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Self-rubbing of psammophiids

Critical remarks on "A Contribution to the understanding of the self-rubbing behaviour in psammophiid snakes (Squamata: Psammophiidae)" by Stphanie de Pury & Wolfgang Bhme Salamandra !"#$%& $'()* #+*$)%
,-. S,//0-12/3 4eeneinde ! +)'$ Weelde Belgium

Corresponding address: March 1, 2014

.otes of the author $5 This article was originally sub itted to Salamandra, but the editors did not want to accept it in this !or !or publication. The reasons they ga"e, were: the article was too long, too aggressi"e and too !ull o! sub#ecti"e state ents. $ did not %stick to the !acts&. $nstead o! utilating y article, $ chose to withdraw it and publish it on the internet. 2. This article is partially based on the critical re"iew o! a thesis o! '. de (ury about the rubbing beha"iour o! psa ophiid snakes, that the author wrote together with 6ichel-7ran8ois 0abersaat, and that is published on the internet too.

$n Salamandra 4)*1+, 201,, -r. 'T.(/01$2 -2 (345 and (ro!. -r. 6789:01: ;</M2 published =0 contribution to the understanding o! the sel!>rubbing beha"iour in psa ophiid snakes *'?ua ata: (sa ophiidae+=, in which they try to clari!y so e aspects o! the sel!>rubbing beha"iour o! psa ophiids snakes and discuss its possible !unctional signi!icance. They conclude that the rubbing o! these snakes @ser"es pri arily the pre"ention o! e"aporati"e water loss@ *p. 2A+. $n that respect, their article see s to consolidate an interpretation o! the sel!>rubbing beha"iour that has been repeated in literature since it was de"eloped by -31'71, -31'71 B C2$T/ *1)DA+. $ think this is a istake.

,he body :ater loss pre;ention interpretation of self-rubbing beha;iour

$n the description o! the hypothesis that the sel!>rubbing o! psa ophiid snakes ser"es to protect the skin against water loss, the authors state: @E se"eral authors considered sel!rubbing as protecting the skin against water loss *;401C/ 1)AA, 80/0F B -M$G28 1))H, 628-71 et al. 200A+@ *p. 1)+. $n this short list o! re!erences, the !ield guide o! ;401C/ *1)AA+ can hardly be called a serious source: he only entions "ery shortly and inaccurately the sel!>rubbing beha"iour in the general paragraph about sand and grass snakes: @'o e species *P. sibilans and P. schokari+ =polish= the sel"es with a nasal gland secretion which reduces skin water>loss@ *p. D0+, without re!erence. The second source, 80/0F B -M$G28 1))H, does surprisingly not e"en ention the rubbing beha"iour. To the contrary, what these authors do write *see below+, can be considered as an argu ent against the water loss hypothesis. 0s !or the last source entioned by -2 (345 B ;</M2, there has also been so e careless re!erring: 628-71 2T 08. *200A+ consider concerning psa ophiids not only the water loss hypothesis: @1asal gland !luids, which dry to !or to a lusterless !il on the skin, are hypothesiIed to retard e"aporati"e water loss or to contain phero ones used to scent> ark conspeci!ics andJor territories@ *p. D4D+. 628-71 2T 08. *200A+ re!er eKclusi"ely to -31'71 2T 08. *1)DA+, including -31'71=' incorrect suggestion that psa ophiids @rub their snout along their dorsal and "entral skin sur!aces@ *p. D4D+, which -31'71 2T 08. on their turn borrow !ro -2 :4$L' *1A)A+, who obser"ed P. sibilans and P. schokari but apparently not "ery closely. 0s we know, they only rub their "entral scales *eKcept !or Ramphiophis that also has been obser"ed to rub lateral scales+, and absolutely not their dorsu *see e.g. -2 /001 1)A2, 1)))M 'T22/73-24 1)ADM -2 (345 2010M "ideos as entioned in the 4e!erence 8ist+. 628-71 2T 08. *1)DA+ re!er to @watery nasal gland secretions@, which is interesting in the conteKt o! a water repelling !unction o! the sa e secretion. They also ention any other !unctions o! che icals in and !ro the skin: @Che icals !ro the integu ent N the epider is and skin glands N also protect reptiles against pathogenic icroorganis s, ectoparasites, including disease "ectors, and predators, in addition to attracting ates and eliciting other phero onal responses.@ $t ay be clear that 628-71 2T 08. *1)DA+ conse?uently can hardly be used as a backing !or the water loss hypothesis. 2"en -31'71, -31'71 B C2$T/ *1)DA+, the original source o! the water loss hypothesis, is doubt!ul, as was elaborately indicated by -2 /001 *1)))+. $ ha"e the i pression all other authors that re!er to this article, only read the abstract. $n the body itsel!, the authors are uch ore care!ul. There is no indication that they the sel"es e"er obser"ed any rubbing

