Noronhomys

Last updated

Noronhomys
Status iucn3.1 EX.svg
Extinct  (1503)  (IUCN 3.1) [1]
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Family: Cricetidae
Subfamily: Sigmodontinae
Genus:† Noronhomys
Carplaton and Olson, 1999
Species:
†N. vespuccii
Binomial name
†Noronhomys vespuccii
Carleton and Olson, 1999
Location Noronha circle.png
Location of Fernando de Noronha, the island group to which Noronhomys was endemic.

Noronhomys vespuccii, also known as Vespucci's rodent, [2] is an extinct rat species from the islands of Fernando de Noronha off northeastern Brazil. Italian explorer Amerigo Vespucci may have seen it on a visit to Fernando de Noronha in 1503, but it subsequently became extinct, perhaps because of the exotic rats and mice introduced by the first explorers of the island. Numerous but fragmentary fossil remains of the animal, of uncertain but probably Holocene age, were discovered in 1973 and described in 1999.

Contents

Noronhomys vespuccii was a fairly large rodent, larger than the black rat (Rattus rattus). A member of the family Cricetidae and subfamily Sigmodontinae, it shares several distinctive characters with Holochilus and related genera within the tribe Oryzomyini, including high-crowned molars with simplified crown features and the presence of several ridges on the skull which help anchor the chewing muscles. Although a suite of traits suggest that Holochilus is its closest relative, it is distinctive in many ways and is therefore classified in a separate genus, Noronhomys. Its close relatives, including Holochilus and Lundomys , are adapted to a semiaquatic lifestyle, spending much of their time in the water, but features of the Noronhomys bones suggest that it lost its semiaquatic lifestyle, after arrival at its remote island.

Discovery and taxonomy

Italian explorer Amerigo Vespucci may have seen this animal on his fourth voyage, which took him to Brazil; [3] the Lettera di Amerigo Vespucci delle Isole Nuovamente in Quattro Suoi Viaggi [4] recorded that he visited an island just south of the equator on August 10, 1503. On this island, identified as Fernando de Noronha, he saw "very big rats and lizards with two tails, and some snakes". [5] The Lettera purports to be an account of Vespucci's voyages, but it is unlikely that he produced it himself and, additionally, his fourth voyage may never have taken place. [6] The biological details given in the Lettera's account of Fernando de Noronha agree with what is known of the natural history of the island, lending weight to the view that it derives from a visit, whether by Vespucci himself or by another explorer. The lizard is probably Trachylepis atlantica and the record of snakes most likely refers to Amphisbaena ridleyi , which is actually an amphisbaenian instead of a snake. [7]

During excavations conducted in 1973, American ornithologist Storrs L. Olson found fossils of a moderately large rat on Fernando de Noronha, which were described as a new genus and species in a 1999 publication by Olson and his colleague, mammalogist Michael D. Carleton. [3] The material is now in the United States National Museum of Natural History in Washington, D.C., and in the museum of the Pontifícia Universidade Católica do Rio Grande do Sul in Porto Alegre, Rio Grande do Sul. The generic name, Noronhomys, combines the name of the island of Fernando de Noronha with the Ancient Greek μῦςmys "mouse" [8] and the specific name, vespuccii, honors Amerigo Vespucci. [9] Noronhomys would have been larger than the black rat (Rattus rattus), which was common on ships and which Vespucci would have been familiar with, consistent with his description of "very large rats". [7]

Suggested relationships of Noronhomys. [10]

Noronhomys is a member of the tribe Oryzomyini, which includes over a hundred species distributed mainly in South America, including nearby islands such as the Galápagos Islands and some of the Antilles. Oryzomyini is one of several tribes recognized within the subfamily Sigmodontinae, which encompasses hundreds of species found across South America and into southern North America. Sigmodontinae itself is the largest subfamily of the family Cricetidae, other members of which include voles, lemmings, hamsters, and deermice, all mainly from Eurasia and North America. [11]

