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FIELDIANA 
Zoology 

Published by Field Museum of Natural History 



New Series, No. 11 



NEOTROPICAL DEER (CERVIDAE) 
PART I. PUDUS, GENUS PUDU GRAY 

PHILIP HERSHKOVITZ 



^^^itli^tff^ 






neio MusEmA imm 



March 22, 1982 
Publication 1330 



NEOTROPICAL DEER (CERVIDAE) 
PART I. PUDUS, GENUS PUDU GRAY 



FIELDIANA 
Zoology 

Published by Field Museum of Natural History 



New Series, No. 11 



NEOTROPICAL DEER (CERVIDAE) 
PART I. PUDUS, GENUS PUDU GRAY 

PHILIP HERSHKOVITZ 

Curator Emeritus 

Division of Mammals 

Department of Zoology 

Field Museum of Natural History 



Accepted for publication May 22, 1980 
March 22, 1982 

Publication 1330 



Library of Congress Catalog Card No.: 81-67141 

ISSN 0015-0754 

PRINTED IN THE UNITED STATES OF AMERICA 



CONTENTS 

I. Introduction 1 

Material 2 

Abbreviations 2 

Measurements 2 

Review of Cervid Classification with Particular Reference to South 

American Genera 2 

Odocoileine Origins and Radiation 8 

Cheiridia 9 

Limb Proportions 11 

II. Genus Pudu Gray 13 

Synonymic History 13 

Conservation 14 

Taxonomy 15 

Generic Characters 15 

Generic Comparisons 22 

Fawnal Spotting Pattern "23 

Ancestral Characters and Relationships 24 

Origin, Dispersal, Spedation 25 

III. The Species of Pudus, Genus Pudu Gray 43 

Key to the Spedes 43 

IV. Pudu mephistophiles de Winton 44 

Synonymic History 44 

Distribution 45 

Conservation 48 

Vernacular Names 50 

Taxonomy 51 

Characters 51 

Comparisons 57 

Spedmens Examined 58 

V. Biology of Pudu mq^histophiles 58 

Habitat 58 

Habits 58 

Maintenance in Captivity 59 

Ungulate Assodations 59 

Reproduction 59 

VI. Pudu puda Molina 60 

Synonymic History 60 

Distribution 62 

Conservation 62 

Taxonomy 64 

Characters 65 

Karyotypes 70 

Comparisons 70 

Spedmens Examined 70 

VII. Biology of Pudu puda 71 

Habitat 71 

Maintenance in Captivity 71 

Ungulate Assodations 73 



vi CONTENTS 

Numbers and Aggregations 73 

Migrations 73 

Locomotion, Play 74 

Enemies 74 

Diseases and Parasites 74 

Food and Water 74 

Earthquake Sensitivity 75 

Temperature Regulation 75 

Reproduction, Growth, Development 75 

Longevity 79 

Vm. Summary 79 

IX. Acknowledgments 79 

X. Gazetteer and Explanation of Maps 80 

XL Literature Qted 83 



LIST OF ILLUSTRATIONS 

1. Dispersal and spedation of pudus 2 

2. Left forefeet and hind feet of brocket, Mazama rufina, and muntjac, 

Muntiacus reevesi 6 

3. Nares in Odocoileinae and Cervinae 7 

4. Artiodactyl hind feet showing gradient of metatarsal bone reduction 10 

5. Size relationship and body form of six South American deer 16 

6. Body form of four South American deer 17 

7. Fadal color pattern of Odocoileinae 18 

8. Head characters of smallest South American deer 19 

9. Fadal markings of five South American deer 20 

10. Front and hind feet with outline of interdigital glands 21 

11. Skulls, left side of adult female pudus and brockets 26 

12. Skulls, dorsal surface of adult female pudus and brockets 27 

13. Skulls, palatal surface of adult female pudus and brockets 28 

14. Skulls, left side of adult male pudus and brockets 29 

15. Skulls, dorsal surface of adult male pudus and brockets 30 

16. Skulls, palatal surface of adult male pudus and brockets; inset, dedduous-like 

pm^ 31 

17. Mandibles of pudus and brockets, lateral surface of left rami 32 

18. Upper cheek teeth of adult pudus and brockets, and juvenal cheek teeth of 
Mazama chunyi 33 

19. Lower cheek teeth of adult pudus and brockets, and juvenal cheek teeth of 
Mazama chunyi 34 

20. Lower front teeth of pudus and brockets, labial surface 35 

21. Lower front teeth of pudus and brockets, lingual surface 36 

22. Permanent and dedduous indsor-canine series in pudus and brockets 37 

23. Upper right and lower left second and third molars of pudus 38 

24. Forefeet and hind feet of pudus with vestiges of metacarpal bones II and V 
(telemetacarpalia) 40 

25. One-week-old southern pudu 41 

26. One-week-old gray brocket 41 

27. Red brocket doe with fawn 42 

28. Northern pudus, adult male and female 43 

29. Northern pudu, adult female 43 

30. Geographic distribution of Pudu mqjhistophiles 46 

31. Pelage and hair types of Pudu mephistophiles 53 

32. Sectioned hair from Pudu mephistophiles 54 

33. Saturation in mammalian tegumentary color 55 

34. Color of hairs of Pudu mephistophiles and P. puda 56 

35. Geographic distribution of Pudu puda 63 

36. Southern pudus in Cologne zoo 67 

37. Southern pudus in Berlin zoo 67 



LIST OF TABLES 

1. Homologies of artiodactyl carpal and tarsal bones 11 

