The turtle shell is a highly complicated shield for the ventral and dorsal parts of turtles, tortoises and terrapins (all classified as "turtles" by zoologists), completely enclosing all the vital organs of the turtle and in some cases even the head. It is constructed of modified bony elements such as the ribs, parts of the pelvis and other bones found in most reptiles. Therefore turtles have no backs. The bone of the shell consists of both skeletal and dermal bone, showing that the complete enclosure of the shell probably evolved by including dermal armor into the rib cage.
The shell of the turtle is an important study, not just because of the obvious protection it provides for the animal, but also as an identification tool, in particular with fossils as the shell is one of the likely parts of a turtle to survive fossilization. Hence understanding the structure of the shell in living species gives us comparable material with fossils.
The shell of the hawksbill turtle, among other species, has been used as a material for a wide range of small decorative and practical items since antiquity, but is normally referred to as tortoiseshell.
The turtle shell is made up of numerous bony elements, generally named after similar bones in other vertebrates, and a series of keratinous scutes which are also uniquely named. The top of the shell is known as the carapace, and the ventral surface is called the plastron. These are joined by an area called the bridge. The actual suture between the bridge and the plastron is called the anterior bridge strut. In Pleurodires the posterior pelvis is also part of the carapace, fully fused with it. This is not the case in Cryptodires which have a floating pelvis. The anterior bridge strut and posterior bridge strut are part of the plastron, on the carapace are the sutures into which they insert, known as the Bridge carapace suture.
The bones of the shell are named for standard vertebrate elements. As such the carapace is made up of 8 pleurals on each side, these are a combination of the ribs and fused dermal bone. Outside of this at the anterior of the shell is the single nuchal bone, a series of 11 paired periphals then extend along each side. At the posterior of the shell is the pygal bone and in front of this nested behind the eighth pleurals is the suprapygal.
Between each of the pleurals are a series of neural bones, which although always present are not always visible, in many species of Pleurodire they are submerged below the pleurals. Beneath the neural bone is the Neural arch which forms the upper half of the encasement for the spinal chord. Below this the rest of the vertebral column. Some species of turtles have some extra bones called mesoplastra, these are located between the carapace and plastron in the bridge area. They are present in most Pelomedusid turtles.
The skeletal elements of the plastron are also largely in pairs. Anteriorly there are two epiplastra, with the hyoplastra behind them. These enclose the singuar entoplastron. These make up the front half of the plastron and the hyoplastron contains the anterior bridge strut. The posterior half is made up of two hypoplastra (containing the posterior bridge strut) and the rear is a pair of xiphiplastra.
Overlying the boney elements are a series of scutes, which are made of keratin and are a lot like horn or nail tissue. In the center of the carapace are 5 vertebral scutes and out from these are 4 pairs of costal scutes. Around the edge of the shell are 12 pairs of marginal scutes. All these scutes are aligned so that for the most part the sutures between the bones are in the middle of the scutes above. At the anterior of the shell there may be a cervical scute (sometimes incorrectly called a nuchal scute) however the presence or absence of this scute is highly variable, even within species.
On the plastron there are two gular scutes at the front, followed by a pair of pectorals, then abdominals, femorals and lastly anals. A particular variation is the Pleurodiran turtles have an intergular scute between the gulars at the front, giving them a total of 13 plastral scutes. Compared to the 12 in all Cryptodiran turtles.
The carapace is the dorsal (back), convex part of the shell structure of a turtle, consisting of the animal's ribcage combined with dermal bone. The spine and ribs are fused to dermal plates beneath the skin which interlock to form a hard shell. Exterior to the skin the shell is covered by scutes, which are horny plates made of keratin that protect the shell from scrapes and bruises. A keel, a ridge that runs from front to the back of the animal is present in some species, these may be single, paired or even three rows of them. In most turtles the shell is relatively uniform in structure, species variation in general shape and color being the main differences. However the soft shell turtles, pig-nose turtles and the leatherback sea turtle have lost the scutes and reduced the ossification of the shell. This leaves the shell covered only by skin. These are all highly aquatic forms.
