Water trek

Water trek
Systematics
without rank: Afroinsectiphilia Order : Tenrecus (Afrosoricida) Family : Tenreks (Tenrecidae) Subfamily : Travel treks (Oryzorictinae) Genre : Kleintenreks ( Microgale ) Type : Water trek
Scientific name
Microgale mergulus
( Major , 1896)

The water tenrek , also water tanrek or occasionally water-bristle hedgehog ( Microgale mergulus , Syn .: Limnogale mergulus ), is a species of mammal from the genus of small tenreks within the tenrek family . It lives in eastern Madagascar and inhabits a relatively small area where it occurs in only a few localities. The animals inhabit the tropical rainforests of the middle and higher mountain regions, but are rarely seen in the wild. They are the largest representatives of the Kleintenreks, have, like their relatives, a spindle-shaped body with powerful limbs and a narrow, pointed head. In contrast to other Kleintenreks, the snout is more blunt, the tail is flattened at the tip and webs are formed between the fingers and toes. The water tenrek is the only member of the tenreks adapted to aquatic life. It needs clear, fast-flowing rivers where it swims in search of food. Prey includes insects , crustaceans and frogs . The animals are solitary and nocturnal, they retire to rest in burrows near the river. Little is known about reproduction, and the way of life is generally considered to be little researched. The species was scientifically introduced in 1896 and predominantly placed in the genus Limnogale in the course of the 20th century . The exact relationships between species and genus have been the subject of scientific discussion in the history of research. Only genetic studies in the transition from the 20th to the 21st century confirmed the close relationship with the Kleintenreks. The population of the water tarnre is threatened.

features

Habitus

The water tenrek is the largest representative of the small tenreks. In general, the animals have a total length of 25.0 to 32.5 cm, a head-torso length of 12.2 to 17.0 cm and a tail length of 11.9 cm, after examining around a dozen individuals from various locations in the distribution area up to 16.1 cm. The weight is given as 60 to 107 g. Two animals from the Ranomafana National Park were 28.3 and 29.3 cm long with a body length of 14.5 and 13.8 cm, a tail length of 13.8 and 15.5 cm and a body weight of 80 and 105 g, respectively. In terms of physique, the water tenrek, with its spindle-shaped body, short and powerful limbs and narrow head with a pointed snout, corresponds to the other small tenreks. However, the snout of the water tenrek is shorter and blunt. The back fur is dark brown to black in color and consists of a mixture of short brownish and longer blackish hair, the underside is lighter. The tail, which is about as long as the rest of the body, is also two-colored. It is also strongly built and conical in shape, at the root it has a diameter of 8 to 10 mm, towards the middle it tapers to around 5 to 6 mm, only in the rear section it is flattened to around 30 mm in length. The top of the snout is covered with a brown-black fur, the conspicuous vibrissae are stiff, around 30 mm long and tinted white. Eyes and ears appear comparatively small. The latter are only 7 to 9 mm long, and short dark gray hairs grow on the outside and inside of the auricle. There is an equally colored spot behind the ears. Hands and feet have five rays on the fingers of the hands and the toes of the feet are blackish tinted webbed linked, as are the hind limbs on to the front and leave only the nails. A short-haired gray to whitish colored fur is formed on the inside and outside. The length of the hind feet is 32 to 36 mm. Females have a pair of teats in the breast, stomach and lumbar region.

Skull and dentition features

The skull of the type specimen is 32 mm long and at the cranium 16.5 mm wide. The nasal bone takes up about 11.5 mm of the length of the skull. The frontal bone is short and narrow, the parietal bones have a broad and elongated shape. The interrupted zygomatic arch is typical , which is a characteristic of all tenreks. The dentition consists of 40 teeth, the dental formula is as follows: . The inner upper incisors resemble a canine tooth ( caniniform ) and are larger than this. The same applies to the lower second incisors. In contrast, the lower anterior incisors are very small, while the second upper incisors are about the size of the canine. The premolars and molars resemble those of the other Kleintenreks. The molars in particular have a zalambdodontic chewing surface pattern with three main cusps. ${\displaystyle {\frac {3.1.3.3}{3.1.3.3}}}$