2. Artiodactyl limb proportions 12 

3. External and cranial measurements of adult Pudu mephistophiles 52 

4. External and cranial measurements of adult Pudu puda 68 

5. Antler cycle in Pudu puda 69 

6. Birth records of Pudu puda 72 

7. Growth and development of a male Pudu puda in the Cologne zoo 76 

8. Chronology of weight increment in Pudu puda 77 

9. Three oldest longevity records for Pudu puda 78 

10. Composition of milk produced by Pudu puda 79 



I. INTRODUCTION 

The diminutive, spike-antlered deer of the genus Pudu are smallest of living 
true deer and smallest of New World ungulates. Of the two known species, the 
northern occurs in the high, equatorial Andes from southern Colombia to central 
Peru between latitudes 5°N and 12°S, and the southern is found in the southern 
half of Chile and in bordering parts of Argentina between latitudes 35°S and 53°S 

(fig. 1). 

This report is the first of a proposed series on the genera of neotropical Cer- 
vidae and part of a monograph begun more than three decades ago with the 
intent of completing a unified work. My preliminary publications in ensuing 
years include attempts to clarify the nomenclature and taxonomic status of some 
neotropical deer (1948, 1958a, 1958b), descriptions of a new subspecies and 
species of brockets of the genus Mazama (1959a, 1959b), and comments on South 
American cervid origin and distribution (1958c, 1966, 1972). Work on the mono- 
graph itself, which had already become a sizable manuscript, suffered delays 
and lengthy interruptions and seemed interminable as a single work. 

The present plan is to update and complete each generic review as a separate 
publication, with particular attention paid to taxonomy, zoogeography, and life 
history of the included species. In this report, the issues of origin, evolution, and 
supergeneric classification are touched upon mainly for orientation and refer- 
ence. Full discussion of these topics necessarily awaits completion of studies of 
both living and fossil forms. Detailed descriptions of more specific subjects, such 
as dental anatomy, hoofs, antlers, glands, genitalia, and other soft parts, will be 
presented in those reports where most appropriate or demanding. For example, 
because simple antlers of pudus and brockets are similar, the attention in this 
account focuses on other characters, such as limb bones, which distinguish 
pudus from all other deer. On the other hand, distinction of brockets {Mazama 
spp.) from their nearest relative, the white-tailed deer {Odocoileus virginianus) , is 
based largely on antler size and shape. It is proper, therefore, to reserve both 
description of the morphology and discussion of the evolution of branched 
antlers from the simple, spike-like form for reports on those particular deer. The 
final number of the proposed series will include a synthesis. 

The study of neotropical deer, particularly in the field, has been largely ne- 
glected. The 11 or so extant species exceed the combined number of spedes of 
the Odocoileinae and tribe Cervini native to other comparable zoogeographic 
regions. 

Man's persecution of deer is persistent and practically uncontrolled. Most of 
the species have already vanished from large areas of their former range. Under 
present conditions of mock conservation, some spedes may disappear com- 
pletely before the end of the century. 




Fig. 1. Dispersal and spedation of pudus. Dashed line at northeastern base of Andes 
indicates presumed temperate zone habitat of ancestral spedes during a gladal period. 
With onset of the postgladal period, pudus presumably moved with their natural temper- 
ate zone habitat to higher altitudes in the north (arrows) and to higher latitudes in the 
south (triangles) in the wake of retreating gladers. The widely separated populations then 
differentiated into Pudu mephistophiles in the north, and the more evolved P. puda in the 
south. Character for character, the southern spedes can be derived from the northern. 
Actual geographic range of P. mqjhistophiles, if not continuous, probably is — or was — less 
fragmented than indicated on the basis of available data. 



HERSHKOVITZ: NEOTROPICAL DEER 3 

Material 

Specimens examined in the preparation of this report include skins, skulls, 
and postcranial skeletons representing 57 pudus, as well as an uncounted 
number of representatives of all other forms of neotropical deer preserved in the 
institutions listed under the "Abbreviations" section. In addition, reference was 
made, as occasion required, to other artiodactyls in the Field Museum collec- 
tions. 

