Diplomys labilis, Bangs, 1901Santamartamys rufodorsalis (J.A.Allen, 1899)Diplomys caniceps (Gunther, 1877)

Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Echimyidae, Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona: Lynx Edicions, pp. 552-604 : 590-591

71.

Central American Rufous Tree-rat

Diplomys labilis

French: Rat-épineux du Panama / German: Panama-Baumstachelratte / Spanish: Rata arboricola de Centroamérica

Other common names: Rufous Soft-furred Spiny-rat, Rufous Tree-rat

Taxonomy. Loncheres labilis Bangs, 1901 , “San Miguel Island, [Archipiélago de las Perlas], Panama.”

Diplomys labilis has been variously placed in the genera Echimys or Phyllomys by early authors; O. Thomas in 1916 was the first to assign it to Diplomys . Two subspecies recognized.

Subspecies and Distribution.

D. I. labilis Bangs, 1901 — San Miguel I, Pearl Is (off SC Panama).

D. l. darlingi Goldman, 1912 — from C Panama to the Choco and Antioquia departments of W Colombia and S along Pacific coast to NW Ecuador.

Descriptive notes. Head-body 340 mm, tail 200 mm; weight up to 492 g. The Central American Rufous Tree-rat is larger than the Colombian Rufous Tree-rat ( D. caniceps ). Its fur is stiff but soft, without bristles or spines. Upper parts are ocherous buff to bright rusty, with some black-tipped hairs. Face has grayish mask. Vibrissae are streaked at bases by distinctive white vertical stripes. Whitish stripe of hairs is present around edge of pinnae and behind ears. Ears are otherwise dark, rounded, and short, divided by large anterior lobe and smaller posterior lobe, edged by small fringe of longer hairs. Vibrissae are long, extending toward shoulders. Ventral colors are quite variable, ranging from dark to pale rufous. Face is marked with small pale spots above eyes, contrasting a black eye ring. Tail is robust, 70-100% of head-body length, unicolored brownish, and covered by short hairs that become longer toward tip. The Central American Rufous Tree-rat has distinct short tuft of hair at tip of tail. Like in the Colombian Rufous Tree-rat,tail scales are not visible beneath hair cover. Hindfeet and front feet are silvery gray above. Skull and teeth of the Central American Rufous Tree-rat are typical of Diplomys but with three major differences compared with the Colombian Rufous Tree-rat: narrower palate, premaxilla sometimes extending anteriorly beyond the nasals, and tooth rows that are parallel.

Habitat. Evergreen and deciduous forests, mangroves, cultivated plantations, and second-growth forests at elevations of 300-1500 m. The arboreal Central American Rufous Tree-rat occurs in lowland primary and secondary rainforests of north-western Colombia, southern Panama and northern Ecuador. A study of ectoparasites based on 77 captured specimens indicated preference for disturbed forests near agricultural plots of corn, yucca, coffee, and citrus fruits. It was mainly captured in trees preserved to provide shade for crops. All individuals were obtained from tree holes well above the ground and in trees mainly near rivers and covered with epiphytes that otherwise housed a large arboreal mammal community.

Food and Feeding. Captive Central America Rufous Tree-rats in Panama have been reported to feed on leaves, seeds, and fruits, thus suggesting a generalized frugivorous and folivorous diet.

Breeding. Female Central American Rufous Tree-rats give birth to precocial large, fully haired, and well-developed young of 30-40 g. One female was found with a near term fetus, weighting 59-5 g, or 12% of her 492g body weight. Pregnant females have been collected in January, March-May, August-September, and November. Recorded litter sizes are 1-2 young (mean 1-2). Nests are built in tree holes.

Activity patterns. The Central American Rufous Tree-ratis strictly nocturnal. During the day, individuals sleep individually or in pairs of adults in their tree hole nests. Movements, Home range and Social organization. The Central American Rufous Tree-rat has been reported to move slowly and avoid detection by remaining motionless for long time periods. Itis an agile climber. Local people comment that individuals may stick out their heads from their tree holes when disturbed. Central American Rufous Tree-rats have been reported to live in pairs that might travel and forage together at night. Captive individuals have been reported to be very aggressive, especially compared with their more gentle terrestrial and sympatric echimyid relatives, Hoplomys and Proechimys . Central American Rufous Tree-rats emit high-pitched calls when disturbed.

