Mazama Rafinesque, 1817a

Jasper, Jonathan G, Lee, Thomas E, Zabel, Carson J, Twohy, Chelsea L, Lane, Kristina K & Robertson, Coral S, 2022, Mazama rufina (Artiodactyla: Cervidae), Mammalian Species 54 (1016), pp. 212-219 : 212-215

Mazama Rafinesque, 1817a

Moschus : Erxleben, 1777:319. Part, not Moschus Linnaeus, 1758 .

Cervus : Fischer, 1814:465. Part, not Cervus Linnaeus, 1758 .

Mazama Rafinesque, 1817a:44 . Type species Mazama pita Rafinesque, 1817b ; (= Cervus rufus Illiger, 1815 ), by subsequent designation (Merriam 1895).

Subulo C. Hamilton Smith, 1827:318 . No type species selected.

Passalites Gloger, 1841:140 . Type species Cervus nemorivagus Cuvier, 1817 , by monotypy.

Coassus J. E. Gray, 1843:174 . No type species selected.

Homelaphus J. E. Gray, 1872:90 . Type species Homelaphus inornatus Gray, 1872 (= Moschus americanus Erxleben, 1777 ), by monotypy.

Doryceros Fitzinger, 1873:360 . No type species selected.

Nanelaphus Fitzinger, 1873:360 . Part, type species Nanelaphus namby Fitzinger 1879: 26 (= Cervus gouazobira G. Fisher, 1814 ), by subsequent designation ( Lydekker 1898).

Cariacus : Brooke, 1878:918. Part, not Cariacus Lesson, 1842 .

Hippocamelus : Elliot, 1907:50. Part, not Hippocamelus Leuckart, 1816 .

Doratoceros Lydekker, 1915:210 . No type species mentioned; attributed to Fitzinger; lapsus evidente for Doryceros Fitzinger, 1873 .

Azarina Larrañaga, 1923:348 . No type species selected.

CONTEXT AND CONTENT. Order Artiodactyla , suborder Ruminantia , family Cervidae , subfamily Capreolinae , tribe Rangiferina ( Prothero and Liter 2007) . The genus Mazama currently contains eight species: Mazama americana , Mazama bororo , Mazama chunyi , Mazama gouazoubira , Mazama nana , Mazama nemorivaga , Mazama rufina , and Mazama temama ( Burgin et al. 2020) . The generic synonymy follows Martín-Varela et al. (2010) and Grubb (2005).

Mazama rufina (Pucheran, 1851) Dwarf Red Brocket

Cervus rufinus Pucheran, 1851:561 . Type locality “la vallée de Lloa, sur le versant occidental de la Cordillière du Pichincha;” (Pichincha, Pichincha Mountains, Lloa valley), Ecuador.

Cariacus [( Coassus )] rufinus : Brooke, 1878:925. Name combination.

Mazama tema : Lydekker, 1898:302. Part, not Mazama tema Rafinesque (1817b) (= Mazama temama Kerr, 1792 ).

Mazama bricenii Thomas, 1908:349 : Type locality “Paramo de la culata, Merida, Venezuela. Altitude 3000 m.”

Mazama rufinus : Thomas, 1908:349. Name combination.

Mazama rufina : Thomas, 1913:586. First use of current name combination.

Mazama bricenii : Lönnberg, 1913:1. Part, not Mazama bricenii Thomas, 1908:349 sensu Czernay (1987) .