distribution

Distribution area of ​​the water tarnre

The water trek is endemic in Madagascar and inhabits a relatively narrow, more or less north-south strip in the eastern part of the country. However, it is only occupied by a total of 10 localities there. The most important sites from north to south include the forest area of ​​Sihanaka and the area around Andekaleka in the Toamasina province , the forest area around the Antsampandrano river on the Ankaratra massif and a region south of Antsirabe in the Antananarivo province and in the forest area of ​​Ranomafana and the Surroundings of Antanifotsy, the Maitso forest area and the upper reaches of the Iantara River in the province of Fianarantsoa , the latter three sites are in the Andringitra Mountains . The species may also be found in other places, but so far there have been too few field studies, and on the other hand, possible habitable habitats are known that do not harbor any populations . The animals live along the banks of clear and fast flowing rivers in the middle of tropical rainforests at altitudes between 450 and 2000 m. Because of this narrowly defined habitat, the actual occurrence of the species may only be around 2000 km². In general, the water trek is very rare. From its discovery at the end of the 19th century to the beginning of the 21st century, only around 40 individuals were observed, including around a dozen in the mid-1960s in the northern and central parts of the distribution area.

Way of life

Territorial behavior

There are only a few studies on the behavior of the aquatic tendre that have been carried out on some wild animals that have been kept briefly in human captivity. As the only representative of the Kleintenreks and the only land-living mammal in Madagascar, it is adapted to a semi-aquatic way of life, which is also reflected in some anatomical features. In addition to the webbed feet and the flattened tail, these include the strikingly large size compared to other small tenreks, the streamlined physique, the small ears and the dense, water-repellent fur. The animals can swim excellently, the rear feet are the main propulsion system, the tail serves as a rudder and the front feet are used to grasp the prey. The semimembranosus muscle is comparatively large on the hind legs . This serves as a straightener for the knee joint and supports a powerful forward movement in the water. The water trek is solitary and nocturnal. Activities begin shortly after sunset around 6:00 p.m. and end around 60 to 90 minutes before sunrise. According to investigations by two individuals equipped with radio transmitters, the distances covered in one night were on average 860 and 1067 m respectively, with the shortest distance covered being 220 and the longest 1550 m. The river sections used for this stretched over 550 and 1160 m in length. Using this data, action areas of 6960 or 7070 m² can be determined. Sometimes the water trek interrupts its nightly activities for shorter (1 hour) or longer (4 to 5 hours) periods. In the latter case, he often returns to his den. A structure examined in the Antsampandrano forest area was located on a river island 10 by 3 m in size and 0.5 m above the water surface. It had a diameter of 10 cm, was about 17 cm deep and had a nest made of plant material inside. Since the river with the observed nest has a strongly fluctuating water level in the course of the year, it is assumed that the water trek changes its structure at least seasonally.

nutrition

The aquatic tendrek is a carnivore and feeds mainly on crayfish , shrimp as well as insects and their larvae . Animals kept in captivity also ate offered fish , but there is no evidence as to whether they catch them themselves due to their nimble and agile nature in the wild. Analyzed droppings in the Ranomafana forest area mainly contained insect remains , including mayflies and caddis flies , dragonflies , butterflies and beetles . Subordinate also were crabs and crayfish, the latter often remains of Astacoides and tadpoles proved. According to the remains from the dung heaps, the prey consists of animals under 10 to 40 mm in size. The animals undertake dives lasting 10 to 15 seconds to catch prey. In the search for food, the vibrissae are especially helpful, the tactile perception is thus well developed. After catching the prey, the water trek leaves the river and eats it with its forefeet. The daily distance covered while foraging in the water varies between 200 and 1550 m. To defecate , the water tendrek leaves the river and climbs on stones protruding from the flowing water or goes ashore in leaf waste. The droppings themselves are 5 to 25 mm long, 3 to 7 mm wide and blackish in color. It is not known whether latrines are communal or territorial in character.

In contrast to other aquatic mammals, the aquatic trek does not show an increased metabolic rate . In the resting state, this is in the range of 79 to 96% of the values ​​to be expected for animals of similar size, but increases when they move or eat. The body temperature is around 32.5 ° C, which is around three degrees above the ambient temperature and corresponds to the values ​​of other small tenreks.