The evolution of the turtle's shell is unique because of how the carapace represents transformed vertebrae and ribs. While other tetrapods have their scapula, or shoulder blades, found outside of the ribcage, the scapula for turtles is found inside the ribcage. The shells of other tetrapods, such as armadillos, are not linked directly to the vertebral column or rib cage allowing the ribs to move freely with the surrounding intercostal muscle. However, analysis of the transitional fossil, Eunotosaurus africanus shows that early ancestors of turtles lost that intercostal muscle usually found between the ribs.
In the figure to the right, the exploded shell view, the roman numerals and other notations are as follows:
(i) Neural 1, (ii) Neural 2, (iii) Neural 3, (iv) Neural 4, (v) Neural 5, (vi) Neural 6, (vii) Neural 7, (viii) Neural 8, (ix) extra neural, divided, (x) suprapygal, (xi) nuchal, (xii) right peripheral 1, (xiii) right peripheral 2,
(xiv) right peripheral 3, (xv) right peripheral 4, (xvi) right peripheral 5, (xvii) right peripheral 6, (xviii) right peripheral 7, (xix) right peripheral 8, (xx) right peripheral 9, (xxi) right peripheral 10, (xxii) right peripheral 11, (xxiii) pygal, (xxiv) left peripheral 11, (xxv) left peripheral 10, (xxvi) left peripheral 9, (xxvii) left peripheral 8, (xxviii) left peripheral 7, (xxix) left peripheral 6, xxx left peripheral 5, xxxi left peripheral 4, (xxxii) left peripheral 3, (xxxiii) left peripheral 2, (xxxiv) left peripheral 1, (xxxv) right 1st rib, (xxxvi) right pleural 1, (xxxvii) right pleural 2, (xxxviii) right pleural 3, (xxxix) right pleural 4, (xl) right pleural 5, (xli) right pleural 6, (xlii) right pleural 7, (xliii) right pleural 8, (xliv) right 10th rib, (xlv) left 1st rib, (xlvi) left pleural 1, (xlvii) left pleural 2, (xlviii) left pleural 3, (xlix) left pleural 4, (l) left pleural 5, (li) left pleural 6, (lii) left pleural 7, (liii) left pleural 8, (liv) left 10th rib, (9-18) centrums.
The plastron is the nearly flat part of the shell structure of a turtle, what one would call the belly or ventral surface of the shell. It also includes within its structure the anterior and posterior bridge struts and the bridge of the shell. The plastron is made up of nine bones and the two epiplastra at the anterior border of the plastron are homologous to the clavicles of other tetrapods. The rest of the plastral bones are homologous to the abdominal ribs of other tetrapods.
In certain families there is a hinge between the pectoral and abdominal scutes allowing the turtle to almost completely enclose itself. In certain species the sex of a testudine can be told by whether the plastron is concave, male or convex, female. This is because of the mating position; the male's concave plastron allows it to more easily mount the female during copulation.
The plastral scutes join along a central seam down the middle of the plastron. The relative lengths of the seam segments can be used to help identify a species of turtle. There are six laterally symmetric pairs of scutes on the plastron: gular, humeral, pectoral, abdominal, femoral, and anal (going from the head to the tail down the seam); the abdominal and gular scute seams are approximately the same length, and the femoral and pectoral seams are approximately the same length.
The gular scute or gular projection on a turtle is the most anterior part of the plastron, the underside of the shell. Some tortoises have paired gular scutes, while others have a single undivided gular scute. The gular scutes may be referred to as a gular projection if they stick out like a trowel.
- Gular (disambiguation), gular anatomical formations in other species
The plastral formula is used to compare the sizes of the individual plastral scutes (measured along the midseam). The following flastral scutes are often distinguished (with their abbreviation):
- intergular = intergul
- gular = gul
- humeral = hum
- pectoral = pect
- abdominal = abd
- femoral = fem
- anal = an
The turtle's shell is covered in scutes that are made of keratin. The individual scutes as shown above have specific names and are generally consistent across the various species of turtles. Terrestrial tortoises do not shed their scutes. New scutes grow by the addition of keratin layers to the base of each scute. Aquatic chelonia shed individual scutes. The scute effectively forms the skin over the underlying bony structures; there is a very thin layer of subcutaneous tissue between the scute and the skeleton. The scutes can be brightly colored in some species, but the basal color is a grey to dark brown color dorsally; the plastral scutes are often white to yellow in base color.