Carleton and Olson performed a detailed comparison of Noronhomys to members of the mainland genera Holochilus and Lundomys on the basis of both general morphology and morphometrical data, concluding that the Fernando de Noronha rat is distinct from both other animals. [12] They used a cladistic analysis to examine its relationships within Oryzomyini, also including two species of Holochilus, Lundomys, and five other oryzomyines. They found that Noronhomys appeared closest to Holochilus, with Lundomys more distantly related. Eighteen shared characters (synapomorphies) supported the grouping of Noronhomys with Holochilus. [10] Another form described as a species of Holochilus, Holochilus primigenus , may also be related, but is likely to fall outside the Holochilus–Noronhomys clade. [13]

In 1998, a fragmentary fossil of another species of this group of oryzomyines was found in eastern Argentina. It was initially identified as a possible second species of Noronhomys on the basis of the presence of a crest on the upper first molar, the mesoloph, [14] but the specimen is different from Noronhomys vespuccii in other respects, and in 2008 it was described as a new genus and species, Carletonomys cailoi , related to Noronhomys and associated genera. [14]

Description

The skull of the living Lundomys molitor is similar to that of Noronhomys. (The illustrated mandible is not of a Lundomys.) Lundomys molitor type.png
The skull of the living Lundomys molitor is similar to that of Noronhomys. (The illustrated mandible is not of a Lundomys.)

Noronhomys vespuccii is known from bone fragments, including five skulls, damaged to various degrees, and many isolated jaws and other bones. [17] This material documents that, with a skull of about 4 cm (1.5 in) (occipitonasal length), Noronhomys was a moderately large oryzomyine, smaller than Lundomys but well within the range of Holochilus. [18] It shares a number of the features that characterize the group of Holochilus and related genera, [19] including a reduction in the complexity of the chewing surface of the molars, [15] simple posterolateral palatal pits (perforations of the palate near the third molar), [20] and a similarly shaped interorbital region of the skull. [21] Noronhomys is distinctive, among other characters, in its lack of a spinous process on the zygomatic plate, the flattened front portion of the zygomatic arch (cheekbone); the short palate, which does not extend behind the third molars; and the presence of an accessory crest, the mesoloph, on the upper molars. [18]

Analysis of morphometrical data from the known material of Noronhomys suggests that growth continued in adults—the older the animal, the larger the depth of the mandible and the size of the lower incisor [22] —and does not provide evidence for a difference in size between males and females. [23] Skull shape is strikingly different from both Holochilus and Lundomys, resulting in a clear separation from both of these taxa in statistical analyses of measurement data. [24] In three individuals measured, the occipitonasal length, a measure of skull length, varies from 38.0 to 39.2 mm (1.50 to 1.54 in), averaging 38.5 mm (1.52 in). The width of the braincase is 13.4 to 14.8 mm (0.53 to 0.58 in), averaging 14.1 mm (0.56 in). Between the first molars, the width of the palate is 8.1 to 8.9 mm (0.32 to 0.35 in) in four specimens, averaging 8.4 mm (0.33 in). [25] The lower molars have a total length of 7.57 to 8.29 mm (0.298 to 0.326 in), averaging 8.00 mm (0.315 in), in 39 specimens with intact molar rows. [26] The femur (upper leg bone) is 32.5 to 41.0 mm (1.28 to 1.61 in) in length in nine specimens, averaging 36.5 mm (1.44 in). [27] Carleton and Olson estimated that body mass in Noronhomys was similar to that of some populations of living Holochilus sciureus at about 200 to 250 g (7.1 to 8.8 oz). [7]

Skull

View of Morro do Pico on Fernando de Noronha. Fernando de Noronha Conceicao Beach.jpg
View of Morro do Pico on Fernando de Noronha.

