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African (Common) Puff Adder - Bitis arietans
Topic Started: Aug 9 2012, 05:50 AM (3,440 Views)
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Shark Toothed Reptile
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African (Common) Puff Adder - Bitis arietans

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Scientific Classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Family: Viperidae
Subfamily: Viperinae
Genus: Bitis
Species: B. arietans

Bitis arietans is a venomous viper species found in savannah and grasslands from Morocco and western Arabia throughout Africa except for the Sahara and rain forest regions. It is responsible for causing the most fatalities in Africa owing to various factors, such as its wide distribution and frequent occurrence in highly populated regions. Two subspecies are currently recognized, including the nominate subspecies described here.

The species is commonly known as puff adder, African puff adder, or common puff adder.

The average size is about 1 m in length and very stout. Large specimens of 190 cm (75 in), weighing over 6.0 kg (13.2 lbs) and with a girth of 40 cm (16 in) have been reported. Specimens from Saudi Arabia are not as large, usually no more than 80 cm in length. Males are usually larger than females and have relatively longer tails. The head has a less than triangular shape with a blunt and rounded snout. Still, it is much wider than the neck. The rostral scale is small. The circumorbital ring consists of 10–16 scales. Across the top of the head, there are 7–11 interocular scales. 3–4 scales separate the suboculars and the supralabials. There are 12–17 supralabials and 13–17 sublabials. The first 3–4 sublabials contact the chin shields. Often, there are two fangs on each maxilla and both can be functional.

Midbody there are 29–41 rows of dorsal scales. These are strongly keeled except for the outermost rows. The ventral scale count is 123–147, the subcaudals 14–38. Females have no more than 24 subcaudals. The anal scale is single.
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The color pattern varies geographically. The head has two well-marked dark bands: one on the crown and the other between the eyes. On the sides of the head, there are two oblique dark bands or bars that run from the eye to the supralabials. Below, the head is yellowish white with scattered dark blotches. Iris color ranges from gold to silver-gray. Dorsally, the ground-color varies from straw yellow, to light brown, to orange or reddish brown. This is overlaid with a pattern of 18–22 backwardly-directed, dark brown to black bands that extend down the back and tail. Usually these bands are roughly chevron-shaped, but may be more U-shaped in some areas. They also form 2–6 light and dark cross-bands on the tail. Some populations are heavily flecked with brown and black, often obscuring other coloration, giving the animal a dusty-brown or blackish appearance. The belly is yellow or white, with a few scattered dark spots. Newborn young have golden head markings with pinkish to reddish ventral plates toward the lateral edges.
One unusual specimen, described by Branch and Farrell (1988), from Summer Pride, East London in South Africa, was striped. The pattern consisted of a narrow (1 scale wide) pale yellowish stripe that ran from the crown of the head to the tip of the tail.
Generally, though, these are relatively dull-looking snakes, except for male specimens from highland east Africa and Cape Province, South Africa, that usually have a striking yellow and black color pattern.

Geographic Range
This species is probably the most common and widespread snake in Africa. It is found in most of sub-Saharan Africa south to the Cape of Good Hope, including southern Morocco, Mauritania, Senegal, Mali, southern Algeria, Guinea, Sierra Leone, Ivory Coast, Ghana, Togo, Benin, Niger, Nigeria, Chad, Sudan, Cameroon, Central African Republic, northern, eastern and southern Democratic Republic of the Congo, Uganda, Kenya, Somalia, Rwanda, Burundi, Tanzania, Angola, Zambia, Malawi, Mozambique, Zimbabwe, Botswana, Namibia and South Africa. It also occurs on the Arabian peninsula, where it is found in southwestern Saudi Arabia and Yemen. The type locality given is "Promontorio bonae spei" (Cape of Good Hope, South Africa).

It is found in all habitats except true deserts, rain forests, and (tropical) alpine habitats. Most often associated with rocky grasslands.

It is not found in rainforest areas, such as along the coast of West Africa and in Central Africa (i.e., central DR Congo); it is also absent from the Mediterranean coastal region of North Africa. On the Arabian peninsula, it is found as far north as Ta'if. It has been reported to be found in the Dhofar region of southern Oman.

