| 词条 | Timber rattlesnake |
| 释义 |
| name = Timber rattlesnake | image = ZollmanTimberRattlesnake.jpg | image_caption = At Zollman Zoo | status = LC | status_system = IUCN3.1 | status_ref = [1] | genus = Crotalus | species = horridus | authority = Linnaeus, 1758 | range_map = | synonyms = * Crotalus horridus Linnaeus, 1758
}} The timber rattlesnake, canebrake rattlesnake or banded rattlesnake (Crotalus horridus),[3] is a species of venomous pit viper endemic to the eastern United States. This is the only rattlesnake species in most of the populous northeastern United States and is second only to its cousins to the west, the prairie rattlesnake, as the most northerly distributed venomous snake in North America.[4][5] No subspecies are currently recognized.[6] TaxonomyThe timber rattlesnake was one of the many reptile species originally described by Linnaeus in the landmark 1758 10th edition of his Systema Naturae, and still bears its original name Crotalus horridus.[7] The subspecies C. h. atricaudatus (Latreille in Sonnini and Latreille, 1802), often referred to as the canebrake rattlesnake,[3] is currently considered invalid.[8] Previously, it was recognized by Gloyd (1936) and Klauber (1936). Based on an analysis of geographic variation, Pisani et al. (1972) concluded no subspecies should be recognized. This was rejected by Conant (1975), but followed by Collins and Knight (1980). Brown and Ernst (1986) found evidence for retaining the two subspecies, but state it is not possible to tell them apart without having more information than usual, including adult size, color pattern, the number of dorsal scale rows and the number of ventral scales. Dundee and Rossman (1989) recognized C. h. atricaudatus, but others take a more neutral point of view.[10] DescriptionAdults usually grow to total length of {{convert|91|–|152|cm|in|round=0.5|abbr=on}}.[4] It was found in Pennsylvania that the smallest size females that could produce viable eggs was {{convert|72.2|cm|in|abbr=on}}.[9] Most adult timber rattlesnakes found measure less than {{convert|100|to|115|cm|in|abbr=on}} in total length and weigh on average between {{convert|500|and|1500|g|lb|abbr=on}}, often being towards the lower end of that weight range.[10][11][12][13] The maximum reported total length is {{convert|189.2|cm|in|abbr=on}} (Klauber, 1956). Holt (1924) mentions a large specimen caught in Montgomery County, Alabama, which had a total length of {{convert|159|cm|in|round=0.5|abbr=on}} and weighed {{convert|2.5|kg|lb|abbr=on}}.[14] Large specimens can reportedly weigh as much as {{convert|4.5|kg|lb|abbr=on}}.[15] The dorsal scales are keeled[16] and arranged in 21–26 scale rows at midbody (usually 25 rows in the southern part of its geographic range, and 23 rows in the northern part). The ventral scales number 158-177 in males and 163–183 in females. Males have 20–30 subcaudal scales, while females have 15–26. The rostral scale is normally a little higher than it is wide. In the internasal-prefrontal area there are 4–22 scales that include 2 large, triangular internasal scales that border the rostral, followed by 2 large, quadrangular prefrontal scales (anterior canthals) that may contact each other along the midline, or may be separated by many small scales. Between the supraocular and internasal, only a single canthal scale is present. There are 5–7 intersupraocular scales. The number of prefoveal scales varies between 2 and 8. Usually the first supralabial scale is in broad contact with the prenasal scale, although slightly to moderately separated along its posteroventral margin by the most anterior prefoveals.[14] Dorsally, they have a pattern of dark brown or black crossbands on a yellowish brown or grayish background. The crossbands have irregular zig-zag edges, and may be V-shaped or M-shaped. Often a rust-colored vertebral stripe is present. Ventrally they are yellowish, uniform or marked with black.