Photographs of skulls and dentition are based on "normal" specimens in good 
condition selected from material at hand. In some cases, it was necessary to 
substitute mandibles or teeth of specimens other than those used for illustrating 
the main parts of the skull. 

Abbreviations 

AMNH — American Museum of Natural History 
ANSP — Academy of Natural Sciences, Philadelphia 
BM — British Museum (Natural History) 
FMNH — Field Museum of Natural History 
MCZ — Museum of Comparative Zoology, Harvard University 
USNM — National Museum of Natural History, Smithsonian Institution 
I take this opportunity to thank the authorities of the sister institutions men- 
tioned here for permission to study the material in their charge. 

Measurements 

Standard external and cranial measurements are used here. Limb bone mea- 
surements are of greatest length, irrespective of the points. Width of metapo- 
dials, however, is measured between widest points across the distal epiphyseal 
union. Combined length of the articulated thoracic and lumbar vertebrae is 
equated with trunk length (TL, table 2), and used as principal reference for 
proportional lengths of limb bones. In most cases, articulations were made man- 
ually and the vertebrae measured in sections. Carpal, tarsal, and phalangeal 
dimensions are omitted here. 

In comparisons of pudu postcranial skeletal bones with those of other ar- 
tiodactyls (table 2), ratios instead of actual measurements are given to eliminate 
or minimize actual size differences between species, sexes, age classes, and 
individuals. 

Review of Cervid Classification 
with Particular Reference to South American Genera 

The first comprehensive and soundly documented arrangement of deer, in- 
cluding the South American forms, is that by Brooke (1878, p. 889). His scheme, 
less a classification than a dichotomous key to the genera or generic groups, is 
abstracted as follows: 

I. Plesiometacarpi (proximal ends of lateral metacarpals persistent, fig. 2). 
A. Posterior portion of nasal cavity not divided into two chambers by 
vomer (fig. 3). Genera: Cervulus i=Muntiacus), Elaphodus, Cervus (sub- 
genera, Cervus, Rusa, Rucervus, Elaphurus, Axis, Pseudaxis, Dama). 

II. Telemetacarpi (distal ends of lateral metacarpals persistent, fig. 2). 



4 FIELDIANA: ZOOLOGY 

A. Posterior portion of nasal cavity not divided into two chambers by 
vomer (fig. 3). Genera: Alces, Hydropotes, Capreolus. 

B. Posterior portion of nasal cavity divided into two chambers by vomer 
(fig. 3). Genera: Cariacus {=Odocoileus, with subgenera Odocoileus, Blas- 
tocerus [includes Ozotoceros=Blastoceros], Furcifer [=Hippocamelus], Coas- 
sus [=Mazama]), Pudua {=Pudu), Rangifer. 

The subfamilies Cervinae and Capreolinae, recognized by Pocock (1911, p. 
971) in his arrangement of cervids, are equivalent to Brooke's Plesiometacarpi 
and Telemetacarpi. In 1923, Pocock (p. 206) elevated each well-marked genus or 
generic group of telemetacarpalid deer to subfanruly rank without indication of 
interrelationships, except through the following artificial key (abstracted): 

A. Telemetacarpalia (fig. 2). 

1. Vomer dividing posterior nares (fig. 3). 

a. Cuboideonavicular and external cuneiform bones united (Pudinae, 
figs. 4, 24). 

b. Cuboideonavicular and external cuneiform bones separated (fig. 2). 
(1). Antlers present in males only (Odocoileinae). 

(2). Antlers present in both sexes (Rangiferinae). 

2. Vomer not dividing posterior nares (fig. 3). 

a. Antlers absent; upper canine hypertrophied in males (Hy- 
dropotinae). 

b. Antlers present; upper canine minute or absent in either sex. 
(1). Muzzle short, not swollen (Capreolinae). 

(2). Muzzle very long, much swollen (Alceinae). 

B. Plesiometacarpalia (fig. 2). 

1. Cuboideonavicular and external nuddle cuneiform bones fused (Mun- 
tiadnae = Cervulinae, fig. 2). 

2. Cuboideonavicular and cuneiform bones discrete (Cervinae). 

Lydekker (1915, p. vii) grouped all deer in the family Cervidae (Section Pec- 
ora). The family, in turn, was divided into subfamilies Moschinae and Cervinae, 
the latter with genera Muntiacus, Elaphodus, Dama, Cervus, Elaphurus, Odocoileus, 
Blastocerus, Hippocamelus, Mazama, Pudu {Pudella, a subgenus), Capreolus, Ran- 
gifer, and Hydropotes. 

