sign in sign up
<<
Home , Colubridae, Colubrinae, Dolichophis, Dolichophis jugularis, Lampropeltini, Little whip snake

Herpetology Notes, volume 14: 735-744 (2021) (published online on 22 April 2021)

Male-male combat in the large whip , jugularis (Serpentes: )

Mohammad A. Abu Baker1, Mohammad Al-Saraireh2, Zuhair S. Amr3,, and Philip J. Senter4,*

Abstract. The male-male combat dance is prevalent among but is previously unreported in . Here, we present data from video footage of 13 instances of the male-male combat dance in D. jugularis. The large number of observations in this sample enables a more detailed study of the parameters of the combat dance in this than the small number of recorded observations in other species has allowed. In this species, the combatants coil around each other and lift their heads high, and each tries to push the other’s head downward. The phylogenetic presence of these behaviours among snakes indicates that coiling, high head-lifting, and downward pushing are ancestral behaviours for male-male combat in macrostomatan snakes that are retained in D. jugularis. The videos used in this study were collected from social media and demonstrate that social media can be useful in collecting data for scientific studies.

Keywords. ; Serpentes; Colubridae; Dolichophis jugularis; combat; behaviour; social media

Introduction distribution of these behaviours shows that male-male combat with “topping” and downward pushing may be The male-male combat ritual is a common behaviour in an ancestral trait for the snake clade Macrostomata, with snakes that involves a sequence of interactions between some clades adding more complex behaviours such as two males in the form of a contest for superiority or neck biting and dorsal body bowing (“body bridging” of dominance. It includes body contact during which Senter et al. (2014)), while in other snake clades (e.g., the males exert pressure by entwining, twisting, and and Natricidae), no male-male combat “topping” (in which each snake attempts to achieve the dance has been reported. Male-male combat with biting more dorsal position) to physically subdue or topple the opponent (Carpenter, 1977, 1984). It generally and pushing the opponent’s head down is also known occurs during the mating season (Capula and Luiselli, in legless of the family (Jablonski, 1997; Schuett, 1997). The ritual is phylogenetically 2018). widespread among snakes and has been reported in The Large Whip Snake, Dolichophis jugularis over 120 species of the families , , (Linnaeus, 1758), is a member of the family Colubridae. Colubridae, , Psammophiidae, and It is distributed through some of the eastern Greek (Shine, 1978, 1994; Senter et al., 2014), although it islands, southwestern and southern , , has not been thoroughly described in all such species. eastern , northern and western , , According to Senter et al. (2014), the phylogenetic , Palestine, northern Sinai, and western (Geniez, 2018), where it is often encountered in suburban and agricultural areas (Amr and Disi, 2011). It is characterised by its black, shiny body and is one 1 Department of Biology, The University of Jordan, Amman, of the largest snakes of the Levant, with a maximum Jordan. total length of up to 3 m. It is a diurnal, terrestrial snake 2 Oncology Department, Royal Medical Services, Amman, associated mostly with mild Mediterranean . It Jordan. is considered a common species in the mountains of 3 Department of Biology, Jordan University of Science & Technology, Irbid, Jordan. Jordan (Amr and Disi, 2011). 4 Department of Biological and Forensic Sciences, Fayetteville Despite its wide geographic distribution, no form of State University, 1200 Murchison Road, Fayetteville, North intrasexual competition has been previously recorded in Carolina 28301, United States. D. jugularis. In this paper, we present the first recorded * Corresponding author. E-mail: [email protected] observations of male-male combat in D. jugularis. © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. We furthermore present an unusually high number of 736 Mohammad A. Abu Baker ������ observations. For most snake species in which male- relative to each other, the sets of behaviours that were male combat has been described, it has been described employed during the combat dance, the in for very few pairs of males, usually only one pair (see which the dance occurred, whether the dance involved references in Shine, 1978, 1994; and Senter et al., 2014). simultaneous locomotion, the number of coils with which Here, we present observations of thirteen pairs of D. each combatant encircled the other, the approximate jugularis engaged in combat. The present observations percentage of body length that was anteriorly raised, the contribute to a fuller understanding of the behaviour approximate percentage of body length that was free of of this species and of the evolution of intrasexual coils anteriorly, the frequency of successful downward competition in snakes. pushes, and the frequency and duration of pauses in the combat dance. Materials and methods Phylogenetic distribution of male-male combat in snakes. To determine the contribution of these data to Videos. With the spread of social media tools, the reconstruction of the evolution of male-male combat “citizen science” has become an important source of in snakes, we mapped the behaviours noted here in D. data by curious amateurs and enthusiasts. Two social jugularis onto a cladogram of snake relationships (Fig. media groups on Facebook (www.facebook.com) were 1). The cladogram uses the phylogeny of Reynolds et founded and administered by M. Al-Saraireh: , al. (2014) for boas and pythons, and that of Zaher et al. Arthropods and Wildlife Amateurs of Jordan; and (2019) for endoglyptodontan snakes. The mapping of Reptiles, Arthropods and Wildlife Amateurs of Syria. behaviours onto the cladogram is based on Figure 2 of The two groups have over 25,000 subscribers apiece Senter et al. (2014), with corrections and with updates who follow and actively contribute their sightings and from additional references (Shaw, 1948, 1951; Saint field observations of reptiles from Jordan, Syria and Girons, 1949; Fleay 1951; Kroll, 1971; Carpenter et al., other countries in the region. It came to our attention that 1976; Martin, 1976; Carpenter and Gillingham, 1977; 13 videos were posted on the groups capturing male- Stickel et al., 1980; Murphy et al., 1981; Bennefield, male combat behaviour of D. jugularis. The videos were 1982; Nishimura et al., 1983; Clark et al. 1984; York, captured during June–July 2020 from several localities 1984; Secor, 1990; Almeida-Santos et al, 1998; Muniz- in Jordan, Syria, the Palestinian Territories, and Turkey, da-Silva and Almeida-Santos, 2013; Missassi et al., and have been posted onto YouTube (www.youtube. 2017; Goodyear and Gilbert, 2018; Valencia and com) (Table 1). Garzon-Tello, 2018; Guedes et al., 2019; Valencia et al., After the videos were collected, each was scrutinised 2020). to record such details as the sizes of the combatants