Normally a sluggish species, it relies on camouflage for protection. Locomotion is primarily rectilinear, using the broad ventral scales in a caterpillar fashion and aided by its own weight for traction. When agitated, it can resort to a typical serpentine movement and move with surprising speed. Although mainly terrestrial, these snakes are good swimmers and can also climb with ease; often they are found basking in low bushes. One specimen was found 4.6 m above the ground in a densely branched tree.

If disturbed, they will hiss loudly and continuously, adopting a tightly coiled defensive posture with the fore part of their body held in a taut "S" shape. At the same time, they may attempt to back away from the threat towards cover. They may strike suddenly and at a high speed, to the side as easily as forwards, before returning quickly to the defensive position, ready to strike again. During a strike, the force of the impact is so strong, and the long fangs penetrate so deeply, that prey items are often killed by the physical trauma alone. The fangs are apparently able to penetrate soft leather.

They can strike to a distance of about one third of their body length, but juveniles will launch their entire bodies forwards in the process. These snakes rarely grip their victims, instead releasing quickly to return to the striking position.
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Mostly nocturnal, they rarely forage actively, preferring instead to ambush prey as it happens by. Their prey includes mammals, birds, amphibians, and lizards

Females produce a pheromone to attract males, which engage in neck-wrestling combat dances. A female in Malindi was followed by seven males. They give birth to large numbers of offspring: litters of over 80 have been reported, while 50–60 is not unusual. Newborns are 12.5–17.5 cm in length. Very large specimens, particularly those from East Africa, give birth to the highest numbers of offspring. A Kenyan female in a Czech zoo gave birth to 156 young, the largest litter for any species of snake.

These snakes do well in captivity, but there are reports of gluttony. Kauffeld (1969) mentions that specimens can be maintained for years on only one mouse a week, but that when offered all they can eat, the result is often death, or at best wholesale regurgitation. They are bad-tempered snakes and some specimens never settle down in captivity, always hissing and puffing when approached.

This species is responsible for more fatalities than any other African snake. This is due to a combination of factors, including its wide distribution, common occurrence, large size, potent venom that is produced in large amounts, long fangs, their habit of basking by footpaths and sitting quietly when approached.

The venom has cytotoxic effects and is one of the most toxic of any vipers based on LD50. The LD50 values in mice vary: 0.4–2.0 mg/kg IV, 0.9–3.7 mg/kg IP, 4.4–7.7 mg/kg SC. Mallow et al. (2003) give an LD50 range of 1.0–7.75 mg/kg SC. Venom yield is typically between 100–350 mg, with a maximum of 750 mg. Brown (1973) mentions a venom yield of 180–750 mg. About 100 mg is thought to be enough to kill a healthy adult human male, with death occurring after 25 hours.

In humans, bites from this species can produce severe local and systemic symptoms. Based on the degree and type of local effect, bites can be divided into two symptomatic categories: those with little or no surface extravasation, and those with hemorrhages evident as ecchymosis, bleeding and swelling. In both cases there is severe pain and tenderness, but in the latter there is widespread superficial or deep necrosis and compartment syndrome. Serious bites cause limbs to become immovably flexed as a result of significant hemorrhage or coagulation in the affected muscles. Residual induration, however, is rare and usually these areas completely resolve.

Other bite symptoms that may occur in humans include edema, which may become extensive, shock, watery blood oozing from the puncture wounds, nausea and vomiting, subcutaneous bruising, blood blisters that may form rapidly, and a painful swelling of the regional lymph nodes. Swelling usually decreases after a few days, except for the area immediately around the bite site. Hypotension, together with weakness, dizziness and periods of semi- or unconsciousness is also reported.

If not treated carefully, necrosis will spread, causing skin, subcutaneous tissue and muscle to separate from healthy tissue and eventually slough with serous exudate. The slough may be superficial or deep, sometimes down to the bone. Gangrene and secondary infections commonly occurs and can result in loss of digits and limbs.

