Dedication
Thiscontributionisrespectfullydedicatedtothefondmemoryof ProfessorShermanA.Minton,MadgeR.Minton,andProfessorAlan W.Bernheimer.Allwereinnatenaturalists,humble,talentedscientists, brilliantteachersandmentors,and,mostimportantly,generous, treasuredfriends.
ShermanA.andMadgeR.Minton,1985. Anoutstanding herpetologistofhistoricalimportance,Sherman(1919 99)wasa pioneerintwentieth-centuryresearchofvenomoussnakesandsnake venoms.Hehadaspecialinterestinthemedicalimportanceofnonfront-fangedsnakesandpioneeredthemoderninvestigationoftheir venoms.AhighlyrespectedfacultymemberintheDepartmentof MicrobiologyandImmunologyattheIndianaUniversitySchoolof Medicine,hewasacompassionatephysicianandaccomplished microbiologist/parasitologist.Madge(1920 2004)wasakeen herpetologist,pilot,ethnologist,andlapidary-gemologist.Sheserved intheWomenAirforceServicePilots(WASP)programduringWorld WarII,whileShermanwasanavalmedicalofficerontheUSS Brooks deployedontheCoralSea.Theirsharedintensepassionfor herpetologyandtoxinologyformedoneofthedeeplinksoftheir greaterthan50-yearlovingpartnership.Theirprofessional collaborationsproducedtwopopularbooksandmultiplescientific contributionsandalsocontributedsignificantlytotheconservationof theherpetologicalfaunaofIndiana.Shermanpublishedover150 papers,includingamajorcomprehensivemonographonthe herpetologyofPakistan,andco-authoredandeditedseveralbookson
snakebiteaswellasregionalherpetologyguides.Todate,hisbook, AmphibiansandReptilesofIndiana,isthemostcomprehensivestudy oftheherpetologyofthatstate.Hisengagingautobiographywas publishedposthumously(fordetailedinformationonShermanA. Minton’slifeandcareer,seeBechtel,1999;Stewart,2000;Karns, 2001;Weinstein,2003,andMinton,2001;photocopyrightto BrianMarianandScottA.Weinstein).
ShermanA.MintonandScottA.Weinstein(1984)discussingthe extractionofvenomsamplesfromapairofmidgetfaded rattlesnakes, Crotalusoreganusconcolor (photocopyrighttoBrianMarian).
GroupphotoattheFirstWorldCongressofHerpetology, Canterbury,UK(1989),afterthesymposiumonvenomoussnakes.
Picturedfromleft:HerbertRosenberg,DavidA.Warrell,ShermanA. Minton,DietrichMebs,JulianWhite,ElazarKochva,andDavidL. Hardy,Sr.(photocopyrighttoDavidA.Warrell).
AlanW.Bernheimer(dateunknown).Oneofthegreat microbiologistsofthetwentiethcentury,Alan(1914 2006)wasone ofthepioneersofmodernbacterialtoxinology.Stronglyinterestedin hemolytictoxins,hisresearchhadnoboundariesandencompassed thestudyoftoxinsfromnumerousmicro-organismsaswellas cnidarians(jellyfish,anemones),hymenopterans(especiallyants),and snakevenoms.HewastherecipientofaNationalInstitutesof Healthcareerawardandatwo-timerecipientoftheEliLillyAward, giventooutstandingmicrobiologistsfortheirnoteworthy accomplishments.Hecontributedover130scientificpapersand editedseveralbooks(seeLinder[2006]forfurtherinformationabout AlanW.Bernheimer’slifeandcareer;photocopyrighttoAlanW. Bernheimer,Jr.).