Table 1. Details on the videos included in this study. JRAWA (Jordanian Reptiles, Arthropods and Wild Life Amateurs Herping) and SRAWA (Syrian Reptiles, Arthropods and Wild Life Amateurs Herping) indicate the Facebook pages to which the videos wereTable 1.originally Details on uploaded.the videos included in this study. JRAWA (Jordanian Reptiles, Arthropods and Wild Life Amateurs Herping) and SRAWA (Syrian Reptiles, Arthropods and Wild Life Amateurs Herping) indicate the Facebook pages to which the videos were originally uploaded.

Video Photo-grapher Location Date of filming YouTube url number 1 Nedal Shriedeh (JRAWA) Tafilah, Jordan 9 July, 2020 https://www.youtube.com/watch?v=8jgi2EkzdYA 2 Mohammad Mousa (JRAWA) Iraq-al-Amir, Jordan 7 July, 2020 https://www.youtube.com/watch?v=WtPMB0khTIE 3 unknown (JRAWA) Palestinian Territories 5 July, 2020 https://www.youtube.com/watch?v=Rq-998bWjX8 4 Mousa Ghanam (JRAWA) Jarash, Jordan 19 June, 2020 https://www.youtube.com/watch?v=i1c7kerDA-w 5 unknown, in Hebron (JRAWA) Palestinian Territories 27 June, 2020 https://www.youtube.com/watch?v=mRz4NraQ9go 6 Sami Hasan (SRAWA) Baniyas, Syria 20 June, 2020 https://www.youtube.com/watch?v=3uObLjGOkdw 7 Sami Hasan (SRAWA) Baniyas, Syria 20 June, 2020 https://www.youtube.com/watch?v=0Iytf2xcLmo 8 Sami Hasan (SRAWA) Baniyas, Syria 3 July, 2020 https://www.youtube.com/watch?v=w3rQAxIhjrw 9 Mesam Houssen (SRAWA) As-Suwayda, Syria 25 June, 2020 https://www.youtube.com/watch?v=yDdSFYffH_Y 10 Ali Qutish (JRAWA) Tafilah, Jordan 20 June, 2020 https://www.youtube.com/watch?v=gpyXty-wMgk 11 unknown (shared to SRAWA from Turkey) Turkey unknown https://www.youtube.com/watch?v=h5PdjjiVkxg 12 unknown (JRAWA) Palestinian Territories unknown https://www.youtube.com/watch?v=q57LNs9tOjw 13 Wadi Ara (JRAWA) Palestinian Territories 2018 https://www.youtube.com/watch?v=gJc_H_DsoG8