Reproduction

Reproduction is hardly documented, but a seasonal restriction is assumed. A single lactating female has been observed in December so far. The discovery of the individual building in the forest area of ​​Antsampandrano took place in January, it hid two almost fully grown young animals.

Systematics

General

Internal systematics of the Kleintenreks according to Everson et al. 2016  Microgale    Microgale pusilla      Microgale majori      Microgale principula      Microgale jenkinsae      Microgale longicaudata      Microgale mergulus      Microgale parvula      Microgale brevicaudata      Microgale grandidieri      Microgale drouhardi      Microgale monticola      Microgale taiva      Microgale gracilis      Microgale thomasi      Microgale cowani      Microgale jobihely      Microgale dryas      Microgale gymnorhyncha      Microgale soricoides      Microgale fotsifotsy      Microgale nasoloi

The web-footed tenrec is a kind from the genus of microgale ( Microgale ) within the family of tenrecs (Tenrecidae). The Kleintenreks also belong to the subfamily of the Reistenreks (Oryzorictinae), which also include the rice burrowers ( Oryzorictes ) and the representatives of the genus Nesogale . They are very varied and represent the numerically strongest member of the Tenreks with more than 20 species. Due to some morphological features, the Kleintenreks are considered to be originally within the family. Molecular genetic analyzes suggest that the genus developed as early as the Lower Miocene , around 16.8 million years ago, and the Kleintenreks diversified to a large extent in the period that followed. Today's representatives have adapted to different ways of life, some species that dig underground, live above ground, or tree-climb and inhabit water. The much larger part of the Kleintenreks inhabit the humid forests of eastern Madagascar, whereas only a few forms are found in the drier landscapes of the western part of the island. Within the genus, different kinship groups can be identified both morphologically and genetically, but the Wassertenrek was originally not considered a member of the Kleintenreks. However, genetic studies have shown that it is deeply embedded in the genus Microgale and represents its largest member. The closest relative was the dwarf lesser rek ( Microgale parvula ), which weighs less than 5 g and is the smallest member.

The first scientific description of the water tenrek comes from Charles Immanuel Forsyth Major in 1896. Major used the binomial Limnogale mergulus , with which he classified the water tenrek as separate from the small tenreks. The generic name Limnogale refers to the water-dwelling way of life of the animals ( λίμνη ( limne ) Greek for "lake" and γαλἑη ( gale ) for "weasel"; the epithet of the species mergulus is derived from the Latin word mergere for "diving"). Major based his new species and genus on two individuals, a male that comes from an area at 1600 m altitude west of Lake Andraykiba in the central-eastern highlands near the Malagasy capital Antananarivo , and a female from Imasindrary in the central-southern highlands. The latter forms the holotype and was collected in March of the year of the first description; its area of ​​origin also represents the terra typica of the species.

On the genus Limnogale - a phylogenetic controversy

The genus Limnogale only included one species, the water trek , and was therefore considered to be monotypical . By referring to its own genus, Major placed the water tenrek outside of the small tenreks. Major defined Limnogale mainly through the trained webbed feet, the strong tail and individual skull and tooth features, the latter clearly linked the genus with the tenreks and, according to Majors, especially with the smaller, soft-haired forms such as the small tenreks. However, in their adaptation to a semi-aquatic way of life, Major recognized similarities with the African otter shrews (Potamogalidae). In general, the otter shrews are classified as closely related to the Malagasy tenreks, which is expressed in similarities in the structure of the teeth and skull, such as the zalambdodontic chewing surface pattern of the molars, the large front, upper incisors or the non-closed zygomatic arch. However, there are also individual significant anatomical differences between the two groups. In contrast to the tenreks, the otter shrews lack the collarbone , on the other hand, the otter shrews have an overgrown second and third toe rays on the hind foot, which again does not occur in the tenreks. In addition to the external morphological similarities of the water tern, especially to the otter shrews, there are also various differences here, as, for example, the presence of webbed shrews is variable in the otter shrews. This in turn results in deviations in the details of the way of life that separate the water tarn from the otter shrews. The latter mainly use their tail to move in the water, while the former mainly uses their hind legs for this.