AlanW.Bernheimer(1967). AlanwasappointedProfessorof MicrobiologyatNewYorkUniversitySchoolofMedicinein1941and remainedProfessorEmeritusuntilhispassingin2006.Inadditionto hisinsightfulapproachtoresearch,hewasatalented,patientteacher andwaspopularamongthemedicalandgraduatestudentswho attendedhisinformation-packedmicrobiologylectures.Hewasalso aneruditelepidopterist(withaparticularinterestinSouthAmerican butterflies)andatalentedcreative/experimental(“reflectographs”) photographer.Hiswife,Harriet(1919 2009),wasanaccomplished microbiologistwhospecializedinthecharacterizationofthe biologicalrolesofthecapsularantigensof Streptococcuspneumoniae (photocopyrighttoAlanW.Bernheimer,Jr.).
Memoriam
Duringthelast2decades,theWorldhasverysadlylostseveral distinguishedherpetologistsandtoxinologistswhocontributed essentialworksaboutnon-front-fangedsnakes,theirglands,dentition, venoms,andevolutionarybiology.Theirlegacieswillcontinueto impactthefieldstowhichtheymadeenduringcontributions.
Prof.KennethV.Kardong(imagecopyrighttothelateKennethV.Kardong,andthanksto Prof.StephenP.Mackessy).
KennethV.Kardong(1943 2018) Abrilliantfunctional morphologist,herpetologist,andvertebrateanatomist,Ken’sethical anddedicatedapproachtoresearchwasmatchedbyhiswarmth, genuineness,andgenerosity.Ken’ssingularcontributionstothe functionalsignificanceoflow-pressureandhigh-pressurevenom glandscompriseonlyaportionofhisstudiesintothebiologyofnonfront-fangedsnakes.Hispassingisnotonlyaprofoundlosstoscience butalsothatofawarm,closefriend(fordetailedinformationabout Ken’slife,career,andreflectionsoffriendsandcolleagues,see Youngetal.,2019).
Prof.ElazarKochva(imagecopyrighttoProf.DavidA.Warrell).
ElazarKochva(1926 2018) Aninsightfulandthorough investigatorofvenomoussnakesandthefunctionalmorphologyofthe venomapparatus,ElazarKochvamethodicallycharacterizedkey aspectsofvenomdeliveryanditsevolutionamongdiverseophidian lineages.Hiscarefullydocumentedresearchisanindicatorofhis depthofcomprehensionespeciallyabouttheevolution,venom apparatus,andvenomsofunusualformssuchastheburrowingasps (Atractaspis spp.).ElazarKochvawasanimportantandinfluential memberoftheIsraeliscientificcommunity(forfurtherdetails aboutElazarKochva’slifeandaccomplishments, seeWeinsteinandWarrell,2019).
William(Bill)R.Branch(1946 2018) Fromthebeginningofhis scientifictraining,BillBranchseemedtobedestinedtobeaforcein Africanherpetology,adestinythatheclearlywascommittedto realize.HisinterestinAfricanherpetofaunadefinedhiscareerand resultedinhisswitchfromapost-doctoralcellbiologiststudying hepatocellularcarcinomatoarenownedherpetologistwho reinvigoratedAfricanherpetologicalresearch.Hehadatalentfor detectingthehiddenfeaturesofsecretiveAfricanfossorialsnakesand hadaparticularinterestinnon-front-fangedsnakes,aswellas unusualspeciessuchastheburrowingaspsormolevipers, Atractaspis spp.,andthequill-snoutedsnakes, Xenocalamus spp.Billwasawarm andgenerouscolleague(forfurtherinformationaboutBill’slifeand hisimportantcontributions,seeBauerandRo¨del,2019,Rogers,2018, andWeinsteinandWarrell,2019).
Dr.William(Bill)RoyBranch(imagecopyrighttoJohanMarais).
DonaldG.Broadley(1932 2016) Oneofthemosthistorically importantherpetologistswhospecializedinAfricanspecies,Donald Broadleyepitomizeddetailed,meticulousresearchandanalysisof Africanherpetofauna.Hisskillfulobservationsandcareinreporting hisvoluminousdatastandasanexampleofthesignificantdifference oneindividualcanmakewithsuchcarefulandmethodicalresearch.In heart-feltnarratives,closecolleaguesandfriendshavevividly describedDonaldBroadley’swarmthandamiablecollaborativespirit (foradditionalinformationaboutDonaldBroadley’slife,career,and hispersonalimpactonthelivesoffriends/colleagues,seeBranch, 1991,BranchandBates,2018).