Male-male combat in the large whip snake, Dolichophis jugularis 737

Figure 1. Phylogenetic distribution of recorded behaviours used in male-male combat in snakes, with black boxes representing the recorded presence of a behaviour. This figure is based on Fig. 1 of Senter et al. (2014), with corrections and with updates from additional references (see Materials and Methods section for details). On the cladogram, white circles represent families, grey circles represent subfamilies, and the grey square is a tribe. Taxonomic abbreviations: B = Boidae, C = Colubridae, c = , cr = Crotalinae, E = Elapidae, la = , P = Pythonidae, Ps = Psammophiidae, V = Viperidae, v = Viperinae, X = Xenopeltidae. For abbreviations of names of behaviours, see Table 2. Mohammad A. Abu Baker ������ 738

Table 2. Behaviours charted in Fig. 2. Abbreviations on the left are those that are used in Fig. 2. Table 2. Behaviours charted in Fig. 2. Abbreviations on the left are those that are used in Fig. 2.

B: Bite.—One snake bites another. Bow: Body bowing.—Formation of a high vertical arc with a section of the body. This behaviour was listed as “body bridging” by Senter et al., 2014, but because some authors use “body bridging” in reference to horizontal bending of the body (e.g. Green and Mason, 2000; Goodyear and Gilbert, 2018; Valencia et al., 2020), it is prudent to avoid the use of the term “body bridging” for body bowing. C: Coil.—One snake forms several coils around another, or both coil around each other. CR: Chin-rub.—One snake draws its chin along the skin of another. CS: Closed-mouth strike.—One snake strikes at another, with its mouth closed. DWP: Downward push.—Both snakes have their anterior ends elevated, and each attempts to push the other toward the ground. HR1: Head raise (type 1).—The snake raises its head, and only very little of its neck, off the substrate. HR2: Head raise (type 2).—The snake raises its head and much of its anterior body off the substrate. J1: Jerk (type 1).—The snake gives its head a sudden, staccato jerk. J2: Jerk (type 2).—The snake gives a large part of its body a sudden, staccato jerk. MG: Mouth gape.—The snake holds its mouth open. MN1: Mounting (type 1).—One snake presses down on another with its head. We called the following five behaviours mounting also, because each type seemed to be a modification of the previous type, with an increase in magnitude or an elaboration added in each successive type. MN2: Mounting (type 2).—One snake lies atop another, conforming to the same bodily bends. MN3: Mounting (type 3).—One snake lies atop another, with S-shaped bends draped over the other’s dorsum. MN4: Mounting (type 4).—One snake lies atop another, with lateral undulations that move anteriorly or posteriorly. SP: Spur poke.—One snake pokes another with its spurs. SR: Spur rub.—One snake rubs another with its spurs. Sw: Sway.—The snake sways its neck back and forth with its anterior portion elevated. TQ: Tail quiver.—The snake vibrates its tail. TW: Tail whip.—The snake rapidly whips its tail back and forth.

The behaviours mapped onto the cladogram are Results described in Table 2, which is based on the table of Videos. All videos commence after combat has descriptions in Senter et al. (2014). Here, the behaviour already begun (Fig. 2). In each case, the two snakes are called body bridging by Senter et al. (2014) is called body approximately equal in size. In each case, the combat bowing (Table 2). This behaviour involves the making dance involves both snakes coiling around each other of a vertical loop with the body. To avoid ambiguity, is and weaving their raised necks left and right and slightly prudent to avoid the use of the term body bridging for dorsally in attempts to “top” each other. In each case, this behaviour, because some authors use the term body the snakes face the same direction, so that they are never bridging in reference to horizontal bending of the body confronted belly-to-belly (facing each other). In most (e.g., Green and Mason, 2000; Goodyear and Gilbert, cases, the movements of the dance carry the snakes 2018; Valencia et al., 2020), whereas others use it in in some direction, but this locomotion is accidental, reference to vertical bending of the body (e.g., Murphy a consequence of the movements of the dance. In all et al., 1978). videos but 11 and 13 (in which the snakes decouple in Senter et al. (2014) mapped coiling onto the phylogeny an apparent attempt to flee the photographer), the two only when the coiling was decoupled from high head- snakes are continuously coupled through the video, raising (head raise type 2). However, that obscured the with their posterior ends coiled around each other. In full phylogenetic distribution of coiling. Here, coiling is all videos, the number of complete turns with which mapped onto the phylogeny in all species that employ it each snake is coiled around the other changes nearly with or without a simultaneous head raise. continuously, as does the percentage of body length that is anteriorly raised and the percentage of body length that Male-male combat in the large whip snake, Dolichophis jugularis 739