Because of these deviations and similarities, the systematic position of Limnogale to the other tenreks and to the otter shrews was the subject of discussion for a long time. In the course of the 20th century, numerous scientists saw a clear connection between the water tenreks and other Malagasy tenreks, especially the soft-haired rice treks, whereby the adaptation to a life in a watery environment was emphasized as a clear convergence to the otter shrews. Christian Guth and research colleagues came to the conclusion after intensive skull studies in 1959 that the water tenrek is only an aquatic form of the small tenreks ( simple "microgale" aquatique ). However, in 1957 it was Serge Frechkop who, despite the existing differences, recognized enough similarities between the water tenrek and the otter shrews to unite the two groups. He also suggested that the common subfamily be renamed Limnogalinae due to naming priority. Three decades later, Louis L. Jacobs and colleagues pointed out the greater similarities between the water tenreks and the African otter shrews than with the other Malagasy tenreks. The phylogenetic analyzes by Robert J. Asher , which, published in 1999 and based on anatomical features, identified the water tenrek as the sister taxon of the otter shrews, were classified as controversial . Asher justified this, among other things, with peculiarities in the skull structure, such as the very short structure of the frontal bone and the reduction of the tear hole (foramen lacrimale), which occur in both forms. As a result, the Malagasy tenreks would be seen as paraphyletic .

Especially in the transition to the 21st century, the emerging molecular genetic studies revealed new views on the relationship between the Tenreks. In a first genetic study from 2002, a research group led by Christophe J. Douady contradicted Asher's results and found the monophyly of the Malagasy tenreks to be confirmed. However, Douady and colleagues examined only a few representatives in total. The RNA analysis by Link E. Olson and Steven M. Goodman , which was published a year later and included around two dozen species of Tenreks, was much more extensive . She showed that the water tenrek on the one hand is much closer to the Malagasy tenreks than the African otter shrews, and on the other hand that Limnogale is deeply embedded in the genus Microgale . Subsequent genetic tests on DNA material were able to reproduce the result by Olson and Goodman and refine it increasingly. From this it could be seen in the end that it is not the Malagasy tenreks in general, but the small tenreks that form a paraphyletic group. For this reason, according to a genetic study from 2016, which took into account all tenrek and otter shrew species recognized up to that point in time, the genus Limnogale was synonymous with Microgale . As a result, the water tenrek now represents an aquatic representative of the small tenreks, which corresponds to the statements made by Guth and colleagues in 1959. The respective semi-aquatic way of life of the water ternect and the otter shrew thus represents a convergent development. In this sense, the common cranial features of the water ternrek and the otter shrew are to be understood as homoplasias . The reduction of the frontal bone and the re-formation of the tear hole go here probably with the reduction of the nasolacrimal duct (lacrimal canal) and the olfactory lobe (olfactory lobe) accompanied. This caused a reduction in the sense of smell, a trait that other smaller, partly aquatic and carnivorous mammals share.

Threat and protection

The water trek lives in a relatively small area in eastern Madagascar with few known find locations and is dependent on clean waters. As a result, the species shows a particular susceptibility to human interference. These include flooding or erosion of the soil as a result of deforestation. This leads not only to the impairment of the aquatic tenrek itself, but also of its benthic prey. In addition, the expansion of the arable land causes a fragmentation of the surrounding area, so that usable habitats are separated from one another. At some of the well-known sites such as Antsampandrano and Andekaleka, where individual animals were still observed in field studies in the 1960s, there has recently been no scientific evidence of the water tarragon. Another threat is fishing. The IUCN therefore lists the water trek as "endangered" ( vulnerable ). It occurs in several nature reserves, for example in the Andringitra National Park , in the Ranomafana National Park or in the Nosy Volo Reserve . For the conservation of the species, more intensive studies on its actual distribution and the influence of the threat factors are necessary. The protection of their prey is also of great importance.