Dr.DonaldBroadley(imagecopyrighttoJohanMarais).
Prof.GarthUnderwood(imagecopyrighttoProf.JudithGobinandtheDepartmentofLife Sciences,St.AugustineCampus,TheUniversityoftheWestIndies).
GarthUnderwood(1919 2002) Aprescientsquamatereptile systematicsbiologistanderuditeherpetologistwithacomprehensive perspective,GarthUnderwoodwasalsoakeenevolutionarybiologist andhighlyrespectedacademician.Hehadanearlyinterestinthe evolutionofsnakecephalicglandsandcontributedsomeofthemost commonlycited(tothisday)investigationsoftheseorgans.Afterhis passing,contributedmemorialsbycolleaguesandfriendshighlighted histalentasateacherandhispreviouslyunknownroleinBritish militaryintelligenceoperationsinWWII(forfurtherinformation aboutGarthUnderwood’slifeandcareer, seeThorpe,2003andBranch,2003).
Contents
Abouttheauthors ..................................................................................xix
Foreword.. .xxiii
Prefacetothesecondedition. ..................................................................xxv
Acknowledgmentsforthesecondedition xxix
Introduction... .................................................................................... xxxiii
CHAPTER1Asummaryoftaxingtaxonomicconsiderations: non-front-fangedsnakesandtheshiftingsands ofsystematics ........................................................1
1.1 Theshiftingsandsoftaxonomicreassignmentsandtheir relevancetotheenvenomedpatient .....................................16
1.2 Briefglimpsesofavastlandscape:thebasicsoftaxonomic reassignmentsanddiscordanceofmorphologicaland molecularsystematicsinvestigations... .................................24
1.3 Outlinedinventoryofarecenthigherphylogenyof Caenophidiansnakes .........................................................33 1.3.1Classification ..........................................................33
1.4 GeneralphylogeneticinventoryofCaenophidiansnakes: commentsandconsiderationofknownmedical significance. ....................................................................34
CHAPTER2Differencesbetweenbuccalglandsecretionand associatedvenomdeliverysystemsoffront-fanged snakesandnon-front-fangedsnakes:Low-versus high-pressureglandfunctionandcanaliculated versussoliddentition .............................................41
2.1 Basicconsiderationsregardingglandstructureand function ..........................................................................42
2.2 Overviewofhypothesesfortheevolutionofvenom-delivery systemsandotheradaptiveglandfunctions ...........................68
2.3 Theoriesconsideringtheevolutionofcanaliculatedfangs andenlargedgroovedteeth ................................................71
2.4 Low-pressure‘Duvernoy’sglands’andhigh-pressure venomglands:aquestionofsemantics? ...............................79
CHAPTER3Abriefoverviewofthehistoryofnon-front-fanged snakevenomresearch ...........................................81
3.1 Backgroundandbeginnings.. .............................................81
3.2 Themodernera:advancesandattributions............................85
3.3 Abriefoverviewofmajorclassesofnon-front-fanged colubroidvenomcomponents .............................................95
3.3.1Three-finger-foldtoxins(3-FFTXs).. .........................95
3.3.2Cysteine-richsecretoryproteins(CRiSPs) ..................97
3.3.3Snakevenommetalloproteases(SVMPs),matrix metalloproteases(MMPs),andsnakevenomserine proteases(SVSPs)..................................................98
3.3.4PhospholipasesA2 andB(PLA2andPLB).. .............102
3.3.5‘Other’components.. ............................................104
3.3.6Acommentonthehypothesizedassignmentofsnake venomtoxinstoclinicaleffectsofenvenoming: causalityandstrengthofassociation ........................105
