mmsp_04_209.1

AMMALIAN

PECIES

No. 758, pp. 1–9, 3 figs.

Chinchilla laniger.

By Angel E. Spotorno, Carlos A. Zuleta, J. Pablo Valladares,

Amy L. Deane, and Jaime E. Jime´nez

Published 15 December 2004 by the American Society of Mammalogists

. 1.

Photograph of an adult male

Chinchilla laniger

from

border of Reserva Nacional Las Chinchillas, Auco´, Illapel, IV Re-
gio´n, Chile (specimen Coleccio´n del Laboratorio de Citogene´tica,
Facultad de Medicina, Universidad de Chile [LCM] 400). Photo-
graph by A. Spotorno O. (1975).

Chinchilla

Bennett, 1829

Mus

: Molina, 1782:301. Part, not

Mus

Linnaeus, 1758.

Lemmus

: Tiedemann, 1808:476. Part, not

Lemmus

Link, 1795.

Cricetus

: E`. Geoffroy St.-Hilaire, 1803:197. Part, not

Cricetus

Les-

ke, 1779.

Chinchilla

Bennett, 1829:1. Type species

Mus laniger

Molina,

1782, by monotypy.

Eriomys

Lichtenstein, 1830:plate 28. Type species

Eriomys chin-

chilla

Lichtenstein, 1830, by monotypy.

Callomys

d’Orbigny and I. Geoffroy Saint-Hilaire, 1830:289. Part,

no type species selected.

Aulacodus

: Kaup, 1832:column 1211. Not

Aulacodus

Temminck,

1827:245.

Lagostomus

: Cuvier, 1830:plate 64. Not

Lagostomus

Brookes,

1828.

CONTEXT AND CONTENT.

Order Rodentia, suborder

Hystricognatha (Caviomorpha), superfamily Chinchilloidea, family
Chinchillidae, subfamily Chinchillinae. Chilean chinchilla has
been treated as 1 of 3 subspecies of

Chinchilla chinchilla

(e.g.,

Osgood 1943), but most recent authorities recognize 2 species of

Chinchilla

C. laniger

and

C. brevicaudata

(Cabrera 1961; Corbet

and Hill 1980; Mun˜oz-Pedreros 2000; Woods 1993), a distinction
that is congruent with recent molecular data (Spotorno et al. 2004).
Valladares and Spotorno (2003) have submitted a petition to the
International Commission on Zoological Nomenclature to conserve
the specific names

Mus laniger

Molina, 1782, and

Eriomys chin-

chilla

Lichtenstein, 1830, for such 2 species but the petition is not

yet published, and the Commission has not rendered an opinion.

Chinchilla laniger

(Molina, 1782)

Chilean Chinchilla

Mus laniger

Molina, 1782:301. Type locality ‘‘Provincie Boreali

del Chili,’’ [northern provinces of Chile]; given as ‘‘Coquimbo’’
by Osgood (1943:134).

Cricetus laniger

: E`. Geoffroy St.-Hilaire, 1803:197. Name combi-

nation.

[

emmus

laniger

: Tiedemann, 1808:476. Name combination.

[

ricetus

chincilla

Fischer, 1814:55. Type locality ‘‘Chile boreal-

ibus,’’ based on

Mus laniger

Molina, 1782.

Chinchilla lanigera

Bennett, 1829:1. First use of current name

combination and incorrect subsequent spelling of

Mus laniger

Molina, 1782.

Callomys laniger

: d’Orbigny and I. Geoffroy Saint-Hilaire, 1830:

289. Name combination.

Lagostomus laniger

: Cuvier, 1830:plate 64. Name combination.

Aulacodus laniger

: Kaup, 1832:column 1211. Name combination.

Chinchilla velligera

Prell, 1934:100. Replacement name for

Mus

laniger

Molina, 1782.

CONTEXT AND CONTENT.

Context as for genus.

Chin-

chilla laniger

is monotypic.

DIAGNOSIS.

Body size of

Chinchilla laniger

(Fig. 1) is

smaller (

260 mm in length) than that of

Chinchilla brevicaudata

(

320 mm), with more rounded ears (

45 mm in length) and a

longer (

130 mm) hairy tail (Redford and Eisenberg 1992) than

those of

C. brevicaudata

(

32 mm and

100 mm, respectively).

Number of caudal vertebrae is 23 in

C. laniger

and 20 in

brevicaudata

(Grau 1986).

GENERAL CHARACTERS.

Wild Chilean chinchilla from

Reserva Nacional Las Chinchillas in Auco´ are smaller and less

sexually dimorphic than domestic animals: males weigh (mean

unless otherwise indicated) 412

9 g (range, 369–493 g;

15) and females weigh 422

7 g (range, 379–450 g;

14—

Jime´nez 1990a). Domestic females weigh up to 800 g, whereas
males reach near 600 g (Neira et al. 1989).

Hair is 2–4 cm long, with gray, white, and black bands; it is

silky, extremely soft, and firmly adhered to skin (Bennett 1835);
thus, it is considered commercially valuable (Corte´s et al. 2000;
Seele 1968). Many hairs (50–75) emerge together from a single skin
pore. Wool hairs, 5–11

m in diameter, are grouped in 2 lateral

clusters. A single guard hair, 10–15

m in diameter, is located at

center of each cluster. Vibrissae are abundant, strong, long (100–
130 mm), and emerge from single follicles (Wilcox 1950).

General color of upper parts is bluish or silvery gray, under

parts are yellowish white. Tail has long, coarse, gray and black
hairs on its dorsal surface that are 30–40 mm long near the body,
50–60 mm long near the tip (Albert 1900), and form a bristly tuft
that exceeds vertebrae by 50 mm (Bennett 1835).

Head is broad, with rounded large ears (range, 45–48 mm;

mean, 46.7;

3) covered by tiny hairs. Skull (Fig. 2) has very

expanded auditory bullae, which protrude over the upper plane.
Forelimbs are much smaller in size than hind legs (Yarrell 1831).
Tibia is longer than femur, and fibula is virtually nonexistent
(Schaeffer and Donnelly 1997). Short forefeet have 5 digits, and
narrow hind feet have 3 digits plus 1 rudimentary having stiff bris-
tles surrounding a small and flat claw. Other claws present but very
short.

