The spiny softshell turtle (Apalone spinifera) is a species of softshell turtle, one of the largest freshwater turtle species in North America. Both the common name, spiny softshell, and the specific name, spinifera (spine-bearing), refer to the spiny, cone-like projections on the leading edge of the carapace, which are not scutes (scales).
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Scientific classification 
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Order: | Testudines |
| Suborder: | Cryptodira |
| Family: | Trionychidae |
| Genus: | Apalone |
| Species: | A. spinifera
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| Binomial name | |
| Apalone spinifera | |
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Description
editThe spiny softshell turtle's scientific name is very descriptive of the animal. Apalone comes from the Greek word apalos, meaning soft or tender, and spinifera is of Latin origin; spina- referring to thorn or spine and -ifer meaning bearing. This species is a member of the family Trionychidae, and one of the most distinguishing features of members in this family is the presence of a leathery, moderately flexible carapace.[3] This is caused by loss of keratinized scutes and some bony tissue loss. Spiny softshell turtles have webbed feet, each with three claws. Another distinguishing feature of softshell turtles is the presence of a fleshy, elongated nose.
The carapace (the upper part of the shell) ranges from brown or yellow-brown to olive in color, while the plastron (lower part of the shell) is lighter, usually white or yellow. Hatchlings usually have dark spots on the carapace, but as females age, they frequently become darker in color, or their carapace becomes splotched. Males tend to maintain the same coloration pattern from birth. Coloration also varies between each subspecies, and the exact coloration can also depend on an individual turtle's environment. Spiny softshell turtles are cryptically colored, meaning that their coloration helps them blend in with their surrounding environment.
Spiny softshell turtles also have pale lines bordered by black lines running from its head down the side of its neck. The carapace length ranges from 18 to 54 cm (7.1–21.3 in), with females growing larger than males. The namesake spines are found along the anterior border of the carapace and are more commonly found in males. The variation in coloration, size, and spine presence indicates that this species exhibits sexual dimorphism.
A. spinifera has ridges on its nasal septum, making its nostrils c-shaped, unlike the nostrils of the similar species A. mutica which are round.[4]
Distribution
editGeographic range
editThis species is the most widely distributed softshell turtle in North America.[5] This could be due to habitat fragmentation caused by humans.[6]The spiny softshell has a wide range, extending throughout much of the United States, as well as north into the Canadian provinces of Ontario and Quebec, and south into the Mexican states of Tamaulipas, Nuevo León, Coahuila and Chihuahua.[1] As of recently, the spiny softshell has expanded its geographic range (through human assistance) into Washington State and California. They were also recently discovered in Lake Champlain despite the historical records.[7]
Ecological range
editSpiny softshell turtles are often referred to as ecological generalists, meaning that they are found in a wide variety of habitats.[8] The spiny softshell can be found in bodies of fresh water including ponds, lakes, rivers, tributaries, and streams. They can persist in more urban environments as they are well adapted to periodic habitat disturbances.[9] They inhabit shallow water less than 1 m (3 ft 3 in) deep, but can also be found as far as 10 m (33 ft) deep. When swimming, a study suggests that they get most of their thrusting power from their forelimbs as opposed to their hindlimbs, which is common in other species.[10] They can be found in areas with varying levels of vegetation, and although they are generally found in more slow-moving waters, this species abundance is greatest in waters with higher visibility and slower water velocity [11] Spiny softshells prefer waters with sandy bottoms and clean, sandy banks. Sandy environments are important for nesting sites, proper juvenile growth and development, and camouflage.[12] Spiny softshells migrate between warm and cold seasons. In each season, turtles generally stay in a single zone, and they move more within their zone during warm months. The spiny softshell will migrate between their zones and their average home range length is 10.8 km (6.7 mi).[13]