beha"iour. Those snakes that were kept ali"e during their research, were kept in eKperi ental conditions that ake it rather i probable they e"er rubbed. The eKperi ents that were carried out to deter ine their e"aporati"e water loss, did not include a co parison between snakes that had rubbed and snakes that had not. 0nd yet, without rubbing, water loss was eKtre ely low. @$ndeed it ay ha"e the lowest rate o! loss e"er easured !or an ophidian@ *p. 4D2+. 0nd: @3nder lab conditions *low acti"ity, 24>2DOC+ M. monspessulanus does not need to drink to aintain its body weight. 'u!!icient water can be obtained !ro a alian prey *lab ice are D,P water+@ *p. 4D0+. They conclude: @$t ight see unlikely that s all a ounts o! a substance spread o"er a snake=s skin could ha"e any signi!icant e!!ect on water per eability. QER 'ince the che ical co position o! the !luid is only partially known, nothing de!inite can be said about its ability to retard passage o! water across the skin. /owe"er the well established property o! certain co pounds, such as cetyl alcohol and long chain !atty acids applied in onolayers, to considerably retard e"aporation !ro water sur!aces pro"ides a use!ul odel@ *p. 4D2+. 0 use!ul odel !or !urther research, that=s all. 0pparently, -31'71, -31'71 B C2$T/ *1)DA+ do not take into consideration any other possible !unction o! the secretion, which see s re arkable. Can this be the reason why -2 (345 !ro the beginning *her 2010 dissertation+ see s to ha"e been !iKated on, !irstly, the presence o! lipids in the secretion and, secondly, on the possible water repellent !unction o! lipids in the secretionS $t is in y opinion ob"ious that -31'71=s hypothesis that s earing a watery secretion in a relati"ely narrow track only on the "entral scales *where it has the largest chance o! being rubbed o!! in a short ti e by contact with the substrate+ would be able to pre"ent water loss through the skin, ust in !act be considered as eKtre ely unlikely, and $ !ind it re arkable that -2 (345 B ;</M2 re ain so persistently !iKed on that hypothesis. -2 (345 B ;</M2 state: @This hypothesis recei"ed !urther support !ro so e anecdotal reports that sel!>rubbing !re?uency was lower when the snakes were eKposed to higher hu idity *'T22/73-24 1))2, ;401-'TTTT24 1))H+ and beca e higher when snakes were eKposed to higher te peratures and drier conditions *-042F'C5 1)UH, 4$2((28 1)D,, -2 /001 1)A2, 'T22/73-24 1)AD, 1))2+@ *p. 1)+. /owe"er, the authors do not consider any other eKplanation. $ ysel! *'T22/73-24 1))1+ !or instance, suggested *assu ing that the rubbing would ser"e to ark en"iron ent+ that less rubbing would take place in non>arid conditions because in a such an en"iron ent scents will e"aporate and disappear at a slower rate, resulting in less need to re!resh the traces. 0lso unpro"en, o! course, but uch ore likely than the oisture loss hypothesis, as could be concluded !ro the study o! 08;24T' *1))2+ and 650TT *200,+. The relation between high te peratures and water loss see s co plicated in psa ophiids snakes. 8$C/T B ;2112TT *1)D2+ de onstrated the general ability o! snake skin to protect against e"aporation. 80/0F B -M$=28 *1))H+ disco"ered eKtre ely low cutaneous water loss in eKperi ents with Psammophis schokari, at low and at high te peratures, the lowest in the stripeless !or o! Psammophis !ro Keric biotopes in the 1ege" desert. The desert !or o! Psammophis appeared able to conspicuously change the "alues o! cutaneous water loss. The authors think it likely that this @re!lects peripheral "aso otor changes, i.e., "asodilatation and "asoconstriction o! the skin blood "essels. -ue to its diurnal acti"ity, the Keric !or o! Psammophis is o!ten eKposed to intense solar radiation, high te peratures and low hu idity co on in deserts. (resu ably, the "aso otor response enables this snake a better and ore i ediate control o! its C68@ *p. 1,A+.