DISTRIBUTION.

Wild populations of

C. laniger

occur in

Auco´, near Illapel, IV Regio´n, Chile (31

S, 71

W), in Re-

serva Nacional Las Chinchillas (Fig. 3) and in La Higuera, ca. 100
km north of Coquimbo (29

S, 71

W—Jime´nez 1996). Chi-

lean chinchillas were reported from Talca (35

S), Chile, reach-

ing north to Peru (Bennett 1835), and eastward from Chilean coastal

MAMMALIAN SPECIES

758—

Chinchilla laniger

. 2.

Dorsal, ventral, and lateral views of cranium and lat-

eral view of mandible of an adult female

Chinchilla laniger

from

Auco´, Illapel, IV Regio´n, Chile (specimen Z65, Universidad de La
Serena Collection, Chile). Greatest length of cranium is 59 mm.

. 3.

Historic (ca. 18th century) geographic distribution of

Chinchilla laniger

(modified from Grau [1986]). Stars indicate

sites of current wild populations (Jime´nez 1996): La Higuera, Co-
quimbo, and Auco´, Illapel, Chile.

hills throughout low mountains. By the mid-19th century, Chilean
chinchillas were not found south of the Choapa river (32

S—Gay

1847; Jime´nez 1990a,b). They were mistakenly reported to occur
far south, at 52

S (Walker 1968). No fossils are known.

FORM.

Epidermis of domestic adults has 3 recognizable lay-

ers: a 2-cell-thick stratum germinativum, a single-cell stratum gran-
ulosum, and a thick stratum corneum. No stratum lucidum is found
between last 2 (Wilcox 1950).

Each group of hairs has 1 arrector pili muscle. Guard hairs

have their own sebaceous glands, whereas 3 or 4 other glands serve
all hairs of a lateral group. Solitary hairs are found as vibrissae or
on tail and pinnae. Follicular layering is found only in vibrissae
(Wilcox 1950).

Females have 2 pairs of thoracic and 1 pair of inguinal mam-

mae (Weir 1974), although latter are not visible in wild females.
Area around mammae is devoid of fur during lactation.

Skull is relatively broad, with little or no ridging. Infraorbital

canal is very large, without a distinct groove for nerve passage; at

its front edge, lacrimal canal has large opening on rostrum side.
Paroccipital process is short and attached to bulla. Jugal approachs
lacrimal (Lawlor 1979). Encephalization quotient for domestic an-
imals (ratio of observed versus expected brain mass for a defined
body mass) is 1.342 (Eisenberg 1981). Pituitary is 3.46 mg/% of
body weight in males and 4.02 mg/% in females (Asdell 1964).

Auditory bullae have 3 large vesicular protuberances; superior

protuberance is nearly hemispherical, posterior is oblong, and in-
ferior is pyriform (Bennett 1835). Mean total volume of middle ear
is 1.52 ml

0.26

(

9—Vrettakos et al. 1988). Tympanic

membrane is nearly parallel to medial wall of external meatus and
anchored to bony annulus; maximum diameter is 8.53 mm

0.18

SD,

and minimum diameter is 8.32

0.27 mm (

5), with a

circumference of 26.47

0.65 mm, and a depth of its cone of 1.78

MAMMALIAN SPECIES

758—

Chinchilla laniger

0.22 mm (

5). Tympanic flat area is 55.64 mm

2.7

SD,

but when its conical form is considered, average area increases 9%,
to a mean of 60.44

2.3 mm

(

5). In the ossicular chain,

incudomalleolar joint is fused, and 2 ossicles form a curved bar
that fit snugly in epitympanic recess. Malleus handle and posterior
margin of incus long process are almost perpendicular to rotational
axis. Footplate of stapes has a toe oriented posteriorly; crura of
stapes are more central than in humans and mice. No stapedial
artery occurs in adult Chilean chinchilla. Overall stapes height from
footplate plate to incudostapedial joint is 1.78 mm, with mean mass
0.4 mg

0.3

(

6). Oval window has diameters of 2.19 and

0.78–0.93 mm. Round window is oval, with diameters of 1.40 and
0.95 mm. Ligamentous attachments occur between short process of
incus and fossa incudis, as well as other 3 ossicular ones. Tensor
tympani and stapedius muscles are well developed (Vrettakos et al.
1988).

Middle ear mucosa consist of columnar, cuboidal, and squa-

mous epithelia. Lining epithelium of eustachian tube is ciliated
pseudostratified columnar. Density of ciliated cells is highest in
columnar epithelium area of transitional zone. Density of secretory
cells is highest in pharyngeal orifice (Hanamure and Lim 1987).

Teeth are hypsodont and evergrowing (rootless). Upper tooth-

rows are convergent anteriorly and crowns of cheekteeth flat, con-
sisting of a series of transverse plates (Bennett 1835). Dental for-
mula is i 1/1, c 0/0, p 1/1, m 3/3, total 20. Incisors grow 5–7.5 cm
per year. Enamel is usually dark yellow in healthy animals (Hillyer
et al. 1997); some inner radial enamel occurs close to dentin–
enamel junction, as well as in portio externa (Von Koenigswald
1985).

Lower jaw has no ridge or groove on lateral surface; angular

process is elongated and not deflected. Condyle is small and lon-
gitudinal, and glenoid cavity is superficial, permitting motion in
anterior–posterior direction.

Vertebral formula is 7 C, 13 T, 6 L, 2 S, 23 Ca, total 51

(Bennett 1835), with 13 pairs of ribs (Yarrell 1831). Radius and
ulna, although distinct, are so closely applied to each other at their
carpal extremity that they appear anchylosed for half their length.
Interarticular cartilage between clavicle and sternum is long, ca. 6
mm.

Masseter is divided in 3 portions. The superficial portion aris-

es by a strong tendon from ventral side of zygoma below infraorbital
foramen and inserts along ventral margin of angle of jaw. The lateral
portion arises from zygoma and inserts on ventral part of masseteric
fossa and ventral margin of mandible. The deep portion also is
subdivided: anterior deep portion arises from dorsal part of max-
illary portion of snout and passes backward and downward through
a large infraorbital foramen; posterior deep portion arises from ven-
tral surface of zygoma and inserts along dorsal half of masseteric
fossa. Gluteus medius is enlarged. Temporalis is reduced and un-
differentiated.

