text:
Commander (Royal Navy)
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For other nations and militaries that use this rank, see Commander.
A Royal Navy commander's rank insignia.
Commander (often abbreviated Cdr) is a senior officer rank of the Royal Navy of the United Kingdom. It is immediately junior to captain and immediately senior to the rank of lieutenant commander. Officers holding the junior rank of lieutenant commander are not considered to be commanders.
History[edit]
The title (originally 'master and commander')[1] originated in around 1670 to describe naval officers who commanded ships of war too large to be commanded by a lieutenant, but too small to warrant the assignment of a post-captain, or (before about 1770) a sailing-master who was in charge of a ship's navigation.[2] These ships were usually unrated sloops-of-war of no more than 20 guns, fireships, hospital ships and store ships.[3]:127 The commanding officer of this type of ship was responsible for both sailing and fighting the ship and was thus its 'master and commander'.
Before 1750 the rank was broadly considered as the limit of advancement for those without patronage, especially those who had been promoted from among a ship's crew.[4] By contrast those with Parliamentary supporters or family connections were more likely spend only a nominal period as master and commander of a sloop before being elevated to post-captain. From 1718 the Navy List began recording an officer's date of appointment to the rank of master and commander, with the intention of establishing seniority as a guide to promotion, but there is little evidence that this carried through into actual appointments.[3]:130
Over the later eighteenth century the rank evolved into a more regular stage of service between lieutenant and captain. The Royal Navy shortened 'master and commander' to 'commander' in 1794;[5] however, the term 'master and commander' remained (unofficially) in common parlance for several years.[6]
The rank was a popular recognition of service during the Napoleonic Wars, resulting in promotion of more commanders than there were commands; in 1812 the Navy List recorded 586 commanders against 168 available vessels. Commanders unable to secure a ship were left ashore on half-pay, with limited prospects for future advancement. This promotions bottleneck was addressed from 1827 with the introduction of commanders as a second-in-command on larger vessels.[7]:98
In the 20th and 21st centuries, the rank has been assigned the NATO rank code of OF-4, matching the army rank of lieutenant colonel.
Seniority and usage[edit]
A commander in the Royal Navy is senior to an officer holding the rank of lieutenant commander but junior to a captain. A commander may command a frigate, destroyer, submarine, aviation squadron or shore installation, or may serve on a staff. Formerly equivalent to the Army rank of major,[8] a commander is now equivalent in rank to a lieutenant colonel in the British Army or a wing commander in the Royal Air Force. The rank of wing commander was derived from the naval rank of commander via the usage in the World War I Royal Naval Air Service.
The rank insignia of a commander features three rings of gold braid with a loop in the upper ring.[9]
References[edit]
Jump up ^ See also: Master and Commander by Patrick O'Brian.
Jump up ^ Rodger, N. A. M. (2001). "Commissioned officers' careers in the Royal Navy, 1690–1815". Journal for Maritime Research. 3 (1): 88–89. doi:10.1080/21533369.2001.9668314.
^ Jump up to: a b Baugh, Daniel A. (1965). British Naval Administration in the Age of Walpole. Princeton University Press. ISBN 978-0691051079. OCLC 729683642.
Jump up ^ Rodger 2001, pp. 88-89.
Jump up ^ Hickox, Rex (2005). All You Wanted to Know about 18th Century Royal Navy: Medical Terms, Expressions and Trivia. Bentonville, Arkansas: Rex Publishing. p. 30. ISBN 978-1-4116-3057-4. Retrieved 7 September 2015.
Jump up ^ "Why is the Colonel called a 'Kernal?'". Naval Historical Center. 1998. Archived from the original on February 24, 2011.
Jump up ^ Lavery, Brian (2012). Nelson's Navy: The Ships, Men and Organisation, 1793-1815. Naval Institute Press. ISBN 9781591146124.
Jump up ^ The Royal Kalendar, and Court and City Register for England, Scotland, Ireland and the Colonies. London: Oxford University. 1835. p. 148. Retrieved 7 September 2015.
Jump up ^ "Uniforms and Badges of Rank at Royal Navy website". Archived from the original on 27 January 2009. Retrieved 7 September 2015.
[hide]
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Commissioned officer ranks of the British Armed Forces
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Student officer
OF-1
OF-2
OF-3
OF-4
OF-5
OF-6
*
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**
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***
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****
OF-10
*****
Royal Navy
O Cdt Mid SLt Lt Lt Cdr Cdr Capt Cdre RAdm
(list) VAdm
(list) Adm
(list) Adm of the Fleet
Royal Marines
2Lt Lt Capt Maj Lt Col Col Brig Maj-Gen Lt-Gen Gen
(list)
Army
O Cdt 2Lt Lt Capt Maj Lt Col Col Brig Maj-Gen
(list) Lt-Gen
(list) Gen
(list) Fd Mshl
Royal Air Force
Off Cdt / SO APO / Plt Off Fg Off Flt Lt Sqn Ldr Wg Cdr Gp Capt Air Cdre AVM Air Mshl Air Chf Mshl
(list) Mshl of the RAF
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Immunoglobulin class switching
From Wikipedia, the free encyclopedia
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Mechanism of class-switch recombination that allows isotype switching in activated B cells.
