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Observations and measurements on the North American Wood Turtle: Glyptemys (Clemmys) insculpta with notation of two different phenotypes.
- A 1985 study
Typical dimensions of Glyptemys insculpta
Based on eight live specimens
Research conducted 1985
Carapace length Carapace width Head width Carapace- plastron length ("Carastron index"
inches / centimeters
Males -Yellowlegs phenotype
7 3/4 19.8 5 9/16 14.2 1 5/16 3.5 11/16 1.6
7 5/16 18.3 5 1/2 13.9 1 7/16 3.6 6/16 1.0
Males -redlegs phenotype
7 1/16 17.8 5 4/16 13.6 1 3/16 2.8 9/16 1.3
7 14/16 19.9 5 12/16 14.4 1 4/16 3.2 1 5/16 2.2
Females -yellowlegs phenotype
6 11/16 17.4 5 9/16 14.3 1 1/16 2.7 -1/16 -0.2
7 3/16 18.3 5 11/16 14.4 1 3/16 2.9 -2/16 -0.3
Females -redlegs phenotype
7 4/16 18.5 5 1/2 13.9 1 2/16 2.8 4/16 0.7
7 4/16 18.5 5 9/16 14.0 1 3/16 2.9 5/16 0.9
Abstract:
Research and observations were conducted in 1985 by Brian L. Schnirel on Glyptemys insculpta.
Live specimens were acquired and it was noted at that time that two distinct phenotypes were in
evidence with this species. The different color aspect of this species has been mentioned by Pope
(1938) and Harding (1997).
Both types of Glyptemys insculpta were measured for carapace and plastron length, head width,
and difference in length between the carapace and plastron ('carastron' index). Two of these
measurements (the head width and the 'carastron index are linked with sexual dimorphism of
Glyptemys insculpta. The two varieties were tested using variations of the Tinklepaugh labyrinth
experiment, Yerkes space reaction experiment, and mirror association experiments. Photographs
were taken at this time on these two phenotypes and were included in this report.
Introduction:
The third suborder of turtles to evolve from the Amphichelydia
are the Cryptodires- which are the most successful group of turtles
today. They possess the ability to pull their heads directly into their
shells without a full sideways motion. The largest family of
Cryptodirians are the Emydidae. Within this family, lies the genus
Glyptemys; which began in the Paleocene Epoch (55 million years
ago). The North American wood turtle (Glyptemys insculpta)- the
sculptured turtle- begins extensively in the Pleistocene
(approximately 1 million years ago). Voorhies (2000) reported a
discovery of a wood turtle shell from the Hemiphillian Miocene (6
million years ago. Pope (1939) mentions the remains of N.A. wood
turtles found in Pennsylvania that were at least ten's of thousands
of years old. As wiil be mentioned in the Geographic Range section,
ice age wood turtles were found in Tennessee and Georgia where
they have never been known from modern times. Since the advent
of Homo sapiens, the population has diminished considerably due to
overcollecting, highway deaths, extensive habitat destruction, etc.
In the past, native americans reduced some of the population for
rituals, shell helmets, rattles, and so on. For the white settler,
Insulptas were a convenient source of food. Indeed, in the early
part of the 20th century, quite a market developed for the 'Redleg'
as Glyptemys insculpta was called in the vernacular. Many
individuals were pulled out of muskrats holes in the winter using
sticks to locate something hard (Pope, 1939). The toughened
individual would then reach into the icy, cold water to claim his
prize.
Relationships:
The wood turtle, as stated belongs to the family Emydidae. The
Genus Clemmys, (as of 1985) consisted of the wood turtle Clemmys
insculpta, the bog turtle Clemmys muhlenbergi, the spotted turtle
Clemmys guttata, and the pacific pond Turtle Clemmys marmorata.
As 0f 2001, Mitochondrial DNA studies have now placed the wood
and bog turtles in the genus: Glyptemys and the pacific pond turtle
in the genus Actinemys. Only the spotted turtle remains in the now
monotypic genus Clemmys. The spotted turtle and the bog turtle
(although widely separated and rare) are also denizens of the
northeast. The pacific pond turtle is the most aquatic member of
this former group and resides along the Pacific coast.
An interesting example of parallel evolution or common ancestral
trait exists between the wood turtle and the semi-box turtle
Emydoidea blandingi , also known as the blanding turtle. The
plastron of both species is amazingly similar. The pattern is a
yellow-white background with black blotches on each of the scutes.