Hearts in domestic animals are more elongated and more

rounded at apex than nearly quadrate heart of

Lagidium

. Internal

carotid artery is absent and brain is supplied by vertebral–basilar
artery system alone, whereas external carotid artery system has as-
sumed entire stapedial area of supply by means of 3 anastomoses;
internal ophthalmic artery aborts (Bugge 1984). Arterial irrigation
of testicles and epididymis proceeds from testicular arteries, which
emerge from renal arteries; accesorial sexual glands and ductus
deferens irrigation is from external iliac artery (Adaro et al. 1998).

Lungs are asymmetric with 3 lobes in left and 4 in right; lower

lobe is the largest, deeply bifid in latter. Trachea is transversely
oval, with rings dorsally imperfect (Bennett 1835).

Vomeronasal organ is a pair of tubular structures, ca. 6 mm

in length, situated bilaterally along base of nasal septum. Its sen-
sory epithelium consists of sensory, supporting, and basal cells.
Acinar cells of Jacobson organ in respiratory epithelium possess 2
types of secretory granules, 1 homogeneous and ca. 1,700 nm in
diameter, and the other ca. 2,200 nm and varying in electron den-
sity (Oikawa et al. 1994).

Oral cavity is small and narrow. Oropharynx communicates

with pharynx through palatal ostium (Hillyer et al. 1997). Stomach
is pyriform, with a length of 63 mm, its greatest breadth on left is
44 mm and in middle is 25 mm (Bennett 1835). Esophagus enters
near middle of cavity, and pyloric portion forms a curve upward,
on which beginning of duodenum makes a sudden turn (Bennett
1835). Liver has 4 lobes, 2 large and 2 small (Yarrell 1831), with

both cystic and lobulus spigelii deeply cleft. Spleen is 25 mm in
length, with a breadth of 22 mm at its lower extremity.

Small and large intestine in 1 adult domestic animal measured

350 cm (Hillyer et al. 1997). In a wild specimen with total length
of nearly 21 cm, total length of small intestine was ca. 117 cm,
and of large intestine, without cecum, was 145 cm. Duodenum, after
descending to right lumbar region, turns upward, crosses spine, and
then becomes free. A relatively large and coiled cecum (Hillyer et
al. 1997) occupies left iliac region and has its blind extremity con-
cealed behind 1st fold (Bennett 1835). Colon curves upward and
toward left side, and then by a 2nd and very acute fold turns upon
itself again and descends to hypogastrium. It reascends to hypo-
chondrium, where it is attached to duodenum by peritoneum, and
then returns on itself to descend in a long, loose, double fold, 23
cm in length. Formation of pellets begins at this long fold (Bennett
1835).

Kidneys are 19 mm long by 13 mm in breadth. Renal capsules

are oblong white bodies, mesiad of upper part of kidney, and mea-
sure near 12 mm in length (Dı´az and Corte´s 2003).

Female reproductive system includes 2 uterine horns and 2

cervices (Weir 1970). Ovary is ovoid and suspended from dorsal
wall of abdomen by mesovarium. Fallopian tubes are suspended by
mesosalpinx mesenterium, which may partly invest ovary, forming
a very thin and tenuous partial bursa ovarica. Fimbria are not prom-
inent. Blood vessels and nerves enter ovary along cranial part of
mesovarium, and hilum is well defined and extensive. Ovary surface
is smooth and follicles may appear translucent but rarely project
as domes on surface. Stigmata are not prominent and corpus lutea
are not visible; therefore, ovulation is difficult to detect a few days
later (Weir and Rowlands 1974). Polyovular follicles are particu-
larly frequent at birth; 61% of animals had 0.4–19% of such fol-
licles.

Testes are relatively large. Those from 4 wild mature males

were 21–29 mm long (mean

26.2), with mass of 3.0–5.1 g (mean

4.3).

C. laniger

lacks a true scrotum; instead, testes are con-

tained within inguinal canal, and caudal epididymis can drop into
small moveable postanal sacs. Inguinal canal is open (Weir 1970).

Accesory reproductive glands of domestic males include large

bulbourethral glands (mean diameter

6.4

0.16 mm;

50) that are surrounded by a capsule of dense connective tissue

and striated muscles, with fibers arranged in 3 dimensions (Cepeda
et al. 1999). Presence of prostate is variable (Adaro et al. 1999;
Weir 1974). Large vesicular glands provide the bulk of accessory
gland secretions, and this fluid gels to form a copulatory plug in
females (Weir 1974).

Penis is separated from anus by bare skin (Weir 1974). Glans

is partially covered with small simple spines directed toward body.
A ventral 7-mm-deep sacculus urethralis or penian intromittent sac
with small spines directed away from body occurs below meatus
urinarium. Two large elastic bands connect to the internal part of
sacculus bottom and, together with directions of spines, suggest that
whole sacculus is everted during copulation. No spikes are present
at external bottom of sacculus urethralis (Spotorno 1979).

Large black eyes have a vertical slit pupil (Walker 1968).

Crystalline lens is large and convex.

FUNCTION.

Middle ear conductive apparatus of domestic

Chilean chinchilla is attached to surrounding cavity by suspensory
ligaments, and thus is free to move independently. The single axis
for the mobile middle ear of Chilean chinchilla passes through the
center of gravity. When ossicles are in motion, no force is generated
that would cause them to move out of axis. Bearings or suspensory
ligaments, because they lie on axis, minimize resistance. Large
mass of anterior process malleus also is distributed evenly, reducing
moment of inertia, and probably contributing to high-frequency re-
sponse (Vrettakos et al. 1988).

Basal metabolic rate of wild

C. laniger

(

6) averages 0.66

ml O

, and skin thermal conductance is 0.0376 ml O

. Thermal conductance in He-O

is 0.089 ml O

. Thus, Chilean chinchilla has the highest thermal insulation

(Corte´s et al. 2000).