Living and fossil South American deer are central to the classification of the 
Cervidae presented by Carette (1922, p. 442). In the following, the two sub- 
families and attributed characters are abstracted, cervine generic names omitted: 

A. Cervinae (Eurasian). 

1. Vomer incompletely ossified, the vertical plate not extending to pos- 
terior portion of nasal cavity. 

2. Plesiometacarpalid or telemetacarpalid. 

3. Metatarsal tuft, when present, on proximal half of metatarsus. 

4. Upper canine tusk-like to absent. 

5. Premaxillary and nasal bones usually articulating. 

6. Antlers simple, dichotomous, branched, or absent. 

B. Neocervinae (New World). 

1. Vomer completely ossified, vertical plate extending to posterior portion 
of nasal cavity. 

2. Telemetacarpalid. 

3. Metatarsal tuft, when present, on distal half of metatarsus. 



HERSHKOVITZ: NEOTROPICAL DEER 5 

4. Upper canine small or absent. 

5. Premaxillary and nasal bones usually not articulating. 

6. Antlers. 

a. Absent {iBlastomeryx, North American Miocene). 

b. Simple {Mazama, Pudu). 

c. Bifurcate {Hippocamelus). 

d. Dichotomous (Blastocerus) . 

e. Branched {Odocoileus, Rangifer). 

Included within the "Neocervinae" are the extinct, branch-antlered Antifer, 
Paraceros, Morenelaphus, Pampaeocervus, and Epieuryceros. "Neocervinae," the 
group name erected by Carette, is not derived from a generic name and has no 
status. 

Kraglievich (1932, pp. 425-429) dismissed Carette's concept of "Neocervinae" 
as contrived and composite. His own arrangement, based primarily on antler 
form, is: 

a. Mazaminae {Mazama, Pudu). 

h. Odocoileinae {Odocoileus, Blastocerus, Hippocamelus). 

c. tAntiferinae {iAntifer). 

d. Aldnae {Alces). 

e. Cervinae {Cervus, Rangifer, fParaceros, fMorenelaphus [iPampaecervus, a 
subgenus]). 

f. Incertae sedis {iEpieuryceros). 

Living and fossil genera of North American deer (Cervidae) are classified in 
key form by Frick (1937, pp. 32, 34, 205) primarily on the basis of teeth and 
antlers. South American genera are included only "tentatively, for 
comparison — unstudied," in "Division C.-Cervini. Pleistocene — Recent," with 
the following subfamilies and their respective genera: 

Subfamily 7. Cervinae {Procoileus, Cervus). 
Subfamily 8. Odocoilenae {Odocoileus, Eucervus). 
Subfamilies 8 and 9a. Alcenae-Rangiferinae {Cervalces, Alces, Rangifer) 
Subfamilies 8 and 9a (continued). Odocoilenae and (?)Rangiferinae {Pudu, 
Hippocamelus, Blastocerus, Mazama, Ozotoceros, fPalaeodocoileus, fParaceros, 
fMorenelaphus, iAntifer). 

Simpson's (1945, p. 152) Cervoidea, with the single family Cervidae, contains 
the following subfamilies, with extant American tribes and genera included 
where pertinent: 

tPalaeomerydnae 
Moschinae 
t Dromomery d nae 
Muntiadnae 
Cervinae 
Odocoileinae 
Odocoileini {f Antifer, iEpieuryceros, fMorenelaphus, Odocoileus, Mazama, Hip- 
pocamelus, Blastocerus, Ozotoceros, Pudu) 
Aldni {iCervalces, Alee) 
Rangiferini {Rangifer) 
Hydropotini {Hydropotes) 
Capreolini {fProcapreolus, Capreolus) 



FRONT LIMB 





Mazama rufina 

70563 



cal caneam 




navicular — >^^^_^m— cuboid 
cuneiform — P33P^~-t5 









DORSAL 



VENTRAL 



Muntiacus reevesi 

15575 
LATERAL DORSAL 



sesamoid 



HIND LIMB 




Fig. 2. Left forefoot and hind foot of brocket, Mazama rufina, with vestiges of metacarpal 
bones II and V distal in position (telemetacarpalia), and of muntjac, Muntiacus reei'esi, with 
vestiges of the same metacarpal bones proximal in position (plesiometacarpalia). Front 
digital bones oi Mazama rufina lacking. 



HERSHKOVITZ: NEOTROPICAL DEER 




Fig. 3. Nares are divided posteriorly by bony extension of vertical plate of vomer 
(arrow) in Odocoileinae, as in Mazama americana, but are undivided posteriorly in Cer- 
vinae, as in Capreolus capreolus. 



Simpson (1945, p. 267) discarded the prior subfamily group name Capreolinae 
for Odocoileinae, "because Odocoileus makes a better type." 

Flerov (1952) criticized Simpson's concept of cervoid relationships and rear- 
ranged the Cervoidea as follows: 

Moschidae 
Cervidae 

tPala eomery d na e 
tDromomery ci na e 
Cervulinae (=Muntiacinae) 

Cervinae {Capreolus, Cervus, Elaphurus, Alces, iPliocervus, iProcapreolus, 
■UZervavitus, iEucladocerus, iPseudalces, iMegaloceros, iCervalces) 
Neocervini {Mazama, Pudu, Odocoileus [with subgenera Odocoileus, Blas- 
tocerus, Hippocamelus, Rangifer, iAntifer, fEpieuryceros, iPampaeocervus, 
iMorenelaphus ]) . 