Immunoglobulin class switching, also known as isotype switching, isotypic commutation or class-switch recombination (CSR), is a biological mechanism that changes a B cell's production of immunoglobulin (antibodies) from one type to another, such as from the isotype IgM to the isotype IgG. During this process, the constant-region portion of the antibody heavy chain is changed, but the variable region of the heavy chain stays the same (the terms "variable" and "constant" refer to changes or lack thereof between antibodies that target different epitopes). Since the variable region does not change, class switching does not affect antigen specificity. Instead, the antibody retains affinity for the same antigens, but can interact with different effector molecules.
Contents
1 Mechanism
2 Cytokines responsible for class switching
3 Gene regulatory sequences responsible for class switching
4 See also
5 References
6 External links
Mechanism[edit]
Class switching occurs after activation of a mature B cell via its membrane-bound antibody molecule (or B cell receptor) to generate the different classes of antibody, all with the same variable domains as the original antibody generated in the immature B cell during the process of V(D)J recombination, but possessing distinct constant domains in their heavy chains.[1]
Naïve mature B cells produce both IgM and IgD, which are the first two heavy chain segments in the immunoglobulin locus. After activation by antigen, these B cells proliferate. If these activated B cells encounter specific signaling molecules via their CD40 and cytokine receptors (both modulated by T helper cells), they undergo antibody class switching to produce IgG, IgA or IgE antibodies. During class switching, the constant region of the immunoglobulin heavy chain changes but the variable regions, and therefore antigenic specificity, stay the same. This allows different daughter cells from the same activated B cell to produce antibodies of different isotypes or subtypes (e.g. IgG1, IgG2 etc.).[2]
The order of the heavy chain exons are as follows:
μ - IgM
δ - IgD
γ3 - IgG3
γ1 - IgG1
α1 - IgA1
γ2 - IgG2
γ4 - IgG4
ε - IgE
α2 - IgA2[3]
Class switching occurs by a mechanism called class switch recombination (CSR) binding. Class switch recombination is a biological mechanism that allows the class of antibody produced by an activated B cell to change during a process known as isotype or class switching. During CSR, portions of the antibody heavy chain locus are removed from the chromosome, and the gene segments surrounding the deleted portion are rejoined to retain a functional antibody gene that produces antibody of a different isotype. Double-stranded breaks are generated in DNA at conserved nucleotide motifs, called switch (S) regions, which are upstream from gene segments that encode the constant regions of antibody heavy chains; these occur adjacent to all heavy chain constant region genes with the exception of the δ-chain. DNA is nicked and broken at two selected S-regions by the activity of a series of enzymes, including Activation-Induced (Cytidine) Deaminase (AID), uracil DNA glycosylase and apyrimidic/apurinic (AP)-endonucleases.[4][5] The intervening DNA between the S-regions is subsequently deleted from the chromosome, removing unwanted μ or δ heavy chain constant region exons and allowing substitution of a γ, α or ε constant region gene segment. The free ends of the DNA are rejoined by a process called non-homologous end joining (NHEJ) to link the variable domain exon to the desired downstream constant domain exon of the antibody heavy chain.[6] In the absence of non-homologous end joining, free ends of DNA may be rejoined by an alternative pathway biased toward microhomology joins.[7] With the exception of the μ and δ genes, only one antibody class is expressed by a B cell at any point in time. While class switch recombination is mostly a deletional process, rearranging a chromosome in "cis", it can also occur (in 10 to 20% of cases, depending upon the Ig class) as an inter-chromosomal translocation mixing immunoglobulin heavy chain genes from both alleles.[8][9]
Cytokines responsible for class switching[edit]
T cell cytokines modulate class switching in mouse (Table 1) and human (Table 2).[10][11] These cytokines may have suppressive effect on production of IgM.
Gene regulatory sequences responsible for class switching[edit]
In addition to the highly repetitive structure of the target S regions, the process of class switching needs S regions to be first transcribed and spliced out of the immunoglobulin heavy chain transcripts (where they lie within introns). Chromatin remodeling, accessibility to transcription and to AID and synapsis of broken S regions are under the control of a large super-enhancer, located downstream the more distal Calpha gene, the 3' regulatory region (3'RR).[12] In some occasions, the 3'RR super-enhancer can itself be targeted by AID and undergo DNA breaks and junction with Sµ, which then deletes the Ig heavy chain locus and defines locus suicide recombination (LSR).[13]
Table 1. Class switching in mouse
T cells
Cytokines
Immunoglobulin classes
IgG1
IgG2a
IgG2b
IgG3
IgG4
IgE
IgA
Th2
IL-4
↑ ↓ ↓ ↓ ↓ ↑
IL-5
↑
Th1
IFNγ
↓ ↑ ↓ ↑ ↓ ↓
Treg
TGFβ
↑ ↓ ↑
IL-10[14]
↑
Table 2. Class switching in human
T cells
Cytokines
Immunoglobulin classes
IgG1
IgG2
IgG3
IgG4
IgA
IgE
Th2
IL-4
↑ ↓ ↑ ↑
IL-5
↑
Th1
IFNγ
↓ ↑ ↓
Treg
TGFβ
↓ ↑
IL-10[15][16]
↑
↑
See also[edit]
Antibody
Immunogenetics
References[edit]
Jump up ^ Eleonora Market, F. Nina Papavasiliou (2003) V(D)J Recombination and the Evolution of the Adaptive Immune System PLoS Biology1(1): e16.