Evaporative water loss of wild Chilean chinchilla (mean

6) is 0.58

0.10 mg H

O g

at ambient temperature

C. At temperatures of 25 and 30

C, evaporative water loss is

0.74

0.09 and 0.89

0.14 mg H

O g

, respectively (Corte´s

et al. 2000).

Chinchilla laniger

produces concentrated urine. In 10 do-

MAMMALIAN SPECIES

758—

Chinchilla laniger

mestic Chilean chinchillas deprived of water for up to 8 days,
mean urine osmolality was 3.505 mOsm/kg (range, 2.345–7.599
mOsm/kg—Weisser et al. 1970). Chilean chinchilla shows elon-
gated renal papilla and high renal indices, with relative medullary
thickness of 6.7

0.8

SD,

and ratio of medulla to cortex of 5.6

0.2 (Dı´az and Corte´s 2003). No reduction of urinary sodium

concentration under food and water deprivation occurred (Gutman
and Beyth 1970).

Feces size (mean

8.6 by 3.9 mm) varies according to size

of animal and is longer and narrower in males (Jime´nez 1990a).
Pellets are dark and can hardly be broken with bare hands. Freshly
defecated feces feel dry to touch.

Rectal body temperature is 35.8

C in domestic males and

36.4

C in females (Kraft 1994). Body temperatures range from 38

to 39

C. Pulse is 100–150 beats/min, and respiratory rate is 40–

80 breaths/min (Webb 1991).

Blood values in captive animals are: erythrocytes, 6.6–10.7

million/

l; hemoglobin, 11.7–13.5 g/dl; mean hematocrit, 38%;

leucocytes, 7.6–11.5

103/

l; monocytes, 1–4%; lymphocytes,

51–73%; neutrophils, 23–45%; eosinophils, 0.5–2.6%; and baso-
phils 0–1%. Number of platelets is 254–298

l (Hillyer et

al. 1997). Chilean chinchillas are as sensitive to ox insulin as they
are to their own, and antigenic determinants and active sites are
intermediate between ox and guinea pig insulins (Neville et al.
1974).

Accessory reproductive glands of domestic males throughout

year follow concentration pattern of blood testosterone, which is
highest between April and August, with a marked but brief descent
in June (Orostegui et al. 1996). Mass and diameter of seminal ves-
icles of males is highest in austral autumn and winter (minimum
mean mass

in February: 0.32

0.05 g; maximum mean

mass

in May: 1.4

0.5 g; minimum mean diameter

in January: 1,488

352.2 mm; maximum mean diameter

June: 4,635.1

1,418.6 mm;

48). Bulbo-urethral glands also

vary (minimum mean mass

in November: 0.11

0.05 g;

maximum mean mass

in May: 0.4

0.06 g; minimum mean

diameter

in February: 3.2

0.52 mm; maximum mean di-

ameter

in June: 6.3

0.19 mm;

50—Cepeda et al.

1999).

Semen ejaculate volume is 0.1–0.5 ml and contains 20–200

million sperm. Sperm can travel 25 mm in 4 min (Asdell 1964).

Follicles in pregnant domestic females are refractory to en-

dogenous gonadotropin, but ovulation may occurs during anestrus.
Ovulation can be induced if coitus occurs early enough to cause a
premature surge of luteinizing hormone. Corpora lutea of pregnancy
are about twice as big as those during nonreproductive cycles; they
do not regress until ovulation recurs. Regressing corpora lutea may
disappear quickly or remain for several cycles. Accessory corpora
lutea are usually formed during pregnancy, but occur at infertile
estrus in some females. Extensive remnants of mesonephric tubules
and rete ovarii exist (Weir and Rowlands 1974). Pregnancy depends
on a progressive increase of progesterone produced only by a com-
plex ovary with accesory corpora lutea and interstitial tissue (Tam
1974).

ONTOGENY AND REPRODUCTION.

Seven (

100 g)

juveniles were caught in the wild between October and December
without a 2nd litter (J. E. Jime´nez, pers. comm.). Domestic

C. lan-

iger

breed from May to November in the southern hemisphere (Nei-

ra et al. 1989), and from November to May in the northern hemi-
sphere (Weir 1973). Usually 2 litters are produced annually.

Long estrous cycles are exhibited by domestic females (mean

SD,

38.1

0.7 days;

323; range, 16–69); estrus lasts 48

h. A postpartum estrus usually occurs at 57.4

2.6 days, whether

or not the litter is lost. Vaginal closure membrane of anestrus may
open and close in 12 h, but usually takes 2–4 days; leucocytes are
present when the membrane is open (Weir 1974). An open vulva,
often with visible mucus, is an external indication of estrus. Vulvar
swelling does not occur during estrus; rather, a change in perineal
color from a dull pink color to a deep red occurs. Color of perineum
increases dramatically at vaginal perforation and remains intense
throughout most of luteal phase (Brookhyser and Aulerich 1980).

Ovulation in captivity is usually spontaneous, sometimes be-

fore vagina opens. A copulatory plug is usually visible after mating.
Single-cell zygotes have been found in fallopian tubes 1–2 days
postcoitum. Fertilization is difficult to detect because some eggs
show no traces of polar bodies, sperm, or pronuclei, although some

show a single polar body and sperm in zona pellucida, or a single
polar body and 2nd maturation spindle at 2.5 days postcoitum
(Roberts and Perry 1974). Polyspermy has not been observed. Pre-
implantation blastocysts have been detected at 3.5 days postcoitum
(Weir and Rowlands 1974).

Implantation occurs at 5.5 days postcoitum and is completely

interstitial. Abembryonal pole of blastocyst is composed of ectopla-
cental trophoblast, which encloses an ectoplacental cavity frequent-
ly filled with extravasated maternal blood. The decidual cavity in-
creases considerably in size, particularly in an antimemetrial re-
gion, where it expands and becomes dumbbell shaped by 10 days
postcoitum. Amniotic cavity is formed at 15 days postcoitum, al-
lantois at 25 days, and chorioallantoic placenta at 30 days after
mating. In 400- to 700-g females, full-term placenta weighs 9 g
with a diameter of 22 mm. Fetal growth rates in domestic Chilean
chinchillas are the lowest observed among hystricomorphs, with a
specific fetal growth velocity of 0.043 (Roberts and Perry 1974).

