African pythons, growing to a maximum length of 182 cm (72 in). The name "ball python" refers to its tendency to curl into a ball when stressed or frightened.
Ophidia is a group of squamate reptiles including modern snakes and reptiles more closely related to snakes than to other living groups of lizards.
Loxocemus bicolor, the sole member of the monotypic family Loxocemidae and commonly known as the Mexican python, Mexican burrowing python and Mexican burrowing snake, is a species of python-like snake found in Mexico and Central America. No subspecies are currently recognized. Analyses of DNA show that Loxocemus is most closely related to the true pythons and the sunbeam snakes.
Henophidia is a former superfamily of the suborder Serpentes (snakes) that contains boas, pythons and numerous other less-well-known snakes.
The green anaconda, also known as the giant anaconda, emerald anaconda, common anaconda, common water boa, or southern green anaconda, is a semi-aquatic boa species found in South America and the Caribbean island of Trinidad. It is the largest, heaviest, and second longest snake in the world, after the reticulated python. No subspecies are currently recognized. Like all boas, it is a non-venomous constrictor. The term "anaconda" often refers to this species, though the term could also apply to other members of the genus Eunectes. Fossils of the snake date back to the Late Pleistocene in the Gruta do Urso locality.
The Indian python is a large python species native to tropical and subtropical regions of the Indian subcontinent and Southeast Asia. It is also known by the common names black-tailed python, Indian rock python, and Asian rock python. Although smaller than its close relative the Burmese python, it is still among the largest snakes in the world. It is generally lighter colored than the Burmese python and reaches usually 3 m. Like all pythons, it is nonvenomous.
A hemipenis is one of a pair of intromittent organs of male squamates. Hemipenes are usually held inverted within the body, and are everted for reproduction via erectile tissue, much like that in the human penis. They come in a variety of shapes, depending on species, with ornamentation such as spikes.
The blue-lipped sea krait, also known as the blue-banded sea krait or common sea krait, is a species of venomous sea snake in the subfamily Laticaudinae of the family Elapidae. It is found in the Indian and Western Pacific Oceans.
Pachyrhachis is an extinct genus of snake with well developed hind legs known from fossils discovered in Ein Yabrud, near Ramallah, in the central West Bank. It is a relatively small snake, measuring more than 1.5 metres (4.9 ft) long at maximum. Pachyrhachis appears to have been an ancient marine snake; the fossils occur in a marine limestone deposit, and the thickened bone of the ribs and vertebrae would have functioned as ballast to decrease the buoyancy of the animal, allowing it to dive beneath the ancient Cretaceous seas that it once inhabited.
Python is a genus of constricting snakes in the Pythonidae family native to the tropics and subtropics of the Eastern Hemisphere.
Madtsoiidae is an extinct family of mostly Gondwanan snakes with a fossil record extending from early Cenomanian to late Pleistocene strata located in South America, Africa, India, Australia and Southern Europe. Madtsoiidae include very primitive snakes, which like extant boas and pythons would likely dispatch their prey by constriction. Genera include some of the longest snakes known such as Vasuki, measuring at least 11–15 metres (36–49 ft) long, and the Australian Wonambi and Yurlunggur. As a grouping of basal forms the composition and even the validity of Madtsoiidae is in a state of flux as new pertinent finds are described, with more recent evidence suggesting that it is paraphyletic as previously defined.
The Alethinophidia are an infraorder of snakes that includes all snakes other than blind snakes and thread snakes. Snakes have long been grouped into families within Alethinophidia based on their morphology, especially that of their teeth. More modern phylogenetic hypotheses using genetic data support the recognition of 19 extant families, although the taxonomy of alethinophidian snakes has long been debated, and ultimately the decision whether to assign a particular clade to a particular Linnaean rank is arbitrary.
Eupodophis is an extinct genus of snake from the Late Cretaceous period. It has two small hind legs and is considered a transitional form between Cretaceous lizards and limbless snakes. The feature, described as vestigial, was most likely useless to Eupodophis. The type species Eupodophis descouensi was named in 2000 and resides now in the paleontology section of the Mim Museum in Beirut, Lebanon. The specific name is dedicated to the French naturalist Didier Descouens.
The boa constrictor, also known as the common boa, is a species of large, non-venomous, heavy-bodied snake that is frequently kept and bred in captivity. The boa constrictor is a member of the family Boidae. The species is native to tropical South America. A staple of private collections and public displays, its color pattern is highly variable yet distinctive. Four subspecies are recognized.
A spur is an outgrowth of bone covered in a sheath of horn found in various anatomical locations in some animals. Unlike claws or nails, which grow from the tip of the toes, spurs form from other parts of the foot, usually in connection with joints where the toes meet the foot or the foot meets the long bones. Spurs are most commonly found on the hindfeet, though some birds possess spurs at the leading edge of the wings.
Malayopython is a genus of constricting snakes in the family Pythonidae. The genus is native to India and Southeast Asia. It contains two species, both of which were previously classified within the genus Python. However, multiple studies recovered these species as distinct. Known as the "reticulatus clade", it was eventually found to be a sister lineage to a lineage giving rise to the Indo-Australian pythons rather than the genus Python.
The Amerophidia, also known as amerophidian snakes, are a clade of snakes that contains two families: Aniliidae and the boa-like Tropidophiidae.