[17] Melanism is common, and some individuals are very dark, almost solid black.[18]Distribution and habitatFound in the eastern United States from southern Minnesota and southern New Hampshire, south to east Texas and north Florida.[19] One hundred and fifteen rattlesnakes have been marked within Brown County State Park in Indiana, one of the only places where they can be found in the state.[20] Its historic range includes southern Ontario and southern Quebec in Canada,[2] but in May 2001, the Canadian Species at Risk Act listed it as extirpated in Canada.[21] A Canadian government sponsored recovery strategy is currently under study to support the reintroducing of this predator of many pests to its former Canadian habitat. Although several experts disagree, many were found in some of the thick forest areas of central and southeastern Iowa, mostly within the Mississippi, Skunk, Iowa, and Des Moines River valleys, in several places in these areas; bites from timber rattlesnakes have been widespread, especially in a localized area of Geode State Park, in southeastern Henry County, along Credit Island Park, in southern Scott County, and in the forested areas of southern Clinton County.{{Citation needed|reason=Cite source of assertion|date=September 2009}} In Pennsylvania, it is not found west of Chestnut Ridge, which is in the Laurel Highlands, nor is it found in the southeastern corner of the state. Thus, its range does not include the areas of Philadelphia and Pittsburgh, the two largest cities in Pennsylvania.[4] C. horridus is extirpated in Maine and Rhode Island and is almost extirpated in New Hampshire. In Massachusetts, the snakes are active from mid-May to mid-October.[22] Early settlers were afraid of the snake, as its population was widespread throughout the state. Since that time their habitat has been reduced to the Blue Hills south of Boston, The Berkshires in Western Massachusetts as well as parts of the Connecticut River Valley, notably in the area of the Holyoke Range.[22] The snake is so rare in the state that it is rarely encountered by people and is considered endangered, making it illegal to harass, kill, collect, or possess.[23] Generally, this species is found in deciduous forests in rugged terrain. During the summer, gravid (pregnant) females seem to prefer open, rocky ledges where the temperatures are higher, while males and nongravid females tend to spend more time in cooler, denser woodland with more closed forest canopy.[24] BehaviorFemale timber rattlers often bask in the sun before giving birth, in open rocky areas known as "basking knolls".[25] During the winter, timber rattlesnakes brumate in dens, in limestone crevices, often together with copperheads and black rat snakes.[18] FeedingTheir prey are mainly small mammals, but may include small birds, frogs, other small animals, including other snakes. Although capable of consuming other rattlesnakes, the most common snake they prey upon are garter snakes.[24] Like most rattlesnakes, timber rattlesnakes are known to utilize chemical cues to find sites to ambush their prey and will often strike their prey and track them until they can be consumed.[26][27] Timber rattlesnakes are known to use fallen logs as a waiting site for prey to pass by, giving them an elevated perch from which to effectively strike their prey, which is almost entirely terrestrial rather than arboreal (even arboreal prey such as squirrels tends to be caught when they come to the ground).