Hairs of 105-day-old fetus are single and independent. Tran-

sition to adult hair occurs in captivity before 2 months of age.
Epidermis of fetus, newborn, and 2-month-old animals resembles
that of adult, but stratum granulosum is quite prominent, and stra-
tum corneum is somewhat thinner (Wilcox 1950).

Fetal reabsorption occurs frequently and may take place at

any stage of pregnancy. Even in late stages, when skeletal tissue
of fetus has formed, reabsortion rather than mummification or spon-
taneous abortion occurs. Neither placental nor fetal tissue can be
recognized in the necrotic mass, but a central blood-filled cavity is
usual (Roberts and Perry 1974).

Gestation in captivity is 111 days (range, 105–118 days). Par-

turition typically occurs early morning, and the placenta usually is
eaten (Hillyer et al. 1997). Litter size is 1–6, usually 2 or 3 (mean,
1.75;

273 litters—Neira et al. 1989). Sex ratio at birth (male :

female) was 1.19 (Galton 1968) in a Northern Hemisphere colony,

and 1.10 in a Southern Hemisphere colony; at weaning in the latter
sex ratio was 1.24 (Vin˜as 1997).

Mean neonatal mass in captivity is 52 g (range, 50–70,

149—Neira et al. 1989). Mean mass of newborn litter is 83.3 g (

211); individual mass is inversely related to litter size. Mean

number born alive is 1.59 (

273), with 10.4% mortality at birth.

Sometimes, 1 newborn is smaller than others from same litter. All
neonates are precocious, fully furred, with erupted teeth, open eyes,
and able to walk within an hour after birth. They begin to take
solid foods at 1 week of age (Hillyer et al. 1997). Growth rate is
3.6 g/day during the 1st month, decreasing to 1.56 g/day from 2 to
6 months, and to 0.65 from 6 to 12 months (Neira et al. 1989).

Lactation normally lasts 6–8 weeks in captivity (mean

days), and the minimal period of suckling necessary for survival is
25 days. Mean litter size at weaning is 1.31 (

273), and mean

mass of litter at weaning is 342.9 g (

193). Mortality from birth

to weaning is 21.3% (Neira et al. 1989).

Sexual maturity in both sexes occurs on average at 8 months,

but may occur as early as 5.5 months (Weir 1974). In Southern
Hemisphere captives, nonprimiparous females are most likely to
become pregnant during summer and spring. Females have a 1st
litter at a mean age of 459 days, with an interbirth interval of 214
days; they weigh 605 g at mating, 742 g at parturition, and 648 g
postpartum (Neira et al. 1989).

Maximum recorded life span in the wild is 6 years (Jime´nez

1990a). A few captives have lived

20, and some have bred at 15

years. Mean lifespan in zoos is 4 years (maximum

7—Eisenberg

1981).

ECOLOGY.

The natural habitat of Chilean chinchilla is the

barren, arid, and rugged areas of transverse mountain chains in
north-central Chile that connect the coastal mountain ranges to
Andes with elevations from 400 to 1,650 m. Many intermittent
streams occur. Channels are dry most of year, although some areas
experience year-round flows. Average annual rainfall from 1974
through 1998 in that region was 165.4 mm

122.6

(Deane

1999). Maximum rainfall usually occurs May–August during austral
winter (Waylen and Caviedes 1990). A prolonged dry season occurs
October–April. Precipitation varies from 15.8 to 513.4 mm/year.

Typical habitat is rocky or sandy with a sparse cover of thorn

shrubs, few herbs and forbs, scattered cacti, and patches of suc-
culent bromeliads toward the coast. These plant communities are a
mixture of species typical of xeric northern desert and mesic south-
ern mediterranean zones (Sarmiento 1975). Chilean chinchillas

MAMMALIAN SPECIES

758—

Chinchilla laniger

shelter in crevices, holes among or below rocks, or within large
agavelike bromeliads (

Puya berteroniana

—Jime´nez 1990a).

Chinchilla laniger

is social. Colonies of ca. 100 individuals

are usual (Albert 1900). Colonies are 1.5–113.5 ha in size, usually
on equatorial slopes (Jime´nez 1995). Two colonies had 450–500
individuals in 1990 (Jime´nez 1990a). Isolated colonies form a me-
tapopulation, with frequent local extinctions and colonizations of
suitable habitat patches. Population density is 4.37 individuals/ha.
Population size does not fluctuate between years of low and high
rainfall (Jime´nez et al. 1992). Predators include culpeo foxes (

Pseu-

dalopex culpaeus

), which take both adults and juveniles, and Ma-

gellan-horned owls (

Bubo magellanicus

), which prey mainly on

juveniles (Jime´nez 1993).

Wild Chilean chinchilla feed on up to 24 plant species, mainly

herbs and grasses. Diet changes between sites both seasonally (Ser-
ra 1979) and across years. They consume succulents in summer
(Corte´s et al. 2002). The primary native plant chosen in cafeteria
tests during autumn and spring was

Nasella chilensis

(coironcillo—

Pizarro 1988), which is also most frequent item in feces collected
in wild (Corte´s et al. 2002). Grass pasto rey (

Stipa plumosa

) is 2nd

preference of

C. laniger

in same season. Chilean chinchillas

choose plants with more fiber and less lignin content (Pizarro 1988).
They appear to drink no water in the wild.

Among diseases, only Chagas disease, caused by spirochete

Trypanosoma cruzi

and transmitted by endemic hemipteran

Tri-

atoma spinolai

, has been reported in wild Chilean chinchilla. At

Auco´, percentages of infection were 20% of 35 individuals (Duran
et al. 1989) and 40% of 20 animals (Jime´nez and Lorca 1990).
From 1939 to 1987, 29.4% of 53 animals were infected with

Try-

panosoma

(Apt and Reyes 1990).

Bacterial flora is abundant in wild individuals. Most species

found in digestive (53 individuals), respiratory (18 individuals), and
male genital (33 individuals) systems are same as in domestic pop-
ulations (Miller and Finegold 1967). Most common is

Listeria gra-

, a nonpathogenic bacteria (Mathieu 1980).