CONTEXT AND CONTENT. Context as for genus. Some authors have cited Mazama nana and Mazama bricenii as subspecies of M. rufina ( Gutiérrez et al. 2015; Heckeberg et al. 2016; Lizcano and Alvarez 2016a), but currently M. rufina is considered monotypic ( Burgin et al. 2020). Historically, M. rufina and M. bricenii were considered separate species; however, compared to the other species, Mazama rufina was seen to be the most morphologically and genetically similar to M. bricenii . Allen (1915) remarked that M. rufina and M. bricenii are so similar in size, color, and pelage character that the two species would be regarded as local forms of the same species if they shared the same geographical range. Initially it was thought that the depth of the lacrima fossa and the shape of the frontal bones could be used to distinguish between M. rufina and M. bricenii ( Hershkovitz 1982) . However, these two characters have proven to be highly variable and an inconsistent morphometric marker for differentiating between the two ( Gutiérrez et al. 2015; Heckeberg 2020; see “Genetics” for more information concerning M. rufina and M. bricenii ). Mazama bricenii is currently considered a synonym of M. rufina and M. nana has been elevated to species status pending revisions of the genus ( Burgin et al. 2020).

NOMENCLATURAL NOTES. Mazama rufina was first documented by Jacques Pucheran (1851) and has many common names but is most well known as the dwarf red brocket ( Lizcano and Alvarez 2016a). An additional common name is little red brocket ( Mattioli 2011). Local names have also been found through personal interviews and descriptions of M. rufina , which are as follows: Venado chonta, venado colorado, cervicabra, soche enano or colorado, ciervo de monte, chivicabra, venado chudingo, venado de páramo, venado soche, corzuela enana, soche de páramo, yamala, and chonto ( Rodríguez-Mahecha et al. 1995; Tirira 2017). Hershkovitz (1982) noted the locals referred to M. rufina as venado chonta (chonta is a type of local palm tree with a dark, spiky bark) in some regions; the same term was used to describe Pudu mephistopheles ( Hershkovitz 1982; Hassanin et al. 2012). The etymology of Mazama is Mexican (Aztec) in origin meaning small deer. The name is derived from “mazame,” “maçame,” or “teuthlamaçame,” names used by Hernandez in 1651 for species of Mexican ungulates ( Palmer 1904). The species name rufina (Latin) means red-haired referring to the red fur of M. rufina .

DIAGNOSIS

Mazama rufina is a small-sized brocket (head–body length, 85–90 cm; height at shoulders, 45 cm; weight, 10–15 kg); pelage is reddish, head and legs are black ( Mattioli 2011). It is similar in size and coloration to other species of Mazama and Pudu . The three species of Mazama that have geographic distributions in close proximity to that of M. rufina are Mazama americana (common red brocket), M. chunyi (common dwarf brocket), and M. nemorivaga (Amazonian brown brocket— Mattioli 2011).

Mazama americana is the largest brocket (head–body length, 90–145 cm; height at shoulders, 60–80 cm; weight, typically 30–35 kg, but up to 65 kg) and notably larger than M. rufina ( Allen 1915; Mattioli 2011). It has cinnamon red fur on the body with face and legs ranging in color from brown to almost black and the neck is gray ( Groves and Grubb 2011; Mattioli 2011). Mazama chunyi is slightly smaller than M. rufina (head–body length, 70–75 cm; shoulder height, 38 cm; weight, 11 kg). Its coat is brown with reddish over the mid-back and flanks. Mazama nemorivaga is similar in size to M. rufina (head–body length, 75–100 cm; height at shoulders, 50 cm; weight 14–16 kg) but its coat is dark brown dorsally with faded-brown flanks.

Mazama rufina has distinctive facial features as described by Hershkovitz (1982) most notably a white mental and white narial patch ( Barrio 2010; Lee et al. 2011). These facial characteristics help distinguish M. rufina from the northern pudu ( Pudu mephistophiles ) which is sympatric. In contrast, the northern pudu has a completely black face, without any white markings around the mental or narial areas ( Fig. 1 ; Lee et al. 2011). Both species have similar preorbital glands and lacrimal fossae ( Hershkovitz 1982; Mattioli 2011; Tirira 2017). Compared to M. rufina , P. mephistophiles is smaller (head–body length, 75 cm; shoulder height, 25–38 cm; weight 5–6 kg — Mattioli 2011) with shorter legs and a darker fur that at no time shows a red color ( Tirira 2017). Pudu puda , the southern pudu, is slightly larger than the northern pudu (head–body length, 80 cm; height at shoulders, 30–40 cm; weight 9–10 kg — Mattioli 2011). Species of Pudu have some of the broadest metapodial shafts among cervids, thus allowing Puda to be distinguished from Mazama ( Hershkovitz 1982) . Mazama rufina is the smallest species of its genus found in Ecuador ( Eisenberg and Redford 1999).