As far as home range, a study of the "eastern spiny" subspecies (which has the largest and most northern-reaching distribution) home range behaviour found that turtles of northern Lake Champlain generally had two annual concentration areas for spring-summer and fall-winter, contributing to a large home range. Mean annual home range was also found, in the same study, to be over ten times larger for female softshells compared to males. It was suggested in this study that the high home range was due to habitat fragmentation.[14]
Behavior
editDiet
editSpiny softshell turtles feed on a variety of food items. They are primary consumers and feed on invertebrates (crayfish and aquatic insects), fish,[15] algal stocks and other plant material, and mussels. They are generally observed as benthic feeders; they can either actively hunt prey or bury themselves in the sand and wait to ambush prey.[16]
Some evidence suggests that spiny softshell turtles exhibit a nuclear-follower foraging association with fish. A study that took place in an urban drainage canal in Louisiana found that when foraging, spiny softshell turtles were observed moving along the creek bottom thrusting their probosces into the substrate which then allowed fish to enter the suspended sediment and capture prey otherwise unavailable to the fish. The fish benefit from increased foraging success, while few benefits accrue to the foraging turtles.[17]
Reproduction
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Spiny softshells begin mating between ages 8 and 10. A large female turtle may live up to 50 years. The turtles mate in mid-to-late spring in deep water. The male will nudge the female's head while swimming, and if she chooses to mate, the male will swim above the female without clasping her with his claws (unlike other turtles). Prior to nesting, the females have been observed to have a 3 stage activity pattern throughout the day. The females start the day by basking and moving around an area, then they spend a majority of their afternoon swimming in a new area, and finally they would stop and search for an ideal nesting spot.[18] A few months later, the female turtle quickly lays her eggs along a sunny sandbar or gravel bank in a flask-shaped cavity she has dug close to the water. This nesting behavior typically begins around July with the females leaving the water and probing the ground with her snout to find the spot to lay her eggs.[19] The turtle nests more than once during a single season. She can lay between 9 and 38 round, calcareous-shelled eggs. The eggs are laid around July and September, and they hatch in the spring. Larger bodied females have been found to lay a second clutch in late June or July.[20]In studies observing nesting behavior, it has been found that the females are more likely to lay eggs on days where there is a small difference between the air and water temperatures.[21] Unlike in some other turtles, in the spiny softshell turtle, the sex of the hatchlings is not determined by temperature variations; it is determined by genetics.[22]
Late-term embryos and hatchlings of spiny softshell turtles have been recently found to make various click- and chirp-type sounds before emergence onto the surface. The function of these sounds is currently unknown.[23]
Respiration
editSpiny softshell turtles are bimodal breathers, meaning they have the ability (to some degree) to perform oxygen and carbon dioxide exchange by breathing air or while breathing underwater. A variety of factors allow for these turtles to perform respiration underwater. They have an increase of cutaneous surface area and blood flow, reduction in lung size, and increase of respiratory epithelium in the cloaca and buccopharynx. This turtle is also known to utilize cutaneous respiration as well.[24] Spiny softshell turtles are more dependent on underwater respiration than other freshwater species. This has led to their low tolerance of hypoxic waters; this becomes especially important during times of hibernation, when these turtles must choose hibernacula that are unlikely to become hypoxic. Underwater dormancy can last up to six months.[25] They are known as an anoxia-intolerant species because of their dependency on underwater respiration. Other turtle species are more adapted to variation in oxygen levels but spiny softshell turtless are not able to regulate it as well, especially during hibernation periods. This makes choosing hibernacula very important to their over-winter survival. While they are susceptible to anoxia, they have the ability to maintain their metabolism completely via aerobic means which gives them a slight advantage when compared to other hibernating aquatic turtles during winter.[26]