The skull is flattened in general shape. The front part is short and broad. The broad interorbital region (located between the eyes) is hourglass-shaped, [20] with squared edges and weakly developed beading. A postorbital ridge is present, obscuring the suture (joint) between the frontal and squamosal bones, [28] a feature shared only with Holochilus among oryzomyines. [29] The braincase is squarish. The interparietal bone is wide, but does not reach the squamosals on the sides. The zygomatic arches, which are well-developed, are furthest apart at the back and converge toward the front. The jugal bone is small. [20]

The back margin of the zygomatic plate is located close to the front of the upper first molar. The incisive foramina do not extend between the molars. [20] Unlike in both Holochilus and Lundomys, [30] the palate is flat, lacking a distinct ridge at the midline. The parapterygoid fossae, which are located behind the palate at the level of the molars, are excavated somewhat above the level of the palate. A strut of the alisphenoid bone is present, separating two openings in the skull, the masticatory–buccinator foramen and the foramen ovale accessorium. [20] The subsquamosal fenestra, an opening at the back of the skull determined by the shape of the squamosal, is present but small. The squamosal probably lacks a suspensory process that contacts the tegmen tympani, the roof of the tympanic cavity, [15] a defining character of oryzomyines. [31]

The mandible (lower jaw) is robust. The two masseteric ridges, which anchor some of the chewing muscles, are joined together as a single crest for a portion of their length and extend forward to a point below the first molar. The capsular process, a raising of the mandibular bone at the back end of the lower incisor, is well developed. [15]

Teeth

In the upper incisors, the chewing edges are located behind the vertical plane of the incisors; thus, they are opisthodont. [15] The microstructure of the enamel of the lower incisor was reported in a 2005 study. The inner portion (portio interna, PI) is much thicker than the outer portion (portio externa, PE). The PI consists of Hunter-Schreger bands, which are uniserial (consisting of a single enamel prism), [32] as in all myomorph rodents. [33] The PE consists of radial enamel, with flattened prisms that are almost parallel to the junction between the enamel and the dentine. [34] The microstructure is similar to that of Holochilus brasiliensis [35] and displays several characters seen only in Myomorpha. [36]

Map of the Fernando de Noronha archipelago, showing the fossil site where remains of Noronhomys have been found (in red). Map of Fernando de Noronha-en-fossil site.png
Map of the Fernando de Noronha archipelago, showing the fossil site where remains of Noronhomys have been found (in red).

The molars are high-crowned (hypsodont) and planar, with the main cusps as high as the crests connecting them, [15] a configuration shared only with Holochilus and Carletonomys among oryzomyines. [38] The first molars are the longest and the third molars are longer but narrower than the second molars. The molars lack many accessory ridges, including the anteroloph on the first upper molar, the posteroloph on the first and second upper molar, and the anterolophid and mesolophid on all lower molars. The first and second upper molar have a short mesoloph and the front cusp of the first lower molar, the anteroconid, encloses a large internal pit. Most of the folds between the cusps and crests are open at the margins of the molars, but two—the posteroflexid on the lower second molar and the entoflexid on the lower third molar—are closed by a wall, or cingulum, at the inner margin of the tooth. As in most oryzomyines, the upper molars all have one root on the inner (lingual) side and two on the outer (labial) side and the lower molars have a single root at the front and back of each molar; in addition, the first upper molar has another labial root and the first lower molar has a small labial and usually also a small lingual root located between the main roots. [15]

Postcranial skeleton

The entepicondylar foramen is absent, [15] as in all members of the Sigmodontinae; if present, as in some other rodents, this foramen (opening) perforates the distal (far) end of the humerus (upper arm bone). [39] The pelvis and the bones of the hindlimbs are heavily built. The femoral tubercle of the acetabulum (part of the pelvis), which anchors the rectum femoris muscle, is reduced relative to Holochilus and Lundomys. [40]

Distribution and origin

The lizard Trachylepis atlantica, which still lives on Fernando de Noronha, is one of the few other animals found on the island. Mabuya maculata-001.png
The lizard Trachylepis atlantica , which still lives on Fernando de Noronha, is one of the few other animals found on the island.