Other studies indicate a close genetic relationship between the two
(See 2003 update in Morphs and Ancestral Relationships). The semi-
box turtle, a cold hardy species, also resides in the northeast;
overlapping the range of the wood turtle. This turtle's range-like so
many others- is slowly shrinking. At one time, it may well indeed
have had the exact same range as the wood turtle.
Behavior and Intelligence:
Tinklepaugh (1932) conducted experiments to
determine the intellegence of the wood turtle. Using a
labyrinth, he concluded Glyptemys (Clemmys) insculpta had
the learning capacity of a rat. In personal experiments
and observations(B. Schnirel,1985,1998) the wood turtle
does indeed show great resourcefulness in problem
solving. One male in particular was quite adroit at
climbing and would systematically probe for weaknesses at
all levels in the outdoor fencing he was kept in. In a
variation to the labyrinth experiment, a three
dimensional approach was conducted in 1985. Ramps were
used to allow choices at 3 different heights. Wood
turtles in the experiment quickly learned the right route
to find food or females placed in a consistent area.
Yerkes (1901) conducted the famous 'space
reaction'experiments. Included on his guest list was
none other than our friend, the wood turtle. The
experiments tested the animal's fear of heights. What
was learned was the more aquatic the species, the more
careless the species is in regards to heights. This is
due to aquatic animals being used to dropping off stones,
logs, or whatever into boyount water. How did the wood
turtle make out? It was found to have less fear of
heights than the more aquatic spotted turtle. In tests
performed (B.Schnirel,1985) at various heights up to 20
feet, great fear and respect of height was demonstrated
by the wood turtle. The turtles would stick their heads
way over to inspect the untouchable surface with a note
of displeasure. More aquatic species would simply walk
straight off the edge. No turtles were hurt in the
experiments and special precautions were taken.
Glyptemys (Clemmys) insculpta appears more cognizant
of it's own reflections when facing mirrors. Experiments
conducted (B. Schnirel, 1985) with wood turtles showed
that individuals would stop, lower their heads and touch
'noses' with their reflections. Other turtles and
tortoises tested (Geocheleone carbonaria, Geocheleone
denticulata,Geochelone chilensis, Testudo horsfieldi,
Kinixys belliana belliana, Gopherus polyphemus, Terrapene
carolina carolina,Terrapene carolina bauri,and
Rhinoclemmys pulcherima showed no interest in their
mirrored image.
Results:
Dimensional Parameters:
The 1985 measurements on the eight insculpta's showed the following pertaining to sexual
dimorphism: Carapace length - males: 7 1/16 - 7 14/16 inches (17.8 - 19.9 centimeters).
Carapace length - females: 6 11/16 - 7 4/16 inches (17.4 - 18.5 centimeters). All test subjects
were mature adults and as such, males showed a larger ultimate size. Carapace width - males:
5 4/16- 5 12/16 inches (13.6 - 14.4 centimeters). Carapace width - females: 5 1/2 - 5 11/16
inches (13.9 - 14.4 centimeters). Again, males were slightly larger. Width of head - males:
1 3/16 - 1 7/16 inches (2.8 - 3.6 centimeters). Width of head - females: 1 1/16 - 1 3/16 inches
(2.7 - 2.9 centimeters). Males show a significant difference. Carapace minus Plastron length
(Carastron index) - Males: 6/16 - 15/16 inches ( 1.0 - 2.2 centimeters). Carapace minus
Plastron length (Carastron index) - females: 5/16 - 2/16 inches ( 0.9- -0.3 centimeters). This
was a sizeable difference with males possessing a large carapace overhang. In subsequent
observations, this is always the case. Female redlegs phenotype insculpta have a narrower
overhang but in the case of the yellowlegs phenotype insculpta, the plastron was actually longer
than the carapace (giving a minus Carastron value). One yellowlegs female that had a Carastron
value of -.06 (not included in parameter table) was unusual in having a completely round
carapace similar to a dinner plate. One thought is that she was a member of a peripheral isolate
population. Another thought was paedomorphism of the carapace. Hatchling
Glyptemys insculpta display a round carapace which develops into a more oval shape during
development towards adulthood.
Results: (Continued)
Tinklepaugh Labyrinth Experiments:
Glyptemys insculpta indeed showed great resourcefulness and speed in problem solving. In
addition to finding escape routes quickly, subjects in variation quickly found correct pathways
to food and male Glyptemys insculpta to females in different sections of the labyrinth. Male
insculpta had been observed in 1985 lying in ambush near pools for females to happen by
looking to soak or a drink. The male would proceed to jump on the back of the female to
mate. A three dimensional approach in the labyrinth experiments did not slow down the
wood turtles in their quest. One male in particular was very adroit in climbing and would
systematically probe for weaknessess. This was done at all levels in the outdoor fencing he
was kept in.