According to Churcher (1966), iEpieuiyceros and iAntifer (in the preceding 
rearrangement) probably are congeneric with Blastocerus. 



8 FIELDIANA: ZOOLOGY 

Cabrera (1%1, pp. x, 324), in his Catdlogo de los mamiferos de America del Sur, 
recognizes the extant South Anierican genera of Cervidae in the following order: 
Odocoileus, Blastocerus, Ozotoceros, Hippocamelus, Mazama, Pudu (with subgenus 
Pudella). 

For convenience of a single reference, a provisional classification of currently 
recognized genera and living spedes of South American cervids is provided 
herewith: 

Cervidae Gray 1821 
Odocoileinae Pocock 1923 (New World deer) 
Pudini (new rank=Pudinae Pocock 1923) 
Pudu Gray 1850 (Pudus) 
P. mephistophiles de Winton 1896 
P. puda Molina 1782 
Odocoileini Simpson 1945 
Mazama Rafinesque 1817 (Brockets) 
M. americana Erxleben 1777 
M. chunyi Hershkovitz 1959 
M. gouazoubira Fischer 1814 
M. rufina Pucheran 1851 
iCharitoceros Hoffstetter 1963 
Hippocamelus Leuckart 1816 (Huemuls) 
H. antisensis D'Orbigny 1834 
H. bisulcus Molina 1782 
Blastoceros Fitzinger 1860 {=Ozotoceros Ameghino 1891 [Pampas deer]) 

B. bezoarticus Linnaeus 1758 
Blastocerus Gray 1850 (?synonyms: iAntifer Ameghino 1889, iParaceros 
Ameghino 1889, iEpieuryceros Ameghino 1889 [Swamp deer]) 
B. dichotomus lUiger 1815 
Odocoileus Rafinesque 1832 (synonyms: iPaleodocoileus Spillmann 1931, 
iProtomazama Spillmann 1931 [White-tailed deer]) 
O. virginianus Zimmermann 1780 
fAgalmaceros Hoffstetter 1952 
iRohnia Castellanos 1957 

iMorenelaphus Carette 1922 (?synbnyms: iPampaeocervus Carette 1922, 
iHabromeryx Cabrera 1929) 

Representatives of all listed genera have been recorded from the Pleistocene of 
South America. In addition, deer fragments from the Uquian or the Pliocene- 
Pleistocene boundary in Argentina have been identified with Blastoceros 
{=Ozotoceros) , Antifer, and Habromeryx (Pascual et al, 1966, p. 21). 

Odocoileine Origins and Radiation 

It has been suggested by Matthew (1908; Flerov, 1952) that the Miocene Blas- 
tomeryx Cope 1877 from South Dakota, Wyoming, and Colorado is a primitive 
generic model from which New World deer can be derived. Blastomeryx, in turn, 
he declared, appeared to be derived from the Oligocene Leptomeryx. Judged by 
descriptions, skeletal parts of the known spedes of Leptomeryx are indeed com- 
paratively primitive, and antlers are lacking. However, the large body size, 
comparable to that of the larger spedes of Mazama, and, above all, the laniary or 



HERSHKOVITZ: NEOTROPICAL DEER 9 

traguline-like upper canine remove Blastomeryx from the line leading to modern 
Odocoileinae, or, for that matter, to the Cervidae. 

Whatever the ancestral stock, it is no less likely that the Odocoileinae origi- 
nated in South America as in North America. The principal theater of radiation, 
however, almost certainly was South America. Pudus, tribe Pudini, with their 
mosaic of primitive, highly specialized and unique characters, appear to repre- 
sent a very early offshoot of the ancestral stock. Brockets and branch-antlered 
deer represent a second offshoot, if not the main stem itself. They have retained 
most of the basic ancestral characters and their evolution has been directed, in 
the main, toward increasing body mass and social complexity. 

Among the branch-antlered deer now living in South America, Odocoileus 
alone may have originated in Middle America, although its ancestral home may 
have been elsewhere. From Middle America, Odocoileus must have spread into 
South and boreal America with major radiation taking place in the latter. Cir- 
cumpolar caribou or reindeer, genus Rangifer, are the northernmost 
odocoileines. They could have diverged from the same stock that gave rise to 
modem Odocoileus. The extinct Argentine Morenelaphus with caribou-type antlers 
may represent a third group, or, perhaps, a subgroup of the second. 