Emmons, 2005

73.

Red Crested Tree-rat

Santamartamys rufodorsalis

French: Rat-épineux a créte rouge / German: Rotschopf-Baumratte / Spanish: Rata arboricola de dorso rojo

Other common names: Red Crested Soft-furred Spiny-rat

Status and Conservation. Classified as Least Concern on The IUCN Red List. Parasitological surveys in Panama suggest that the Central American Rufous Tree-rat is abundant near disturbed habitat. Because much of its distribution has been logged, additional ecological studies of the Central American Rufous Tree-rat are needed because of its rarity in collections and lack offield studies.

Taxonomy. Isothrix rufodorsalis J. A. Allen, 1899 , “Onaca, Santa Marta District,”

Magdalena, Colombia.

Santamartamys rufodorsalis was formerly classified in the genera Echimys and Diplomys . Monotypic.

Distribution. N Colombia (Sierra Nevada de Santa Marta); it might occur in NW Venezuela.

Bibliography. Alberico et al. (2000), Allen (1916b), Bangs (1901), Eisenberg (1989), Emmons (1990, 1997a, 2005), Goldman (1912a, 1912b), Hall (1981), Handley (1966), Méndez (1967), Patton et al. (2015), Paynter (1993), Tesh (1970), Thomas (1916a, 1916b), Tirira (2007), Trouessart (1904), Woods (1993), Woods & Kilpatrick (2005)

72.

Colombian Rufous Tree-rat

Diplomys caniceps

French: Rat-épineux a téte grise / German: Graukopf-Baumstachelratte / Spanish: Rata arboricola de Bolivia

Other common names: Arboreal Soft-furred Spiny-rat, Colombian Soft-furred Spiny-rat

Taxonomy. Loncheres caniceps Gunther, 1877 , “Medellin,” Antioquia, Colombia .

Diplomys caniceps has been variously placed in the genera Loncheres or Phyllomys by 19" century authors; first assigned to Diplomys by O. Thomas in 1916. Monotypic.

Distribution. The Caribe Biogeographical Region of NW Colombia, in the Cauca Valley near Medellin, and possibly the Ser-rania de San Lucas in the Bolivar State.

Descriptive notes. Head-body 212-390 mm, tail 178-267 mm. No specific data are available for body weight. The Colombian Rufous Tree-rat is larger than its congener, the Central American Rufous Tree-rat ( D. labilis ). Its fur is stiff but soft, without bristles or spines. Overall upper parts are dull rusty, red-brown slightly streaked with dark brown hairs, but bright red on back one-half of body; forequarters are fulvous in some Colombian Rufous Tree-rats. Nose is grayish brown, and pre-auricular grayish patch extends partially to eyes. Face is marked with small whitish spots over eyes and at bases of vibrissae. A few whitish or yellowish spots are present below ear notches and behind ears. Ears are short, rounded, and divided by small anterior lobe and larger posterior lobe. Vibrissae are long, extending backward to shoulders. Belly and chin are whitish, yellowish, or pale orange; some individuals have rich orange venters. Dark rusty fur often occurs on midline of abdomen, extending posteriorly toward base of tail. Robusttail is 90-105% of head-body length and is covered sparsely by dark brown, bristle-like hairs;tail scales are not well exposed beneath hair cover. Forefeet are whitish-brown or gray-brown. Hindfeet are broad, yellow-brown to brown or gray above. Skulls of both species of Diplomys are characteristically convex in outline when viewed laterally. Rostrum is short and narrow; bullae are not inflated but possess well-developed auditory tube thatis oriented anteriorly; and auditory meatus is very close to squamosal bone. Jugal becomes narrow below post-orbital process of zygoma and exhibits an inconspicuous ventral process posterior to dorsal one. Alisphenoid arterial foramen pattern is unique in Diplomys , with short distance between buccinator foramen and foramen ovale. Hamular process of pterygoid bone has developed anteriorly oriented spur. Mandible has large masseteric ridge and characteristic shallow angle of sigmoid notch between condyloid and angular processes. Mandibular foramenis located anteriorly to condyloid ridge and not in a fossa. Cheekteeth are large and long and are highly convergent in occlusal morphology with those of species of Phyllomys . Incisors are strongly curved. Maxillary and mandibular tooth rows are strongly divergent antero-posteriorly, with outward curving of last two molars causing disjunction with straight edge formed by molars M' and M*. Habitat. Caribe Biogeographical Region rainforest of Colombia. Few data on specific habitats of the highly arboreal Colombian Rufous Tree-rat are available. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Additional ecologicalstudies of this rare Colombian endemic are needed. Bibliography. Alberico et al. (2000), Allen (1916b), Cuervo et al. (1986), Eisenberg (1989), Emmons (1990, 1997a, 2005), Goldman (1916), Gunther (1877a), Mares (1982), Patton et al. (2015), Thomas (1916a, 1916b), Trouessart (1880, 1904), Woods (1993), Woods & Kilpatrick (2005).