Jump up ^ Stavnezer J, Amemiya CT (2004). "Evolution of isotype switching". Semin. Immunol. 16 (4): 257–75. doi:10.1016/j.smim.2004.08.005. PMID 15522624.
Jump up ^ Parham, Peter (2015). The Immune System (4th ed.). Garland Science.
Jump up ^ Durandy A (2003). "Activation-induced cytidine deaminase: a dual role in class-switch recombination and somatic hypermutation". Eur. J. Immunol. 33 (8): 2069–73. doi:10.1002/eji.200324133. PMID 12884279.
Jump up ^ Casali P, Zan H (2004). "Class switching and Myc translocation: how does DNA break?". Nat. Immunol. 5 (11): 1101–3. doi:10.1038/ni1104-1101. PMC 4625794  . PMID 15496946.
Jump up ^ Lieber MR, Yu K, Raghavan SC (2006). "Roles of nonhomologous DNA end joining, V(D)J recombination, and class switch recombination in chromosomal translocations". DNA Repair (Amst.). 5 (9–10): 1234–45. doi:10.1016/j.dnarep.2006.05.013. PMID 16793349.
Jump up ^ Yan CT, Boboila C, Souza EK, Franco S, Hickernell TR, Murphy M, Gumaste S, Geyer M, Zarrin AA, Manis JP, Rajewsky K, Alt FW (2007). "IgH class switching and translocations use a robust non-classical end-joining pathway". Nature. 449 (7161): 478–82. doi:10.1038/nature06020. PMID 17713479.
Jump up ^ Reynaud S, Delpy L, Fleury L, Dougier HL, Sirac C, Cogné M (May 2005). "Interallelic class switch recombination contributes significantly to class switching in mouse B cells". Journal of Immunology. 174 (10): 6176–83. PMID 15879114.
Jump up ^ Laffleur B, Bardet SM, Garot A, Brousse M, Baylet A, Cogné M (2014). "Immunoglobulin genes undergo legitimate repair in human B cells not only after cis- but also frequent trans-class switch recombination". Genes and Immunity. 15 (5): 341–6. doi:10.1038/gene.2014.25. PMID 24848929.
Jump up ^ Janeway CA Jr.; Travers P; Walport M; Shlomchik MJ (2001). Immunobiology (5th ed.). Garland Publishing. (via NCBI Bookshelf) ISBN 0-8153-3642-X.
Jump up ^ Male D, Brostoff J, Roth DB, Roitt I (2006). Immunology, 7th ed. Philadelphia: Mosby Elsevier, ISBN 978-0-323-03399-2 (pbk.)
Jump up ^ Pinaud E, Marquet M, Fiancette R, Péron S, Vincent-Fabert C, Denizot Y, Cogné M (2011). "The IgH locus 3' regulatory region: pulling the strings from behind". Advances in Immunology. 110: 27–70. doi:10.1016/B978-0-12-387663-8.00002-8. PMID 21762815.
Jump up ^ Péron S, Laffleur B, Denis-Lagache N, Cook-Moreau J, Tinguely A, Delpy L, Denizot Y, Pinaud E, Cogné M (May 2012). "AID-driven deletion causes immunoglobulin heavy chain locus suicide recombination in B cells". Science. 336 (6083): 931–4. doi:10.1126/science.1218692. PMID 22539552.
Jump up ^ Shparago, N.; Zelazowski, P.; Jin, L.; McIntyre, T. M.; Stuber, E.; Peçanha, L. M.; Kehry, M. R.; Mond, J. J.; Max, E. E. (1996-05-01). "IL-10 selectively regulates murine Ig isotype switching". International Immunology. 8 (5): 781–790. doi:10.1093/intimm/8.5.781. ISSN 0953-8178. PMID 8671667.
Jump up ^ Brière F, Servet-Delprat C, Bridon JM, Saint-Remy JM, Banchereau J (February 1994). "Human interleukin 10 induces naive surface immunoglobulin D+ (sIgD+) B cells to secrete IgG1 and IgG3". The Journal of Experimental Medicine. 179 (2): 757–62. doi:10.1084/jem.179.2.757. PMC 2191366  . PMID 8294883.