Noronhomys is known only from Fernando de Noronha, a small archipelago of volcanic origin off northeastern Brazil, consisting of a main island and several associated smaller islands. The formation of the archipelago, which has never been connected to the mainland, began about 11 million years ago; active vulcanism ceased about 2 million years ago. [43] Remains of Noronhomys were found in association with remains of various reptiles, birds, and snails, several of which are also restricted to the archipelago, [18] in sand dunes near the northeastern tip of the main island. [44] The age of the deposits is unknown, but is likely late Holocene, at most a few thousand years old. [3]

The ancestor of Noronhomys may have been a semiaquatic animal, similar to living Holochilus or Lundomys, that arrived on Fernando de Noronha by chance on a floating log. [45] The morphology of the limb bones in Noronhomys suggests that the animal was not semiaquatic like its relatives, but terrestrial, consistent with its occurrence on a small island, where streams and pools are rare or absent. [46]

Extinction

Vespucci's account suggests that Noronhomys was common when the island was first visited, but it was not encountered by the first biological explorers of the island, who conducted their research in the late 19th century. The introduced black rat and house mouse (Mus musculus), which became very common on the island, may have driven it to extinction by directly competing for food, preying on young Noronhomys, or transmitting diseases. Other factors that may have played a role include modification of its habitat, introduction of predators such as cats (Felis catus), and predation by visiting sailors. These extinction mechanisms are common for island endemic species. [47] As early as 1888, Henry Nicholas Ridley suggested that the rat Vespucci had seen had been driven to extinction by the introduced black rat. [48] The International Union for Conservation of Nature lists it as "extinct". [1]

Related Research Articles

<i>Oryzomys nelsoni</i> Extinct species of rodent

Oryzomys nelsoni, also known as the Nelson’s Rice Rat, is an extinct rodent of María Madre Island, Nayarit, Mexico. Within the genus Oryzomys of the family Cricetidae, it may have been most closely related to the mainland species O. albiventer. Since its first description in 1898, most authors have regarded it as a distinct species, but it has also been classified as a mere subspecies of the marsh rice rat (O. palustris).

<i>Oryzomys</i> Genus of semiaquatic rodents

Oryzomys is a genus of semiaquatic rodents in the tribe Oryzomyini living in southern North America and far northern South America. It includes eight species, two of which—the marsh rice rat (O. palustris) of the United States and O. couesi of Mexico and Central America—are widespread; the six others have more restricted distributions. The species have had eventful taxonomic histories, and most species were at one time included in the marsh rice rat; additional species may be recognized in the future. The name Oryzomys was established in 1857 by Spencer Fullerton Baird for the marsh rice rat and was soon applied to over a hundred species of American rodents. Subsequently, the genus gradually became more narrowly defined until its current contents were established in 2006, when ten new genera were established for species previously placed in Oryzomys.

<i>Euryoryzomys emmonsae</i> Amazonian rodent

Euryoryzomys emmonsae, also known as Emmons' rice rat or Emmons' oryzomys, is a rodent from the Amazon rainforest of Brazil in the genus Euryoryzomys of the family Cricetidae. Initially misidentified as E. macconnelli or E. nitidus, it was formally described in 1998. A rainforest species, it may be scansorial, climbing but also spending time on the ground. It lives only in a limited area south of the Amazon River in the state of Pará, a distribution that is apparently unique among the muroid rodents of the region.

<i>Lundomys</i> A semiaquatic rat species from southeastern South America.

Lundomys molitor, also known as Lund's amphibious rat or the greater marsh rat, is a semiaquatic rat species from southeastern South America.

<i>Pseudoryzomys</i> Genus of rodent from South America with one species

Pseudoryzomys simplex, also known as the Brazilian false rice rat or false oryzomys, is a species of rodent in the family Cricetidae from south-central South America. It is found in lowland palm savanna and thorn scrub habitats. It is a medium-sized species, weighing about 50 grams (1.8 oz), with gray–brown fur, long and narrow hindfeet, and a tail that is about as long as the head and body. The IUCN has assessed its conservation status as being of least concern, although almost nothing is known about its diet or reproduction.