Descriptive notes. Head—body 190 mm, tail 267 mm; weight 492 g. The Red Crested Tree-rat is bright rustred, with legs and sides pale yellow-orange, gray underfur, and white belly. Distinct crest of long bright red hairs is present on crown between

.

629

FAMILY ECHIMYIDAE

Hutias, South American Spiny-rats and Coypu

Plate 38

Species Accounts

ears. Sides and upper part of muzzle tip are yellow-gray, streaked with black. Mask of pale yellow-gray occurs between eyes, ears, and on rostrum, delimited dorsally by dark stripe of fur. Chest and throat are pale orange, and tip of chin is white. Its fur is soft, without any bristles or spines, having long, lax dorsal guard hairs. Dense woolly fur covers legs, ankles, and wrists. Vibrissae are short and do not extend beyond ears. Ears are short and naked, with tufts of long hairs emerging from inner surfaces. Like in species of Diplomys , the Red Crested Tree-rat has two pairs of lateral abdominal mammae along ventro-lateral side of body. Robusttail is c¢.140% of head-body length and densely clothed with 20 mm of rusty basal hairs, followed by large band of black and brown hairs covering scales for three-fifths of tail length and ending with pure white part. Hindfeet and front feet are brown above, washed with silvery gray color. There are no palmar or plantar tubercles, and pollex has a nail. Skull of the Red Crested Tree-rat is curved in dorsal profile, as in species of Diplomys , rostrum is short and broad. Similar to species of Diplomys , small auditory meatusis placed close to squamosal bone but has short auditory tube, anteriorly oriented. Auditory bullae are flattened and relatively small compared with skull length and merged with alisphenoid bone at shallow angles. Lateral process of supraoccipital is extremely short. Long bony bridge is present between foramen ovale and masticatory foramen. Condyloid process ofjaw is deep; shallow angle is present between sigmoid notch and angular and condyloid processes. Mandibular foramina are located in fossa downward on condyloid ridge. Pterygoid shelf is small. Lower incisors are strongly curved. Distinct maxillary and mandibular cheekteeth are rectangular and longer than wide. Tooth rows are long and bowed inward. Teeth differ from species of Diplomys , with only one separation between first two lophs of fourth upper premolar. Lophidsare still connected on lower molars and are very distinct compared with species of Diplomys .

Habitat. Endemic in montane primary and secondary rainforests of the Sierra Nevada de Santa Marta, north-eastern Colombia at elevations of 680-2000 m.

Food and Feeding. There is no specific information available for this species, but the Red-Crested Tree-rat is likely folivorous.

Breeding. There is no information available for this species.

Activity patterns. There is no information available for this species.

Movements, Home range and Social organization. There is no information available for this species.

Status and Conservation. Classified as Critically Endangered on The IUCN Red List. The Red-Crested Tree-rat was previously classified as Vulnerable, but the Critically Endangered status is a consequence of the decline known in the extent of occurrence, important fragmentation of the population, small area of occupancy, and degradation of habitat in this region. Additional ecological and field studies are needed to have a better estimate ofits distribution and area of occupancy. There are no data on abundance and survival capacities of the Red-Crested Tree-rat, but available records indicate that it is rare. It is only known from three specimens in the American Museum of Natural History and a potential unrecorded specimen in the Instituto de Investigacion Alexander von Humboldt in Colombia. One individual was observed and photographed at the El Dorado Eco Lodge after more than a century without confirmed records.