Septicemic cutaneous ulcerative disease (SCUD)
Originally described by Kaplan (1957) is often known by its acronym SCUD. This disease is known to be caused by a variety of bacteria entering through some sort of abrasion, combined with poor animal husbandry. The disease is identified by its progression and what starts as ulcerative lesions of the plastron leads to a septacemic infection causing the degradation of the liver and other organs. Without treatment, this will lead to death.
Ulcers of the shell may be superficial or deep, and may be termed "shell rot." Ulcers are generally a result of poor husbandry. Turtles with ulcerative shell lesions should be examined and treated by a veterinarian, as the ulcers may become infected and penetrate through the shell. The shell will need to be cleaned daily, and dead tissue removed. Topical and/or injectable antibiotics are required in the case of bacterial infections. Deep ulcers may need to be repaired through surgery and the application of acrylic or fiberglass material. In a disease called "septicemic cutaneous ulcerative disease," or SCUD, ulcers may be seen on both the shell and legs. This condition is often associated with the bacteria, Citrobacter freundii.
Pyramiding is a shell deformity commonly found in captive turtles, in which the shell grows unevenly resulting in a "pyramid" shape underlying each scute. This deformity can vary in severity from barely noticeable to life-threatening. Note that Indian star tortoises and some other species display this morphology naturally.
Several causes can lead to pyramiding, including excessive animal or vegetable protein; inadequate calcium, sunlight, and/or vitamin D3; and excessive moisture / poor nutrition foods. Once pyramiding has occurred, it cannot be reversed, though if the underlying problem is corrected, the animal may still lead a long, healthy life.
Further information on pyramiding:
- Romer, A.S. (1956) Osteology of the Reptiles. Univ. of Chicago Press.
- Zangerl, R. 1969. The turtle shell. In: Gans, C., Bellairs, D.d'A. and Parsons, T.A. (Eds). Biology of the Reptilia, Vol 1, Morphology A. London: Academic Press. pp. 311–340
- Thomson, S., White, A. & Georges, A (1997). "Re-Evaluation of Emydura lavarackorum: Identification of a Living Fossil". Memoirs of the Queensland Museum 42 (1): 327–336.
- Pritchard, P.C.H. (1988). "A survey of neural bone variation among recent chelonian species, with functional interpretations". Acta Zoologica Cracoviensa 31 (26): 625–686.
- Thomson, S. and Georges, A. (1996). "Neural bones in chelid turtles". Chelonian Conservation and Biology 2: 82–86.
- Rhodin, A.G.J. and Mittermeier, R.A. (1977). "Neural bones in chelid turtles from Australia and New Guinea". Copeia 1977 (2): 370–372. doi:10.2307/1443917. JSTOR 1443917.
- Pritchard, P.C.H., and P. Trebbau. 1984. The Turtles of Venezuela. SSAR Contributions to Herpetology 2:.
- Bojanus, L. H. 1819. Anatome testudinis Europaeae. 178pp, 31 plates
- Nagashima, H., F. Sugahara, M. Takechi, R. Ericsson, Y. Kawashima-Ohya, Y. Narita, and S. Kuratani. 2009. Evolution of the turtle body plan by the folding and creation of new muscle connections. Science 325:193-196.
- Wang, Z., J. Pascual-Anaya, A. Zadissa, W. Q. Li, Y. Niimura, Z. Y. Huang, C. Y. Li et al. 2013. The draft genomes of soft-shell turtle and green sea turtle yield insights into the development and evolution of the turtle-specific body plan. Nature Genetics 45:701-+.
- Hirasawa, T., H. Nagashima, and S. Kuratani. 2013. The endoskeletal origin of the turtle carapace. Nature Communications 4.
- Lee, M. S. Y. 2013. Palaeontology: Turtles in transition. Current Biology 23:R513-R515.
- Gilbert, S. F., G. A. Loredo, A. Brukman, and A. C. Burke. 2001. Morphogenesis of the turtle shell: The development of a novel structure in tetrapod evolution. Evolution & Development 3:47-58.
- C.H. Ernst, R.G.M. Altenburg and R.W. Barbour. "Terrapene carolina". Netherlands Biodiversity Information Facility. Retrieved 12 February 2011.
- Kaplan, H. M. (1957). "Septicemic, cutaneous ulcerative disease of turtles". Proc. Animal Care Panel 7: 273–277.
- Mader, D. (2006) Reptile Medicine and Surgery, 2nd ed., Saunders, ISBN 072169327X.