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Painted turtle
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==Behavior== ===Daily routine and basking=== [[File:Painted Turtle on a log.jpg|thumb|right|alt=A painted turtle standing on a floating log|Basking for warmth]] A [[Ectotherm|cold-blooded]] reptile, the painted turtle regulates its temperature through its environment, notably by basking. All ages bask for warmth, often alongside other species of turtle. Sometimes more than 50 individuals are seen on one log together.{{sfn|Ernst|Barbour|Lovich|1994|p=283}} Turtles bask on a variety of objects, often logs, but have even been seen basking on top of [[common loon]]s that were covering eggs.{{sfn|Gervais et al.|2009|p=13}} The turtle [[diurnality|starts its day at sunrise]], emerging from the water to bask for several hours. Warmed for activity, it returns to the water to forage.{{sfn|Ernst|Barbour|Lovich|1994|p=282}} After becoming chilled, the turtle re-emerges for one to two more cycles of basking and feeding.{{sfn|Ernst|Barbour|Lovich|1994|pp=282β283}} At night, the turtle drops to the bottom of its water body or perches on an underwater object and sleeps.{{sfn|Ernst|Barbour|Lovich|1994|p=282}} To be active, the turtle must maintain an internal body temperature between {{convert|17|-|23|C|F|abbr=on|0}}. When fighting infection, it manipulates its temperature up to {{convert|5|C-change|abbr=on}} higher than normal.{{sfn|Ernst|Barbour|Lovich|1994|p=283}} ===Seasonal routine and hibernation=== <!-- [[File:Muskrat lodge.jpg|thumb|right|alt=mound of sticks in the water about 10 feet from shore|Muskrat burrowβpossible painted turtle hibernation spot]] not enough room--> In the spring, when the water reaches {{convert|15|-|18|C|F|abbr=on|0}}, the turtle begins actively foraging. However, if the water temperature exceeds {{nowrap|{{convert|30|C|F|abbr=on|0}}}}, the turtle will not feed. In fall, the turtle stops foraging when temperatures drop below the spring set-point.{{sfn|Ernst|Lovich|2009|p=293}} During the winter, the turtle hibernates. In the north, the inactive season may be as long as from October to March, while the southernmost populations may not hibernate at all.{{sfn|Carr|1952|p=217}} While hibernating, the body temperature of the painted turtle averages {{nowrap|{{convert|6|C|F|abbr=on|0}}}}.{{sfn|Ernst|Barbour|Lovich|1994|p=284}} Periods of warm weather bring the turtle out of hibernation, and even in the north, individuals have been seen basking in February.{{sfn|Ernst|Barbour|Lovich|1994|p=281}} The painted turtle hibernates by burying itself, either on the bottom of a body of water, near water in the shore-bank or the burrow of a [[muskrat]], or in woods or pastures. When hibernating underwater, the turtle prefers shallow depths, no more than {{nowrap|{{convert|2|m|ft|abbr=on|0}}}}. Within the mud, it may dig down an additional {{nowrap|{{convert|1|m|ft|abbr=on|0}}}}.{{sfn|Ernst|Barbour|Lovich|1994|p=284}} In this state, the turtle does not breathe, although if surroundings allow, it may get some oxygen through its skin.<ref name="Jackson">{{cite journal |last=Jackson |first=D. C. |author2=Rauer, E. M. |author3=Feldman, R. A. |author4= Reese, S. A. |title=Avenues of extrapulmonary oxygen uptake in western painted turtles (''Chrysemys picta belli'') at 10 Β°C |journal=Comparative Biochemistry and Physiology A |date=August 2004 |volume=139 |issue=2 |pages=221β227 |pmid=15528171 |doi=10.1016/j.cbpb.2004.09.005 }}</ref> The species is one of the best-studied [[vertebrate]]s able to survive long periods [[Hypoxia (medical)|without oxygen]]. Adaptations of its blood chemistry, brain, heart, and particularly its shell allow the turtle to survive extreme [[lactic acid]] buildup while oxygen-deprived.<ref name="Jackson 2002">{{cite journal|last=Jackson|first=Donald C.|date=2002|title=Hibernating without oxygen: physiological adaptations of the painted turtle|journal=The Journal of Physiology|volume=543|issue=3|pages=731β737|doi=10.1113/jphysiol.2002.024729|pmc=2290531|pmid=12231634}}<!