3.3.7Illuminationofbiologicalfunction:prey-specificity andresearchtrends... ............................................130
3.3.8Immunologyofnon-front-fangedsnakevenoms. ........146
3.3.9Commercialantivenomsagainstmedically importantelapidsandviperids:possibleusefor envenomingbynon-front-fangedsnakes;utilityor futility?... ............................................................149
3.3.10Futureresearch.....................................................152 CHAPTER4Medicallysignificantbitesbynon-front-fanged snakes(NFFCs) ....................................................155
4.1 Typicalfeaturesofdocumentedcasesandevidence-based risk ..............................................................................160
4.1.1Publishedandcommunicatedormanagedcasesof medicallysignificantnon-front-fangedcolubroid bites:summariesandstrengthofevidence ..................161
4.1.2Epidemiologyofbitesfromnon-front-fangedsnakes....320
4.1.3Circumstancesassociatedwithspeciescapableof inflictinglife-threateningenvenoming ........................345
4.2 Somerepresentativegenera:typicalfeaturesofbitesand anoverviewoftheirnaturalhistoryandtoxinology.. ............347 Colubridae,Ahaetuliinae. .................................................347
4.2.1Genus Chrysopelea (Boie,1826):backgroundand generalfeaturesofdocumentedbites. ........................347
Colubridae,Colubrinae ....................................................350
4.2.2Genera Boiga (Fitzinger,1826),and Toxicodryas (Hallowell,1857):backgroundandgeneralfeatures ofdocumentedbites. ...............................................350
4.2.3Genera Hemorrhois (Boie,1826), Platyceps (Blyth,1860), Hierophis (Fitzinger in Bonaparte,1834), Dolichophis (Gistel,1868),and Coluber (Linnaeus,1758):backgroundandgeneralfeatures ofdocumentedbites. ...............................................363
Dipsadidae ....................................................................369
4.2.4Genus Philodryas (Wagler,1830):backgroundand generalfeaturesofdocumentedbites... ......................369
4.2.5Genus Heterodon (Latreille,1801):backgroundand generalfeaturesofdocumentedbites... ......................385
4.2.6Genera Borikenophis and Cubophis (Hedgesetal.,2009):backgroundandgeneral featuresofdocumentedbites ....................................395
Natricidae.. ....................................................................401
4.2.7Genus Thamnophis (Fitzinger,1843)andcomments ongenus Nerodia (BairdandGirard,1853): backgroundandgeneralfeaturesofdocumented bites. ....................................................................401
Psammophiidae ..............................................................407
4.2.8Genus Psammophis (Boie,1825):backgroundand generalfeaturesofdocumentedbites... ......................407
4.2.9Summarizedconsiderationofrepresentative specieswithlimitedevidence... ................................410
4.3 Life-threateningandfatalcases:non-front-fangedsnakes withlethalpotentialandassessmentof evidence-basedrisk. ........................................................417
4.3.1 Dispholidustypus (Smith,1828), Thelotornis capensis (Smith,1849),and Thelotorniskirtlandii (Hallowell,1844);Colubridae,Colubrinae: backgroundandgeneralfeaturesofdocumented bites. ....................................................................417
4.3.2 Rhabdophissubminiatus (Schlegel,1837), Rhabdophistigrinus (Boie,1826),and Rhabdophis ceylonensis (Gunther,1858);Natricidae:background andgeneralfeaturesofdocumentedbites.. .................435
4.3.3Comparisonofmajorfeaturesofhazardlevel1 non-front-fangedsnakes(Dispholidustypus, Thelotornis capensis, Thelotorniskirtlandii, Rhabdophistigrinus, Rhabdophissubminiatus,and Rhabdophis ceylonensis)...........................................................457