Figure 2. Stills (cropped and with image quality enhanced in most cases) from the 13 videos used in this study. Note the presence of coiling and high head-raising during the combat dance. 740 Mohammad A. Abu Baker ������ is free of coils anteriorly. Below, body length percentage turns with which each snake is coiled around the other estimates are based on measurements taken by PJS varies from one to four. The percentage of body length with a ruler held up to a computer screen. Admittedly, that is anteriorly raised varies from approximately 15% that is not a precise way to measure the length of a to approximately 30%. The percentage of body length moving in video footage, and therefore all such that is free of coils anteriorly varies from almost none measurements are approximate. In all videos, after a to approximately 40%. A successful downward push successful downward push (with success defined as a occurs at 0:11. There are no pauses in the dance. case in which one snake succeeds in pushing the other’s In video 4 (23 s long), the dance begins amid a head and neck onto the ground), both snakes re-raise collection of boulders that are surrounded by high, dry their necks immediately, and the dance continues. grass. The dance movements carry the snakes forward Below, the reported percentages of raised body length and into the grass. Obscuration of view by boulders do not include the moment in which a downward push and grass prevents a coil count. The percentage of body is successful. length that is anteriorly raised varies from approximately In video 1, the longest video (2 m 38 s), the dance 20% to approximately 50%. The percentage of body takes place in a wide, rocky gulley that lacks vegetation length that is free of coils anteriorly varies from almost but is surrounded by low vegetation. The movements none to approximately 20%. No pauses or successful of the dance carry the snakes slowly forward. During downward pushes occur. a stretch of 22 s, from 1:12 to 1:33, their forward In video 5 (1 m 40 s long), the dance takes place in a progress has brought them so far across the gulley that cobble-filled ditch next to the wall of a building. The vegetation obstructs the view of their heads, but then snakes are too far away to be clearly visible until 0:12, the photographer moves and their heads become visible when the zoom feature is employed, and the camera again. Through the video, the number of complete turns is moved so that the snakes are out of the frame from with which each snake is coiled around its opponent 1:27 to 1:36. The dance is stationary until the snakes varies from two to at least five. The percentage of turn completely around (180º) at 0:48, and the dance body length that is anteriorly raised varies from none then carries them forward. Obscuration of view by to approximately 30%. The percentage of body length cobbles prevents a coil count until 0:59, soon after that is free of coils anteriorly varies from none (i.e. the which the number of complete turns is five and then first coil begins immediately posterior to the head) to varies from three to five. The percentage of body length approximately 40%. Successful downward pushes occur that is anteriorly raised varies from approximately at 0:10, 0:18, 0:32, 0:54, and 2:13. The dance pauses at 15% to approximately 35%. The percentage of body 0:20 (for 2 s), at 0:38 (for 7 s), at 1:01 (for 7 s), at 1:22 length that is free of coils anteriorly varies from none (for 20 s), at 2:15 (for 2 s), and 2:31 (for at least 7 s, to approximately 30%. Successful downward pushes through the end of the video). occur at 1:00, 1:14, 1:16, and 1:24. The dance is paused In video 2 (17 s long), the dance takes place in a from 0:12 until 0:38 and pauses again at 1:36 through narrow, dirt-floored gulley that is surrounded by low the end of the video. vegetation. The dance movements carry the snakes a In video 6 (1 m 51 s long, although the video is little rearward. The tails of the snakes are obscured by taken from too far away to make out details until the vegetation, preventing a count of coils. The percentage zoom feature is employed at 0:31), the dance takes of body length that is anteriorly raised varies from none place on a patch of rocky ground at the base of a wall, to approximately 50%. The percentage of body length surrounded by grass on most sides, with low trees close that is free of coils anteriorly varies from approximately to the wall. The dance is stationary until 1:20, when 15% to approximately 40%. In video 2, there are no dance movements begin to carry the snakes sideways, pauses and no successful downward pushes, although to their left. The number of complete turns with which one snake loses its balance and falls at 0:01. each snake is coiled around the other varies from zero In video 3 (20 s long), the dance takes place on dirt to five. The percentage of body length that is anteriorly with a shallow covering of dry, dead vegetation, between raised varies from approximately 15% to approximately a patch of low vegetation next to a human-made wall 40%. The percentage of body length that is free of coils and a pile of what appear to be insulated cables and anteriorly varies from none to 100% (briefly, from 1:00 an overturned bucket. The dance movements carry the to 1:01). Successful downward pushes occur at 0:35 and snakes sideways, to their left. The number of complete 1:01. No pauses in the dance occur. Male-male combat in the large whip snake, Dolichophis jugularis 741