--|access-date=2010-12-13 --></ref> The painted turtle, like many other turtles, has the ability to breathe through its anus, or cloaca. This unusual adaptation, known as cloacal respiration, allows turtles to hibernate overwinter in colder climates where water surfaces may freeze over. Painted turtles also have specialized skin cells that absorb oxygen from the water. This process is known as cutaneous respiration. These two respiratory strategies, along with their other resilient traits, are vital to this reptile's success in colder climates. === Anoxia tolerance === During the winter months, painted turtles become ice-locked and spend their time in either [[Hypoxia (environmental)|hypoxic]] (low oxygen) or anoxic (no oxygen) regions of the pond or lake.<ref>{{Cite journal |last=Jackson |first=D. C. |date=2002 |title=Hibernating without Oxygen: Physiological Adaptations of the Painted Turtle |journal=The Journal of Physiology |volume=543 |issue=3 |pages=731β737 |doi=10.1113/jphysiol.2002.024729 |pmid=12231634 |pmc=2290531 }}</ref> Painted turtles essentially hold their breath until the following spring when the ice melts. As a result, painted turtles rely on [[anaerobic respiration]], which leads to the production of lactic acid.<ref name=":02">{{Cite journal |last=Jackson |first=D. C. |date=2000 |title=Living without oxygen: lessons from the freshwater turtle |journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology |volume=125 |issue=3 |pages=299β315 |doi=10.1016/s1095-6433(00)00160-4 |pmid=10794959 }}</ref> However, painted turtles can tolerate long periods of anoxia due to three factors: a depressed [[metabolic rate]], large [[glycogen]] stores in the liver, and sequestering lactate in the shell and releasing carbonate [[Buffer solution|buffers]] to the extracellular fluid.<ref name=":02" /> The shell of an adult painted turtle has the largest concentration of carbonate content recorded among animals.<ref>{{Cite journal |last1=Keenan |first1=S. W. |last2=Pasteris |first2=J. D. |last3=Wang |first3=A. |last4=Warren |first4=D. E. |date=2019 |title=Heterogeneous bioapatite carbonation in western painted turtles is unchanged after anoxia |journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology |volume=233 |pages=74β83 |doi=10.1016/j.cbpa.2019.03.019 |pmid=30930203 |s2cid=89619510 }}</ref> This large carbonate content helps the painted turtle buffer the accumulation of lactic acid during anoxia. Both the shell and skeleton release calcium and magnesium carbonates to buffer extracellular lactic acid.<ref>{{Cite journal |last1=Jackson |first1=D. C. |last2=Crocker |first2=C. E. |last3=Ultsch |first3=G. R. |date=2000 |title=Bone and shell contribution to lactic acid buffering of submerged turtles''Chrysemys picta bellii''at 3Β°C |journal=American Journal of Physiology. Regulatory, Integrative and Comparative Physiology |volume=278 |issue=6 |pages=R1564βR1571 |doi=10.1152/ajpregu.2000.278.6.r1564 |pmid=10848524 |s2cid=27785024 }}</ref><ref>{{Cite journal |last=Jackson |first=D. C. |date=2004 |title=Surviving extreme lactic acidosis: the role of calcium lactate formation in the anoxic turtle |journal=Respiratory Physiology & Neurobiology |volume=144 |issue=2β3 |pages=173β178 |doi=10.1016/j.resp.2004.06.020 |pmid=15556100 |s2cid=33342583 }}</ref> A painted turtle can also sequester 44% of total body lactate in their shell.<ref>{{Cite journal |last=Jackson |first=D |date=1997 |title=Lactate accumulation in the shell of the turtle Chrysemys picta bellii during anoxia at 3Β°C and 10Β°C |journal=Journal of Experimental Biology |volume=200 |issue=17 |pages=2295β2300 |doi=10.1242/jeb.200.17.2295 |pmid=9320212 }}</ref> Despite the shell's large buffering contribution, it does not experience any significant decrease in mechanical properties under natural conditions.