'o e psa ophiids inhabit en"iron ents that are not so dry a!ter all, !or instance Psammophis (Dromophis) lineatus that typically inhabits wetlands *;iodi"ersity o! the Va beIi ;asin 6etlands 2000+. 'till, these also eKhibit rubbing beha"iour. $t would be worth considering this aspect too. $! -2 (345 B ;</M2 want to seriously de!end the so unlikely water e"aporation hypothesis, against all odds, one would eKpect a "ery serious research. $ was rather disappointed when $ read their paper. 9or instance: they analysed the ultrastructure o! dorsal and lateral scales. $n the article in Salamandra they don=t tell us why, but reading the 2010 dissertation o! -2 (345 akes clear that this ser"es to in"estigate i! the ultrastructure o! psa ophiids scales di!!ers !ro those o! other snakes in a certain aspect, that is the absence o! pores. @E i! psa ophiids ha"e a porous structure to allow the eKit o! lipids *E+, why should they additionally s ear a secretion containing lipids on their bodyS@ *par. 1.Da+. This is a rather strange oti"ation, as it already assu es what yet has to be pro"en: that the secretion contains lipids that can and do protect the snake=s skin against water loss. 0nd what is ore: i! there would ha"e been pores, this would not necessarily ha"e eant that the *hypothetical+ lipids would 17T ha"e acted as an additional protection. 0nd thirdly: i! there were pores, would they ser"e as secreting ducts !or lipidsS 0nd !inally: what proo! would there be that the !ound lipids would ser"e to pre"ent water loss and not ha"e any other !unction, !or instance phero onalS ;ut y ost ob"ious ob#ection is: why didn=t the authors analyse the ventral scales, as these are the scales that are rubbed and s earedS $nstead, they analysed "ertebral and lateral scales > that are not rubbed at all. 7nly Rhamphiophis sp. see to rub lateral scales, all other species only rub the "entral scales. 4eaders that ne"er ha"e obser"ed the rubbing beha"iour o! these snakes, can easily see what $ ean i! they look at the "ideos $ placed in y 5outube Channel *address added to the list o! re!erences+. $t is a pity that the "ideos that were used in the research o! -2 (345 B ;</M2 are not a"ailable !or public use. Considering the abo"e, the analysis o! the scales as described in the paper o! -2 (345 B ;</M2, see s interesting but in this conteKt rather irrele"ant, as is conse?uently e"erything the authors state about it in their discussion section. 9ro the 2010 dissertation o! -e (ury, where the research is ore elaborately described, $ learned that the lateral scales ha"e been le!t out o! !urther research !ro the beginning, as there appeared to be "ery little structure on their sur!ace *-2 (345 2010, par. ,.1+. The "entral scales *which, $ repeat, are the only scales that are rubbed+ al ost co pletely lack sur!ace structure, eKcept !or a short period a!ter shedding *;401-'TTTT24 1))U+. The authors state on p. 2H: @ 6ith regard to rubbing beha"iour, the delicately sculptured icro>orna entation ight ser"e to retain the nasal glandsG secretion on the scale sur!ace@. 6hat is there to retain i! the secretion is not e"en applied to these dorsal scalesS 7nly i! this retaining would concern secretion that was rubbed o!! !ro the belly o! a snake to ark another snake, would the re ark be sensible, but that see s not to be the intention. $ can be short about the analysis o! the secretion that is described and discussed in the article o! -2 (345 B ;</M2: absolutely nothing can be concluded. @The presence o! lipids in the nasal secretion could be clearly deter ined in the two species studied. /owe"er it could not be clearly identi!ied which lipids eKactly they are@ *p. 2D+. 1either was there any clue to what the !unction o! the unknown lipids could be.