<i>Oryzomys gorgasi</i> Rodent from the family Cricetidae from northwestern Colombia and Venezuela

Oryzomys gorgasi, also known as Gorgas's oryzomys or Gorgas's rice rat, is a rodent in the genus Oryzomys of family Cricetidae. First recorded in 1967, it is known from only a few localities, including a freshwater swamp in the lowlands of northwestern Colombia and a mangrove islet in northwestern Venezuela. It reportedly formerly occurred on the island of Curaçao off northwestern Venezuela; this extinct population has been described as a separate species, Oryzomys curasoae, but does not differ morphologically from mainland populations.

<span class="mw-page-title-main">Hammond's rice rat</span> Species of rodent in the family Cricetidae from Ecuador

Mindomys hammondi, also known as Hammond's rice rat or Hammond's oryzomys, is an endangered species of rodent in the tribe Oryzomyini of family Cricetidae. Formerly considered to be related with Nectomys, Sigmodontomys, Megalomys, or Oryzomys, it is now placed in then genus Mindomys, but its relationships remain obscure; some evidence supports a placement near Oecomys or as a basal member of Oryzomyini.

<i>Eremoryzomys</i> Rodent species in the family Cricetidae from central Peru

Eremoryzomys polius, also known as the gray rice rat or the Marañon oryzomys, is a rodent species in the tribe Oryzomyini of the family Cricetidae. Discovered in 1912 and first described in 1913 by Wilfred Osgood, it was originally placed in Oryzomys and named Oryzomys polius. In 2006, a cladistic analysis found that it was not closely related to Oryzomys in the strict sense or to any other oryzomyine then known, so that it is now placed in its own genus, Eremoryzomys. The Brazilian genus Drymoreomys, named in 2011, is probably the closest relative of Eremoryzomys. Eremoryzomys has a limited distribution in the dry upper valley of the Marañón River in central Peru, but may yet contain more than one species.

<span class="mw-page-title-main">Oryzomyini</span> Tribe of rodents

Oryzomyini is a tribe of rodents in the subfamily Sigmodontinae of the family Cricetidae. It includes about 120 species in about thirty genera, distributed from the eastern United States to the southernmost parts of South America, including many offshore islands. It is part of the clade Oryzomyalia, which includes most of the South American Sigmodontinae.

<i>Carletonomys</i> Extinct genus of rodents

Carletonomys cailoi is an extinct rodent from the Pleistocene (Ensenadan) of Buenos Aires Province, Argentina. Although known only from a single maxilla with the first molar, its features are so distinctive that it is placed in its own genus, Carletonomys. Discovered in 1998 and formally described in 2008, it is part of a well-defined group of oryzomyine rodents that also includes Holochilus, Noronhomys, Lundomys, and Pseudoryzomys. This group is characterized by progressive semiaquatic specializations and a reduction in the complexity of molar morphology.

Reigomys primigenus is an extinct oryzomyine rodent known from Pleistocene deposits in Tarija Department, southeastern Bolivia. It is known from a number of isolated jaws and molars which show that its molars were almost identical to those of the living Lundomys. On the other hand, the animal possesses a number of derived traits of the palate which document a closer relationship to living Holochilus, the genus of South American marsh rats, and for this reason it was placed in the genus Holochilus when it was first described in 1996. The subsequent discoveries of Noronhomys and Carletonomys, which may be more closely related to extant Holochilus than H. primigenus is, have cast its placement in Holochilus into doubt, and it was ultimately made the type species of a separate genus, Reigomys.

Nephelomys moerex is a species of rodent in the genus Nephelomys of family Cricetidae. The type locality is at Mindo in western Ecuador, where it has been recorded together with three other rodents of the oryzomyine group, Sigmodontomys aphrastus, Mindomys hammondi, and Handleyomys alfaroi, as well as three opossums, Chironectes minimus and unidentified species of Didelphis and Marmosa. Mindo is a "tiny agricultural community" located at 0°02'S, 78°48'W and 1,264 metres (4,150 ft) above sea level. It was originally described by Oldfield Thomas as a subspecies of Oryzomys albigularis. It remained synonymized under this species until it was recognized as a separate species when the genus Nephelomys was established for Oryzomys albigularis and related species in 2006.