The fatality rate highly depends on the severity of the bites and some other factors. Deaths can be exceptional and probably occur in less than 10% of all untreated cases (usually in 2–4 days from complications following blood volume deficit and a disseminated intravascular coagulopathy), although some reports show that severe envenomations have a 52% mortality rate. Most fatalities are associated with bad clinical management and neglect.

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Edited by Taipan, Aug 18 2012, 02:33 PM.
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Puff Adders are Masters of Disguise and Smell
The puff adder (Bitis arietans) is a master of camouflage but did you know that it can also mask its scent to make it nearly invisible to animals that rely on scent to find prey items? That is right, the venomous puff adder is a master of scent camouflage.
Researchers with the University of the Witwatersrand’s School of Animal, Plant and Environmental Sciences in Johannesburg, South Africa, trained dogs and meerkats to determine whether these animals could detect puff adders via olfaction, or via sense of smell. They also trained the animals to detect via smell, five species of active foraging snakes versus ambush snakes like the puff adder.

What their research determined was the dogs and meerkats could easily detect the foraging snakes--their accuracy in locating them was better than 80 percent. They were unable, however to detect either the captive or wild puff adders.
The puff adder hides itself scent-wise via what scientists call chemical crypsis, which basically means it can mask its scent so as to go undetectable. The researchers speculate that other ambush snakes may also mask their scent to remain undetected. The puff adder is found in nearly all of sub Saharan Africa, South Africa, and on the Arabian Peninsula into Saudi Arabia and Yemen. It is found in most habitats with the exception of true deserts, rain forests and tropical alpine habitats. They feed on small mammals, birds, amphibians and lizards. It bites more people in Africa than any other snake. The venom of this species is one of the most toxic of viper based on LD 50 values. Symptoms can include bleeding and swelling, hemorrhages, and deep necrosis.
An ambusher's arsenal: chemical crypsis in the puff adder (Bitis arietans)
Ambush foragers use a hunting strategy that places them at risk of predation by both visual and olfaction-oriented predators. Resulting selective pressures have driven the evolution of impressive visual crypsis in many ambushing species, and may have led to the development of chemical crypsis. However, unlike for visual crypsis, few studies have attempted to demonstrate chemical crypsis. Field observations of puff adders (Bitis arietans) going undetected by several scent-orientated predator and prey species led us to investigate chemical crypsis in this ambushing species. We trained dogs (Canis familiaris) and meerkats (Suricata suricatta) to test whether a canid and a herpestid predator could detect B. arietans using olfaction. We also tested for chemical crypsis in five species of active foraging snakes, predicted to be easily detectable. Dogs and meerkats unambiguously indicated active foraging species, but failed to correctly indicate puff adder, confirming that B. arietans employs chemical crypsis. This is the first demonstration of chemical crypsis anti-predatory behaviour, though the phenomenon may be widespread among ambushers, especially those that experience high mortality rates owing to predation. Our study provides additional evidence for the existence of an ongoing chemically mediated arms race between predator and prey species.
Edited by Ceratodromeus, Dec 24 2015, 03:15 AM.
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account of a puff adder biting and killing unfortunate donkey foals,
Animal life in africa(By James Stevenson-hamilton)