Yerkes Space Reaction Experiments:
At all levels up to twenty feet, great fear and respect of heights was demonstrated by
Geochelone carbonaria. Glyptemys insculpta and Terrapene carolina carolina showed
respect for all distances but did not show fear at the two foot height. Clemmys guttata
showed respect and fear at ten and twenty feet, but did not so at two and five feet. Indeed, at
the latter two heights, the subjects tried to drop off as aquatic species would be expected to
into buoyant water.
Results: (Continued)
Mirror Experiment:
All test species with the exception of Glyptemys insculpta, showed no interest in their reflected
images. Glyptemys insculpta appeared more cognizant of it's own reflections when facing
mirrors. test subjects would stop, lower their heads, and touch 'noses' with their reflections.
This behavior has been noted between encounters between two Glyptemys insculpta. In addition
to the head lowering and nose tipping, individuals will often tip their heads and sniff each other
in the manner of dogs. They seem to treat each other on an individual basis. Knowlton describes
the behavior of insculpta in an encounter in the following manner: "Their heads were slowly
lowered when the turtles were with eight inches of each other. Next, their heads were
swung from side to side for up to an hour without cessation. Mating occurred in a nearby
pool. " Harding also mentions that Glyptemys insculpta conduct a mating dance initiated by
both parties. They position themselves in front of one another and swing their heads in a
sideways swing.
Text (Introduction):
The North American wood turtle belongs to the Family Emydidae. The Genus Clemmys
(as of 1985) consisted of the wood turtle Clemmys insculpta, the bog turtle
Clemmys muhlenbergi, the spotted turtle Clemmys guttata, and the Pacific pond turtle
Clemmys marmorata. As of 2002, mitochrondrial DNA studies have now placed the wood
and bog turtles in the Genus: Glyptemys and the pacific pond turtle in the Genus: Actinemys.
Only the spotted turtle remains in the now monotypic Genus: Clemmys.
In 1985, several live Glyptemys insculpa were acquired for study. Individuals were selected
from extreme ends of their natural range. Some individuals came from Wisconsin
(Oneida county). This represented the western end of the range. Other individuals hailed
from Northern Virginia. This represented the eastern as well as the southern part of their range.
It immediately became apparent that there were consistent differences in coloration between the
geographically separated groups. The eastern group consisted of the standard reddish orange
color phase known in the vernacular as the "redlegs" (Pope, 1939). This standard morph also is
characteristic of having a yellow-ringed iris. The western group of Glyptemys insculpta was
different in that the skin color was yellow instead of reddish-orange and the iris was entirely
black with no yellow ring. This phenotype will be referred to as the "yellowlegs" for the duration
of this paper. Photographs of both phenotypes were taken and accompany this report. In the
early twentieth century, Clifford Pope mentions having specimens of the yellowlegs phenotype
which he describes as follows: " The soft parts lack the normal salmon red color of the adult,
Text (Introduction) - Continued:
the plastron is dark except for a narrow light margin and a little light mottling ".
Harding (1997) also mentions the color variation aspect and geographical distinctiveness of
Glyptemys insculpta.
Measurements were recorded on eight selected individuals (4 males, 4 females- divided by 2
from each region). The data recorded was significant in regards to sexual dimorphism.
Typically, male Glyptemys insculpta are noted for having wider heads, carapace's significantly
longer than the plastron (always), longer claws, concave plastron, and thicker tails with the vent
further out from the shell than the female. Female Glyptemys insculpta posses narrower heads,
a carapace slightly longer than the plastron or in some cases, the plastron is longer than the
carapace, shorter claws, flat plastron, and thinner tails with the vent close to the edge of the
shell.
Tinklepaugh (1932) conducted experiments to determine the intelligence of Glyptemys insculpta.
He concluded that this species had the learning capacity of a rat. The 1985 experiments of the
same were done with a labyrinth constructed of 2 x 4's. The width of the corridors were ten
inches wide with area covering twenty feet by twenty feet. In a variation of the original
experiment, a three dimensional approach was conducted. Ramps were used to allow choices
at three different levels. Incentives included food and females placed at specific locations.
Text (Introduction) - Continued:
Yerkes (1901) conducted the famous 'space' reaction experiments. Included on his guest list was
Glyptemys insculpta. The experiment tested the animal's fear of heights. What was learned was
the more aquatic the species, the more careless the species is in regards to heights. This behavior
has evolved due to aquatic turtles being used to dropping off stones, logs, or whatever into
buoyant water. The following test heights were chosen: two feet, five feet, ten feet, and twenty
feet. In addition to Glyptemys insculpta, other species tested were the spotted turtle:
Clemmys guttata, the redfoot tortoise: Geochelone carbonaria, and the Eastern box turtle,
Terrapene carolina carolina.