[27][28] The primary foods by genera of timber rattlesnakes were as follows: Peromyscus (33.3%), Microtus (10.9%), Tamias (qv) (10.6%), Sylvilagus (10.4%), Sigmodon (5.3%) and Sciurus (4.2%). Based on examination of the snout-to-vent length, it was found that juvenile timber rattlesnakes differed slightly in dietary preferences from adult rattlesnakes, being more likely to consume smaller prey such as shrews (averaging {{convert|8|g|oz|abbr=on}} and unable to attack subadult eastern cottontail rabbits (averaging {{convert|500|-|1000|g|lb|abbr=on}} but Peromyscus was the number one prey item for both young and adult rattlesnakes. Several birds, although always secondary to mammals, are also known to be hunted, mainly ground-dwelling species such as bobwhites, but also a surprising number of passerines.[29] VenomPotentially, this is one of North America's most dangerous snakes, due to its long fangs, impressive size, and high venom yield. This is to some degree offset by its relatively mild disposition[30] and long brumation period. Before striking, they often perform a good deal of preliminary rattling and feinting.[31] Cist (1845) described how he lived in western Pennsylvania for many years, and the species was quite common there, but in all that time, he heard of only a single death resulting from its bite.[3] Considerable geographic and ontogenetic variation occurs regarding the toxicity of the venom, which can be said for many rattlesnake species. Four venom patterns have been described for this species: Type A is largely neurotoxic, and is found in various parts of the southern range. One effect of the toxin can be generalized myokymia.[32] Type B is hemorrhagic and proteolytic, and is found consistently in the north and in parts of the southeast. Type A + B is found in areas where the aforementioned types apparently intergrade in southwestern Arkansas and northern Louisiana. Type C venom has none of the above components and is relatively weak.[30] The neurotoxic component of the type A venom is referred to as canebrake toxin, and is a phospholipase A2. It is analogous to the neurotoxins found in the venoms of several other rattlesnake species, and when present, contributes significantly to the overall toxicity. Other components found in the venom include a small basic peptide that works as a myotoxin, a fibrinogen-clotting enzyme that can produce defibrination syndrome, and a bradykinin-releasing enzyme.[30] CroFab antivenom, while not specific for C. horridus, is used to treat envenomations from this species.[33]SymbolThe timber rattlesnake was designated the state reptile of West Virginia in 2008.[34] That state's legislature praised "...a proud contribution by the eighth grade class at Romney Middle School, from West Virginia's oldest county, in West Virginia's oldest town, to have been instrumental in making the timber rattlesnake the state reptile..."[35] This snake became a prominent symbol of American anger and resolve during the American Revolution due to its fearsome reputation. In the 18th century, European-trained doctors and scientists had little firsthand experience with or information on timber rattlesnakes,[36] and treatment of their bites was poorly effective. The motto Nemo me impune lacesset (with the verb in the future tense) appears above a Crotalus horridus on a 1778 $20 bill from Georgia as an early example of the colonial use of the coiled rattlesnake symbol, which later became famous on the Gadsden flag. Conservation statusThis species is classified as least concern on the IUCN Red List (assessed in 2007).[1] Species are listed as such due to their wide distribution, presumed large population, or because they are unlikely to be declining fast enough to qualify for listing in a more threatened category.[37] The timber rattlesnake is listed as endangered in New Jersey, Vermont, Massachusetts[38] (along with the copperhead viper), Virginia, New Hampshire, Indiana,[39] and Ohio, and it is threatened in New York, Connecticut, Illinois, Minnesota, and Texas. {{Citation needed|date=May 2011}} Timber Rattlesnakes have already gone extinct in Maine and Rhode Island and only one population remains in New Hampshire. They are protected in many of the Appalachian states but their populations continue to decline.[40] See also