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3D reconstruction of fang replacement in the venomous snakes Dendroaspis jamesoni (Elapidae) and Bitis arietans (Viperidae)
"Venomous snakes use highly specialized teeth, so-called
fangs, to kill living prey items. The evolution of these fangs
and that of the associated venom-producing and -delivery
system has been the subject of continuous research (e.g.,
Kochva 1978, Kardong 1982, Knight & Mindell 1994,
Jackson 2003, Fry et al. 2008, Vonk et al. 2008). In addition
to the evolutionary origin of tubular fangs, their ontogenetic
formation has been studied as well (e.g., Tomes
1874, Bogert 1943, Klauber 1972, Lake & Trevor-Jones
1987, Lake & Trevor-Jones 1995, Jackson 2002). While
our knowledge has significantly increased thanks to the
integrated evaluation of palaeontological, morphological,
physiological, molecular and other data, some details still
remain unresolved. One such remaining problem concerns
the timing and regulation of fang replacement, i.e.,
when and how a functional fang is replaced by a new fang.
This topic has been briefly touched by Jackson (2007).
Snakes replace their teeth, including the fangs, regularly
and continuously, therefore there is always a number of
replacement fangs posterior to the functional one. The
maxilla typically has sockets for two fangs in lateral versus
medial positions. At any given time, one would expect
one fang to be solidly fused (ankylosed) to the socket and
the other more or less loose and in the process of either
attaching or being shed. However, as a functional safety
factor, the phase when both fangs are ankylosed should
overlap to some degree, avoiding a time window where
both the replacement fang and the one in the process of
being shed would not be fixed to the upper jaw. According
to Klauber (1972), this period of overlap must be short,
since two functional fangs on one side (i.e., both are firmly
set in neighbouring sockets) are observed very infrequently."
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"Puff adders (Bitis arietans) are heavy ambush predators
in Africa, and they have very large fangs that may exceed
30  mm in length, fused to short but wide maxillae. The
anatomy of the skull and the mass of attached muscles facilitate
a powerful strike, and the amount of injected venom
can be very high compared to that in other venomous
snakes (see also Fig. 3, left). Our specimen was fixed and
preserved with the mouth closed, and therefore its fangs
remained in a resting position during the reconstruction.
On the right-hand side, two fangs are visible in a parallel
position, resulting in equal striking angles (Figures 3–4).
From a mechanical point of view, both fangs would be fully
efficient in penetrating a prey animal. At first sight, both of
these fangs seem to be fully functional. They are roughly
equal in size and positioned very close to each other. To
have a better view, pieces of bones were digitally cut away
around the fangs to make close-up views possible (Fig. 5).
These show that fang No. 1, fixed in a lateral socket of the
maxilla is the functional fang, while the one less firmly ankylosed
in the medial socket is the next fang in the replacement
series. The alternate possible interpretation would
have been that the medial fang is older than the lateral one
and in the process of loosening prior to being shed. However,
based on a comparison of the angles of the shafts of
fangs 1 and 2 (# 1 is in a closer-to-vertical position, and
# 2 lies at an angle intermediate between 1 and replacement
fang 3), we interpret fang # 2 as being in the process of attaching
to the maxilla rather than in the process of detaching.
According to Klauber’s (1972) explanation of the sequence
of fang replacement, fang No. 3 will replace whichever
in-place fang (lateral or medial) is shed first"
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Figure 5. Frontal (left) and lateral views (right) of the fangs of Bitis arietans. Numbers indicate the sequence of fangs in the replacement