The last series of experiments involved reaction to reflections in mirrors. The mirror used was
twenty four inches wide. In addition to the North American wood turtle: Glyptemys insculpta,
the following chelonian species were tested: the redfoot tortoise: Geochelone carbonaria, the
yellowfoot tortoise: Geochelone denticulata, The Chaco tortoise: Geochelone chilensis, the
Afghan tortoise: Testudo horsefieldi, the hingeback tortoise: Kinixys belliana, The Eastern
gopher tortoise: Gopherus polyphemus, the Eastern box turtle: Terrapene carolina carolina,
the Florida box turtle: Terrapene carolina bauri, and the Central American wood turtle:
Rhinoclemmys pulcherima. Discussion:
The differences between the redlegs and yellowlegs phenotypes of Glyptemys insculpta are suttle
but distinctive. Present evidence suggests the yellowlegs phenotype resides in the western part
of the biozone. The yellowlegs form may exist as peripheral isolates evolving eventually into a
new subspecies if not disturbed by Homo sapiens. However, an alternative view may have the
yellowlegs as ancestral stock. Harding (1999) mentions hybrids between Emydoidea blandingi
and Glyptemys insculpta. The range of these two species overlaps with the greatest core
concentration (as of 1985) remaining in the Great Lakes region. During the 1985 research,
individuals of Emydoidea blandingi were present and studied with similarities noted with
Glyptemys insculpta. The plastron of both species is very similar with the same arrangement
of black spots on the plastral scutes. The skin color is yellowish on blandingii as is with the
yellowlegs insculpta. It might be possible that a common emydid ancestor to both species
resided in the same region and the ancestral color trait being yellow in regards to skin color.
This would be a pre-Miocene event as the earliest known wood turtle (an adult male) found
(Voorhies, 2000) was in the Hemphillian Miocene. If this is the case, the redlegs phenotype
would be a derived apomorphic trait.
Ernest, Lovitch and Barborn in 1994 reported that in the cooler climes of
the ice age, the wood turtle had a more southern distribution. The evidence
suggested that the species could be found as far south as Tennessee and
Georgia.
The noted differences in the phenotypes with dimensional measurements were the females with
the Carastron values. Plastrons seem to be longer than the carapace with the yellowlegs
phenotype. However, more measurements on more yellowlegs and redlegs individuals would
be needed to draw an absolute conclusion.
Discussion: (Continued)
In behavior, both phenotypes acted along similar lines and showed no differences in intelligence.
This was true for the Tinklepaugh labyrinth experiment, the Yerkes space reaction experiment
and the mirror reflection experiment.
References:
Behler/King. 1979. Audubon Society field guide to North American reptiles and amphibians. Knopf, New York. 744 pp.
Bickman, J. 1996. Molecular systematics of the genus Clemmys and the intergeneric Lamb, T. Minx, P. relationships of Emydid turtles. Herpetologica. Bainbridge, Ga. Patton, J. 521 (1) pp.89-97.
Burton, M. 1973. The world of reptiles and amphibians. Bounty Books, London-New York. 128 pp.
Carr, A.F. 1952. Handbook of turtles. Comstock Publ. Associates, Cornell University Press. Ithaca, New York. 542 pp.
Conant, R. 1958. A field guide to reptiles and amphibians of Eastern North America. Houghton Mifflin Co., Boston Massachusetts. 366 pp.
Ernest, C. H./ 1972. Turtles of the United States. University of Kentucky Press. 347 pp. Barbour, R.W.
Harding, James 2003. Clemmys Insculpta. Animal Diversity Web - Michigan State University Museum of Zoology 7 pp.
Lanworn, R.A. 1972. The book of reptiles. Hamlin, London. 127 pp.
Pope, C. H. 1939. Turtles of the United States and Canada. Alfred A. Knolpf, New York. 343 pp.
Pritchard, P. 1979. Encyclopedia of turtles. T.F.H. Publications, Neptune, New Jersey. 895 pp.
Roberts, M. 1980. Turtles. T.F.H. Publications, Neptune, New Jersey. 93 pp.
Tinklepaugh, 1932. Maze learning of a turtle. Journal Comprehensive Psychology 13: O. pp. 210-206.
Zim, H. 1953. Reptiles and amphibians (North America). Golden Press, New
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