$n the study o! -31'71, -31'71 B C2$T/ *1)DA+ lipids were also !ound in the secretion, but only in s all ?uantities: @The lipid content o! the secretion appears to be a "ery inor co ponent@ *p. 4HA+. There is no indication !or the relati"e ?uantity o! lipids -2 (345 B ;</M2 ha"e !ound. The rest in -31'71=s sa ples were ainly proteins, which is according to the !indings o! -2 /001 *pers. co .+ in the 1)A0=s and e"en in the late 1)H0=s, who had sa ples o! dried secretion that was scraped o!! a !ront glass panel *see abo"e+ gas chro atically analysed. (roteins do o!ten ha"e a phero one !unction > as do lipids and !atty acids *650TT 200,+. -31'71, -31'71 B C2$T/ *1)DA+ stated: @ The task that re ains is to collect the natural secretion and to directly test its e!!ect on skin water per eability@. This was apparently not possible within the boundaries o! the research o! -2 (345 B ;</M2. The re!erence to the rubbing beha"iour o! so e !rogs *p. 2A+ is not "ery con"incing. -31'71, -31'71, -31'71 B C2$T/ *1)DA+ already pointed to the super!icial si ilarity with the rubbing o! Malpolon, but ad it that in the !rog secretion lipids were do inantly present, but @that the nasal secretion o! M. monspessulanus contains !atty acids and the skin sur!ace in snakes is considerably ore keratiniIed@ *p. 4D2+. $n short: the analysis o! the scale structure and the secretion in"ol"ed, as described in the article o! -2 (345 B ;</M2, pro"es interesting but in y opinion co pletely !ruitless. The ain conclusion see s to be that nothing could be concluded. 9ollowing 8$C/T B ;2112TT *1)D2+ who concluded that the presence o! scales has nothing to do with the e"aporati"e water loss through the skin, -2 (345 B ;</M2 state: @$ndeed, the protecti"e echanis o! the skin against water loss lies in the epider al layer o! the skin and not on its sur!ace@ *p. 2H+. 0pparently the authors and $ agree that the analysis o! the scale structure and o! the secretion did not support the water loss hypothesis in any way. 8ipids play a role in the pre"ention o! water loss in snakes, but they see to do so in the skin tissue, not applied on its sur!ace > at least, there is not a single indication that this would be the case, neither in the literature cited by the authors, neither in their own research.

,he chemical marking interpretation of rubbing beha;iour

Considering y !oregoing ob#ections, it ay be clear that $ think that the hypothesis that the sel!>rubbing ser"es as pre"ention o! water loss will probably ne"er be con!ir ed, and that another eKplanation can only be ore plausible. 'uch an other interpretation o! the rubbing beha"iour is that sel!>rubbing is used to ark substrates, territories and conspeci!ics, as is eKtensi"ely argued by -2 /001 *1)A2, 1))), 200,+ and -2 /001 B C83C/$24 *200H+. This hypothesis is also the one that is the ost likely. The analyses o! -042F'C5 *1)UH+, -31'T71, -31'T71 B C2$T/ *1)DA+ and o! -2 (345 B ;</M2 *201,+ all show the presence o! che icals that are co only used in or as phero ones *see 650TT 200,+. 0s !ar as lipids are concerned: @The signal li!e o! a alian phero ones used as territory arkers was increased by carrier co ponents in secretions such as sebu , a lipid>rich, oily substance produced by sebaceous glands@ *650TT 200,+. $n the section o! the 201, article by -2 (345 B ;</M2 in which the beha"ioural eKperi ents are described and discussed, the authors try to !ind indications that would !alsi!y or con!ir -2 /001=s arking interpretation. The original dissertation that has been the basis o! the 201,

article, e"en had a working hypothesis that =4ubbing beha"iour is not a *par. 1.D o! the dissertation+.

