Eastern montpellier snake | Malpolon insignitus | Greek herpetofauna

Introduction

The Eastern Montpellier snake – Malpolon insignitus; where does one even begin to describe this remarkable reptile? These fiery, lightning-fast, beautiful, snake-eating, omnipresent and charismatic rear-fanged snakes are possibly the favorite Greek snake of several members of our team, and understandably so. In fact, the sheer force and might of this snake is a fact so well-established that even its genus name “Malpolon” hints at it. It is derived from the Greek “μάλας” meaning “grand/strong” and “πολύς”, with roughly the same translation in this context. The sole extant members of Psammophiidae in Europe, Montpellier snakes possess numerous unique features that clearly set them apart from all other European snakes, from their unusual scalation, to their exceptional intelligence and associated one-of-a-kind behaviors. Members of the genus Malpolon in Europe were somewhat recently split into two species, M. monspessulanus and M. insignitus, the latter being the species that occurs in Greece (with all Greek populations assigned under the subspecies M. i. fuscus) while M. monspessulanus is restricted to Western Europe and Western North Africa (Maghreb) across the Gibraltar strait. The epithet “insignitus” means “distinguished/separate” in Latin, denoting its split from M. monspessulanus. Eastern Montpellier snakes are widely distributed in Greece, from Thrace to the Peloponnese and from the Ionian to the Anatolian islands and they seem to persist even in the most inhospitable and disturbed habitats. Regrettably, they have a terrible reputation as a dangerous and aggressive snake not only among locals but also some herpers, which is entirely undeserved. Their Greek common name “sapíti” (σαπίτης) also quite literally translates to “rotter”, which is a misnomer, as despite their tough behavior, these snakes are harmless to people, their venom certainly does not cause rotting, and as with all Greek snakes, they avoid us at all cost.

Description

Very easily identifiable due to the characteristic scalation on the head; the supraocular, preocular and prefrontal scales on either side are raised, forming two ridges/arches, which give the en face view of the snake’s head an unmistakable V-shape. The dorsal scales are arranged in 17 rows (occasionally 19), lack keels and are concave (grooved), which gives the snake a satin or matte appearance instead of the glossy texture of a typical smooth-scaled snake. Maximum TL very likely over 200 cm, though a more usual adult size in most areas of the mainland ranges between 90-160 centimeters TL. Nevertheless, larger individuals are not uncommon and adults can also be smaller. Females are typically considerably smaller than males, though there are rare exceptions to the rule, with females over 150 centimeters TL having been recorded. A fairly robust snake, with mature individuals being proportionately more thick-bodied than juveniles. Large adults can reach a kilogram in weight, if not surpass it. The tail is thin, not particularly long in relation to the rest of the body and it tapers gradually. The eyes are large and the pupil is round.  Coloration and pattern vary immensely according to age, sex, as well as individual variation, and broadly speaking, no two Montpellier snakes ever look the same. Dorsal coloration of juveniles can range from cream, to tan, to light bronze, to greyish olive, to reddish brown/gray, to dark brown/gray with varying amounts, size and intensity of darker spots that may form vague rhombus/triangle-like shapes with light-colored edges. The ventral is a pinkish-salmon orange, sometimes with rows of small dark dots and semi-defined light (yellowish) longitudinal lines. Females retain juvenile dorsal coloration and patterns into adulthood to a great extent, though, at the same time, there are obvious differences between the two. The ventral of adult females can sometimes be incredibly ornate, consisting of many well-defined rows of dots with several intense and vibrant colors such as reddish/bronze, orange and pink that are most intense at the height of the neck. Males, on the other hand, lose their dorsal markings and intense ventral colors/patterns as they mature, and the dorsal coloration can range from greenish/olive-gray, blueish green-gray, dark gray and brown-gray to almost solid black. The ventral can be pale or with a poorly defined lighter stripe in down the middle. While light spots exist on the labial scales of adults of both sexes, they are highly defined in females (sometimes with almost black/white contrast and intense orange), as opposed to males, the spots of which can be characterized as faded and relatively indistinct. The color/pattern transition from juvenile to adult is very gradual, giving rise to a vast spectrum of intermediate forms, and the sex of a snake can be discerned via coloration and pattern usually only when the animal is well into adulthood. The three dorsal scale rows closest to each side of the ventral have patches of white or at least very pale coloration in their edges, which gives most snakes the appearance of two or three very faint light longitudinal lines near the ventral. Melanism has been recorded at least once in Croatia, though it is extremely rare and melanistic individuals do not exhibit a jet-black appearance. Situated approximately in the middle of the upper jaw is a pair of enlarged, fixed, grooved fangs. Malpolon insignitus also possess the Duvernoy’s gland, which produces a low-grade neurotoxic venom, which is harmless to humans.