Jump up ^ Malisan F, Brière F, Bridon JM, Harindranath N, Mills FC, Max EE, Banchereau J, Martinez-Valdez H (March 1996). "Interleukin-10 induces immunoglobulin G isotype switch recombination in human CD40-activated naive B lymphocytes". The Journal of Experimental Medicine. 183 (3): 937–47. doi:10.1084/jem.183.3.937. PMC 2192363  . PMID 8642297.
External links[edit]
Immunoglobulin+class+switching at the US National Library of Medicine Medical Subject Headings (MeSH)
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Immunology: lymphocytic adaptive immune system and complement
Lymphoid
Antigens
Antigen
Superantigen
Allergen
Hapten
Epitope
Linear
Conformational
Mimotope
Antigen presentation/Professional APCs: Dendritic cell
Macrophage
B cell
Immunogen
Antibodies
Antibody
Monoclonal antibodies
Polyclonal antibodies
Autoantibody
Microantibody
Polyclonal B cell response
Allotype
Isotype
Idiotype
Immune complex
Paratope
Immunity vs.
tolerance
action: Immunity
Autoimmunity
Alloimmunity
Allergy
Hypersensitivity
Inflammation
Cross-reactivity
inaction: Tolerance
Central
Peripheral
Clonal anergy
Clonal deletion
Tolerance in pregnancy
Immunodeficiency
Immunogenetics
Affinity maturation
Somatic hypermutation
Clonal selection
V(D)J recombination
Junctional diversity
Immunoglobulin class switching
MHC/HLA
Lymphocytes
Cellular
T cell
Humoral
B cell
NK cell
Substances
Cytokines
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Cytolysin
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text:
Groundhog
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For other uses, see Groundhog (disambiguation) and Woodchuck (disambiguation).
Groundhog
Groundhog at Laval University campus, Quebec, Canada
Conservation status

Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Family: Sciuridae
Genus: Marmota
Species: M. monax
Binomial name
Marmota monax
(Linnaeus 1758;
Trouessart, Catal. Mamm. viv. foss., suppl., p. 344)
Subspecies
M. m. monax Linnaeus, 1758
M. m. canadensis Erxleben, 1777
M. m. ignava Bangs, 1899
M. m. rufescens A. H. Howell, 1914
Groundhog range
Synonyms
Mus monax Linnaeus 1758
Arctomys monax, True, Proc. U. S. Nat. Mus., vol. 7 (1884), p. 593. 1885.
A groundhog burrow hole showing size.
An adult female groundhog walking through snow in March
A groundhog has climbed up to reach water in a bird bath.
Juvenile groundhog siblings on a morning in early June.
Mated groundhogs in March. Female on the left, male on the right.
Juvenile groundhog siblings play-fighting
Groundhog prepares to feast on an acorn
Emerged from hibernation in February, groundhog takes leaves to line the burrow nest or toilet chamber.
The groundhog (Marmota monax), also known as a woodchuck, is a rodent of the family Sciuridae, belonging to the group of large ground squirrels known as marmots.[2] It was first scientifically described by Carl Linnaeus in 1758.[3] The groundhog is also referred to as a chuck, wood-shock, groundpig, whistlepig,[4][5] whistler, thickwood badger, Canada marmot, monax, moonack, weenusk, red monk[6] and, among French Canadians in eastern Canada, siffleur.[7] The name "thickwood badger" was given in the Northwest to distinguish the animal from the prairie badger. Monax was a Native American name of the woodchuck, which meant "the digger".[8] Young groundhogs may be called chucklings.[9]:66 Other marmots, such as the yellow-bellied and hoary marmots, live in rocky and mountainous areas, but the groundhog is a lowland creature. It is found through much of the eastern United States across Canada and into Alaska[10]
Contents
[hide]
1 Description
2 Etymology
3 Distribution and habitat
4 Survival
5 Behavior
5.1 Diet
5.2 Burrows
5.3 Hibernation
5.4 Reproduction
6 Relationship with humans
7 References
8 Further reading
9 External links
Description[edit]
The groundhog is the largest sciurid in its geographical range. Adults are 16 to 20 inches long, including a six inch tail.[11] A large woodchuck thought to weigh twenty pounds when carried was exactly half that weight when weighed by scale.[12] Woodchuck weight ranges from five to twelve pounds.[13] Extremely large individuals may weigh up to 15 pounds.[14] Seasonal weight changes indicate circannual deposition and use of fat. Progressive higher weights are attained each year for the first 2–3 years after which weights plateau.[15] Groundhogs have four incisor teeth which grow 1/16" per week. Constant usage wears them down again by about that much each week.[16] Unlike the incisors of other rodents, the incisors of groundhogs are white to ivory white.[17][18] Groundhogs are well adapted for digging, with short, powerful limbs and curved, thick claws. Unlike other sciurids, the groundhog's spine is curved, more like that of a mole,[citation needed] and the tail is comparably shorter as well — only about one-fourth of body length. Suited to their temperate habitat, groundhogs are covered with two coats of fur: a dense grey undercoat and a longer coat of banded guard hairs that gives the groundhog its distinctive "frosted" appearance.[citation needed]
Etymology[edit]
The etymology of the name woodchuck is unrelated to wood or chucking. It stems from an Algonquian (possibly Narragansett) name for the animal, wuchak.[19] The similarity between the words has led to the popular tongue-twister:
How much wood would a woodchuck chuck
if a woodchuck could chuck wood?