literature

• Jonathan Benstead, Kevin H. Barnes and Catherine M. Pringle: Diet, acvtivity patterns, foraging movements and response to deforestation of the aquatic tenrec Limnogale mergulus (Lipotyphla: Tenrecidae) in eastern Madagascar. Journal of Zoology 254, 2001, pp. 119-129
• Paulina D. Jenkins: Tenrecidae (Tenrecs and Shrew tenrecs). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 134–172 (p. 171) ISBN 978-84-16728-08-4
• Charles Immanuel Forsyth Major: Diagnoses of new mammals from Madagascar. The Annals and magazine of natural history 18, 1896, pp. 318–321 ( [2] )
• P. Malzy: Un mammifere aquatique de Madagascar: Le Limnogale. Mammalia 29 (3), 1965, pp. 399-411
• Ronald M. Nowak: Walker's Mammals of the World . The Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9

Individual evidence

1. ↑ ^{a b c d e f g h i} P. Malzy: Un mammifere aquatique de Madagascar: Le Limnogale. Mammalia 29 (3), 1965, pp. 400-411
2. ↑ ^{a b c d e f g} J. F. Eisenberg and Edwin Gould: The Tenrecs: A Study in Mammalian Behavior and Evolution. Smithsonian Institution Press, 1970, pp. 1-138
3. ↑ ^{a b c d e} Kathryn M. Everson, Voahangy Soarimalala, Steven M. Goodman and Link E. Olson: Multiple loci and complete taxonomic sampling resolve the phylogeny and biogeographic history of tenrecs (Mammalia: Tenrecidae) and reveal higher speciation rates in Madagascar's humid forests. Systematic Biology 65 (5), 2016, pp. 890-909 doi: 10.1093 / sysbio / syw034
4. ↑ ^{a b c d e f g} Jonathan Benstead, Kevin H. Barnes and Catherine M. Pringle: Diet, acvtivity patterns, foraging movements and response to deforestation of the aquatic tenrec Limnogale mergulus (Lipotyphla: Tenrecidae) in eastern Madagascar. Journal of Zoology 254, 2001, pp. 119-129
5. ↑ ^{a b c d} Charles Immanuel Forsyth Major: Diagnoses of new mammals from Madagascar. The Annals and magazine of natural history 18, 1896, pp. 318-321
6. ↑ ^{a b c d e} Paulina D. Jenkins: Tenrecidae (Tenrecs and Shrew tenrecs). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 134–172 (p. 171) ISBN 978-84-16728-08-4
7. ^ ^{A b} P. J. Stephenson, Voahangy Soarimalala and Steven M. Goodman: Limnogale mergulus. The IUCN Red List of Threatened Species 2016. e.T11979A97189690 ( [1] ); last accessed on October 10, 2016
8. ↑ Hideko Endo, Takahiro Yonezawa, Felix Rakotondraparany, Motoki Sasaki and Masami Hasegawa: The adaptional strategies of the hindlimb muscles in the Tenrecidae species including the aquatic web-footed tenrec (Limnogale mergulus). Annals of Anatomy 188, 2006, pp. 383-390
9. ^ ^{A b c} Edwin Gould and John F. Eisenberg: Notes on the biology of the Tenrecidae. Journal of Mammalogy 47 (4), 1966, pp. 660-686
10. ^ Peter J. Stephenson, Paul A. Racey, and Félix Rakotondraparany: Maintenance and reproduction of tenrecs (Tenrecidae) at Parc Tsimbazaza, Madagascar. International Zoo Yearbook 33, 1994, pp. 194-201
11. ↑ Peter J. Stephenson: Resting metabolic rate and body temperature in the aquatic tenrec Limnogale mergulus (Insectivora: Tenrecidae). Acta Theriologica 39 (1), 1994, pp. 89-92
12. ^ Paul A. Racey and Peter J. Stephenson: Reproductive and energetic differentiation of the Tenrecidae of Madagascar. In: W. R Lourenço (Ed.): Biogéographie de Madagascar. Paris, 1996, pp. 307-319
13. ↑ ^{a b} R. DE MacPhee: The Shrew Tenrecs of Madagascar: Systematic revision and distribution of Holocene Microgale (Tenrecidae, Insectivora). American Museum Novitates 2889, 1987, pp. 1-45
14. ^ ^{A b} Robert J. Asher and Robert Hofreiter: Tenrec Phylogeny and the Noninvasive Extraction of Nuclear DNA. Systematic Biology 55 (2), 2006, pp. 181-194