In video 7 (1 m 50 s long, with the snakes out of the is coiled around the other varies from one to four. The frame due to camera movement from 0:05 to 0:17, percentage of body length that is anteriorly raised varies 0:53 to 1:01), the dance takes place in an area of low from approximately 20% to approximately 30%. The vegetation with boulders and cobble amid the vegetation. percentage of body length that is free of coils anteriorly The dance movements carry the snakes sideways, to varies from none to approximately 40%. No successful their left. Obscuration of view by vegetation prevents downward pushes occur. The dance pauses at 0:14, and a coil count, approximation of anteriorly raised body at 0:22 the snake whose anterior body is beneath the length, and anteriorly coil-free body length. Successful other’s suddenly slithers swiftly to the snakes’ right and downward pushes occur at 0:26, 0:36, and 1:25. No dives under the flattened box, its tail disappearing from pauses in the dance occur. view at 0:24. Whether it is acknowledging defeat or In video 8 (14 s long), the dance takes place on a dirt fleeing the photographer is unclear. The snake that had road and is stationary. The number of complete turns been on top follows the other snake’s path to the edge with which each snake is coiled around the other varies of the cardboard box, pauses with its head and neck at from three to at least five. The percentage of body length the edge of the box at 0:26, tongue-flicks once at 0:31, that is anteriorly raised varies from approximately then slithers over the edge of the box at 0:32, apparently 20% to approximately 35%. The two snakes’ bodies following the path of the other snake. Whether it is are coiled even immediately behind the neck through pursuing the other snake or fleeing the photographer is much of the video. There are no pauses or successful unclear. downward pushes. In video 12 (1 m 37 s long), the dance takes place In video 9 (12 s long), the dance takes place on a beneath an olive tree on a patch of ground with short paved road and is stationary. The number of complete grass and dry, dead vegetation. The dance movements turns with which each snake is coiled around the other cause the snakes to make a left turn at 0:16, then the varies from two to five. The percentage of body length movements carry them forward. Obscuration of the that is anteriorly raised varies from nearly none to posterior ends of the snakes prevents a coil count through approximately 25%. The percentage of body length most of the video, but the number of complete turns that is free of coils anteriorly varies from none to with which each snake is coiled around the other is at approximately 15%. There are no pauses or successful least three at several points in the video. The percentage downward pushes. of body length that is anteriorly raised varies from less In video 10 (1 m 37 s long, although the snakes are than 10% to approximately 30%. The percentage of out of the frame through much of the video: 0:08 – 0:12, body length that is free of coils anteriorly varies from 0:49 – 0:51, 0:54 – 0:58, 1:26 – 1:37), the dance takes none to approximately 50%. Successful downward place in a field of high grass that is surrounded by trees. pushes occur at 0:16 and 0:46. There are no pauses in The dance movements carry the snakes forward. At the the dance. beginning of the video, the dance carries the snakes In video 13 (46 s long), the dance takes place in a dirt over part of a downed branch that projects horizontally footpath with vegetation on one side of it. The dance perhaps about 20 cm above the ground (Fig. 2). Low is stationary. The number of complete turns with which resolution, camera movement, and obscuration of view each snake is coiled around the other varies from three by grass prevent coil counts and estimations of anteriorly to four. The percentage of body length that is anteriorly raised and anteriorly uncoiled body length through most raised varies from approximately 10% to approximately of the video. But at 1:02 the number of complete turns 30%. The percentage of body length that is free of coils with which each snake is coiled around the other appears anteriorly varies from none to approximately 40%. A to be four or five, and approximately 15 – 20% of the successful downward push occurs at 0:30. The dance body is anteriorly raised from 1:01 to 1:08. A successful pauses at 0:17 (for 10 s). At 0:36 the snake whose neck downward push occurs at 0:20. No pauses in the dance is dorsal to the other’s appears to notice the forward occur when the snakes are in frame. progress of the photographer and suddenly turns rearward In video 11 (37 s long), the dance takes place on a and flees into the vegetation. The other snake pauses for flattened cardboard box amid other discarded cardboard 4 s, then also appears to notice the photographer and and plastic items, all next to a chain-link fence with flees in the same direction as its opponent. low vegetation at its base. The dance is stationary. In summary, the combat dance of D. jugularis involves The number of complete turns with which each snake mutual coiling, high head-raising, and downward 742 Mohammad A. Abu