arking beha"iour@

The obser"ations in the beha"ioural eKperi ents as described in the dissertation and in the 201, article, are "ery interesting. $t is a pity that not ore results o! the obser"ations o! rubbing beha"iour !ro the 2010 dissertation ha"e been included in the 201, article. /owe"er, a a#or ob#ection is, that all ani als in"ol"ed were obser"ed in conditions that were totally di!!erent !ro their natural: in s all enclosures that were te porarily inhabited, solitary, so eti es placed in a new enclosure and re o"ed within so e hours. $t is not at all clear to what eKtent obser"ations ade in such circu stances can be applied to nor al beha"iour in natural circu stances. -2 /001 *1))), 200,+ obser"ed Malpolon in se i>natural circu stances, in wide open enclosures in an area that belongs to their natural habitat. Co parisons should thus be "ery care!ully ade. The description o! the eKperi ents o! -2 (345 B ;</M2 is un!ortunately inco plete. 9or instance: it is stated that continuous recording was chosen so that the ani als would not be disturbed *p. 20+, but eKactly how undisturbed were theyS -2 /001 *pers. co .+, /0;24'00T *pers. co .+, 'T22/73-24 *unpubl.+, they all obser"ed that the rubbing beha"iour was strongly in!luenced by the presence o! obser"ers. The ani als that were kept in enclosures in a roo al ost only rub when they !eel sa!e and are not disturbed by the sight o! people o"ing through the roo or watching the . 6hen they ha"e started a rubbing session, they al ost always stop when an obser"er co es in sight. The 201, paper o! -2 (345 B ;</M2 and the 2010 dissertation do not !ully describe the circu stances under which the ani als were obser"ed. 6hat we do know is, that in part o! the eKperi ent there has been a lot o! anipulating o! the ani als in"ol"ed. That decreases the reliability o! the obser"ations. -2 (345 B ;</M2 tried to deter ine whether there would be a di!!erence in !re?uency in rubbing beha"iour, assu ing that arking beha"iour would be acti"ated i! the ani als were put in a clean, new en"iron ent. Their data did not con!ir this assu ption. @4ubbing !re?uency was not signi!icantly higher on the !irst day than during the !ollowing days in the 1U psa ophiids tested@ *p. 24+. 0necdotal data !ro 'C/0$C/ *pers. co .+ and ysel! see to contradict the obser"ation o! -2 (345 B ;</M2. 0nd what to do with the re ark o! the authors the sel"es on p. 21: @7ur preli inary eKperi ents had shown that 4; is ainly per!or ed during the !irst H0N)0 inutes a!ter being placed into a new terrariu @S The authors do not try to eKplain the de"iation between eKpected and obser"ed beha"iour. 7n the other hand, the assu ption in itsel! is also disputable, as it is based on a narrow interpretation o! the hypothetical arking beha"iour, "iI. that it ser"es to ark a territory, and that without a spatial, en"iron ental or social conteKt. $t is not sure that being placed in a absolutely clean and new enclosure, without traces o! conspeci!ics or other snakes would speci!ically elicit rubbing beha"iour. -2 /001 *200,+: @ $n Malpolon monspessulanus, a species showing pronounced seKual di orphis QER $ obser"ed che ical arking with "entrally scraped o!! nasal secretion particles Q1U,1)R: o! hunting routes traced on substrate by all indi"iduals, irrespecti"e o! seK and ageM o! conspeci!ics in holes and other dark places, where the arking a!terwards ser"es !or utual recognition, pre"ents panic and !a"ours clan !or ation, but ight also discourage aggregation since gi"en indi"iduals risk beco ing the prey o! opportunist conspeci!icsM