Bibliography. Allen (1899b), Eisenberg (1989), Emmons (1990, 1997a, 2005), Emmons & Patton (2015e), Woods & Kilpatrick (2005).

73.

Red Crested Tree-rat

Santamartamys rufodorsalis

French: Rat-épineux a créte rouge / German: Rotschopf-Baumratte / Spanish: Rata arboricola de dorso rojo

Other common names: Red Crested Soft-furred Spiny-rat

Status and Conservation. Classified as Least Concern on The IUCN Red List. Parasitological surveys in Panama suggest that the Central American Rufous Tree-rat is abundant near disturbed habitat. Because much of its distribution has been logged, additional ecological studies of the Central American Rufous Tree-rat are needed because of its rarity in collections and lack offield studies.

Taxonomy. Isothrix rufodorsalis J. A. Allen, 1899 , “Onaca, Santa Marta District,”

Magdalena, Colombia.

Santamartamys rufodorsalis was formerly classified in the genera Echimys and Diplomys . Monotypic.

Distribution. N Colombia (Sierra Nevada de Santa Marta); it might occur in NW Venezuela.

Bibliography. Alberico et al. (2000), Allen (1916b), Bangs (1901), Eisenberg (1989), Emmons (1990, 1997a, 2005), Goldman (1912a, 1912b), Hall (1981), Handley (1966), Méndez (1967), Patton et al. (2015), Paynter (1993), Tesh (1970), Thomas (1916a, 1916b), Tirira (2007), Trouessart (1904), Woods (1993), Woods & Kilpatrick (2005)

72.

Colombian Rufous Tree-rat

Diplomys caniceps

French: Rat-épineux a téte grise / German: Graukopf-Baumstachelratte / Spanish: Rata arboricola de Bolivia

Other common names: Arboreal Soft-furred Spiny-rat, Colombian Soft-furred Spiny-rat

Taxonomy. Loncheres caniceps Gunther, 1877 , “Medellin,” Antioquia, Colombia .

Diplomys caniceps has been variously placed in the genera Loncheres or Phyllomys by 19" century authors; first assigned to Diplomys by O. Thomas in 1916. Monotypic.

Distribution. The Caribe Biogeographical Region of NW Colombia, in the Cauca Valley near Medellin, and possibly the Ser-rania de San Lucas in the Bolivar State.

Descriptive notes. Head-body 212-390 mm, tail 178-267 mm. No specific data are available for body weight. The Colombian Rufous Tree-rat is larger than its congener, the Central American Rufous Tree-rat ( D. labilis ). Its fur is stiff but soft, without bristles or spines. Overall upper parts are dull rusty, red-brown slightly streaked with dark brown hairs, but bright red on back one-half of body; forequarters are fulvous in some Colombian Rufous Tree-rats. Nose is grayish brown, and pre-auricular grayish patch extends partially to eyes. Face is marked with small whitish spots over eyes and at bases of vibrissae. A few whitish or yellowish spots are present below ear notches and behind ears. Ears are short, rounded, and divided by small anterior lobe and larger posterior lobe. Vibrissae are long, extending backward to shoulders. Belly and chin are whitish, yellowish, or pale orange; some individuals have rich orange venters. Dark rusty fur often occurs on midline of abdomen, extending posteriorly toward base of tail. Robusttail is 90-105% of head-body length and is covered sparsely by dark brown, bristle-like hairs;tail scales are not well exposed beneath hair cover. Forefeet are whitish-brown or gray-brown. Hindfeet are broad, yellow-brown to brown or gray above. Skulls of both species of Diplomys are characteristically convex in outline when viewed laterally. Rostrum is short and narrow; bullae are not inflated but possess well-developed auditory tube thatis oriented anteriorly; and auditory meatus is very close to squamosal bone. Jugal becomes narrow below post-orbital process of zygoma and exhibits an inconspicuous ventral process posterior to dorsal one. Alisphenoid arterial foramen pattern is unique in Diplomys , with short distance between buccinator foramen and foramen ovale. Hamular process of pterygoid bone has developed anteriorly oriented spur. Mandible has large masseteric ridge and characteristic shallow angle of sigmoid notch between condyloid and angular processes. Mandibular foramenis located anteriorly to condyloid ridge and not in a fossa. Cheekteeth are large and long and are highly convergent in occlusal morphology with those of species of Phyllomys . Incisors are strongly curved. Maxillary and mandibular tooth rows are strongly divergent antero-posteriorly, with outward curving of last two molars causing disjunction with straight edge formed by molars M' and M*. Habitat. Caribe Biogeographical Region rainforest of Colombia. Few data on specific habitats of the highly arboreal Colombian Rufous Tree-rat are available. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Additional ecologicalstudies of this rare Colombian endemic are needed. Bibliography. Alberico et al. (2000), Allen (1916b), Cuervo et al. (1986), Eisenberg (1989), Emmons (1990, 1997a, 2005), Goldman (1916), Gunther (1877a), Mares (1982), Patton et al. (2015), Thomas (1916a, 1916b), Trouessart (1880, 1904), Woods (1993), Woods & Kilpatrick (2005).