Baker ������ pushes. It takes place in a variety of habitats, including combatants and provides a more robust picture of the exposed ground and vegetation-filled areas. It is parameters of combat within the species. However, it sometimes stationary, and at other times its movements revealed that little variation exists in the combat dance produce accidental locomotion that carries the snakes of D. jugularis. On the other hand, our data suggest that forward, rearward, or sideways. The number of coils the onset of the breeding season, starting with male-male continuously changes during the dance and often combat, begins in June and extends through mid-July. reaches as many as five. The percentage of the body Comparison of specific aspects of the behaviours that is anteriorly raised during the dance continuously involved in the combat dance between D. jugularis and changes, frequently reaches approximately 30%, and other snake species (e.g. frequency per minute of pauses occasionally reaches approximately 50%. The coiling and successful downward pushes) is hampered by a lack is nearly continuous in the posterior half of the body of such details in previous reports. However, published and frequently continues up to the back of the head. pictures enable comparison of the magnitude of coiling During the dance, the anterior body (up to 40% of total between species. In our sample, the number of complete length) often briefly escapes the coils of the opponent. turns of each snake’s body during coiling frequently Downward pushes are frequently successful and occur reached five turns. Higher numbers of turns may have at a rate of zero to four per minute. Pauses in the dance occurred but could not be confirmed. In other species, occur at a rate of zero to three per minute. The durations pictorial evidence from drawings and photographs of the pauses range from two to twenty seconds and reveals that the maximum recorded number of complete often last between seven and ten seconds. turns during coiling can be as few as one in Crotalus Phylogenetic distribution of male-male combat in ruber (Shaw 1948); as few as two in Vipera ammodytes snakes. The addition of D. jugularis to the cladogram (Carpenter, 1977), Agkistrodon contortrix (Schuett and fills a gap in colubrid phylogeny and helps to confirm Gillingham, 1989), and Crotalus lepidus (Carpenter et the widespread occurrence of coiling and the downward al., 1976); as many as three in Vipera berus (Andrén, push in male-male combat in the Colubridae, and of the 1986), Protobothrops flavoviridis (Nishimura et al., high head raise (head raise type 2) in male-male combat 1983), Micrurus lemniscatus (Missassi et al., 2017), in the Colubridae outside the clade Lampropeltini. Austrelaps superbus (Shine and Allen, 1980), These behaviours are also widespread both in other bicarinatus (Almeida-Santos and Marques, 2002), and endoglyptodontan snakes and in the pythons and melanoleucus (Shaw, 1951); four in Micrurus boas. Their presence in the combat dance is therefore frontalis (Almeida-Santos et al., 1998), pullatus highly likely to be an ancestral behaviour in the clade (Muniz-da-Silva et al., 2013), Macrostomata. (Bogert and Roth, 1966), and spiloides Most of the other behaviours mapped onto the (Rigley, 1971); five in Epicrates assisi (Guedes et al., phylogeny have a known phylogenetic distribution that 2019), Pseudechis porphyriacus (Fleay, 1951), and is too limited or too erratic to determine evolutionary Rhinoplocephalus flagellum (Turner, 1992); and nine patterns with certainty. Exceptions include biting and in Micrurus mipartitus (Valencia et al., 2020). The body bowing, which are sufficiently prevalent in the frequent achievement of five turns in D. jugularis Lampropeltini to support the hypothesis that they are therefore appears to be typical of a general tendency ancestral behaviours for that clade. Another exception is among snakes to achieve three or more complete turns swaying, which is sufficiently prevalent in the Crotalinae during coiling. It is interesting that only one or two to support the hypothesis that it is an ancestral behaviour turns, if any, are recorded in published illustrations of for that clade. New World crotalines. This may be an artefact, or it may represent a real reduction in the tendency to coil around Discussion one’s opponent in the New World crotaline clade. For most snake species in which the combat dance has In addition to contributing to greater documentation and been described, it has been described for only one pair or a better understanding of the behaviour of D. jugularis, a small number of pairs of combatants. In this study, the this study contributes to a better understanding of the availability of a large number of videos of the combat evolution of male-male combat in snakes. The data dance enabled a more in-depth study than can usually reported here, in combination with previously reported be produced. Our study, therefore, allows comparison data and with an improvement upon the understanding of details of the combat dance among several pairs of of coiling with respect to the study of Senter et al. Male-male combat in the large whip snake, Dolichophis jugularis 743