4.3.4Guiltywithoutatrial:threeadditionalunsupported reportsoffataloutcomesafterbitesby Tachymenis peruviana (Wiegmann,1834)(Dipsadidae), Oligodon arnensis (Shaw,1802)(Colubridae,Colubrinae),and Xenodonseverus (Linnaeus,1758)(Dipsadidae).. ........458
4.4 Aberrantcasesandrepresentativecaseswithoutclear etiology:acriticalassessmentofrisk. ................................464
4.4.1 Boigairregularis (Bechstein,1802)(Colubridae, Colubrinae) ...........................................................464
4.4.2 Malpolonmonspessulanus (Hermann,1804) (Psammophiidae)... .................................................475
4.4.3 Hydrodynastesgigas (Dume ´ riletal.,1854), Dipsadidae ............................................................481
4.4.4 Philodryasolfersiilatirostris (Cope,1863), Dipsadidae ............................................................486
4.4.5 Platycepsnajadum (Eichwald,1831)[Colubridae, Colubrinae]:afatalcaseofprogressiveneuropathy fromacolubridbite?.. ............................................492
4.4.6 Hierophisviridiflavus (Lace ´ pe ` de,1789)[Colubridae, Colubrinae]:acaseofneurotoxiccolubrinebiteor amorecommonetiology?. .......................................497
4.4.7“Venomous”bitesbynon-venomouslizards?. .............503
4.5 Pitfallsnotedindocumentedcases:Differentiating PerceivedversusEvidence-BasedRisk ...............................528
4.5.1Examplesofpitfallsnotedindocumentedcases.. ........528
4.5.2Perceivedversusevidence-basedrisk:human responsetotraumaandsomatosensory amplification... ......................................................546
4.6 Recommendationsformanagementofmedically significantnon-front-fangedsnakebites... ...........................550
4.6.1General.................................................................550
4.6.2Specific ................................................................552
CHAPTER5Summaryandconclusions
AppendixARepresentativeunverifiedcasesofmedicallysignificant non-front-fangedsnakebitespostedontheinternet ...................601
AppendixBRepresentativelethalpotencyrangesandyieldsofvenoms andotheroralproductsfromselectednon-front-fanged colubroidsnakes ..................................................................609
AppendixCStrategiesformanagementofgram-negativesepticemia: aretherelessonstobelearnedformanagingvenom-induced coagulopathies?. ..................................................................615
AppendixDLegalconsiderationsregardingprivateownershipof venomoussnakes(includinghazardlevel1non-front fangedsnakes):anopinionatedessay... ...................................619
AppendixEListofexaminedosteologicalspecimens... ..............................623
Additionalrecommendedreading .............................................................625
Bibliography.. .......................................................................................633
Glossary... ............................................................................................719
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Abouttheauthors
SCOTTA.WEINSTEIN,BA,MSc,PhD,MBBS,MD,DIP,ABFM, FAAFP
ScottWeinsteinwasconsumedfromearliestchildhoodbyaninterestinreptilesand amphibians.AmemberoftheNewYorkHerpetologicalSocietybyage11,hewas oneofaluckycadreofyoungmemberswhoweregentlymentoredbythe well-knowncuratoroftheStatenIslandZoo,thelateCarlKauffeld.Hisstudies of“rear-fangedcolubrids”startedinJuniorHighSchoolwherehestartedtocompare thesparseinformationintheavailableherpetologyliteraturewithcollectedliving specimens.Theseinterestsrapidlyfocusedonthebiologyofvenomoussnakes andherpetologicaltoxinology.ThisledtohisstudiesinherpetologywithProf. EdmundD.Brodie,Jr.atAdelphiUniversity(GardenCity,NY),whereheearned