References1. ^1 Hammerson, G.A. (2007). Crotalus horridus. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. 2. ^1 McDiarmid RW, Campbell JA, Touré T. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Herpetologists' League. {{ISBN|1-893777-00-6}} (series). {{ISBN|1-893777-01-4}} (volume). 3. ^1 2 Wright AH, Wright AA. 1957. Handbook of Snakes of the United States and Canada. Comstock Publishing Associates. Ithaca and London. (7th printing, 1985). {{ISBN|0-8014-0463-0}}. (Crotalus horridus, pp. 956–966.) 4. ^1 2 Conant R. 1975. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Second Edition. First published in 1958. Houghton Mifflin Company. Boston. {{ISBN|0-395-19979-4}}. (Crotalus horridus, pp. 233–235 + Plate 35 + Map 178.) 5. ^Brown, W. S. (1991). Female reproductive ecology in a northern population of the timber rattlesnake, Crotalus horridus. Herpetologica, 101-115. 6. ^{{ITIS |id=174306 |taxon=Crotalus horridus |accessdate=8 February 2007}} 7. ^{{cite book |last= Linnaeus |first=Carl |authorlink= Carl Linnaeus |title= Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis |publisher= (Laurentii Salvii) |location= Holmiae |volume= Vol. I |edition= 10th revised |language= Latin |year= 1758 |page= 280 |url= https://archive.org/stream/carolilinnaeisy00gesegoog#page/n223/mode/1up |via= The Internet Archive}} 8. ^{{ITIS |id=174307 |taxon=Crotalus horridus atricaudatus |accessdate=27 September 2006}} 9. ^Galligan, J. H., & Dunson, W. A. (1979). Biology and status of timber rattlesnake (Crotalus horridus) populations in Pennsylvania. Biological Conservation, 15(1), 13-58. 10. ^Timber Rattlesnake (Crotalus horridus). Tpwd.state.tx.us. Retrieved on 2013-01-05. 11. ^Fitch, H. S., Pisani, G. R., & Greene, H. W. (2004). A FIELD STUDY OF THE TIMBER RATTLESNAKE lN LEAVENWORTH COUNTY, KANSAS. Journal of Kansas Herpetology Number, 1, 18. 12. ^Brown, W. S., Kéry, M., & Hines, J. E. (2007). Survival of timber rattlesnakes (Crotalus horridus) estimated by capture-recapture models in relation to age, sex, color morph, time, and birthplace. Copeia, 2007(3), 656-671. 13. ^Clark, R. W. (2006). Fixed videography to study predation behavior of an ambush foraging snake, Crotalus horridus. Copeia, 2006(2), 181-187. 14. ^1 2 {{cite book|authors=Campbell JA & Lamar WW|title=The Venomous Reptiles of the Western Hemisphere (2 volumes)|publisher=Comstock Publishing Associates|year=2004|isbn=0-8014-4141-2}} {{Page needed|date=May 2011}} 15. ^ANIMAL BYTES – Canebrake Rattlesnake. Seaworld.org. Retrieved on 2013-01-05. 16. ^{{cite book|authors=Behler JL & King FW|year=1979|title=The Audubon Society Field Guide to North American Reptiles and Amphibians|publisher=New York: Alfred A. Knopf. 743 pp.|isbn=0-394-50824-6}} (Crotalus horridus, pp. 688-689 + Plates 619, 620, 653.) 17. ^Boulenger, G.A.. 1896. Catalogue of the Snakes in the British Museum (Natural History), Volume III., Containing the...Viperidæ. Trustees of the British Museum (Natural History). (Taylor and Francis, Printers.) London. xiv + 727 pp. + Plates I.- XXV. (Crotalus horridus, pp. 578–580.) 18. ^1 Schmidt, K.P., and D.D. Davis. 1941. Field Book of Snakes of the United States and Canada. G.P. Putnam's Sons. New York. 365 pp. (Crotalus horridus horridus, pp. 301-302 + Plate 33; Crotalus horridus atricaudatus, p. 302.) 19. ^Conant, Roger & Collins, Joseph T. (1998). Reptiles and Amphibians of Eastern/Central North America. New York: Houghton Mifflin. {{ISBN|0-395-90452-8}} 20. ^{{cite web|url=https://www.in.gov/dnr/parklake/files/sp-Timber_brown-RattlesnakesFAQ2.pdf|title=The Timber Rattlesnake (Crotalus horridus) in Brown County State Park|publisher=Indiana Department of Natural Resources|accessdate=August 8, 2017}} 21. ^Crotalus horridus at Species at Risk Public Registry. Accessed 23 June 2008. 