(2014), support the hypothesis that the ancestral version 1976: 764–780. of the male-male combat dance in the snake clade Clark, D.L., Gillingham, J.C., Rebischke, A. (1984): Notes on Macrostomata included coiling, the downward push, the combat behavior of the California , Lampropetis getulus californiae, in captivity. British Journal of Herpetology and the high head raise (head raise type 2). 6: 380–382. This study also demonstrates how useful social Fleay, D. (1951): Savage battles between snakes. Walkabout 17(1 media can be in providing scientists with data. Indeed, May): 10–13. previous studies have also corroborated the role of Geniez, P. (2018): Snakes of Europe, North Africa and the Middle social media in documenting behavioural observations East. A Photographic Guide. Princeton, Princeton University that may provide the scientific community with useful Press. data on little known or secretive species. Thus, social Goodyear, J., Gilbert, E. (2018): First record of male-male combat in unicolor. Herpetological Bulletin 143: 45–56. media proved to be a reliable and faster source for field Green, M.J., Mason, R.T. (2000): Courtship, mating, and male observations relative to a classic literature review (e.g., combat of the , irregularis. Herpetologica Maritz and Maritz, 2020). 56: 166–175. Guedes, T., Guedes, A., Almeida Santos, S.M. (2019): Male-male Acknowledgments. Many thanks to the photographers (listed in fighting, dominance, and mating in Epicrates assisi (Boidae) in Table 1) of the 13 videos of D. jugularis in combat. Many thanks captivity. Phyllomedusa 18: 131–135. also to reviewer Daniel Jablonski, who provided constructive Jablonski, D. (2018): Male-male combat in Pseudopus apodus suggestions that improved this paper. (Reptilia: Anguidae). Russian Journal of Herpetology 25: 293– 298. References Kroll, J.C. (1971): Combat behavior in male Great Plains Ground Snakes ( episcopa episcopa). Texas Journal of Science Almeida-Santos, S.A., Marques, O.A.V. (2002): Male-male ritual 23: 300. combat in the colubrid snake Chironius bicarinatus from the Linnaeus, C. (1758): Systema Naturae per Regna Tria Naturae, Atlantic forest, southeastern Brazil. Amphibia-Reptilia 23: Secundum Classes, Ordines, Genera, Species, cum Characteribus, 528–533. Differentiis, Synonymis, Locis. Stockholm, Laurentius Salvius. Almeida-Santos, S.A., Schmidt de Aguiar, L.F., Lucchesi Maritz, R.A., Maritz, B. (2020): Sharing for science: high- Balestrin, R. (1998). Micrurus frontalis (coral snake). Combat. resolution trophic interactions revealed rapidly by social media. Herpetological Review 29: 242. PeerJ 8(e9485): 1–21. Amr, Z.S., Disi, A. (2011): Systematics, distribution and ecology of Martin, B.E. (1976): Notes on breeding behavior in a captive pair the snakes of Jordan. Vertebrate Zoology 61: 179–266. of Sonora Mountain (). Andrén, C. (1986): Courtship, mating and agonistic behaviour in a Bulletin of the Maryland Herpetological Society 12: 23–24. free-living population of adders, Vipera berus (L.). Amphibia- Missassi, A.F.R., Coeti, R.Z., Germano, V.J., Almeida-Santos, Reptilia 7: 353–383. S.M. (2017): Micrurus lemniscatus carvalhoi (Coralsnake). Bennefield, B.L. (1982): Combat and mating in the vine Reproduction / male-male combat. Herpetological Review 48: snake Thelotornis capensis mossambicana. Journal of the 214–215. Herpetological Association of Africa 28: 13–14. Muniz-da-Silva, D.F., Almeida-Santos, S.M. (2013): Male-male Bogert, C.M., Roth, V.D. (1966): Ritualistic combat of male gopher ritual combat in (Serpentes: Colubrinae). snakes, affinis (Reptilia, Colubridae). Herpetological Bulletin 126: 25–29. American Museum Novitates 2245: 1–27. Murphy, J.B., Tryon, B.W., Brecke, B.J. (1978): An inventory Capula, M., Luiselli, L. (1997): A tentative review of sexual of reproduction and social behavior in captive gray-banded behavior and alternative reproductive strategies of the Italian kingsnakes, alterna (Brown). colubrid snakes (Squamata: Serpentes: Colubridae). Herpetozoa Herpetologica 34: 84–93. 10: 107–119. Murphy, J.B., Lamoreaux, W.E., Barker, D.G. (1981): Miscellaneous Carpenter, C.C. (1977): Communication and display of Snakes. notes on the reproductive biology of reptiles. 4. Eight species American Zoologist 17: 217–223. of the family Boidae, genera Acrantophis, Aspidites, Candoia, Carpenter, C.C. (1984): Dominance in snakes. In: Vertebrate Liasis and Python. Transactions of the Kansas Academy of Ecology and Systematics. A Tribute to Henry S. Fitch. University Sciences 84: 39–49. of Kansas Museum of Natural History Special Publication 10, Nishimura, M., Otani, T., Nakamotu, E. (1983): Mating and combat p. 195–202. Seigel R.A., Hunt, LE., Knight, J.E., Malert, L., dance of the habu, Trimeresurus flavoviridis. Japanese Journal Zuschlag, N.L., Eds., Lawrence, University of Kansas. of Herpetology 10: 42–46. Carpenter, C.C., Gillingham, J.C. (1977): A combat ritual between Reynolds, R.G., Niemiller, M.L., Revell, L.J. (2014): Toward a two male speckled kingsnakes (Lampropeltis getulus holbrooki: Tree-of-Life for the boas and pythons: Multilocus species-level Colubridae: Serpentes) with indications of dominance. phylogeny with unprecedented taxon sampling. Molecular Southwestern Naturalist 22: 517–254. Phylogenetics and Evolution 71: 201–213. Carpenter, C.C., Gillingham, J.C., Murphy, J.B. (1976): The Rigley, L. (1971): “Combat dance” of the black , o. combat ritual of the rock rattlesnake (Crotalus lepidus). Copeia obsoleta. Journal of Herpetology 5: 65–66. 744 Mohammad A. Abu Baker ������

Saint Girons, H. (1949): Les moeurs nuptiales de la vipère aspic. Shine, R., Allen, S. (1980): Ritual combat in the Australian Terre et Vie 96: 103–113. copperhead, Austrelaps superbus (Serpentes, Elapidae). Schuett, G.W. (1997): Body size and agonistic experience affect Victorian Naturalist 97: 188–190. dominance and mating success in male copperheads. Animal Stickel, L.F., Stickel, W.H., Schmid, F.C. (1980): Ecology of a Behavior 54: 213–224. Maryland population of black rat snakes (Elaphe o. obsoleta). Schuett, G.W., Gillingham, J.C. (1989): Male-male agonistic Herpetologica 103: 1–14. behaviour of the copperhead, Agkistrodon contortrix. Amphibia- Turner, G. (1992): Courtship behaviour and male combat in the Reptilia 10: 243–266. little whip snake Rhinoplocephalus flagellum (Elapidae). Secor, S.M. (1990): Reproductive and combat behavior of the Herpetofauna 22: 14–21. Mexican Kingsnake, Lampropeltis mexicana. Journal of Valencia, J.H., Garzón-Tello, K., Cogălniceanu. (2020): Male- Herpetology 24: 217–221. male combat in the coralsnake Micrurus mipartitus decussatus Senter, P., Harris, S.M., Kent, D.L. (2014): Phylogeny of courtship (Duméril et al., 1854) (Squamata: Elapidae). Herpetology Notes and male-male combat behavior in snakes. PLoS ONE 9(9: 13: 329–332. e107528): 1–10. York, D.S. (1984): The combat ritual of the Malayan pit viper Shaw, C.E. (1948): The male combat “dance” of some crotalid (Calloselasma rhodostoma). Copeia 1984: 770–772. snakes. Herpetologica 4: 137–145. Zaher, H., Murphy, R.W., Arredondo, J.C., Graboski, R., Machado- Shaw, C.E. (1951): Male combat in American colubrid snakes Filho, P.R., Mahlow, K., et al. (2019): Large-scale molecular with remarks on combat in other colubrid and elapid snakes. phylogeny, morphology, divergence-time estimation, and the Herpetologica 7: 149–168. fossil record of advanced caenophidian snakes (Squamata: Shine, R. (1978): Sexual size dimorphism and male combat in Serpentes). PLoS ONE 14(5:e0216148): 1–82. snakes. Oecologia 33: 269–277. Shine, R. (1994): Sexual size dimorphism in snakes revisited. Copeia 1994: 326–346.

Accepted by Jiri Smid