Conservation
editOverall, the spiny softshell turtle is widespread, common and not threatened,[1] but some local populations are under pressure. In Canada, which is at the northern margin of its range and where the species is quite local, it is considered endangered.[27] The very rare black spiny softshell turtle, a subspecies found only in Mexico's Cuatro Ciénegas Basin, is considered critically endangered.[28]
Taxonomy
editThe species was first described by Charles Alexandre Lesueur in 1827.[1] It has been redescribed numerous times, leading to some confusion in its taxonomy.[29]
Subspecies
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The recognized subspecies differ in the markings on their carapaces, on the sides of their heads, and on their feet, although there are also considerable individual (not related to subspecies) variations in the appearance. Their markings, which are distinct as hatchlings, fade as the turtles grow larger. Adult females of the various subspecies, which grow larger than males, are not easily distinguishable from one another, and sometimes can only be assigned to a particular subspecies based on geography.[30][4]
Six subspecies of A. spinifera are recognized, including the nominate subspecies:[31]
- Northern spiny softshell turtle or eastern spiny softshell,[32] A. s. spinifera (Lesueur, 1827)[1]
- Gulf Coast spiny softshell turtle, A. s. aspera (Agassiz, 1857)[1]
- Black spiny softshell turtle or Cuatro Cienegas softshell turtle, A. s. atra (Webb & Legler, 1960)[1]
- Texas spiny softshell turtle, A. s. emoryi (Agassiz, 1857)[1]
- Guadalupe spiny softshell turtle, A. s. guadalupensis (Webb, 1962)[1]
- Pallid spiny softshell turtle, A. s. pallida (Webb, 1962)[1]
A previously recognized subspecies, Apalone spinifera hartwegi (Conant & Goin, 1941), has been synonymized to A. s. spinifera as of 2011.[1]
Genomics
editA rough-draft assembly of the A. spinifera aspera genome was completed in 2013 by the Genome Institute at Washington University in St. Louis. The assembly ASM38561v1 can be accessed via its Genbank accession ID APJP00000000.1 [1]
References
edit- ^ a b c d e f g h i j k l van Dijk et al. (2011), p. 000.206.
- ^ Fritz & Havaš (2007), pp. 306–310.
- ^ Engstrom, T. (2004). "Multiple data sets, high homoplasy, and the phylogeny of softshell turtles (Testudines: Trionychidae)". Systematic Biology. 53 (5): 693–710. doi:10.1080/10635150490503053. PMID 15545250.
- ^ a b Powell, Robert; Conant, Roger; Collins, Joseph T. (2016). A Field Guide to Reptiles and Amphibians of Eastern and Central North America, Fourth Edition. Boston and New York: Houghton Mifflin Harcourt. ISBN 978-0-544-12997-9. xiv + 494 pp., 47 plates, 207 figures. (Apalone spinifera, pp. 232–233 + Plates 20, 22 + Figures 85, 105, 106).
- ^ Lazure, Louis; Pare, Patrick; Bouthillier, Lyne; Galois, Patrick (2019). Nesting biology and conservation of a northern population of spiny softshell turtles (Apalone spinifera) (PDF) (3 ed.). Herpetological Conservation and Biology.
- ^ Galois, Patrick, et al. “Movement Patterns, Activity, and Home Range of the Eastern Spiny Softshell Turtle (Apalone Spinifera) in Northern Lake Champlain, Québec, Vermont.” Journal of Herpetology, vol. 36, no. 3, Sept. 2002, p. 402, https://doi.org/10.2307/1566184.
- ^ Galois, Patrick; Léveillé, Martin; Bouthillier, Lyne; Daigle, Claude; Parren, Steve; Leveille, Martin (September 2002). "Movement Patterns, Activity, and Home Range of the Eastern Spiny Softshell Turtle (Apalone spinifera) in Northern Lake Champlain, Quebec, Vermont". Journal of Herpetology. 36 (3): 402. doi:10.2307/1566184. ISSN 0022-1511. JSTOR 1566184.
- ^ Plummer, Michael; Mills, Nathan (2015). Growth and maturity of spiny softshell turtles (Apalone spinifera) in a small urban stream (PDF) (2 ed.). Herpetological Conservation and Biology.
- ^ Plummer, Michael V.; Krementz, David G.; Powell, Larkin A.; Mills, Nathan E. (September 2008). "Effects of Habitat Disturbance on Survival Rates of Softshell Turtles (Apalone spinifera) in an Urban Stream". Journal of Herpetology. 42 (3): 555–563. doi:10.1670/07-217.1. ISSN 0022-1511. S2CID 51770178.