Descriptive notes. Head—body 190 mm, tail 267 mm; weight 492 g. The Red Crested Tree-rat is bright rustred, with legs and sides pale yellow-orange, gray underfur, and white belly. Distinct crest of long bright red hairs is present on crown between

.

629

FAMILY ECHIMYIDAE

Hutias, South American Spiny-rats and Coypu

Plate 38

Species Accounts

ears. Sides and upper part of muzzle tip are yellow-gray, streaked with black. Mask of pale yellow-gray occurs between eyes, ears, and on rostrum, delimited dorsally by dark stripe of fur. Chest and throat are pale orange, and tip of chin is white. Its fur is soft, without any bristles or spines, having long, lax dorsal guard hairs. Dense woolly fur covers legs, ankles, and wrists. Vibrissae are short and do not extend beyond ears. Ears are short and naked, with tufts of long hairs emerging from inner surfaces. Like in species of Diplomys , the Red Crested Tree-rat has two pairs of lateral abdominal mammae along ventro-lateral side of body. Robusttail is c¢.140% of head-body length and densely clothed with 20 mm of rusty basal hairs, followed by large band of black and brown hairs covering scales for three-fifths of tail length and ending with pure white part. Hindfeet and front feet are brown above, washed with silvery gray color. There are no palmar or plantar tubercles, and pollex has a nail. Skull of the Red Crested Tree-rat is curved in dorsal profile, as in species of Diplomys , rostrum is short and broad. Similar to species of Diplomys , small auditory meatusis placed close to squamosal bone but has short auditory tube, anteriorly oriented. Auditory bullae are flattened and relatively small compared with skull length and merged with alisphenoid bone at shallow angles. Lateral process of supraoccipital is extremely short. Long bony bridge is present between foramen ovale and masticatory foramen. Condyloid process ofjaw is deep; shallow angle is present between sigmoid notch and angular and condyloid processes. Mandibular foramina are located in fossa downward on condyloid ridge. Pterygoid shelf is small. Lower incisors are strongly curved. Distinct maxillary and mandibular cheekteeth are rectangular and longer than wide. Tooth rows are long and bowed inward. Teeth differ from species of Diplomys , with only one separation between first two lophs of fourth upper premolar. Lophidsare still connected on lower molars and are very distinct compared with species of Diplomys .

Habitat. Endemic in montane primary and secondary rainforests of the Sierra Nevada de Santa Marta, north-eastern Colombia at elevations of 680-2000 m.

Food and Feeding. There is no specific information available for this species, but the Red-Crested Tree-rat is likely folivorous.

Breeding. There is no information available for this species.

Activity patterns. There is no information available for this species.

Movements, Home range and Social organization. There is no information available for this species.