A woodchuck would chuck all the wood he could
if a woodchuck could chuck wood![20]
Distribution and habitat[edit]
The groundhog prefers open country and the edges of woodland, and is rarely far from a burrow entrance.[21] Marmota monax has a wide geographic range. It is typically found in low elevation forests, small woodlots, fields, pastures and hedgerows. It constructs dens in well drained soil, and most have summer and winter dens. Human activity has increased food access and abundance allowing M. monax to thrive.[22] As a consequence, the groundhog is a familiar animal to many people in the United States and Canada.[citation needed]
Survival[edit]
Groundhogs can climb trees to escape predators
In the wild, groundhogs can live up to six years with two or three being average. In captivity, groundhogs reportedly live up to 14 years. Humans, dogs, coyotes, and foxes are about the only predators that can kill adult groundhogs although young may also be taken by owls and hawks.[23] The red fox is the major predator of Marmota monax.[24][25] Occasionally woodchucks may suffer from parasitism and a woodchuck may die from infestation or from bacteria transmitted by vectors.[26] In areas of intensive agriculture and dairying regions of the state of Wisconsin, particularly the southern parts, the woodchuck by 1950 had been almost extirpated.[27]:124 Jackson (1961) suggested the amount of damage done by the woodchuck had been exaggerated and that excessive persecution by people substantially reduced its numbers in Wisconsin. In some areas marmots are important game animals and are killed regularly for sport, food, or fur. In Kentucky an estimated 267,500 M. monax were taken annually from 1964 to 1971 (Barbour and Davis 1974)[28] Woodchucks had protected status in the state of Wisconsin[29] until 2017. [30] Woodchuck numbers appear to have decreased in Illinois.[31]
Behavior[edit]
Motionless individual, alert to danger, will whistle when alarmed to warn other groundhogs
The time spent observing groundhogs by field biologists represents only a small fraction of time devoted to the field research.[32] W.J. Schoonmaker reports that groundhogs may hide when they see, smell or hear the observer.[9]:41–43 Ken Armitage, marmot researcher, states that the social biology of the groundhog is well understudied.[33] Despite their heavy-bodied appearance, groundhogs are accomplished swimmers and occasionally climb trees when escaping predators or when they want to survey their surroundings.[34] They prefer to retreat to their burrows when threatened; if the burrow is invaded, the groundhog tenaciously defends itself with its two large incisors and front claws. Groundhogs are generally agonistic and territorial among their own species, and may skirmish to establish dominance.[35] Outside their burrow, individuals are alert when not actively feeding. It is common to see one or more nearly-motionless individuals standing erect on their hind feet watching for danger. When alarmed, they use a high-pitched whistle to warn the rest of the colony, hence the name "whistle-pig".[36][37] Groundhogs may squeal when fighting, seriously injured, or caught by a predator.[37] Other sounds groundhogs may make are low barks and a sound produced by grinding their teeth.[37] When groundhogs are frightened, the hairs of the tail stand straight up, giving the tail the appearance of a hair brush.[citation needed] David P. Barash wrote he witnessed only two occasions of upright play-fighting among woodchucks and that the upright posture of play-fighting involves sustained physical contact between individuals and may require a degree of social tolerance virtually unknown in M. monax. He said it was possible to conclude, alternatively, that upright play-fighting is part of the woodchuck's behavioral repertory but rarely shown because of physical spacing and/or low social tolerance.[38]
Clover is a preferred food source for groundhogs.