- ^ Cinnamon M. Pace, Richard W. Blob, Mark W. Westneat; Comparative kinematics of the forelimb during swimming in red-eared slider (Trachemys scripta) and spiny softshell (Apalone spinifera) turtles. J Exp Biol 1 October 2001; 204 (19): 3261–3271. doi: https://doi.org/10.1242/jeb.204.19.3261
- ^ Barko, Valerie (2006). "Midland smooth softshell (Apalone Mutica) and Spiny Softshell (Apalone Spinifera) Turtles in the Middle Mississippi River: Habitat Associations, Population Structure, and Implications for Conservation". Chelonian Conservation and Biology. 5 (2): 225–31. doi:10.2744/1071-8443(2006)5[225:MSSAMA]2.0.CO;2. S2CID 85571504.
- ^ Plummer, M. (2008). "Effects of habitat disturbance on survival rates of softshell turtles (Apalone spinifera) in an urban streatm". Journal of Herpetology. 42 (3): 555–563. doi:10.1670/07-217.1. S2CID 51770178.
- ^ "Environment and climate change". Canada.ca. / Gouvernement du Canada. 21 December 2018.
- ^ Patrick Galois, Martin Léveillé, Lyne Bouthillier, Claude Daigle, Steve Parren "Movement Patterns, Activity, and Home Range of the Eastern Spiny Softshell Turtle (Apalone spinifera) in Northern Lake Champlain, Québec, Vermont," Journal of Herpetology, 36(3), 402-411, (1 September 2002)
- ^ Anderson, Noah. "Biomechanics of Feeding and Neck Motion in the Softshell Turtle, Apalone Spinifera, Rafinesque". ProQuest 305062583 – via ProQuest.
- ^ Williams, T. (1981). "The niches of two sympatric softshell turtles, Trionyx muticus and Trionyx spiniferus, in Iowa". Journal of Herpetology. 15 (3): 303–308. doi:10.2307/1563433. JSTOR 1563433.
- ^ Platt, Steven G.; Rainwater, Thomas R. (November 2021). "Observations of a Nuclear–follower Foraging Association between Spiny Softshell Turtles (Apalone spinifera) and Fish in an Urban Drainage Canal in Louisiana". Southeastern Naturalist. 20 (4): N108–N114. doi:10.1656/058.020.0406. ISSN 1528-7092. S2CID 244674573.
- ^ Daigle, Claude; Daigle, Claude; Galois, Patrick; Chagnon, Yves (2002). "Nesting activities of an Eastern Spiny Softshell Turtle, Apalone spinifera". The Canadian Field-Naturalist. 116 (1): 104––107. doi:10.5962/p.363405.
- ^ Tornabene; Bramblett, R.G; Zale, A.V; Leathe, S.A (2018). "Factors Affecting Nesting Ecology of Apalone spinifera in a Northwestern Great Plains River of the United States". Chelonian Conservation and Biology. 17 (1): 63–77. doi:10.2744/CCB-1298.1. S2CID 90056294.
- ^ Robinson, Keith M., and George G. Murphy. “The Reproductive Cycle of the Eastern Spiny Softshell Turtle (Trionyx Spiniferus Spiniferus).” Herpetologica, vol. 34, no. 2, 1978, pp. 137–140, www.jstor.org/stable/3891665?seq=1
- ^ Lazure, L., Pare, P., Bouthillier, L., & Galois, P. (2019). Nesting biology and conservation of a northern population of spiny softshell turtles (Apalone spinifera). Herpetological Conservation and Biology, 14(3), 659-667.
- ^ Greenbaum, Eli; Carr, John L (2001). "Sexual Differentiation in the Spiny Softshell Turtle (Apalone spinifera), a Species With Genetic Sex Determination". Journal of Experimental Biology. 290 (2): 190–200. Bibcode:2001JEZ...290..190G. doi:10.1002/jez.1049. PMID 11471149.
- ^ Geller, G.A. and G.S. Casper. 2023. Sound production by late-term embryos and hatchlings of the Spiny Softshell Turtle, Apalone spinifera (LeSueur, 1827) before nest exit. Herpetology Notes 16:483-491.)