Small mammals that coexist with Chilean chinchillas in arid

lands of north central Chile are a marsupial (

Thylamys elegans

murid rodents (

Akodon olivaceus

A. longipilis

, and

Phyllotis dar-

wini

), and hystricomorph rodents (

Abrocoma bennetti

and

Octodon

degus

—Osgood 1943).

Chilean chinchillas are not trapped easily. The best trap for

live capture is made of wire mesh (Jime´nez 1987) and baited with
rolled oats (Jime´nez 1989). Pre-Columbian Amerindian hunted

laniger

for their wool, meat, and pelt (Molina 1782). The pelts

reached high prices at international markets in the 19th century,
and modern trappers harvested them almost to extinction (Jime´nez
1994).

Domestication of

C. laniger

was initially for pelts, but now

Chilean chinchillas are also raised as clean, odorless, and friendly
pets (Webb 1991), and as laboratory animals. They are used for
studies on hearing and behavior; they are easily trained, and they
have a highly evolved middle ear (Vrettakos et al. 1988). Their
middle ear and cochlea (Robles and Ruggero 2001) have easy ac-
cess for recording electrodes. They are less subject to ear infections
than laboratory rats, and their audibility curve is more like that of
humans than common laboratory species (Heffner and Heffner
1991).

ANIMAL HUSBANDRY.

Chinchilla laniger

has been do-

mesticated several times, but most if not all present captive colo-
nies derive from 12 wild Chilean chinchillas captured around Po-
trerillos and taken to California by U.S. engineer Matthew Chapman
in 1923 (Parker 1975). Some breeders refer to these domestic in-
dividuals as hybrids but this does not necessarily imply that they
are true interspecific hybrids.

Chilean chinchillas are territorial and acrobatic animals. They

need to be housed either as family units or individually in large
spaces. Because they are nocturnal, cages should be placed in a
draft-free area that permits daytime sleeping.

Individuals should be kept in cages no smaller than 0.0251

in volume (0.35 m wide, 0.40–0.50 m long, and 0.30 m high).

In breeding systems used in the pelt industry, with 1 male having
access to various collared females maintained in separate cages,
minimum cage size for reproductive females is 0.036 m

. For small

colonies, cages no smaller than 1 m in length and 0.5 m high and
wide should contain shelves, which allow for ample exercise. Tubes,
10–13 cm in diameter, hanging from top of cage, add locomotive

opportunities, and places to hide (Hillyer et al. 1997). Front, back,
and top of cages should be wire mesh that allows fresh air to pass
freely; suggested air change in large colonies are 8 volumes/h in
winter and 15 volumes/h in summer (G. Lara, in litt).

Temperatures in housing areas should be between 17 and 25

(Grau 1986). Chilean chinchillas are prone to heat stroke, and
should never be exposed to temperature above 30

C (Hillyer et al.

1997). Length and density of hair increases when maintained at 5–
12

C (G. Lara, in litt.), and with a relative humidity under 50%

(Webb 1991).

Commercial pellets for Chilean chinchillas are ideal food

when available; accepted formula is 10–20% protein, 2–5% fat,
and 15–35% bulk fiber (Webb 1991). Ca. 30 g of pellets should
be given daily, along with a handful of dried timothy hay, which is
preferred in the United States over higher protein content of alfalfa;
the latter is preferred in South America. Dried apples, figs, grains,
hazelnuts, raisins, and sunflower seeds should be limited to not

large spoon a day. Changes in diet should be gradual because
abrupt dietary change can cause serious illnesses (Hillyer et al.
1997).

Individuals consume ca. 25–50 ml of water daily. Water bottles

should be hung outside of cage, and they should be hard if made
of plastic, with protection to prevent punctures. A metal wheel
should be hung inside cage.

Because incisors are constantly growing, a piece of wood (not

cedar) allows gnawing that results in trimming teeth. A clinical sign
of malocclusion is excessive drooling with wet fur around chin,
chest, and forepaws; roots of molars also begin to extend into orbit,
causing lachrymation (Webb 1991).

Dust baths to remove moisture and oils from fur should be

available daily, or at least several times per week. Sanitized chin-
chilla dust is available commercially; beach or playground sand is
not suitable. Dust is placed at a depth of 2–3 cm in a pan that is
large enough for the animal to roll around; dust should be kept
clean and free of feces (Hillyer et al. 1997). Pets should be allowed
to play nightly in an area that allows for jumping and running.

Although Chilean chinchillas are shy, handling is not difficult,

particularly if they are habituated from early age. They may be
held at shoulders, but grasping fur or rough handling will result in
shedding fur patches. They also may be picked up by the base of
tail; body is then supported by other hand (Webb 1991). Biting is
rare, except in extreme distress.

A good indication of an animal’s health is the state of its feces

(Kluhner 1987). Firm, dark feces are ca. 1 cm in length. Wild
individuals produce ca. 223 fecal pellets daily (Jime´nez 1990a);
domestic animals produce a mean of 209 pellets (

8—G. Lara,

in litt). Chilean chinchillas practice coprophagy or reconsumption
of fecal pellets; this aids in maintenance of microorganisms in di-
gestive system. Thus, oral penicillin should not be administered.

Main reasons for culling in domestic colonies are infertility

(44%), age (24%), illness (20%), and fights (12%—Neira et al.
1989). For 2 breeding colonies in Chile, general rate of natality is
69 per 100 chinchillas, and general fecundity rates are 3.06 new-
borns for every male and 1.73 for every female. Specific fecundity
per age is highest at 1–6 years in females and 1–6.5 in males;
then, fecundity decreases progressively. During reproductive life, a
female produces a mean of 20.8 live newborns, and a male sires
51.1 (Vin˜as 1997).

Information on care and pathology (Bickel 1987), and on vet-

erinary practices (Hillyer et al. 1997; Webb 1991) are available.
Guidelines for adequate and humane animal husbandry of Chilean
chinchillas, including a specific questionnaire for evaluation are
available from Council of the Chinchilla Industry (1992), an orga-
nization that includes representatives from the pelt industry.

BEHAVIOR.