Diet[edit]
Mostly herbivorous, groundhogs eat primarily wild grasses and other vegetation, including berries and agricultural crops, when available.[35] In early spring, dandelion and coltsfoot are important groundhog food items. Some additional foods include: sheep sorrel, timothy, buttercup, tear thumb, agrimony, red and black raspberries, buckwheat, plantain, wild lettuce, all varieties of clover, and alfalfa.[39] Groundhogs also occasionally eat grubs, grasshoppers, insects, snails and other small animals, but are not as omnivorous as many other Sciuridae. Like squirrels, they also have been observed sitting up eating nuts such as shagbark hickory, but unlike squirrels, do not bury them for future use.[citation needed]
Groundhogs will occasionally eat snails, insects and baby birds they come upon by accident.[40] An adult groundhog will eat more than a pound of vegetation daily.[41] In early June, woodchucks' metabolism slows, food intake decreases, their weight increases by as much as 100% and fat deposits are produced upon which they will live during hibernation and late winter.[42] Instead of storing food, groundhogs stuff themselves to survive the winter without eating.[43] Thought not to drink water, groundhogs are reported to obtain needed liquids from the juices of food-plants, aided by their sprinkling with rain or dew.[44][45][46]
Burrows[edit]
Groundhogs are excellent burrowers, using burrows for sleeping, rearing young, and hibernating. W. J. Schoonmaker excavated 11 dens finding the earth removed from these averaged six cubic feet or four and eight-tenths bushels per den. The longest burrow was 24 feet plus 2 short side galleries. The amount of soil taken from this den was eight bushels and it weighed 640 pounds. The average weight of the earth taken from all eleven dens was 384 pounds.[47] Though groundhogs are the most solitary of the marmots, several individuals may occupy the same burrow. Groundhog burrows usually have two to five entrances, providing groundhogs their primary means of escape from predators. Burrows are particularly large, with up to 14 metres (46 ft) of tunnels buried up to 1.5 metres (5 ft) underground, and can pose a serious threat to agricultural and residential development by damaging farm machinery and even undermining building foundations.[36] In a June 7, 2009 Humane Society of the United States article, "How to Humanely Chuck a Woodchuck Out of Your Yard", John Griffin, director of Humane Wildlife Services, stated you would have to have a lot of woodchucks working over a lot of years to create tunnel systems that would pose any risk to a structure.
The burrow is used for safety, retreat in bad weather, hibernating, sleeping, love nest, and nursery. In addition to the nest, there is an excrement chamber. The nest chamber may be about twenty inches to three feet below ground surface. It is about sixteen inches wide and fourteen inches high. There are typically two burrow openings or holes. One is the main entrance, the other a spy hole. Description of the length of the burrow often includes the side galleries of the burrow. Excluding the side galleries, Schoonmaker reports the longest was twenty-four feet, with the average length of eleven dens dug out to be fourteen feet.[48] W. H. Fisher investigated nine burrows, finding the deepest point to be forty-nine inches down. The longest, including side galleries, was forty-seven feet eleven and one half inches.[49] Numbers of burrows per individual groundhog decreases with urbanization.[50][51]
Bachman mentioned that when the young groundhogs are a few months old, they prepare for separation, digging a number of holes in the area of their early home. Some of these holes were only a few feet deep and never occupied but the numerous burrows gave the impression that groundhogs live in communities.[52]
Hibernation[edit]
Groundhogs are one of the few species that enter into true hibernation, and often build a separate "winter burrow" for this purpose. This burrow is usually in a wooded or brushy area and is dug below the frost line and remains at a stable temperature well above freezing during the winter months. In most areas, groundhogs hibernate from October to March or April, but in more temperate areas, they may hibernate as little as three months.[53] Groundhogs hibernate longer in northern latitudes than southern latitudes.[54][55] To survive the winter, they are at their maximum weight shortly before entering hibernation.[56] When the groundhog enters hibernation, there is a drop in body temperature to as low as 35 degrees fahrenheit, heart rate falls to 4-10 beats per minute and breathing rate falls to one breath every six minutes.[57] During hibernation, they experience periods of torpor and arousal.[58] They emerge from hibernation with some remaining body fat to live on until the warmer spring weather produces abundant plant materials for food.[56] Males emerge from hibernation before females.[59][60] Groundhogs are mostly diurnal, and are often active early in the morning or late afternoon.[61]
Reproduction[edit]
A groundhog mother and her cubs (kits) in a suburban yard
Usually groundhogs breed in their second year, but a small proportion may breed in their first. The breeding season extends from early March to mid- or late April, after hibernation. A mated pair remains in the same den throughout the 31- to 32-day[62] gestation period. As birth of the young approaches in April or May, the male leaves the den. One litter is produced annually, usually containing two to six blind, hairless and helpless young. Groundhog mothers introduce their young to the wild once their fur is grown in and they can see. At this time, if at all, the father groundhog comes back to the family.[63]:316 They encourage their young to copy their behaviors and during this time may differ from usual routines.[citation needed] By the end of August, the family breaks up; or at least, the larger number scatter, to burrow on their own.[64]
Relationship with humans[edit]
Both their diet and habit of burrowing make them serious nuisance animals around farms and gardens. They will eat many commonly grown vegetables, and their burrows can destroy farm ponds and undermine foundations. Consequently, there is a thriving business exterminating them. Their preferred habitat of grassy areas near woods also makes them abundant along roads and highways where they often become the victims of passing cars.[citation needed]
Very often the dens of groundhogs provide homes for other animals including skunks, red foxes, and cottontail rabbits. The fox and skunk feed upon field mice, grasshoppers, beetles and other creatures that destroy farm crops. In aiding these animals, the groundhog indirectly helps the farmer. In addition to providing homes for itself and other animals, the groundhog aids in soil improvement by bringing subsoil to the surface. The groundhog is also a valuable game animal and is considered a difficult sport when hunted in a fair manner.[65] In some parts of Appalachia, they are eaten.[66]
A report in 1883 by the New Hampshire Legislative Woodchuck Committee illustrates the attitude of some people towards this animal. In part, the report states:[67]
The woodchuck, despite its deformities both of mind and body, possess some of the amenities of a higher civilization. It cleans its face after the manner of the squirrels, and licks its fur after the manner of a cat. Your committee is too wise, however, to be deceived by this purely superficial observation of better habits. Contemporaneous with the ark, the woodchuck has not made any material progress in social science, and it is now too late to reform the wayward sinner. The average age of the woodchuck is too long to please your committee.... The woodchuck is not only a nuisance, but also a bore. It burrows beneath the soil, and then chuckles to see a mowing machine, man and all, slump into one of these holes and disappear....