o! indi"idual territory li its traced upon and between optical land arks by certain adult ales, li its which they guard and de!end as well as their respecti"e !e ales against ost ale conspeci!ics and di"erse allospeci!ic intruders during HNA weeks in MayJLuneM by a territorial ale, arking his !e ale and one or two "assal> ales, which also N i! not ainly N ark the sel"es dorsally by !re?uently creeping under the territorial do inant.@ $! -2 /001 is right, the arking beha"iour is uch ore co pleK than si ple arking o! a territory. There could also be a connection with a neglected ele ent in the study o! -31'71, -31'71 B C2$T/ *1)DA+, who !ound that @-i!!erent sa ples showed considerable "ariation@ *p. 4HD+. -31'71, -31'71 B C2$T/ *1)DA+: @0 a#or portion o! the central area o! the gland is an eKtensi"ely branched syste o! ducts. $n the only pre"ious study o! the gland, there was no ention o! this peculiarity *-are"ski, =UH+. The need !or such an enor ous syste o! ducts re ains unknown, but it is likely that in addition to storing and passing the !luid released by the tubules, the ducts ha"e an acti"e role in deter ining the !inal co position o! the secretory !luid *p. 4D1+.@ 'o it is not clear i! the co position o! the secretion is always the sa e, and it would be possible that it di!!ers according to the actual !unction o! the rubbing. 0nother part o! the eKperi ent o! -2 (345 B ;</M2 consisted in the use o! glass plates o"er which the ani als crawled. /oriIontal glass plates were used, o"er which the ani als were intended to crawl, rubbing o!! the secretion that had been applied in a rubbing session. @1one o! the indi"iduals o! both species studied had le!t "isible traces o! secretion on the glass plate@ *p. 24+. $ think it would ha"e been "ery unlikely i! they had. The article does not describe eKactly how the glass plates were obser"ed: with or without a icroscopeS =Fisible= suggests that it has been done without. ;ut e"en with the help o! a "ery good icroscope, it would ha"e been "ery hard to !ind particles *presu edly scraped o!!, in this case on s ooth glass+ !or ing a scent trace. This part o! the eKperi ent see s to assu e that large ?uantities o! the secretion, "isibly obser"able, would be needed to ark a trace. 0 "ery strange idea, i! we only re e ber how a dog is able to !ollow a trace or is able to detect a corpse !ro a distance. Lust as an eKa ple: dogs are able to detect A0P o! so e aterials in densities o! 100 particles per *'2TT82' 200U+. 8ike a needle in a haystackW 6hy would be assu e that these snakes would need such enor ous ?uantities o! particles that we would be able to see the e"en without a icroscopeS Maybe there has been a con!usion with the "ery "isible traces o! secretion these snakes o!ten apply acti"ely on the !ront glass panels o! their enclosures by rubbing their nose against the panel * see picture 1+. $t is not clear i! this is a deliberate andJor !unctional action. They can easily be scraped o!! !or in"estigation, and then !or a sticky aterial. The wipes can "ery easily be re o"ed with a oist towel or paper. The water resistance see s low.

7igure $& ;ertical <ig<ag streaks of nasal gland secretion on the front glass panel of an enclosure taken out and laid flat5

,emperature dependent=
$n the last part o! their article, the authors state: @The o"erall analysis o! the 1D psa ophiids in"estigated in the present study support the hypothesis that their rubbing beha"iour depends on te perature E@ *p. 2A+. The background o! this hypothesis is o! course, that higher te peratures ean ore e"aporation and, as we know, the authors wanted to !ind a connection between e"aporation and !re?uency o! sel!>rubbing. /owe"er, the relation between te perature and sel!>rubbing !re?uency in their data is not as i pressi"e as they suggest. $n the original dissertation o! -2 (345 *2010+ ore details are gi"en. Tables ,.U and ,.H in that dissertation show that in one Scutophis *Rhagerhis+ moilensis there was a correlation, in three others notM in !i"e Psammophis mossambicus there was, in one notM in two Psammophis schokari there was a correlation, but in two other ones not. Though there was an o"er>all correlation, the outco e o! this eKperi ent re ains thus unclear, and the cases in which there was no correlation, should ha"e been eKplained. ;esides, other eKplanations would ha"e been possible, apart !ro the one $ entioned in the abo"e. $n general, !or instance, the ani als beco e ore acti"e when te perature rises. $t would be plausible that rubbing beha"iour would be part o! nor al acti"ity and beco e ore

!re?uent. $! one would want to !ind out i! the !re?uency increase o! rubbing beha"iour is independent o! a ore general increase o! acti"ity, ore data are needed.

-2 (345 B ;</M2 *201,+ state: @The o"erall analysis o! the 1D psa ophiids in"estigated in the present study support the hypothesis that their rubbing beha"iour depends on te perature and ser"es pri arily the pre"ention o! e"aporati"e water loss@*p. 2A+. 0s !or the !irst part o! that conclusion: that re ains to be in"estigated, and i! so, there cannot si ply be concluded that this has anything to do with e"aporation. 0s !or the second part o! the conclusion, $ honestly ask ysel! and the readers: where in the world is it based onS 8et us not !orget how "ery unlikely the water loss hypothesis is in the !irst place. 'uch a hypothesis would ask !or a uch stronger support than gi"en in the paper o! -2 (345 B ;</M2, in which $ !ind not e"en a shadow o! proo!. 6e ha"e seen that since 1)DA *uncare!ul+ reading o! the study o! -31'71, -31'71 B C2$T/ *1)DA+ has led authors to #u ping to the conclusion that the rubbing beha"iour o! psa ophiid snakes ser"es to pre"ent body water loss. The article o! -2 (345 B ;</M2 *201,+ see s to perpetuate this incorrect conclusion. $ hope y contribution has ade su!!iciently clear that this has to be considered a istake. The hypothesis o! -2 /001 *1))), 200,, c!. -2 /001 B C83C/$24 200H+ that the psa ophiid sel!>rubbing beha"iour @ser"es as a preli inary to subse?uent che ical arking o! conspeci!ics, nest and hunting routes@ *-2 /001 200H+ see s uch ore pro ising. -2 /001 see s con"inced that this is at least the case !or the 2astern and 6estern Montpellier snakes *resp. Malpolon insignitus and M. monspessulanus+, ost notably !or certain ales in May> Lune, but it could be hypothesised as a odel to eKplain the sel!>rubbing beha"iour o! the other e bers o! the psa ophiid group.