<ref name=":1">{{Cite journal |last1=Odegard |first1=D. T. |last2=Sonnenfelt |first2=M. A. |last3=Bledsoe |first3=J. G. |last4=Keenan |first4=S. W. |last5=Hill |first5=C. A. |last6=Warren |first6=D. E. |date=2018 |title=Changes in the material properties of the shell during simulated aquatic hibernation in the anoxia-tolerant painted turtle |journal=Journal of Experimental Biology |volume=221 |issue=Pt 18 |doi=10.1242/jeb.176990 |pmid=30065038 |s2cid=51890031 |doi-access=free }}</ref> The duration of anoxia tolerance varies depending on the sub-species of painted turtle. The western painted turtle (''C. picta bellii'') can survive 170 days of anoxia, followed by the midland painted turtle (''C. picta marginata'') which can survive 150 days, and finally the eastern painted turtle (''C. picta picta''), which can survive 125 days.<ref name=":1" /><ref name=":3">{{Cite journal |last1=Reese |first1=S. A. |last2=Stewart |first2=E. R. |last3=Crocker |first3=C. E. |last4=Jackson |first4=D. C. |last5=Ultsch |first5=G. R. |date=2004 |title=Geographic variation of the physiological response to overwintering in the painted turtle (''Chrysemys picta'') |journal=Physiological and Biochemical Zoology |volume=77 |issue=4 |pages=619β630 |doi=10.1086/383514 |pmid=15449233 |s2cid=24651637 }}</ref><ref>{{Cite journal |last1=Reese |first1=S. A. |last2=Crocker |first2=C. E. |last3=Jackson |first3=D. C. |last4=Ultsch |first4=G. R. |date=2000 |title=The physiology of hibernation among painted turtles: the midland painted turtle (Chrysemys picta marginata) |journal=Respiration Physiology |volume=124 |issue=1 |pages=43β50 |doi=10.1016/S0034-5687(00)00188-2 |pmid=11084202 }}</ref><ref name=":4">{{Cite journal |last1=Ultsch |first1=G. R. |last2=Hanley |first2=R. W. |last3=Bauman |first3=T. R. |date=1985 |title=Responses to Anoxia during Simulated Hibernation in Northern and Southern Painted Turtles |journal=Ecology |volume=66 |issue=2 |pages=388β395 |doi=10.2307/1940388 |jstor=1940388 |bibcode=1985Ecol...66..388U }}</ref> Differences in anoxia tolerance are partially attributed to the rate of lactate production and buffering capability in painted turtles.<ref name=":3" /> Furthermore, northern populations of painted turtles have a higher anoxia tolerance than southern populations.<ref name=":3" /> Other anoxia tolerant freshwater turtles include: the southern painted turtle (''Chrysemys dorsalis''), which can survive 75β86 days of anoxia, the snapping turtle ([[Common snapping turtle|Chelydra serpentina]]), which can survive 100 days under anoxia, and the map turtle ([[Northern map turtle|Graptemys geographica]]), which can survive 50 days of anoxia.<ref name=":3" /><ref name=":4" /><ref>{{Cite journal |last1=Reese |first1=S. A. |last2=Jackson |first2=D.C. |last3=Ultsch |first3=G. R. |date=2002 |title=The Physiology of Overwintering in a Turtle That Occupies Multiple Habitats, the Common Snapping Turtle (Chelydra serpentina) |journal=Physiological and Biochemical Zoology |volume=75 |issue=5 |pages=432β438 |doi=10.1086/342802 |pmid=12529844 |s2cid=32401168 }}</ref><ref>{{Cite journal |last1=Reese |first1=S. A. |last2=Crocker |first2=C. E. |last3=Carwile |first3=M. E. |last4=Jackson |first4=D. C. |last5=Ultsch |first5=G. R. |date=2001 |title=The physiology of hibernation in common map turtles (Graptemys geographica) |journal=Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology |volume=130 |issue=2 |pages=331β340 |doi=10.1016/S1095-6433(01)00398-1 |pmid=11544078 }}</ref> One reason for the difference in duration between more anoxia-tolerant species and less anoxia-tolerant species is the turtle's ability to buffer lactic acid accumulation during anoxia.<ref>{{Cite journal |last1=Jackson |first1=D. C. |last2=Taylor |first2=S. E. |last3=Asare |first3=V. S. |last4=Villarnovo |first4=D. |last5=Gall |first5=J. M. |last6=Reese |first6=S. A. |date=2007 |title=Comparative shell buffering properties correlate with anoxia tolerance in freshwater turtles |journal=American Journal of Physiology. Regulatory, Integrative and Comparative Physiology |volume=292 |issue=2 |pages=R1008βR1015 |doi=10.1152/ajpregu.00519.2006 |pmid=17008457 |s2cid=6911719 }}</ref> Unlike adult painted turtles, hatchlings can survive only 40 days, but still exhibit high anoxia tolerance and freeze tolerance compared to other hatchling species (30 days for ''Chelydra serpentina'', and 15 days for ''Graptemys geographica'') due to cold winters.