Status and Conservation. Classified as Critically Endangered on The IUCN Red List. The Red-Crested Tree-rat was previously classified as Vulnerable, but the Critically Endangered status is a consequence of the decline known in the extent of occurrence, important fragmentation of the population, small area of occupancy, and degradation of habitat in this region. Additional ecological and field studies are needed to have a better estimate ofits distribution and area of occupancy. There are no data on abundance and survival capacities of the Red-Crested Tree-rat, but available records indicate that it is rare. It is only known from three specimens in the American Museum of Natural History and a potential unrecorded specimen in the Instituto de Investigacion Alexander von Humboldt in Colombia. One individual was observed and photographed at the El Dorado Eco Lodge after more than a century without confirmed records.

Bibliography. Allen (1899b), Eisenberg (1989), Emmons (1990, 1997a, 2005), Emmons & Patton (2015e), Woods & Kilpatrick (2005).

72.

Colombian Rufous Tree-rat

Diplomys caniceps

French: Rat-épineux a téte grise / German: Graukopf-Baumstachelratte / Spanish: Rata arboricola de Bolivia

Other common names: Arboreal Soft-furred Spiny-rat, Colombian Soft-furred Spiny-rat

Taxonomy. Loncheres caniceps Gunther, 1877 , “Medellin,” Antioquia, Colombia .

Diplomys caniceps has been variously placed in the genera Loncheres or Phyllomys by 19" century authors; first assigned to Diplomys by O. Thomas in 1916. Monotypic.

Distribution. The Caribe Biogeographical Region of NW Colombia, in the Cauca Valley near Medellin, and possibly the Ser-rania de San Lucas in the Bolivar State.

Descriptive notes. Head-body 212-390 mm, tail 178-267 mm. No specific data are available for body weight. The Colombian Rufous Tree-rat is larger than its congener, the Central American Rufous Tree-rat ( D. labilis ). Its fur is stiff but soft, without bristles or spines. Overall upper parts are dull rusty, red-brown slightly streaked with dark brown hairs, but bright red on back one-half of body; forequarters are fulvous in some Colombian Rufous Tree-rats. Nose is grayish brown, and pre-auricular grayish patch extends partially to eyes. Face is marked with small whitish spots over eyes and at bases of vibrissae. A few whitish or yellowish spots are present below ear notches and behind ears. Ears are short, rounded, and divided by small anterior lobe and larger posterior lobe. Vibrissae are long, extending backward to shoulders. Belly and chin are whitish, yellowish, or pale orange; some individuals have rich orange venters. Dark rusty fur often occurs on midline of abdomen, extending posteriorly toward base of tail. Robusttail is 90-105% of head-body length and is covered sparsely by dark brown, bristle-like hairs;tail scales are not well exposed beneath hair cover. Forefeet are whitish-brown or gray-brown. Hindfeet are broad, yellow-brown to brown or gray above. Skulls of both species of Diplomys are characteristically convex in outline when viewed laterally. Rostrum is short and narrow; bullae are not inflated but possess well-developed auditory tube thatis oriented anteriorly; and auditory meatus is very close to squamosal bone. Jugal becomes narrow below post-orbital process of zygoma and exhibits an inconspicuous ventral process posterior to dorsal one. Alisphenoid arterial foramen pattern is unique in Diplomys , with short distance between buccinator foramen and foramen ovale. Hamular process of pterygoid bone has developed anteriorly oriented spur. Mandible has large masseteric ridge and characteristic shallow angle of sigmoid notch between condyloid and angular processes. Mandibular foramenis located anteriorly to condyloid ridge and not in a fossa. Cheekteeth are large and long and are highly convergent in occlusal morphology with those of species of Phyllomys . Incisors are strongly curved. Maxillary and mandibular tooth rows are strongly divergent antero-posteriorly, with outward curving of last two molars causing disjunction with straight edge formed by molars M' and M*. Habitat. Caribe Biogeographical Region rainforest of Colombia. Few data on specific habitats of the highly arboreal Colombian Rufous Tree-rat are available. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Additional ecologicalstudies of this rare Colombian endemic are needed. Bibliography. Alberico et al. (2000), Allen (1916b), Cuervo et al. (1986), Eisenberg (1989), Emmons (1990, 1997a, 2005), Goldman (1916), Gunther (1877a), Mares (1982), Patton et al. (2015), Thomas (1916a, 1916b), Trouessart (1880, 1904), Woods (1993), Woods & Kilpatrick (2005).