XXXXXX Fideos o! rubbing psa ophiids can be !ound on the internet, in the 5outube Channel o! Ton 'teehouder, JuserJTon1)4H2J!eaturedS"iewYasZpublic, or through the link http:JJ(sa ophis,nlJindeK!il . XXXXXX

08;24T', 088$'71 C. *1))2+: Constraints on the -esign o! Che ical Co unication 'yste s in Terrestrial Fertebrates. The 0 erican 1aturalist, Fol. 1,), 'upple ent: 'ensory -ri"e. -oes 'ensory -ri"e ;iology ;ias or Constrain the -irection o! 2"olutionS *Mar., 1))2+, pp. 'H2>'A). ;iodi"ersity o! the Va beIi ;asin 6etlands: (hase $$. 2000. (ublished by the Va beIi 'ociety in partnership with the ;iodi"ersity 9oundation !or 0!rica. This pro"ides a detailed e"aluation o! the biodi"ersity o! the Va beIi wetland areas. The !ull docu ent is published in two parts, containing !our "olu es. 4e!. !ro http:JJwww.Ia soc.orgJwp> contentJuploadsJ2012J0,J6etlands>(hase>2>Folu e>$$c>Chap>H>/erps>(art>2.pd! ;401C/, ;. *1)AA+: 9ield guide to the snakes and other reptiles o! 'outhern 0!rica. 'truik, Cape Town. ;401-'TTTT24, 9401C *1))U+: 2ine 4e"ision der :attung Psammophis it ;er[cksichtigung der 'chwestergattungen innerhalb der Tribus (sa ophiini *ColubridaeM 8ycodontinae+. Teil $$: 4asterelektronen ikroskopische 3ntersuchungen Iur 'chuppenultrastruktur bei den 0rten der Tribus (sa ophiini it besonderer ;er[cksichtigung der 0rten der :attung Psammophis. -issertation 3ni". des 'aarlandes, 'aarbr[cken. ;401-'TTTT24, 9. *1))H+: -ie 'andrennattern. N 'pektru 0kade ischer Ferlag, /eidelberg.

-042F'C5, $. '. *1)UH+: 7n the structure and !unction o! the nasal gland o! the liIard snake Malpolon monspessulanus */er .+ N Voological Lournal Moscow, ,U: ,12N,14 *in 4ussian+. :4$L', (. -2 *1A)A+: ;eobachtungen an 4eptilien in der :e!angenscha!t. Vool. :arten, ,): 2,,>24D -2 /001, C. *1)A2+: -escription du co porte ent de \!rotte ent] et notes sur la reproduction et la !onction aKillaire de la Couleu"re de Montpellier Malpolon monspessulanus. 4e ar?ues co parati"es a"ec Malpolon moilensis et Psammophis spp. N ;ulletin de la 'oci^t^ /erp^tologi?ue de 9rance, 2,: ,UN4). -2 (345, 'T.(/01$2 *2010+: 0nalysis o! the 4ubbing ;eha"iour o! (sa opphiids: 0 Methodological 0pproach. -issertation ;onn, 'epte ber 2010. 5ear o! publication: 2011. -2 (345, 'T.(/01$2 B 6789:01: ;</M2 *201,+: 0 contribution to the understanding o! the sel!>rubbing beha"iour in psa ophiid snakes *'?ua ata: (sa ophiidae+. > 'ala andra 4)*1+ 1AN,0. -31'71, 6. 0., M. C. -31'71 B 0. -. C2$T/ *1)DA+: The nasal gland o! the Montpellier 'nake Malpolon monspessulanus: 9ine structure, secretion co position, and a possible role in reduction o! der al water loss. N Lournal o! 2Kperi ental Voology, 20,: 4H1N4D4. -2 /001, C. *1)A2+: -escription du co porte ent de \!rotte ent ] et notes sur la reproduction et la !onction aKillaire de la Couleu"re de Montpellier Malpolon