Chilean chinchillas are quiet, shy, but active

animals. They ricochet on their long hind legs across rocks in their
usual habitat with remarkable agility. Animals are generally phil-
opatric, remaining in a small area for as many as 6 years. They use
conspicuous rocks for resting and observing their range. Chilean
chinchillas urinate and defecate on rocks, forming latrines that ad-
vertise their presence (Jime´nez 1990b). Domestic animals defecate
between 0300 and 0600 h, and consume cecotropes between 0800
and 1400 h (Holtenius and Bjornhag 1985). When eating, Chilean
chinchillas sit erect and hold food in their forepaws.

Infants often lie on their backs during suckling (Weir 1970).

Fathers are tolerant and fairly friendly, sitting with mother and

MAMMALIAN SPECIES

758—

Chinchilla laniger

young in a protective manner (Kleiman 1974). Mothers usually feed
their young with small pieces of food (Albert 1900). Juveniles dis-
play frisky-hop playing, which includes vertical leaps, body twist-
ing, head tossing, racing and pivoting, and prancing with kicking
back of hind feet (Kleiman 1974).

When stressed, captive Chilean chinchillas shed fur. Females

are very aggressive toward each other and toward males, even when
in estrus. Threats are expressed by growling, chattering teeth, or
urinating (only in captive, unreported in wild). Females are less
aggressive at postpartum estrus than at other times (Healey and
Weir 1970). Courtship rituals include mutual grooming and bipedal
approach by males (Kleiman 1974). Copulatory patterns consist of
multiple intromissions (median

5) of 5 s each, and multiple ejac-

ulations (range, 1 or 2), with ejaculatory intromissions lasting 8–
10 s each. Thrusting occurs during intromission. Females remain
receptive for several hours (Bignami and Beach 1968).

Individuals produce various sounds. When frightened or

seized, an ‘‘eek-eek’’ is produced. Isolated males may produce a
‘‘nyak-nyak’’ call (Eisenberg 1974).

Small burrows are built by Chilean chinchillas under or among

rocks. Below the entrance, a straight corridor runs and turns, open-
ing into a wider room, which serves as a dormitory, and probably
as a nest when covered by dried hay or soft materials. Astringent
sheaths and sweet seeds of algarrobilla (

Balsamocarpon brevifol-

ium

) may be stored (Albert 1900). Sometimes, watery fruits of cacti

also are stored within burrows (Opazo 1911).

Chilean chinchillas clean their fine fur by sand or soil bathing

(Stern and Merari 1969). In wild, they repeatedly bathe in the same
place, thus leaving a barren circle of fine dust ca. 40 cm in di-
ameter, near the entrance of their burrows.

GENETICS.

All chromosomes of Chilean chinchilla are bi-

armed with 2n

64 (Hsu and Benirschke 1967) and FN

128

(Makino 1953; Nes 1963). The metacentric X chromosome is the
largest of complement at a 9% of the haploid autosome set (Galton
et al. 1965); it is of a duplicated type (Ohno et al. 1964), with a
large late replication region on its short arm (Galton et al. 1965)
that corresponds to a large C-band in position and size. In addition
to the duplicate X chromosome is a very small Y chromosome,
which was misidentified as a large chromosome (Makino 1953).
Association of X and Y chromosomes is always end-to-end in mei-
osis, denoting absence of complete synapsis. XX bivalents in fe-
males have 4 chiasmata (Galton et al. 1965). Nucleolar organizing
region is on the secondary constriction of the long arm of a single
submetacentric chromosome pair (George and Weir 1974). A single
nucleolus occurs in the subcentral nuclear space of spermatocytes
(Berrı´os et al. 2004).

Cytochrome-

gene sequences from mitochondrial DNA of

wild and domestic

C. laniger

share 3 character states not present

in other chinchillid species: a C base at site 11 (2nd codon posi-
tion), a G at site 93, and a T at site 150. The 2 chinchilla species
differ at 22 sites, among which 3 are 1st or 2nd codon positions,
with Kimura 2-parameter genetic distances near 6% (Spotorno et
al. 2004; Valladares 2002). All wild

C. laniger

have a character-

istic A base at site 366 (Spotorno et al. 2004). Wild samples from
Auco´ have 4 characteristic bases at sites 47, 198, 234, and 495.
A single specimen from La Higuera had 5 autoapomorph mutations
at sites 147, 264, 265, 266, and 429. Five domestic

C. laniger

had 2 characteristic nucleotides: a T base at site 33, and A at site
85 (2nd codon position) and shared with the single wild specimen
from La Higuera, Coquimbo, a unique silent G base at site 63,
concordant with documented historical origin of present domestic

C. laniger

(Parker 1975). No traces of

C. brevicaudata

variants,

wild or domestic, were found in any of 5 domestic

C. laniger

(Spo-

torno et al. 2004). Microsatellite DNA markers have been amplified
from feces (Walker et al. 2000). Many mutations resulting in dif-
ferent pelage colors have been described in domestic colonies
(Grau 1986).

Hybrids obtained from crosses with

C. brevicaudata

are in-

termediate in gestation length. Hybrid males are infertile. Hybrid
females are fertile, but when backcrossed ca. two-thirds of the 2nd-
generation hybrids are sterile (Grau 1986).

CONSERVATION STATUS.

The Chilean chinchilla is en-

dangered (Glade 1988; Thornback and Jenkins, 1982), with the 2nd
highest conservation priority among Chilean mammals (Cofre´ and
Marquet 1999). Wild populations are listed in CITES Appendix 1

(CITES 1991). As a result of overharvesting for its fur (Iriarte and
Jaksic 1986), the entire species was almost extirpated during early
1900s (Jime´nez 1994). Despite protection provided by an agree-
ment signed between governments of Argentina, Bolivia, Chile, and
Peru´ (Grau 1986), and under Chilean law since 1929 (Lagos and
Valverde 1998), poaching continued. By 1950s, the Chilean chin-
chilla was considered extinct (Mann 1978). Wild populations were
rediscovered in 1978, and in 1983 Reserva Nacional Las Chin-
chillas was created (Lagos and Valverde 1998). This reserve com-
prises 4,227 ha, of which 556 ha had colonies before 1983, but
only 264 ha had colonies in 1989 (Jime´nez 1990b). The former
distribution has been reduced primarily to only 2 areas. Only 19
of 42 known colonies in Reserva Nacional Las Chinchillas currently
are protected (Jime´nez 1996). The total wild population is estimated
as 2,500–11,700 individuals and is declining, as indicated by re-
duction in area covered by present colonies and their fragmentation
(Jime´nez 1993). Although protected inside the reserve from human
activities since 1987, population size continues to decline (Jime´nez
1996). A conservation plan has been in place since 1990 (Lagos
and Valverde 1998), but no particular conservation action is being
taken for the small genetically rich northern population of La Hig-
uera.