Link to paper
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Not a Snake I would want to be tagged by

Edited by tigerburningbright, Apr 25 2016, 04:03 PM.
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Effects of a Puff Adder Bite on a Black Rhino

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A lure at both ends: Puff adders leave nothing to chance

Date: February 1, 2017
Source: University of the Witwatersrand
By capturing and analysing thousands of hours of video footage of puff adders hunting in the wild, researchers have shown that puff adders use what is termed 'lingual luring' to attract amphibian prey closer and increase the odds of catching it.

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One of Africa's most iconic snakes, the puff adder.
Credit: Wits University

The Puff Adder (Bitis arietans) is one of Africa's deadliest snake species, not only due to its deadly venom, but also because of its stealthy behaviour in the way that it hunts by ambushing prey.

It has now emerged, however, that it has another deadly weapon in its arsenal, as it not only lies in wait for prey, but that it actively lures prey into striking range.

By capturing and analysing thousands of hours of video footage of puff adders hunting in the wild, Wits University researchers, Xavier Glaudas and Graham Alexander, have shown that puff adders use what is termed "lingual luring" to attract amphibian prey closer, and increase the odds of catching it.

"A puff adder's strike is typically no longer than one or two head lengths (5-10cm) in distance, so it needs a strategy to attract potential prey to come within that striking range in order to catch it," says Glaudas, a herpetologist and Post Doctorate Fellow at the Alexander Herpetology Laboratory at Wits University. "We have found that puff adders use their tongues that resemble an invertebrate that frogs feed on to increase prey capture rate."

Funded by the National Geographic Committee for Research and Exploration, Glaudas and Alexander tracked 86 puff adders over three years at the Dinokeng Game Reserve, about 100km north of Johannesburg in South Africa. Glaudas captured wild snakes and tracked them by surgically implanting radio transmitters into the snakes and releasing them at their place of capture.

"We really wanted to have a closer look into the secretive lives of these fascinating animals, and specifically study their foraging ecology," says Glaudas.

To aid his research, Glaudas and Alexander made use of video cameras, which they set up in front of puff adders that were lying in ambush position.

"We placed our cameras mounted on a tripod about 70cm away from the snake, and the camera continuously recorded what was going on. We came back the next morning to get the memory cards and reviewed everything that happened during the night," says Glaudas. "We gathered over 4600 hours of video footage of snake foraging. So, that is 193 days of continuous footage -- over half a year."

What Glaudas and Alexander saw surprised them. "It was complete luck," says Glaudas. "We know that snakes use their tongues to pick up scent cues in their environment, but these snakes were extending their tongues out of their mouths for up to 30 seconds, which is dramatically longer to what they do when they are just using their tongues to 'smell' their environment. We know of several species that use tongue luring to attract prey. Some wading birds, like egrets do it, as well as alligator snapping turtles and some aquatic snakes, but this is the first time that it is reported in a terrestrial snake," says Glaudas.

Even more surprising was the fact that the snakes only used lingual luring to attract amphibian prey.

"All the cases of lingual luring that we have observed, occurred with frogs, which suggests that puff adders are able to distinguish between amphibian prey and other prey like small mammals."

Glaudas and Alexander also witnessed puff adders waving their tails, suggesting the use of their tails as lures as well. However, none of the tail luring behaviour attracted prey within the camera's field of view, and more data on this are needed.

"We suspect that this behaviour is also used to attract prey, as it is pretty common in snakes, including adders, but we weren't able to observe prey capture with the videos" says Glaudas.

Story Source: University of the Witwatersrand. "A lure at both ends: Puff adders leave nothing to chance." ScienceDaily. www.sciencedaily.com/releases/2017/02/170201092618.htm (accessed February 2, 2017).

Journal Reference:
X. Glaudas, G.J. Alexander. Food supplementation affects the foraging ecology of a low-energy, ambush-foraging snake. Behavioral Ecology and Sociobiology, 2016; 71 (1) DOI: 10.1007/s00265-016-2239-3

The effect of food availability on the spatial ecology of animals varies within and across study systems because a multitude of factors can affect the spatial activity of organisms. Low-energy specialists, such as ambush-foraging snakes, feed infrequently and can endure long periods without food. Because they have low-energy requirements, one possible tactic for feeding may be to simply ambush for longer periods when prey availability is low, thereby decreasing the potential costs associated with locating new ambush sites. We used radiotelemetry, supplemental feeding, and remote video cameras on free-ranging male puff adders (Bitis arietans) in South Africa to test the hypothesis that food intake affects the foraging ecology of extreme low-energy, ambush foragers and to quantify their natural feeding rates. Supplementally fed puff adders improved their body condition, spent less time foraging, and decreased distance traveled compared to control snakes. However, movement frequency and home range size did not differ between the two groups. These findings indicate that control snakes traveled farther within similar-sized home ranges compared to fed snakes and did so at no survival cost. Further, naturally foraging puff adders successfully caught a prey of small size once every 10 days on average. Hence, despite their “sit-and-wait” foraging strategy and their low-energy intake/requirements, underfed puff adders travel widely to presumably find appropriate ambush sites that maximize prey capture. Our research provides the first strong evidence that the spatial activity of a terrestrial vertebrate species with extremely low energetic demands is significantly affected by food intake.

Edited by Taipan, Feb 2 2017, 04:51 PM.
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