monspessulanus. 4e ar?ues co parati"es a"ec Malpolon moilensis et Psammophis spp. N ;ulletin de la 'oci^t^ /erp^tologi?ue de 9rance, 2,: ,UN4). -2 /001, C. *1)))+: -ie 2urop_ische 2idechsennatter Malpolon monspessulanus */er an, 1A04+. N pp. HH1NDUH in: ;`h e, 6. *ed.+: /andbuch der 4eptilien und 0 phibien 2uropas, Fol ,J $$0, 'erpentes $$: Colubridae 2. N 6iebelshei *0380+, :er any. -2 /001, C. *200,+: 2Ktrabuccal in!ralabial secretion outlets in Dromophis, Mimophis and Psammophis species *'erpentes, Colubridae, (sa ophiini+. 0 probable substitute !or %sel!rubbing& and cloacal scent gland !unctions, and a cue !or a taKono ic account. N Co ptes 4endus de ;iologie, ,2H: 2DUN2AH. -2 /001, C. B 0. C83C/$24 *200H+: Che ical arking beha"iour in the (sa ophiine snakes Malpolon monspessulanus and Psammophis phillipsi. N (roceedings. o! the 1,th Congress o! 'ocietas 2uropea /erpetologica: 211N212. 80/0F, '. B 4. -M$G28 *1))H+: 'kin resistance to water loss in colubrid snakes: ecological and taKono ical correlations. N 2coscience, ,: 1,UN1,). 8$C/T, (038 01- ;2112TT, 08;24T 9. *1)D2+. 0 'caleless 'nake: Tests o! the 4ole o! 4eptilian 'cales in 6ater 8oss and /eat Trans!er. Copeia 1)D2, 1o. 4, -ece ber 2), pp. D02>D0D. 4$2((28, 7. *1)D,+: Vu %(utI"erhalten& "on Malpolon monspessulanus */er . 1A04+. N 0?uaterra, 10: 2,N24. '2TT82', :045 '. *200U+: 'ni!!ers: 9luid>-yna ic 'a pling !or 7l!actory Trace -etection in 1ature and /o eland 'ecurityaThe 2004 9ree an. 'cholar 8ecture. Lournal o! 9luids 2ngineering, M04C/ 200U, Fol. 12D, p. 1A)>21A. 'T22/73-24, 0.M. *1)A4+: 4epeated success!ul breeding o! the red striped sand snake Psammophis subtaeniatus sudanensis, and so e re arks on the =polishing beha"iour= o! this species. 8itt. 'erp., Fol. 4 *,J4+: )0>10, J -utch 2d.: )4>10A. 'T22/73-24, T. *1)AD+: 'o e eKperiences with the 'potted 'kaapsteker, Psammophylax rhombeatus. N 8itteratura 'erpentiu , D: 1DUN1A4. 'T22/73-24, T71 *1))1+: =7"er poetsen en #agen.= /et Terrariu A*D+:1,)>142.

'T22/73-24, T. *1))2+: 0 success!ul cross>breeding with Psammophis sibilans and Psammophis subtaeniatus in the terrariu . N 8iteratura 'erpentiu , 12: 1,4N144. 'T22/73-24, T71 *1))H+: 7"er het poetsen "an Iandrenslangen, hagedisslangen en skaapstekers. /et Terrariu 1UJH *dece ber 1))H+, p. 2)>,2. 628-71, (. L., ;. 980C/';04T/ B '. 'C/38V *200A+: 1atural products !ro nona"ian reptiles. N 1atural (roduct 4eports, 2U: D,ANDUH. 650TT, T4$'T40M -. *200,+: (hero ones and ani al beha"iour. Co taste. Ca bridge 3ni"ersity (ress. the integu ent o!

unication by s ell and