<i>Amphisbaena ridleyi</i> Species of lizard

Amphisbaena ridleyi, known by the common names Ridley's worm lizard or the Noronha worm lizard, is a species of amphisbaenian in the family Amphisbaenidae. The species is endemic to the island of Fernando de Noronha off the coast of Brazil. It is one of two indigenous reptiles on the island.

In anatomy, posterolateral palatal pits are gaps at the sides of the back of the bony palate, near the last molars. Posterolateral palatal pits are present, in various degrees of development, in several members of the rodent family Cricetidae. Many members of the family lack them or have only simple pits, but Arvicolinae and Oryzomyini have more highly developed posterolateral palatal pits. Posterolateral palatal pits are also present in some other rodents, including Glis, Jaculus, Hystrix, Abrocoma, Ctenomys, Chinchilla, and Lagidium.

<i>Oryzomys albiventer</i> Species of rodent

Oryzomys albiventer, commonly known as the White-bellied Rice Rat, is a rodent in the genus Oryzomys of family Cricetidae from interior western Mexico, in the states of Jalisco, Guanajuato, and Michoacán. First described in 1901 as a separate species, it was later lumped under O. couesi and the marsh rice rat (O. palustris) until it was reinstated as a species in 2009. It differs from neighboring Oryzomys populations in size and measurements and is a large, brightly colored species with a long tail and robust skull and molars. Its range has been much impacted by agricultural development, but isolated populations are thought to persist.

In mammals, ungual tufts are tufts of hairs at the base of claws of the forefeet and hindfeet. Their presence has been used as a character in cladistic studies of the Cricetidae, a large family of rodents.

Natatory fringes are rows of stiff hairs that occur along the margins of the hindfeet in some rodents. They occur along the plantar margins and in some cases also between the toes. Among sigmodontines, a mostly South American groups, natatory fringes are present in Ichthyomyini and some Oryzomyini. Among ichthyomyines, the fringes are poorly developed in Neusticomys but well-developed in other genera, and in Rheomys mexicanus the hairs of the fringes may exceed 3 millimetres (0.12 in) in length. Amphinectomys, Holochilus, Lundomys, and Nectomys are the only oryzomyines with natatory fringes, but have them only weakly developed; one study also records them in Oryzomys. In oryzomyines, the fringes are an adaptation for a semiaquatic lifestyle that appeared convergently in the Holochilus-Lundomys and Nectomys-Amphinectomys lineages. The term was introduced in 1993 by Voss and Carleton in describing Lundomys.

<i>Agathaeromys</i> Extinct genus of rodents

Agathaeromys is an extinct genus of oryzomyine rodents from the Pleistocene of Bonaire, Netherlands Antilles. Two species are known, which differ in size and some details of tooth morphology. The larger A. donovani, the type species, is known from hundreds of teeth that are probably 900,000 to 540,000 years old, found in four localities. A. praeuniversitatis, the smaller species, is known from 35 teeth found in a single fossil site, which is probably 540,000 to 230,000 years old.

<i>Pennatomys</i> Extinct oryzomyine rodent in the Lesser Antilles

Pennatomys nivalis is an extinct oryzomyine rodent from the islands of Sint Eustatius, Saint Kitts, and Nevis in the Lesser Antilles. The only species in the genus Pennatomys, it is known from skeletal remains found in Amerindian archeological sites on all three islands, with dates ranging from 790–520 BCE to 900–1200 CE. No live specimens are known, but there are several historical records of rodents from Saint Kitts and Nevis that could conceivably refer to Pennatomys. The animal apparently belongs to a group within the tribe Oryzomyini that includes many other island-dwelling species.