The committee concludes that "a small bounty will prove of incalculable good; at all events, even as an experiment, it is certainly worth trying; therefore your committee would respectfully recommend that the accompanying bill be passed."[68]
Groundhogs may be raised in captivity, but their aggressive nature can pose problems. Doug Schwartz, a zookeeper and groundhog trainer at the Staten Island Zoo, has been quoted as saying "They’re known for their aggression, so you’re starting from a hard place. His natural impulse is to kill ’em all and let God sort ’em out. You have to work to produce the sweet and cuddly."[69] Wildlife rehabilitation is the treatment and care of injured, orphaned, or sick wild animals so that they can be released back to the wild. In the event an injured, orphaned, or sick groundhog is found, contact a wildlife rescue center or rehabilitator. Groundhogs cared for by a rehabilitator that survive but can't be returned to the wild may remain with their caregivers and become educational ambassadors.[70][71][72]
In the United States and Canada, the yearly February 2 Groundhog Day celebration has given the groundhog recognition and popularity. The most popularly known of these groundhogs are Punxsutawney Phil, Wiarton Willie, Jimmy the Groundhog, Dunkirk Dave, and Staten Island Chuck kept as part of Groundhog Day festivities in Punxsutawney, Pennsylvania;, Wiarton, Ontario; Sun Prairie, Wisconsin, Dunkirk, New York, and Staten Island respectively. The 1993 comedy film Groundhog Day references several events related to Groundhog Day, and portrays both Punxsutawney Phil himself, and the annual Groundhog Day ceremony. Famous Southern groundhogs include Smith Lake Jake from Graysville, Alabama and General Beauregard Lee, based at the Yellow River Game Ranch outside Atlanta, Georgia.[citation needed]
Groundhogs are used in medical research on hepatitis B-induced liver cancer. A percentage of the woodchuck population is infected with the woodchuck hepatitis virus (WHV), similar to human hepatitis B virus. Humans do not receive hepatitis from woodchucks with WHV but the virus and its effects on the liver make the woodchuck the best available animal for the study of viral hepatitis in humans. The only other animal model for hepatitis B virus studies is the chimpanzee, an endangered species.[73] Woodchucks are also used in biomedical research investigating metabolic function, obesity, energy balance, the endocrine system, reproduction, neurology, cardiovascular disease, cerebrovascular disease, and neoplastic disease.[74]
Groundhog burrows have been known to reveal at least one archaeological site, the Ufferman Site in the U.S. state of Ohio.[75] Although archaeologists have never excavated the Ufferman Site, numerous artifacts have been found because of the activities of local groundhogs. They favor the loose soil of the esker upon which the site lies, and their many diggings for their burrows have brought to the surface significant numbers of human and animal bones, pottery, and bits of stone.[75] Woodchuck remains were found in the Indian mounds at Aztalan, Jefferson County, Wisconsin.[76]
Robert Frost's poem "A Drumlin Woodchuck" uses the imagery of a groundhog dug into a small ridge as a metaphor for his emotional reticence.[citation needed]
References[edit]
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Further reading[edit]
Bezuidenhout, A. J. (Abraham Johannes) and Evans, Howard E. (Howard Edward). Anatomy of the woodchuck (Marmota monax). Lawrence, KS: American Society of Mammalogists, 2005. ISBN 9781891276439. Online at doi:10.5962/bhl.title.61270
External links[edit]
Wikimedia Commons has media related to Marmota monax.