Cheiridia 

The artiodactyl manus and pes evolved for support of the limbs on enlarged 
third and fourth ungual digits. The modification, doubtless beginning at a 
preancestral stage, involved loss of metapodial I and digit 1, disappearance of a 
discrete, carpal central bone, and, usually, the trapezium, reduction of digits 2 
and 5 together with the corresponding metapodial bones, and elongation and 
coalescence of metapodials III and IV into a single, functional element — the 
cannon bone. Fusion of the middle metapodial bones progresses from midshaft 
toward both ends, with union of the distal portions lagging behind the proximal. 
A graded series of metapodials III and IV from separate to entirely fused, ex- 
cepting the distal articular facets, can be formed with species of suborder Suina, 
followed by the tragulid Hyemosc/iMS and selected forms of camelids, bovids, and 
cervids in the order named (fig. 4). Locomotorially, the front limb is specialized 
mainly for pulling, the hind limb mainly for pushing. 

As already noted, in nearly all living Old World cervids, only the proximal 
ends of the disappearing lateral metacarpals persist (plesiometacarpalid, fig. 2). 
In all living cervids of New World origin, the distal ends persist (telemetacar- 
palid, fig. 2). The latter condition may be more primitive because the metacarpal 
vestiges occur at the digital or functional end. However, functional difference 
between the two limb types is not apparent. Carpal bones, on the other hand, 
remained fairly constant throughout the order Artiodactyla. 

Individual forelimb and hind limb bones are serially homologous (table 1), but 
evolution of the homologues does not always proceed at the same rate or in the 
same direction. Metatarsi III and IV and corresponding digits do resemble their 
forelimb homologues. In contrast, metatarsi II and V have disappeared except 
sometimes for vestiges, particularly in pudus, without consistent orientation 
proximally or distally. The corresponding digits do not articulate with the rest of 
the skeleton, thus resembling the condition in plesiometacarpalid deer. The 
tarsal central or navicular bone, unlike the carpal central bone, is present and 
well developed. In Pecora and Tragulina, it is integrated with the cuboid to form 




c=^%r 





u b '- o 




^' = 2 .| 

n "5 t« C 



10 



HERSHKOVITZ: NEOTROPICAL DEER 



11 



Table 1. Homologies of artiodactyl carpal and tarsal bones are shown horizontally. 
Names in columns marked "1" are used in comparative anatomy, those in columns 
marked "2" are the customary English names employed in this work. Names connected by 
a brace are of bones fused in all artiodactyls except as noted; those followed by a slash (/) 
are further fused with the next listed bone in suborders Tragulina and Pecora. Sesamoid 
bones, except the pisiform, are not named. Names and homologizations are adopted from 
Flower (1885, pp. 281-311; 339-360). 



Carpalia 



1 

Radiale 

Intermedium 

Ulnare 



Carpale 1 
Carpale 2/ 
Carpale 3 
Carpale 4 
Carpale 5 

Centrale 



=Scaphoid 
=Lunar' 
=Triquetrum or 

cuneiform 
=Trapezium^ 
=Trapezoici/ 
=Magnum 

=Undform 
= Central' 



1 

Tibiale 

Intermedius 

Fibulare 



Tarsale 1 
Tarsale 2 
Tarsale 3 
Tarsale 4 
Tarsale 5 

Centrale 



Tarsalia 

2 

=Astragalus or Talus 
=Calcaneum 

=Internal cuneiform 
=Middle cuneiform 
=External cuneiform 



=Cunei- 
form^ 



=Cuboid/ 



=Cuboideonavicular3 



= Navicular 



'Lunar and central fused in Artiodactyla. ^Presumed lost in Artiodactyla except in 
Hippopotamus. ^Cuneiform and cuboideonavicular bones always fused in Tragulina, giraf- 
fid Okapia, and cervids Muntiacus and Pudu; middle and external cuneiforms (tarsales 2, 3) 
are distinct in Suina. 

the cuboideonavicular bone. External and middle cuneiform bones (cuneiforms 2 
and 3) of the tarsus are fused, except in Suina, and the internal cuneiform is 
small but discrete in all artiodactyls. In Pudu and Cervulus {=Muntiacus), the 
cuboideonavicular bone is further united with outer and middle cuneiform 
bones into a single element, as in the chevrotains Tragulus and Hyemoschus 
(Tragulidae, Tragulina, Artiodactyla), and giraffids.' 

Fusion of cuboid, navicular, and outer and middle cuneiform bones into a 
single unit obviously evolved independently in the four disparate artiodactyl 
groups just dted. Functional difference between simplified and primitive ar- 
tiodactyl tarsi is questionable, particularly in Giraffa, where both types may occur 
in the same animal. Seemingly, the tragulid and pecorid tarsus operates as a unit 
irrespective of articular relationships between its components. 