Adult male (C) Stamatis Kalogiannis

Adult female (C) Stamatis Kalogiannis

Adult female (C) Stamatis Kalogiannis

Hatchling (C) Stamatis Kalogiannis

Adult female (C) Charis Kouelis

Young male, TL approx. 70 cm (C) Plato Stefanopoulos 

Juvenile, TL approx. 35 cm (C) Plato Stefanopoulos

Adult male (C) Plato Stefanopoulos

Juvenile (C) Plato Stefanopoulos

Large adult female, TL >115 cm (C) Plato Stefanopoulos

Young female (C) Plato Stefanopoulos

Young, nearing the end of a shed cycle (C) Plato Stefanopoulos

Young adult, unsexed (C) Plato Stefanopoulos

Young adult, unsexed (C) Plato Stefanopoulos

Adult female (C) Charis Kouelis

Large adult male, TL 145 cm (C) Plato Stefanopoulos

Adult female (C) Plato Stefanopoulos

Young (C) Plato Stefanopoulos

Juvenile, under 3 months old (C) Plato Stefanopoulos

Adult male (C) Stamatis Kalogiannis

Habitat 

Inhabits almost all Mediterranean habitats from sea level to 2000 m asl, though it is said to become rare above mid elevations (approx. 600-800 m asl), though this is by no means a rule that applies to all places. It can be argued that it is one of the ultimate habitat generalists among the Greek snakes. Its preferred habitat seems to be maquis, phrygana and open hardwood/mixed/conifer forests, though it is by no means limited to the above. It also inhabits meadows, coastal dune systems, marshes, floodplains and coastal woodlands, forest edges, open olive groves, other orchards and plantations and even severely degraded habitat such as agricultural, industrial and even suburban areas. It can also withstand significant drought, and it is often found in the driest, most xeric and inhospitable habitats such as corroded hillsides and pine forests near city limits. It may also be found in bushy grasslands and open cultivations, though, while it can thrive in surprisingly open habitat, it does not seem to tolerate it to a degree as extreme as that of Dolichophis caspius.

Cultivated olive grove with stonewalls – habitat of Malpolon insignitus

Montane oak forest – habitat of Malpolon insignitus

Grassland bordering a lakeside forest – habitat of Malpolon insignitus

Mixed conifer-hardwood forest – habitat of Malpolon insignitus

Wet meadow – habitat of Malpolon insignitus

Maquis and phrygana hillside – habitat of Malpolon insignitus

Maquis, pastures and streamside plane forests – habitat of Malpolon insignitus

Phrygana bordering riparian area – habitat of Malpolon insignitus

Riparian area – habitat of Malpolon insignitus

Open, humid grassland bordering streams – habitat of Malpolon insignitus

Montane forests interrupted by maquis and pastures at approx.1000 m asl. – habitat of Malpolon insignitus

Coastal dune system – habitat of  Malpolon insignitus

Suburban area in a village with Eucalyptus and other plantations – habitat of Malpolon insignitus

Adult female in her open, degraded vineyard and olive grove habitat (C) Plato Stefanopoulos