15. ↑ Peter Vogel: Family Tenrecidae Tenrecs, Otter-shrews. In: Jonathan Kingdon, David Happold, Michael Hoffmann, Thomas Butynski, Meredith Happold and Jan Kalina (eds.): Mammals of Africa Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, 2013, pp. 216-222
16. ↑ George Gaylord Simpson: The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85, 1945, pp. 1–350 (pp. 48–49)
17. ↑ Malcolm C. McKenna and Susan K. Bell: Classification of mammals above the species level. Columbia University Press, New York, 1997, pp. 1-631 (pp. 293-295)
18. ^ Christian Guth, Henri Heim de Balsac and M. Lamotte: Recherches sur la morphologie de Micropotamogale lamottei et l'évolution des Potamogalinae: écologie, denture, anatomie crânienne. Mammalia 23, 1959, pp. 423-447
19. ↑ Serge Frechkop: A propos de nouvelles espèces de Potamogalines. Mammalia 21, 1957, pp. 226-234
20. ↑ Louis L. Jacobs, William Anyonge, and John C. Barry: A Giant Tenrecid from the Miocene of Kenya. Journal of Mammalogy 68 (1), 1987, pp. 10-16
21. ^ Robert J. Asher: A morphological basis for assessing the phylogeny of the "Tenrecoidea" (Mammalia, Lipotyphla). Cladistics 15, 1999, pp. 231-252
22. ↑ ^{a b} Link E. Olson and Steven M. Goodman: Phylogeny and biogeography of tenrecs. In: Steven M. Goodman and Jonathan P. Benstead (Eds.): The natural history of Madagascar. University of Chicago Press, 2003, pp. 1235-1242
23. ↑ Christophe J. Douady, Francois Catzeflis, Diana J. Kao, Mark S. Springer and Michael J. Stanhope: Molecular Evidence for the Monophyly of Tenrecidae (Mammalia) and the Timing of the Colonization of Madagascar by Malagasy Tenrecs. Molecular Phylogenetics and Evolution 22 (3), 2002, pp. 357-363
24. ↑ Christophe J. Douady, Francois Catzeflis, Mark S. Springer and Michael J. Stanhope: Molecular evidence for the monophyly of Tenrecidae: a reply to Asher. Molecular Phylogenetics and Evolution 26, 2003, pp. 331-332
25. ↑ Céline Poux, Ole Madesen, Elisabeth Marquard, David R. Vieites, Wilfried W. de Jong and Miguel Vences: Asynchronous Colonization of Madagascar by the Four endemic clades of Primates, Tenrecs, Carnivores, and Rodents as Inferred from Nuclear Genes. Systematic Biology 54 (5), 2005, pp. 719-730
26. ↑ Céline Poux, Ole Madesen, Julian Glos, Wilfried W. de Jong and Miguel Vences: Molecular phylogenetic and divergence times of Malagasy tenrecs: Influence of data partitioning and taxon sampling on dating analyzes. BMC Evolutionary Biology, 2008, p. 102
27. ^ Matjaž Kuntner, Laura J. May-Collado and Ingi Agnarsson: Phylogeny and conservation priorities of afrotherian mammals (Afrotheria, Mammalia). Zoologica Scripta 40 (1), 2011, pp. 1-15
28. ^ MR Sanchez-Villagra and RJ Asher: Cranio-sensory adaptations in small faunivorous semiaquatic mammals, with special reference to olfaction and the trigeminal system. Mammalia 66 (1), 2002, pp. 93-109
29. ↑ Robert J. Asher: Tenrecoidea. In: Lars Werdelin and William Joseph Sanders (eds.): Cenozoic Mammals of Africa. University of California Press, Berkeley, London, New York, 2010, pp. 99-106
30. ↑ Martin Nicoll and Nanie Ratsifandrihamanana: The growth of Madagascar's protected areas system and its implications for tenrecs (Afrosoricida, Tenrecidae). Afrotherian Conservation 10, 2014, p. 4-8

Web links

Commons : Wassertenrek ( Microgale mergulus )  - Collection of images, videos and audio files

• Information on Animal Diversity Web
• Limnogale mergulus in the endangered Red List species the IUCN 2016 Posted by: PJ Stephenson, Voahangy Soarimalala and Steven M. Goodman, 2014. Accessed on 10/10/2016.