Tarsal similarities among the four groups in question do not imply metatarsal 
similarities. Limb proportions differ widely among them. The tragulid metatar- 
sus is most primitive in retention of complete, albeit slender and elongate, 
metatarsal bones II and V; whereas the same bones are absent in giraffids, 
muntjacs, and deer, except pudus and possibly a few other spedes in which they 
may persist as vestiges. Furthermore, pudu metapodials, as represented by the 
cannon bone (III -I- IV), are relatively shorter and broader than those of other 
artiodactyls, and thus nearer the primitive condition still exhibited by living 
members of the suborder Suina. 



Limb Proportions 

Forelimb length, based on combined length of humerus, radius, and metacar- 
pus (H + R + MC, table 2) in pudus and the great majority of quadrupedal 

'Consistently in Okapia (FMNH 26066, 104733, 108987) and variably in Giraffa (fused in 
both tarsi of FMNH 34930, fused in left, separate in right of FMNH 34424); aU zoo animals. 



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12 



HERSHKOVITZ: NEOTROPICAL DEER 13 

mammals, is less than hind limb length, based on combined length of femur, 
tibia, and metatarsus (F + T + MT). As shown in Table 2, length of forelimb 
(H + R + MC) to trunk length (TL), based on combined length of articulated 
thoradc and lumbar vertebrae, ranges from about 73% (Pudu) to about 104% 
(Mazama) in adult artiodactyls sampled. The average length of hind limb to trunk 
(TL) ranges from about 89% (Bison, followed by about 105% in Pudu and Oreo- 
tragus) to about 139% (Mazama gouazoubira) . Individual variation is considerable, 
and present ratios based on the sum of straight-line lengths of long limb bones 
are greater than — but probably directly proportional to — standing shoulder and 
rump heights in which limb bones articulate at angles. 

The cervid humerus is usually the longest thoracic limb bone. In pudus, it is 
relatively longer than in all other cervids and bovids measured. The humerus, 
however, contributes least to standing shoulder height. The cervid radius may 
be nearly as long as the humerus, sometimes slightly longer. The metacarpus is 
the shortest of the long thoradc limb bones, but nearly all its length enters into 
standing shoulder height. In most Bovidae, the radius is the longest forelimb 
bone, but in others, notably gazelles, the metacarpus is longest. In both cervid 
and bovid hind limbs, the tibia is usually the longest bone, the metatarsus being 
shortest of the three long bones. As in the forelimb, almost the entire length of 
the metatarsus enters into total standing rump height. 

Metapodials of pudus are not only relatively shortest among cervids, but also 
broadest. Ratio of metapodial width to length of the small Andean brockets, 
Mazama rufina and M. chunyi, appears to be nearly the same when greatest width 
is measured across the distal epiphysis. However, metapodial shafts of all 
brockets, particularly those of the metacarpus, are relatively narrower than those 
of pudus. 

II. GENUS Pudu GRAY 
Synonymic History 

Pudu Gray, 1850, Gleanings Knowlsely Menagerie, p. 69 — subgenus of Coassus Gray. 
Gray, 1852, Proc. Zool. Soc. London, 1850: 242— subgenus of Coassus Gray. Gray, 1852, 
Cat. Mamm. Brit. Mus., Ungulata, p. 240 — genus. Sclater, 1870, Proc. Zool. Soc. Lon- 
don, 1870: 116 — classification. Lydekker, 1915, Cat. Ungulate Mammals Brit. Mus., 4: 
214— characters. Knottnerus-Meyer, 1907, Arch. Naturg., (73) 1: 15, 16, 97, 108— cranial 
characters. Carette, 1922, Rev. Mus. La Plata, 26: 412, 420, 422, 425, 428, 445— history; 
characters; classification. Pocock, 1923, Proc. Zool. Soc. London, 1923: 181, 207— 
characters; classification (genus of subfamily Pudinae). Kraglievich, 1932, Anal. Mus. 
Hist. Nat., Montevideo, (3) 2: 265 — characters; phylogeny. Pocock, 1932, Proc. Zool. 
Soc. London, 1932: 395^antlers; comparisons. Flerov, 1952, Fauna U.S.S.R., n.s. no. 
56, Mammals, 1, (2): 218— description. Hershkovitz, 1958, Fieldiana, Zool., 36(6): 
619 — distribution; Pudella a synonym. Cabrera, 1961, Rev. Mus. Argentino Cienc. Nat. 
"Bernardino Rivadavia," 4(2): 343 — subgenus of Pudu Gray; included species: P. pudu 
Molina. Haltenorth, 1963, Handb. Zool., 8(32): 4, 9, 42, 47, 48— subgenus of Mazama; 
characters; classification; Pudella Thomas a synonym. Hershkovitz, 1972, in Keast, Erk, 
and Glass (eds.). Evolution, mammals and southern continents, State U. of New York Press 
(Albany), pp. 362, 371, 392 — history, distribution, ecology, adaptation. Whitehead, 
1972, Deer of the world. Viking Press, New York, pp. 64, 112, 169— spedes, P. pudu, P. 
mephistophiles; classification. 