Juvenile in its traditional orchard habitat (C) Plato Stefanopoulos

Adult male in its riparian habitat (C) Plato Stefanopoulos

Biology

Eastern Montpellier snakes overwinter underground, and while healthy adults will rarely emerge from their hibernacula on mid-winter warm spells, juveniles and old individuals are very prone to winter activity and basking individuals have been recorded on sunny December days where the air temperature was as low as 14 degrees Celsius. That being said, the aforementioned hesitation is not particularly apparent in the species’ southernmost localities in Greece such as lowland Messenia or Laconia. The last few winters, there seemed to be a distinct activity spurt of Malpolon in the Balkans and some individuals were found DOR on the warmest days of January. The spring emergence seems to occur very early on, especially in lowland areas, and healthy adults can usually be reliably found basking on sunny days in February with the air temperature as low as 12.5 degrees Celsius. Their spring emergence is very slow and gradual and oftentimes, in the earliest months, the snakes will only expose their head and neck from the hibernaculum while the rest of the body remains in the refuge, a behavior which we have recorded directly and we have seen apparent evidence of, meaning that before their first shed of the year, adult individuals are often muddy on their lower half, while the upper half is clean. The early-mid spring is a time of preparation for the mating season and an important feeding period, especially for younger snakes. During that time, the active snakes will generally remain close to their hibernacula. The feeding habits of Malpolon insignitus could perhaps best be described as “opportunistic euryphagy”, as they will generally consume whatever prey they can catch. While their preferred prey seems to be medium-large lizards (especially Lacerta), they are also known to eat small mammals, amphibians, birds, insects (especially juveniles), as well as other snakes, including vipers and other Montpellier snakes. Malpolon insignitus has excellent eyesight and is a very visual snake. It is also lightning fast, arguably one of the fastest-moving snakes in Greece. It is ground-dwelling, though it will often climb on dense shrubs and stone walls. When active, the snakes will often stop and rear up the front part of their body in a “periscoping” position to spot prey or assess potential threats from a distance. When a prey item is located, the snake may chase it over a considerable distance before intercepting it. It is also an excellent swimmer and it has been recorded successfully hunting dice snakes (Natrix tessellata) in the water. Once a prey item is caught, it is not killed by constriction, though the snake may use coils of its body to restrain it. Prior to swallowing, the prey is envenomated by chewing, so that as much venom as possible can be transferred via the grooves of the fangs. This action serves to paralyze and ultimately kill the prey so that the risk of injury to the snake while swallowing is minimized. Contrary to most other Greek snakes, Malpolon are not secretive in the slightest, and they’re arguably one of the easiest snakes to observe in Greece. They are very active in general, and they tend to frequently move over considerable distances for various reasons, especially in search of prey and mates, which makes them especially vulnerable to being ran over by cars, and regrettably, they are one of the most common roadkills in Greece. They can easily be seen basking outside refuges on cooler spring days, though, paradoxically, this seems to be a somewhat reliable way of finding them year-round. They have been found on roads in a very wide range of temperatures, from an air temperature of 19.5 C to (allegedly) a ground temperature of 48 C. When it comes to our own observations, we have seen a massive adult active next to a stream in 35 C, in midday, on a clear, sunny day, and routinely see them crossing roads in the summer in air temperatures between 30-32 C, temperatures often too high to observe any other snakes on the road. Apart from being incredibly heat-resistant, they also have a remarkable tolerance for drought. This resistance to drought is related to one of the most intriguing aspects of Eastern Montpellier snakes and Psammophiids in general; a rubbing behavior where the snake will rub its nose against the rest of its body and its ventral side so that it can cover itself with a nasal secretion that prevents water loss. There is a distinct peak in the number of DOR adult males in May-June, suggesting that this is most likely when most of the mating takes place. Montpellier snakes have a very unique mating ritual. Firstly, each male establishes a territory and vigorously defends it against rival males. Male-male combat is very strange, namely, both snakes will bite each other until one asserts dominance over the other. Perhaps the most intriguing part of the mating process is courting, during which the male will kill a prey item and present it to the female as a “gift”. The precise reason for this behavior remains unclear, though it has been theorized to clarify the male’s interest in mating as well as to potentially discourage the female from consuming the male if she happens to be larger than him. When faced with a potential predator (this includes humans), the snake will always rely on its speed to escape. It is very cautious and alert, and therefore it usually sees us from several meters away and instantly disappears into thick vegetation or any other kind of refuge before we can even see it. When cornered or discovered in the open, it will stand its ground, hiss extremely loudly (it has the loudest hiss of all the Greek snakes) and raise the front part of its body in a remarkable defensive display. If approached within less than a meter in this state, it will strike vigorously and potentially bite and then attempt to escape. If caught, it will thrash violently, bite repeatedly and potentially musk and defecate. While Montpellier snakes will readily bite when harassed, the aim of these bites does not seem to be envenomation, as they do not use force and release immediately. If handled for several minutes, only then they may bite, hold on and chew in an attempt to envenomate.