- ^ Plummer, Michael V., and Caleb S. O’Neal. “Aerobic Pushups: Cutaneous Ventilation in Overwintering Smooth Softshell Turtles, Apalone Mutica.” Journal of Herpetology, vol. 53, no. 1, 2019, pp. 27–31, www.jstor.org/stable/26908244.
- ^ Plummer, Michael V., and Caleb S. O’Neal. “Aerobic Pushups: Cutaneous Ventilation in Overwintering Smooth Softshell Turtles, Apalone Mutica.” Journal of Herpetology, vol. 53, no. 1, 2019, pp. 27–31, www.jstor.org/stable/26908244.
- ^ Reese, Jackson, D. C., & Ultsch, G. R. (2003). Hibernation in freshwater turtles: softshell turtles (Apalone spinifera) are the most intolerant of anoxia among North American species. Journal of Comparative Physiology. B, Biochemical, Systemic, and Environmental Physiology, 173(3), 263–268. https://doi.org/10.1007/s00360-003-0332-1
- ^ "Spiny Softshell". OntarioNature. Retrieved 14 October 2023.
- ^ Tortoise & Freshwater Turtle Specialist Group (2016). "Apalone spinifera ssp. atra". IUCN Red List of Threatened Species. 2016: e.T1820A97408016. doi:10.2305/IUCN.UK.2016-1.RLTS.T1820A97408016.en. Retrieved 14 October 2023.
- ^ California Turtle & Tortoise Club: Softshell Turtles.
- ^ Conant R (1975). A Field Guide to Reptiles and Amphibians of Eastern and Central North America, Second Edition. Boston: Houghton Mifflin. 429 pp. (Softshell Turtles: Family Trionychidae, pp. 76-77).
- ^ "Apalone spinifera". Integrated Taxonomic Information System. Retrieved 31 May 2012.
- ^ Species Apalone spinifera at The Reptile Database .
- Bibliography
- van Dijk, Peter Paul; Iverson, John B.; Shaffer, H. Bradly; Bour, Roger; Rhodin, Anders G. J. (31 December 2011). "Turtles of the world, 2011 update: Annotated checklist of taxonomy, synonymy, distribution and conservation status". Conservation Biology of Freshwater Turtles and Tortoises (PDF). Chelonian Research Monographs. Vol. 5.
- Fritz, Uwe; Havaš, Peter (2007). "Checklist of Chelonians of the World". Vertebrate Zoology. 57 (2): 149–368. doi:10.3897/vz.57.e30895. S2CID 87809001.
Further reading
edit- Behler JL, King FW (1979). The Audubon Society Field Guide to North American Reptiles and Amphibians. New York: Alfred A. Knopf. 743 pp. ISBN 0-394-50824-6. (Trionyx spiniferus, pp. 485–486 + Plates 270, 271).
- Boulenger GA (1889). Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). New Edition. London: Trustees of the British Museum (Natural History). (Taylor and Francis, printers). x + 311 pp. + Plates I-III. (Trionyx spinifer, pp. 259–260).
- Lesueur CA (1827). "Note sur deux espèces de tortues, du genre Trionyx de M[onsieur]. Geoffroy-Saint-Hilaire ". Mémoires du Muséum d'Histoire Naturelle, Paris 15: 257-268 + Plates 6–7. (Trionyx spiniferus, new species, pp. 258–263 + Plate 6). (in French).
- Smith HM, Brodie ED Jr (1982). Reptiles of North America: A Guide to Field Identification. New York: Golden Press. 240 pp. ISBN 0-307-13666-3. (Trionyx spiniferus, pp. 31–33).
- Stejneger L, Barbour T (1917). A Check List of North American Amphibians and Reptiles. Cambridge Massachusetts: Harvard University Press. 125 pp. (Amyda spinifera, p. 125).
- Kominek, Emma; Cornies, Olivia; McCurdy-Adams, Hannah; Mooers, Arne Ø (2023). "Evolutionary isolation of Canadian terrestrial vertebrate species". The Canadian Field-Naturalist. 137 (4): 367–380. doi:10.22621/cfn.v137i4.2673. ISSN 0008-3550.