GENERAL CHARACTERS

Mazama rufina is a small deer with a reddish-brown pelage that darkens in color as it transitions from the dorsum to ventral surface until it reaches the face and legs, where it appears black ( Fig. 1 ). Lehmann (1959) has reported completely albino M. rufina taken from the Páramo de las Delicias, Colombia. Some of the notable characteristics of M. rufina are the black ears, which have a frosted snowy-white outline around the edge and white tufts of hair on the inner side of the pinnae. Reddish burgundy hair covers the rest of the body, and the head has a black mask that extends from the nose to the nape ( Barrio 2010). This mask includes the chin but not the cheeks as the border of the mask rises above the buccal cavity under the eyes toward the ears. The buccal patch is dark red similar to the rest of the body ( Lee et al. 2011). The black fur begins at the legs and reaches down toward the hooves ( Barrio 2010). Mazama rufina also has defining white mental and narial patches on an otherwise dark face ( Hershkovitz 1982). It has extremely large preorbital glands, and a round preorbital fossa ( Hershkovitz 1982). Its antlers are in the shape of short, simple spikes ( Hershkovitz 1982; Goss 1983; Heckeberg 2020) which erupt from short and strongly inclined pedicles ( Heckeberg 2020). The small body size and simplicity of antlers of M. rufina are most likely adaptations to allow easier navigation of dense vegetation ( Tirira 2017; Heckeberg 2020).

Mean external measurements (cm or kg) for M. rufina recorded by Eisenberg and Redford (1999) were: head–body length, 85.3; length of hind foot, 26; tail length, 7.8; ear length, 5.3; shoulder height, 45; and weight 8.2. External measurements (cm or kg) as reported by Mattioli (2011) were: head–body length, 85–90; tail length, 8; shoulder height, 45; and weight 10–15 ( Mattioli 2011). Measurement ranges (cm or kg) from adult specimens in Ecuador were: head–body length, 77.5–98.5; tail length, 7.8–9.3; length of hind foot, 26; ear length, 8.3–9.4; shoulder height, 45; weight, 10–15 ( Tirira 2017). Gutiérrez et al. (2015) measured the typical body length between 60–140 cm, and shoulder height 34–80 cm. The antlers are unbranched, in the shape of small spikes; the length averages 6.9 cm and is rarely more than 8.0 cm ( Fig. 2 ; Goss 1983; Groves and Grubb 2011; Mattioli 2011). An old female collected on 15 May 1913 from Vallee de Lloa, Mt. Pichincha had the following skull measurements (cm), as collected by W. B. Richardson: total length of skull, 16.15; condylobasal length, 15.1; occipitonasal length, 14.25; zygomatic breadth, 7.2; interorbital breadth, 3.6; mastoid breadth, 5.55; breadth of braincase, 5.1; and maxillary toothrow 4.8 ( Allen 1915).