Periscoping position (C) Stamatis Kalogiannis

Periscoping position (C) Plato Stefanopoulos

Adult female only exposing her head and neck out of the hibernaculum. February. (C) Charis Kouelis

Various defensive posturing of adult snakes. Rearing of the head and neck, horizontal and vertical neck-flattening (C) Plato Stefanopoulos

Venom and bite symptoms 

The venom of Montpellier snakes is described as a low-grade neurotoxin that serves to paralyze and ultimately kill prey items. It is produced in the Duvernoy’s gland, and it exists in the saliva of the snake. While Malpolon are harmless snakes, we believe that envenomation should still be taken seriously, as symptoms are very likely to arise. While envenomation by M. insignitus remains virtually completely unstudied, there have been scientific publications regarding hospitalizations related to extreme cases of envenomation by the closely related M. monspessulanus, which resulted in symptoms such as headache, vomiting, localized paralysis, pain and localized swelling, among others of similar severity. That being said, these symptoms not only subsided over the course of some hours or a few days at most, but they were the result the snake channeling venom in the patients for over a minute, something that can only possibly happen if the snake is handled or otherwise mistreated for several minutes. No human deaths have been caused by either species, and we do not think it’s possible. When it comes to our personal experience, the symptoms we have recorded from bites where the enlarged, grooved teeth managed to pierce the skin have been very localized swelling and itching (e.g., a single finger) and very mild pain that completely subside within an hour or two. Nevertheless, it is important to consider that we have not allowed the snakes to envenomate for more than a few seconds. Though it may be somewhat of an arbitrary conclusion, the severity of bite symptoms generally seems to be positively correlated with the time that the snake is allowed to chew, i.e., the amount of venom channeled into us, especially considering that bites where the grooved teeth pierce the skin but snake immediately releases do not show any symptoms whatsoever, and all severe symptoms have been the result of bites that lasted for minutes. In conclusion, while we confidently label M. insignitus as a harmless species, we believe that handling these snakes should only be done by people who are experienced and have considered all the probable results of envenomation.

Distribution 

Being one of the most widespread snakes on the Greek mainland, it occurs throughout all of its extent; from the southernmost point of the Peloponnese peninsula, to the northernmost point of Thrace. It is also found on most of the Ionian islands (Corfu, Lefkada, Kefallonia, Ithaca, Zakynthos), as well as on some of the Aegean islands, particularly those closer to the mainland of Greece and Turkey; Skiathos, Skopelos, Thasos, Samothrace, Limnos, Lesvos, Chios, Samos, Rhodes, Salamina, Hydra and Elafonisos. While looking at the distribution, it quickly becomes evident that the species is only absent from islands that are known to have split from the mainland earlier than the ones where it's present - the most likely explanation for this phenomenon is that since Malpolon insignitus is a species of North African origin, it seems to have spread to Anatolia and the Balkan peninsula relatively recently, that is, after a lot of islands (e.g. Crete, Kythera and the Cyclades archipelago) had already been disconnected from the mainland.

Distribution of Malpolon insignitus in Greece