<ref name=":2">{{Cite journal |last1=Reese |first1=S. A. |last2=Ultsch |first2=G. R. |last3=Jackson |first3=D. C. |date=2004 |title=Lactate accumulation, glycogen depletion, and shell composition of hatchling turtles during simulated aquatic hibernation |journal=Journal of Experimental Biology |volume=207 |issue=17 |pages=3123 |doi=10.1242/jeb.207.17.3123 }}</ref><ref>{{Cite journal |last1=Dinkelacker |first1=S. A. |last2=Costanzo |first2=J. P. |last3=Lee |first3=R. E. |date=2005 |title=Anoxia tolerance and freeze tolerance in hatchling turtles |journal=Journal of Comparative Physiology B |volume=175 |issue=3 |pages=209β217 |doi=10.1007/s00360-005-0478-0 |pmid=15739066 |s2cid=12647870 }}</ref>{{Clear}} ===Movement=== [[File:Painted turtle California.jpg|thumb|right|alt=Painted turtle with green slime on its shell, on pebbles, with a couple of leaves on its back. Sun shining.|Moving on land]] Searching for water, food, or mates, the painted turtles travel up to several kilometers at a time.{{sfn|Ernst|Barbour|Lovich|1994|p=286}} During summer, in response to heat and water-clogging vegetation, the turtles may vacate shallow marshes for more permanent waters.{{sfn|Ernst|Barbour|Lovich|1994|p=286}} Short overland migrations may involve hundreds of turtles together.{{sfn|Carr|1952|p=217}} If heat and drought are prolonged, the turtles will [[Estivation|bury themselves]] and, in extreme cases, die.{{sfn|Ernst|Lovich|2009|p=195}} Foraging turtles frequently cross lakes or travel linearly down creeks.<ref>{{cite journal|last1=MacCulloch, R.D. and D.M. Secoy|title=Movement in a river population of ''Chrysemys picta bellii'' in southern Saskatchewan|journal=Journal of Herpetology |date=1983|volume=17|issue=3|pages=283β285|doi=10.2307/1563834|jstor=1563834}}</ref> Daily crossings of large ponds have been observed.{{sfn|Ernst|Lovich|2009|p=195}} [[Tracking animal migration|Tag and release]] studies show that sex also drives turtle movement. Males travel the most, up to {{convert|26|km|mi|abbr=on|0}}, between captures; females the second most, up to {{convert|8|km|mi|abbr=on|0}}, between captures; and juveniles the least, less than {{convert|2|km|mi|abbr=on}}, between captures.{{sfn|Ernst|Barbour|Lovich|1994|p=286}} Males move the most and are most likely to change wetlands because they seek mates.{{sfn|Ernst|Lovich|2009|p=195}} The painted turtles, through visual recognition, have [[Homing (biology)|homing]] capabilities.{{sfn|Ernst|Barbour|Lovich|1994|p=286}} Many individuals can return to their collection points after being released elsewhere, trips that may require them to traverse land. One experiment placed 98 turtles varying several-kilometer distances from their home wetland; 41 returned. When living in a single large body of water, the painted turtles can home from up to {{convert|6|km|mi|abbr=on|0}} away. Another experiment found that if placed far enough away from water the turtles will just walk in straight paths and not orient towards water or in any specific direction which indicates a lack of homing ability.<ref name="Caldwell Nams">{{cite journal |last1=Caldwell |first1=I.R. |last2=Nams |first2=V.O. |title=A compass without a map: tortuosity and orientation of eastern painted turtles (''Chrysemys picta picta'') released in unfamiliar territory |journal=Canadian Journal of Zoology |date=August 2006 |volume=84 |issue=8 |pages=1129β1137 |doi=10.1139/z06-102 }}</ref> Females may use homing to help locate suitable nesting sites.{{sfn|Ernst|Barbour|Lovich|1994|p=286}} Eastern painted turtle movements may contribute to aquatic plant seed dispersal. A study done in Massachusetts found that the quantity of intact macrophyte seeds defecated by Eastern painted turtles can be high and that the seeds of specifically ''Nymphaea ordorata'' that were found in feces were capable of moderate to high level germination. As turtles move between ponds and habitats, they carry seeds along with them to new locations.<ref name="Caldwell Nams"/> {{Clear}}
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