Wikispecies has information related to Marmota monax
Woodchuck, Hinterland Who's Who
Woodchuck (Groundhog), Missouri Conservation Commission
[7] Breeding and Experimental Facility for Woodchucks
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Living species of tribe Marmotini (ground squirrels)
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Suborder: Sciuromorpha
Family: Sciuridae
Subfamily: Xerinae
Ammospermophilus
(antelope squirrels)
Harris's antelope squirrel (A. harrisii)
Espíritu Santo antelope squirrel (A. insularis)
Texas antelope squirrel (A. interpres)
White-tailed antelope squirrel (A. leucurus)
San Joaquin antelope squirrel (A. nelsoni)
Callospermophilus
(golden-mantled ground squirrels)
Golden-mantled ground squirrel (C. lateralis)
Sierra Madre ground squirrel (C. madrensis)
Cascade golden-mantled ground squirrel (C. saturatus)
Cynomys
(prairie dogs)
Gunnison's prairie dog (C. gunnisoni)
White-tailed prairie dog (C. leucurus)
Black-tailed prairie dog (C. ludovicianus)
Mexican prairie dog (C. mexicanus)
Utah prairie dog (C. parvidens)
Eutamias
Siberian chipmunk (E. sibiricus)
Ictidomys
(little ground squirrels)
Mexican ground squirrel (I. mexicanus)
I. parvidens
Thirteen-lined ground squirrel (I. tridecemlineatus)
Marmota
(marmots)
Subgenus Marmota: Gray marmot (M. baibacina)
Bobak marmot (M. bobak)
Alaska marmot (M. broweri)
Black-capped marmot (M. camtschatica)
Long-tailed marmot (M. caudata)
Himalayan marmot (M. himalayana)
Alpine marmot (M. marmota)
Menzbier's marmot (M. menzbieri)
Groundhog or woodchuck (M. monax)
Tarbagan marmot (M. sibirica)
Subgenus Petromarmota: Hoary marmot (M. caligata)
Yellow-bellied marmot (M. flaviventris)
Olympic marmot (M. olympus)
Vancouver Island marmot (M. vancouverensis)
Neotamias
(western chipmunks)
Alpine chipmunk (N. alpinus)
Yellow-pine chipmunk (N. amoenus)
Buller's chipmunk (N. bulleri)
Gray-footed chipmunk (N. canipes)
Gray-collared chipmunk (N. cinereicollis)
Cliff chipmunk (N. dorsalis)
Durango chipmunk (N. durangae)
Merriam's chipmunk (N. merriami)
Least chipmunk (N. minimus)
California chipmunk (N. obscurus)
Yellow-cheeked chipmunk (N. ochrogenys)
Palmer's chipmunk (N. palmeri)
Panamint chipmunk (N. panamintinus)
Long-eared chipmunk (N. quadrimaculatus)
Colorado chipmunk (N. quadrivittatus)
Red-tailed chipmunk (N. ruficaudus)
Hopi chipmunk (N. rufus)
Allen's chipmunk (N. senex)
Siskiyou chipmunk (N. siskiyou)
Sonoma chipmunk (N. sonomae)
Lodgepole chipmunk (N. speciosus)
Townsend's chipmunk (N. townsendii)
Uinta chipmunk (N. umbrinus)
Notocitellus
Tropical ground squirrel (N. adocetus)
Ring-tailed ground squirrel (N. annulatus)
Otospermophilus
(rock squirrels)
Baja California rock squirrel (O. atricapillus)
California ground squirrel (O. beecheyi)
Rock squirrel (O. variegatus)
Poliocitellus
Franklin's ground squirrel (P. franklinii)
Sciurotamias
(Asian rock squirrels)
Père David's rock squirrel (S. davidianus)
Forrest's rock squirrel (S. forresti)
Spermophilus sensu stricto
(Old World ground squirrels)
Alashan ground squirrel (S. alashanicus)
S. brevicauda
European ground squirrel (S. citellus)
Daurian ground squirrel (S. dauricus)
Red-cheeked ground squirrel (S. erythrogenys)
Yellow ground squirrel (S. fulvus)
Russet ground squirrel (S. major)
S. pallicauda
Little ground squirrel (S. pygmaeus)
S. ralli
S. relictus
Speckled ground squirrel (Spermophilus suslicus)
Taurus ground squirrel (Spermophilus taurensis)
Asia Minor ground squirrel (Spermophilus xanthoprymnus)
Tamias
Eastern chipmunk (T. striatus)
Urocitellus
(Holarctic ground squirrels)
Uinta ground squirrel (U. armatus)
Belding's ground squirrel (U. beldingi)
Northern Idaho ground squirrel (U. brunneus)
Southern Idaho ground squirrel (U. endemicus)
Merriam's ground squirrel (U. canus)
Columbian ground squirrel (U. columbianus)
Wyoming ground squirrel (U. elegans)
Piute ground squirrel (U. mollis)
Arctic ground squirrel (U. parryii)
Richardson's ground squirrel (U. richardsonii)
Townsend's ground squirrel (U. townsendii)
Washington ground squirrel (U. washingtoni)
Long-tailed ground squirrel (U. undulatus)
Xerospermophilus
(pygmy ground squirrels)
Mohave ground squirrel (X. mohavensis)
Perote ground squirrel (X. perotensis)
Spotted ground squirrel (X. spilosoma)
Round-tailed ground squirrel (X. tereticaudus)
Category
Taxon identifiers
Wd: Q221612
ADW: Marmota_monax
EoL: 327979
EPPO: MAROMO
Fossilworks: 47902
GBIF: 2437368
iNaturalist: 46095
ITIS: 180137
IUCN: 42458
MSW: 12400961
NCBI: 9995
Retrieved from "https://en.wikipedia.org/w/index.php?title=Groundhog&oldid=841448913"
Categories:
IUCN Red List least concern species
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Mammals described in 1758
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