22. ^1 {{cite web|title=Timber Rattlesnake: Crotalus Horridus|url=http://www.mass.gov/eea/docs/dfg/nhesp/species-and-conservation/nhfacts/crotalus-horridus.pdf|website=Massachusetts Division of Fisheries and Wildlife}} 23. ^{{cite web|title=Timber Rattlesnake|url=http://www.masnakes.org/snakes/timber_rattle/|website=Snakes of Massachusetts}} 24. ^1 Timber Rattlesnake Fact Sheet at NY State Dept. of Environmental Conservation. Accessed 8 February 2007. 25. ^{{cite book|author=Furman, Jon|title=Timber rattlesnakes in Vermont and New York: biology, history, and the fate of an endangered species|publisher=UPNE|year=2007|isbn=978-1-58465-656-2|url=https://books.google.com/books?id=S5bM83jWl9YC&pg=PA133|page=133}} 26. ^Clark, R. W. (2004). Timber rattlesnakes (Crotalus horridus) use chemical cues to select ambush sites. Journal of Chemical Ecology, 30(3), 607-617. 27. ^1 Reinert, H. K., Cundall, D., & Bushar, L. M. (1984). Foraging behavior of the timber rattlesnake, Crotalus horridus. Copeia, 976-981. 28. ^Platt, S. G., Hawkes, A. W., & Rainwater, T. R. (2001). Diet of the canebrake rattlesnake (Crotalus horridus atricaudatus): An additional record and review. Texas Journal of Science, 53(2), 115-120. 29. ^Clark, R. W. (2002). Diet of the timber rattlesnake, Crotalus horridus. Journal of Herpetology, 36(3), 494-499. 30. ^1 2 Norris R. 2004. Venom Poisoning in North American Reptiles. In Campbell JA, Lamar WW. 2004. The Venomous Reptiles of the Western Hemisphere. Comstock Publishing Associates. Ithaca and London. 870 pp. 1500 plates. {{ISBN|0-8014-4141-2}}. 31. ^U.S. Navy. 1991. Poisonous Snakes of the World. US Govt. New York: Dover Publications Inc. 203 pp. {{ISBN|0-486-26629-X}}. 32. ^{{cite web|url=http://www.medscape.com/viewarticle/487281_5 |title=Snake Venoms and the Neuromuscular Junction: Spontaneous Activity |publisher=Medscape.com |date=2004-08-16 |accessdate=2014-07-29}} 33. ^{{cite web|url=http://www.toxinfo.org/antivenoms/indication/CROTALUS_HORRIDUS_HORRIDUS.html |title=MAVIN 2013-05-14, Crotalus horridus horridus |publisher=Toxinfo.org |date= |accessdate=2014-07-29}} 34. ^{{cite web | url = http://www.legis.state.wv.us/Bill_Status/Resolution_History.cfm?year=2008&sessiontype=RS&input4=28&billtype=cr&houseorig=s&btype=res | title = Senate concurrent resolution 28 (bill status 2008 regular session) | accessdate = February 22, 2011 | publisher = West Virginia Legislature}} 35. ^{{cite web|title=Senate concurrent resolution no. 28|url=http://www.legis.state.wv.us/Bill_Status/resolution.cfm?billdoc=scr28%20amended.htm&yr=2008&sesstype=RS&i=28|work=1st session of the 80th legislature|publisher=West Virginia Legislature|accessdate=February 25, 2011|year=2008}} 36. ^Bryson, Bill. 2004. A Short History of Nearly Everything. New York: Broadway Books. p. 81. 37. ^{{Redlist CC2001|date=13 September|year=2007}} 38. ^{{cite web|url=http://www.mass.gov/eea/agencies/dfg/dfw/natural-heritage/species-information-and-conservation/mesa-list/list-of-rare-species-in-massachusetts.html |title=Massachusetts List of Endangered, Threatened and Special Concern Speci |publisher=Mass.gov |date= |accessdate=2014-07-29}} 39. ^{{citation|author=Indiana Legislative Services Agency|year=2011|title=312 IAC 9-5-4: Endangered species of reptiles and amphibians|work=Indiana Administrative Code|accessdate=28 April 2012|url=http://www.in.gov/legislative/iac/}} 40. ^{{cite web|title=Timber Rattlesnake |url=http://www.oriannesociety.org/timber-rattlesnake|publisher=Orianne Society|accessdate=6 August 2015}} Further reading{{refbegin}}
External links
9 : Crotalus|Crotalinae by common name|Crotalinae by taxonomic synonyms|Reptiles of the United States|Reptiles described in 1758|Fauna of the Eastern United States|Reptiles of Ontario|Symbols of West Virginia|Taxa named by Carl Linnaeus |
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