hisBAinbiologicalsciencesandcomparativereligion.Shortlythereafter,he becamethelateProf.ShermanA.Minton’slaststudentattheIndianaUniversity SchoolofMedicine,whereheearnedanMScinMedicalMicrobiologyandImmunology.HisPhDinMedicalMicrobiologyandImmunologywasearnedwiththelate Prof.AlanW.BernheimerattheSacklerInstituteofGraduateBiomedicalSciences ofNewYorkUniversitySchoolofMedicine(NewYork,NY),hisMBBSwasearned atFlindersUniversitySchoolofMedicine,Adelaide,Australia,andtheBoardof RegentsoftheUniversityoftheStateofNewYorkconferredhisMD.Hecompleted familymedicineresidencyatSouthNassauCommunitiesHospital,Oceanside,NY andservedaschiefresident.Healsocompletedtwopostdoctoralresearch
fellowships:oneatNYUMedicalCenterandtheotherattheUSArmyMedical ResearchInstituteofInfectiousDiseases(FortDetrick,Frederick,MD).His researchhasincludedpurificationandcharacterizationofnovelsnakevenomneurotoxins;elucidationofcomponentsandthebiomedicalpropertiesofvenomsand othersecretionsfromnon-front-fangedcolubroids;venomresistanceinophiophagousnon-venomoussnakes;andfieldstudiesofreptilesandamphibians.Herecently servedasaclinicaltoxinologistattheWomen’sandChildren’sHospitalinAdelaide, SouthAustralia,wherehewasaconsultantinthemanagementofenvenomationsand practicestraditionalfamilymedicineinhisnativeNewYorkCityandintermittently inthenorthernsuburbsofAdelaide.Inadditiontohislifelonginterestsintoxinology andherpetology,Dr.Weinsteinhasastronginterestinthemedicalmanagement ofspecial-needspopulations(e.g.,thosewithdisabilities),infectiousdiseases (especiallyacademicandclinicalvenereology),andsubstancedependencymedicine. Hestillparticipatesinfieldherpetologywheneverpossibleandhasalwaysremained activeinthecaptivepropagationofawidevarietyoflivingspecimens.Asiscommon withmanyherpetologistsandtoxinologists,healsohastraveledextensively.Hehas contributedmorethan100peer-reviewedpublicationsintoxinology,herpetology, andmedicine,aswellassixbooks.
DAVIDA.WARRELL,MA,DM,DSc,FRCP,FRCPE,HONFZS, FMEDSCI
DavidAlanWarrellisEmeritusProfessorofTropicalMedicineandHonorary FellowofStCrossCollege,UniversityofOxford,UK.AftertrainingatOxford, StThomas’Hospital,andtheRoyalPostgraduateMedicalSchoolinLondon,he lived,worked,researched,andtraveledinEthiopia,Nigeria,Kenya,SouthAfrica,
Thailand,Burma,SriLanka,Bangladesh,PapuaNewGuinea,Brazil,Ecuador,Peru, theCaribbean,andinothertropicalcountries,foundingtheOxfordUniversity-based TropicalMedicineResearchProgramme,whoseunitsareinThailand(since1979) andelsewheretostudymalariaandothermajortropicaldiseases.HebecametheDirectoroftheOxfordTropicalNetworkin1986andlater,headofTheNuffield DepartmentofClinicalMedicine,UniversityofOxford.HehasbeentheDelegate forMedicineandMusicattheOxfordUniversityPressandwassenioreditorof the OxfordTextbookofMedicine (fifthedition,2010)aswellasthe OxfordHandbook ofExpeditionandWildernessMedicine (firstedition,2008).Hehaspublishedmore than500researchpapers,articles,reviews,andtextbookchaptersonmalaria,rabies, relapsingfevers,meningococcalmeningitis,cryptococcalmeningitis,HIV,other tropicalandinfectiousdiseases,comparativerespiratoryphysiology,respiratorydiseases,herpetology,venomousanimals,envenoming,andplantandchemical poisoning.HeisaconsultanttotheWorldHealthOrganization(WHO)onsnakebite, rabies,andmalaria,theBritishArmy,theUKMedicalResearchCouncil,theForeign andCommonwealthOffice,theEarthWatchInternational(conservation),the ZoologicalSocietyofLondon,theRoyalGeographicalSociety,andtheToxBase UK.HealsoservedasthepastpresidentoftheRoyalSocietyofTropicalMedicine andHygieneandtheInternationalFederationforTropicalMedicine.Hisprincipal researchinterestremainsthepathophysiologyandtreatmentofenvenoming.In 2019,hewasawardedtheprestigiousSirPatrickMansonMedalbytheRoyalSociety ofTropicalMedicineandHygiene.