<i>Drymoreomys</i> A rodent genus with one species in the family Cricetidae from the Atlantic Forest of Brazil

Drymoreomys is a rodent genus in the tribe Oryzomyini that lives in the Atlantic Forest of Brazil. The single species, D. albimaculatus, is known only from the states of São Paulo and Santa Catarina and was not named until 2011. It lives in the humid forest on the eastern slopes of the Serra do Mar and perhaps reproduces year-round. Although its range is relatively large and includes some protected areas, it is patchy and threatened, and the discoverers recommend that the animal be considered "Near Threatened" on the IUCN Red List. Within Oryzomyini, Drymoreomys appears to be most closely related to Eremoryzomys from the Andes of Peru, a biogeographically unusual relationship, in that the two populations are widely separated and each is adapted to an arid or a moist environment.

References

  1. 1 2 Weksler, M. & Costa , L.M. 2019. Noronhomys vespuccii. The IUCN Red List of Threatened Species 2019: e.T136692A22333193. https://dx.doi.org/10.2305/IUCN.UK.2019-1.RLTS.T136692A22333193.en. Accessed on 01 May 2023.
  2. ↑ Musser and Carleton, 2005, p. 1136
  3. 1 2 3 Carleton and Olson, 1999, p. 2
  4. ↑ Carleton and Olson, 1999, p. 44
  5. ↑ Translation from the Italian by Branner, 1888, p. 869, cited in Carleton and Olson, 1999, p. 2
  6. ↑ Carleton and Olson, 1999, p. 46
  7. 1 2 3 Carleton and Olson, 1999, p. 48
  8. ↑ Carleton and Olson, 1999, p. 10
  9. ↑ Carleton and Olson, 1999, p. 12
  10. 1 2 Carleton and Olson, 1999, fig. 22
  11. ↑ Musser and Carleton, 2005
  12. ↑ Carleton and Olson, 1999, pp. 16–32
  13. ↑ Carleton and Olson, 1999, p. 50
  14. 1 2 Pardiñas, 2008, p. 1271
  15. 1 2 3 4 5 6 7 8 Carleton and Olson, 1999, p. 16
  16. ↑ Voss and Carleton, 1993, p. 6
  17. ↑ Carleton and Olson, 1999, pp. 10–11
  18. 1 2 3 Carleton and Olson, 1999, p. 9
  19. ↑ Weksler, 2006, p. 131
  20. 1 2 3 4 5 Carleton and Olson, 1999, p. 13
  21. ↑ Carleton and Olson, 1999, pp. 12–13
  22. ↑ Carleton and Olson, 1999, p. 27; table 5
  23. ↑ Carleton and Olson, 1999, p. 28
  24. ↑ Carleton and Olson, 1999, pp. 29–32
  25. ↑ Carleton and Olson, 1999, table 1
  26. ↑ Carleton and Olson, 1999, table 2
  27. ↑ Carleton and Olson, 1999, table 3
  28. ↑ Carleton and Olson, 1999, p. 14
  29. ↑ Weksler, 2006, p. 30
  30. ↑ Weksler, 2006, p. 35
  31. ↑ Weksler, 2006, p. 40
  32. ↑ Weiss and Malabarba, 2005, pp. 153–154
  33. ↑ Kalthoff, 2000, p. 16
  34. ↑ Weiss and Malabarba, 2005, pp. 153–154; Kalthoff, 2000, p. 17
  35. ↑ Kalthoff, 2000, p. 55
  36. ↑ Weise and Malabarba, 2005, p. 159
  37. ↑ Carleton and Olson, 1999, fig. 2
  38. ↑ Weksler, 2006, pp. 43–44; Pardiñas, 2008, table 2
  39. ↑ Weksler, 2006, p. 55
  40. ↑ Carleton and Olson, 1999, p. 21
  41. ↑ Miralles et al., 2009, p. 57
  42. ↑ Carleton and Olson, 1999, p. 55
  43. ↑ Carleton and Olson, 1999, pp. 5–7
  44. ↑ Carleton and Olson, 1999, p. 8
  45. ↑ Carleton and Olson, 1999, pp. 55–56
  46. ↑ Carleton and Olson, 1999, p. 54
  47. ↑ Carleton and Olson, 1999, pp. 48–49
  48. ↑ Ridley, 1888, p. 474

Bibliography

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.