DISTRIBUTION

Mazama rufina can be found in small populations in the northwestern Andes above 1,000 m ( Fig. 3 ; Lizcano and Alvarez 2016b). The range of the montane forest (typical habitat of M. rufina ) reaches the northern limits of northern Colombia and western Venezuela ( Lizcano and Alvarez 2016b). In Venezuela, M. rufina can be found along the Zulia, Tachira, Apure, Merida, and Trujillo mountains ( Linares 1998). It is mostly found in Venezuelan parks due to forest distributions, primarily including Guaramacal National Park and Dinira National Park ( Lizcano and Alvarez 2016b). It can be found in the Colombian mountains of Antioquia Boyacá, Caldas, Cauca, Cesar, Huila, La Guajira, Nariño, Norte de Santander, Santander, Valle del Cauca, Tolima, Risaralda, and Quindío ( Linares 1998; Alberico et al. 2000; Lizcano and Alvarez 2016b). It is found in Tama national park in Norte de Santander ( Lizcano and Alvarez 2016b). Mazama rufina remains have been found on the eastern slopes of the Períja mountains in Cerro Viruela near Pico Tetari at 3,100 m ( Linares 1998), and it most likely inhabits the western slopes of these mountains as well ( Lizcano and Alvarez 2016b). In Ecuador its range is centered around the eastern and western Andes ( Tirira 2001). A partial skull of a male with antlers was found in temperate cloud forest (3,340 m) in the Guandera Biological Reserve in 2014 ( Lee et al. 2015) in an area where no other cervids are known to inhabit ( Tirira 2007). In Peru it can be found along the eastern slopes of the Andes in Amazon river watersheds ( Lizcano and Alvarez 2016a). The southern limit of the species is the Huancabamba depression ( Barrio 2010). Mazama rufina has been documented from 3,340 to 3,650 m in the Carchi Province of Ecuador ( Lee et al. 2015). The Carchi population was originally misidentified as Mazama nana by Siegfried ( Czernay 1987; Grubb 2005). The distribution was changed to follow revised species restrictions with addition of M. bricenii as a conspecific ( Burgin et al. 2020). The distribution M. rufina in Peru seems to be restricted to the montane forests of the Equatorial Yungas, in northern Cajamarca, and along the ridge between Piura and Cajamarca departments ( Fig. 3 ; Barrio 2010). No fossils are known.

FORM AND FUNCTION

Mazama rufina is shown to have a compressed ratio of metapodial width to length ( Hershkovitz 1982). This ratio is due to the narrow metapodial shafts that are generally found in brockets. Mazama rufina is also lacking its front digital bones and the vestigial II and V metacarpal bones are in the distal position ( Hershkovitz 1982). Metatarsals III and IV are fused and II and V have been lost ( Hershkovitz 1982).

Photos from Hershkovitz (1982) show that M. rufina has the dental formula i 0/4, c 0/0, p 3/3, m 3/3, total 32. This is congruent with the specimen ANMH (American Museum of Natural History) 66742 ( Fig. 2 ). The first lower incisors (i1) are spatulate in M. rufina , a feature shared in the genus ( Hershkovitz 1982; Heckeberg 2020). The upper molars have short crista, rising from the inner distal enamel border of the hypocone ( Hershkovitz 1982). The antlers of M. rufina grow for a considerable amount of time before shedding ( Czernay 1987).

ONTOGENY AND REPRODUCTION

Mazama rufina is not restricted to a mating season and is able to mate, give birth, and produce antlers year-round ( Goss 1983). It has an average gestation period of 210 days. At the end of this period, the female gives birth to a single offspring. The young can be identified by the white spots on their fur ( Hershkovitz 1982; Lizcano and Alvarez 2016a); they are hidden in vegetation and wait for their mother, who returns for feeding and protection. The fawn will continue to hide until it is older and strong enough to follow its mother. Young M. rufina are weaned at 6 months and can enter sexual maturity at 1 year of age ( Lizcano et al. 2010).

Figures

Fig. 1.—An adult female Mazama rufina from Sangay National Park, Morona Santiago Province, Ecuador: 2°11′03″S, 78°29′50″W, 3,123 m. Photographed by TEL.

Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view of mandible of an adult male Mazama rufina (AMNH [American Museum of Natural History] 66742) from 2,960 m, Rio Pita, Pichincha Province. The skull is 171 mm long, with a zygomatic breadth of 78 mm. Photograph by Nil Santana used with permission.

Fig. 3.—Geographic distribution of Mazama rufina. Map redrawn from Lizcano and Alvarez (2016a, 2016b) with modifications (ESRI 2020).