REMARKS.

Molina (1782:301) described ‘‘La Chinchilla,

Mus laniger

(I. Mus cauda mediocri, palmis tetradactylis, plantis

pentadactilis, corpore cinereo lanato.) e` unaltra sorta di sorcio cam-
pestre stimabili assai per la finissima lana, di cui e` coperto in vece
di pelo, la quale e` tanto morbida, quanto la seta, che producono i
ragni dei giardini: essa e` di color cenerino, e assi lunga per potersi
filari.’’. Bennett (1829) proposed the new generic name

Chinchilla

for such Molina’s

Mus laniger

; he also emended

laniger

to femi-

nine

lanigera

, assuming that

laniger

was an adjective. Subsequent

authors have followed this action (Woods 1993), unaware that

lan-

iger

is perfectly acceptable as a noun in apposition; others have

retained original spelling (i.e., Gray 1830). Prell (1934) formally
rejected Molina’s description as ambiguous, unidentifiable, and
even applicable to animals of genus

Abrocoma

undescribed in Mo-

lina’s time. However, Molina (1782) described the tail as dressed
with soft hair (‘‘vestita di morbido pelo’’), in contrast with the tail
of

Abrocoma

, which has only very short hairs

1 mm long and

looks naked. Also, Molina (1782) reported the color as ‘‘cenerino’’,
which usually refers to a brilliant gray that is almost cold silver in
appearance, unlike color of

Abrocoma bennetti

from central Chile,

which is ashy gray mixed with warm brown (Spotorno et al. 1998).
Molina (1782) noted that

Mus laniger

does not have a bad odor,

whereas wild

Abrocoma

has a strong organic odor of plant oil.

Prell’s (1934) rejection of

laniger

led him to propose new specific

name

velligera

, an action followed by Osgood (1941) 1st in con-

ditional terms, and in formal terms (Osgood 1943). Prell’s opinions
were analyzed and found to be without serious basis (Cabrera
1960). The International Commission on Zoological Nomenclature
has been requested to incorporate

C. laniger

to the official list of

scientific names (Valladares and Spotorno 2003).

The common name, chinchilla, may derive from Quechua

words

chin

meaning silent,

sinchi

meaning strong or courageous

(Grau 1986), and a diminutive Quechua

lla.

Together,

chinchilla

would mean the strong, silent little (Aleandri 1998). The name

laniger

is from Latin, meaning woolly. The common name Chilean

chinchilla was used by Osgood in 1941, and also in the 1st modern
revision of Chilean mammals (Osgood 1943) for this Chilean en-
demic and most probable autochtonous species. Cabrera (1960:195)
stated ‘‘Durante siglo y medio,

Mus laniger

Molina, 1782, fue univ-

ersalmente identificado como la chinchilla chilena (o ‘‘costina’’, si
se prefiere el calificativo adoptado en peleterı´a) . . . ).’’ [During a
century and a half,

Mus laniger

Molina, 1782, was universally

identified as the Chilean chinchilla (or ‘‘coastal’’, if the qualification
adopted in the pelt industry is preferred . . . ).] Other common
names for this species are chinchilla, chinchilla chilena, coastal
chinchilla, chinchilla costina, chinchilla velligera, chinchilla cola
larga, long-tailed chinchilla, chinchilla chica, and small chinchilla.

Our work was supported by a Fondo Nacional de Ciencia y

Tecnologı´a grant (FONDECYT 198 1122, Chile to A. E. Spotorno).
FONDECYT Sectorial Project (596 0017 to A. E. Spotorno) allowed
initial field and laboratory work. We thank Servicio Agrı´cola y Gan-
adero and Corporacio´n Nacional Forestal, Ministerio de Agricul-
tura, Chile (collection permits 2273, August 1997; 1454, May
1999; and 3042, October 1999), particularly Agustı´n Iriarte, for

MAMMALIAN SPECIES

758—

Chinchilla laniger

granting collection permits; Gonzalo Lara (former president of In-
stituto de la Chinchilla A.G., Chile) for his kind donation of do-
mestic specimens, some literature, discussion, and revision of ‘‘An-
imal Husbandry’’; Angel S. Spotorno L. for drawing Fig. 3; Juan
Oyarce for technical assistance in collecting and care of animals;
A. Gardner for assistance with the synonymy; and 2 anonymous
reviewers.

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AIL

ICHENER

, P

AMELA

WEN

, and K

RIS

ELLGREN

. Editor was V

IRGINIA

AYSSEN

NGEL

E. S

POTORNO AND

J. P

ABLO

ALLADARES

, L

ABORATORIO DE

ENO

´ MICA

VOLUTIVA

, P

ROGRAMA DE

ENE

´ TICA

UMANA

, I

NSTI

TUTO DE

IENCIAS

IOME

´ DICAS

ICBM, F

ACULTAD DE

EDICINA

NIVERSIDAD DE

HILE

, C

ASILLA

70061, S

ANTIAGO

7, C

HILE

. C

LOS

A. Z

ULETA

, D

EPARTAMENTO DE

IOLOG

ı´

, F

ACULTAD DE

IEN

CIAS

, U

NIVERSIDAD DE

ERENA

, C

ASILLA

599, L

ERENA

HILE

. A

L. D

EANE

, C

ASILLA

302, I

LLAPEL

, C

HILE

. J

AIME

IME

´ NEZ

, L

ABORATORIO DE

COLOG

ı´

, U

NIVERSIDAD DE

AGOS

ASILLA

933, O

SORNO

, C

HILE