DANIELE.KEYLER,BS,BSPharmacy,PharmD,FAACT
Danhashadaninterestinvenomoussnakessincehisearlyyearsofgrowingupalong theTippecanoeRiverinIndiana.Thisenthusiasmwasfurtherintensifiedduringhis highschoolyearswhenhecaredforthesnakecollectionoftheeminentherpetologist,
ProfessorShermanA.Minton,Jr.,MD,whilehewaslivinginPakistanwithhisfamily. HemaintainedarelationshipwithProf.Mintonviaenvenomationconsultationsuntil hispassingin1999.Dan’sinterestcontinuedtoexpandintotherealmoftoxicology, andhehascompleteduniversitycoursesinEnvironmentalToxicology,Industrial Toxicology,VeterinaryToxicology,ClinicalToxicology,GeneralToxicology, AdvancedToxicology,andResearchToxicology.HeholdsaBSdegreefromPurdue UniversityinScienceandaBSinPharmacyandDoctorofPharmacydegreesfrom theUniversityofMinnesota.Continuedinvolvementwiththedisciplineoftoxicology followedasDanwasontheMedicalFaculty,DivisionofClinicalPharmacologyand Toxicology,DepartmentofMedicine,HennepinCountyMedicalCenterandtheCoDirectorofToxicologyResearchwiththeMinneapolisMedicalResearchFoundation 1985 2011.HeservedasPresidentoftheNorthAmericanSocietyofToxinology 2016 18andisaFellowoftheAmericanAcademyofClinicalToxicologyand Professor,DepartmentofExperimental&ClinicalPharmacology,Universityof Minnesota,CollegeofPharmacy(wherehedevelopedthefirstcourseinClinical Toxinology),andwasrecipientofthedistinguishedMelendyLectureawardin2010. Danhasconsultedonhundredsofsnakebitecasesandhasobjectivelycontributedto thefieldsoftoxicologyandtoxinologybyhavingauthored/coauthoredover100scientificpublicationsand45bookchaptersinvolvingimmunotherapeutics(antibodyand vaccinedevelopment),immunotoxicology,toxicology,animaltoxins,andvenomous snakebites.Hehasbeenacoauthorandeditorfortwobooks,thefirsteditionofthis book, VenomousBitesfromNon-VenomousSnakes:ACriticalAnalysisofRiskManagementofColubridSnakeBites,and VenomousSnakebiteintheWesternUnited States.HeservedasChairoftheEnvenomationSectionwiththeAmericanAcademy ofClinicalToxicology2002 07andisamemberoftheMedicalAdvisoryCommittee fortheOnlineAntivenomIndex.Heisareviewerformultiplemedicalandscientific journals,consultanttomultiplezoos,andamemberoftheInternationalSocietyon Toxinology,theAmericanAcademyofClinicalToxicology,theNorthAmericanSocietyofToxinology,theSocietyfortheStudyofAmphibiansandReptiles,andthe Chicago,Indiana,andMinnesotaHerpetologicalSocieties.Additionally,heserves asaConsultingSeniorClinicalToxicologisttoSafetyCallInternational. Prof.KeylerhashadalifelongpassionforTimberRattlesnakes(Crotalushorridus) andhashadmultipleresearchgrantstostudythisspeciesintheUpperMississippi RiverValley.Danisafoundingmemberofacommitteeinvolvedinthedevelopment ofanationalConservationActionPlanregarding C.horridus andhasanongoing interestinthenaturalhistory,conservation,biology,andvenomof C.horridus. Prof.Keylercontinueshisenjoymentoffieldworkwithvenomoussnakespeciesin temperateandtropicalgeographicregionsoftheworld.
Foreword
Prof.DietrichMebs InstituteofLegalMedicine, GoetheUniversityofFrankfurt,Frankfurt,Germany
WhenErikaSchirner,thesecretaryofProf.RobertMertens,calledme:“TheProfessorwasbittenbya Thelotorniskirtlandii”(Kirtland’stwigorvinesnake),Iwas scared.IknewD.G.Broadley’spaper(1957)onfatalbitesof Dispholidus (boomslang) and Thelotornis, andMertens,oneoftheleadingherpetologistsandformerDirectorof thefamousSenckenbergMuseuminFrankfurt,alsomusthavebeenawareofit.I rushedtotheMuseum,whereIwasgreetedbyMertens:“Don’tworry,it’snotserious. Ihavebeenbittenbythatsnakeseveralmonthsagoandnoenvenomingsymptomshad occurred.”Andsmilingheadded:“Isupposeyouarejustinterestedinwritingacase report.”
Mertenskeptanexquisitecollectionofamphibiansandreptilesinhismagnificentvivariumathomeand,asusual,intheearlymorningofAugust5,1975,he wasfeedinghisherps.Whenheofferedasmalllizardtoa Thelotorniskirtlandii of120cmbodylength,thesnakemissedthelizardandbitthethumbofMertens’ righthand.Heremovedthesnakeimmediatelyandputitbackintheterrarium. Tworowsofslightlybleedingteethandfangmarkswereseenonhisthumb.
ItwasawastedefforttoconvinceMertenstocomewithmetotheUniversity Hospitalforacheck-up.BeforeIleft,Iurgentlyaskedhimtocallmewhenever heobservesanysymptoms.
Atabout4p.m.,Mertenscalledmefromhishome:“Bloodisstilloozingfrom theteethmarks.”Iimmediatelywenttohishome,metastillrelaxedMertens,and drovewithhimtotheUniversityHospital,wherehewasadmittedtotheintensive careunit.Firstlaboratorytestsrevealedafibrinogenemiaandmassivefibrinolysis. Dispholidustypus-antivenomwasathandprovidedbytheFrankfurtZoobutwas notadministeredafteracalltoSAIMR(South-AfricanInstituteofMedical Research)confirmingthattheantivenomshowsnocross-reactivitywith Thelotornis venom.
Thefollowingday,Mertens’conditiondeteriorated,helostconsciousnessand after18daysofunsuccessfulattemptstostopbleedingandtoreverseorganfailure, hedied.Hewas80yearsold.
Thissadepisodeprovidesanimpressiveexampleofthelargelyunderestimated toxicandevenlethalpotencythatvenomsfromnon-front-fangedandso-called “non-venomous”snakes(whichinfactmanyarevenomous)maypossess.Their experimentallethalpotencyand,insomecases,clinicaleffectsmaycompetewith venomsfromfront-fanged,“real”venomoussnakessuchasfromRussell’s(Daboia russelii)andsaw-scaledvipers(Echiscarinatus).Exceptfor D.typus and T.kirtlandii.whichattractedtheinterestoftoxinologists,becausetheirbitecauses
life-threateningsymptoms,othermembersofthelargeColubridaesnakefamily wereconsideredtobeofminorimportance,andmostoftheireffectswereclassified as“mild.”However,byapplyingmodernbiochemicaltechniques,studiesonthese venomsorglandextractsledtosurprisingdiscoveries,suchastheidentification of“three-finger”neurotoxinsintheDuvernoy’sglandsecretionofthemangrove snake Boigadendrophila.Thesetoxinsarecloselyrelatedtothosepresentinvenoms ofcobras(Naja spp.)andkraits(Bungarus spp.).
Thefirsteditionofthisbook,publishedin2011wasanimportantstepforwardto drawattentiontothe“non-venomoussnakes,”notjustthroughsummarizingcurrent knowledge,butalsobycriticallyevaluatingcasereports,definitions(themeaningof “toxiceffects”),etc.Thenewupdatededitioncoveringrecentdevelopmentsand providingnewdataandinformationinthatfieldishighlywelcomeandwillserve asanimportantreferencebookforanyone,toxinologists,clinicians,and naturalistsaswell.Chapeautotheauthors,ScottWeinstein,DavidWarrelland DanielKeyler,forcompletingthatmission!
Prof.DietrichMebs,PhD
FranfurtamMain
February2020