Pudua Garrod, 1877, Proc. Zool. Soc. London, 1877: 18— emendation of Pudu Gray; 
spedes, Pudua humilis Bennett. Brooke, 1878, Proc. Zool. Soc. London, 1878: 926— 
characters. De Winton, 1896, Proc. Zool. Soc. London, 1896: 508— characters. Lydekker, 
1898, Deer of all lands, p. 307— characters. 



14 FIELDIANA: ZOOLOGY 

Pudella Thomas, 1913, Ann. and Mag. Nat. Hist., (8)11: 588 — type, Pudua mephistophiles de 
Winton. Lydekker, 1915, Cat. Ungulate Mammals Brit. Mus., 4: 217 — subgenus oiPudu 
Molina. Pocock, 1923, Proc. Zool. Soc. London, 1923: 207 — genus of subfamily Pudinae. 
Cabrera, 1%1, Rev. Mus. Argentino Cienc. Nat. "Bernardino Rivadavia," 4(2): 344 — 
subgenus of Pudu Molina. Brokx, 1972, J. Mammal., 53(2): 260 — may he Mazama. 

Mazama, Flerov (part, not Rafinesque), 1952 — Fauna U.S.S.R., n.s. no. 56, Mammals, 1, 
(2): 218 — part, M. mq)histopheles [sic] de Winton only. Haltenorth, 1963, Handb. Zool., 
8(32): 47, 48 — part, Pudu Gray, a subgenus. 

Mezama [sic] Cabrera, 1%1, Rev. Mus. Argentino Cienc. Nat. "Bernardino Rivadavia," 
4(2): 344 — misprint for Mazama in synonymy of subgenus Pudu. 

Coessus [sic], Cabrera, 1%1, Rev. Mus. Argentino Cienc. Nat. "Bernardino Rivadavia," 
4(2): 344 — misprint for Coassus in synonymy of subgenus Pudu. 

Type species. — Coassus pudu [=Capra puda Molina], by virtual tautonymy. 

Included species. — Pudu mephistophiles de Winton, P. puda Molina. 

Distribution (fig. 1). — Temperate zone forests and fringing grasslands of 
southern Colombia, Ecuador, north central Peru, southern Chile, and adjoining 
parts of Argentina. Altitudinal range from sea level to about 1,000 m above in 
Chile and Argentina, and between 2,(X)0 and 4,000 m in Colombia, Ecuador, and 
Peru. 

Conservation 

A program for the study and management of deer threatened with extinction 
throughout the world was launched by the International Union for Conservation 
of Nature and Natural Resources (lUCN). The first of three reports presented by 
Cowan & Holloway (1973; 1974; 1975) lists 25 threatened forms, including the 
South American pampas deer {Blastoceros bezoarticus) , marsh deer {Blastocerus 
dichotomus), and the northern and southern huemuls {Hippocamelus antisensis and 
H. bisulcus). Certainly, the objectives and rationale of this conservation program 
apply no less to pudus than to any other deer spedes threatened with extinction. 

The geographic distribution of pudus given here (figs. 1, 30, 35), based on 
museum specimens and published records, does not reflect the present status of 
the animal in nature. In the years intervening since 1782, when the first pudu 
was made known to science, there has been enormous shrinkage in the 
peripheral and internal ranges of the genus. Hunting by man is a much greater 
menace to the survival of pudus than is habitat destruction. Pudus readily adapt 
as individuals to the marginal and even seemingly improbable habitats to which 
they are being driven. It is unlikely, however, that the spedes can remain viable 
for long under such drcumstances. Present conservation measures, based on 
establishment of national parks and nature reserves, are inadequate, because in 
virtually all cases, the legally protected animals are defended by nothing more 
potent than the mere letter of the law. 

As yet, Peru has no national park or nature reserve where pudus are definitely 
known to exist. Colombia has three parks that certainly harbor the northern 
pudu and three that probably do. Ecuador has one. Chile has four or five parks, 
and possibly more, where the southern pudu still may occur. Argentina can 
boast of at least three national parks where the southern pudu may still be 
found, if only as a vanishing spedes, unless a successful program for breeding 
the animal in semicaptivity is put into effect immediately. 

Information regarding past and recent status of each of the two kinds of pudus 
in the wild is included under various headings in the spedes accounts. 



HERSHKOVITZ: NEOTROPICAL DEER 15 

Taxonomy 

The name Pudu was proposed to contain Capra puda Molina {Cervus humilis 
Bennett, a synonym) by Gray (1850; 1852a [1850], p. 242), as a subgenus of 
Coassus Gray 1843 [=Mazama Rafinesque 1817]. Soon after. Gray (1852b, pp. x, 
240) raised Pudu to full generic rank. Characterization of the genus, however, 
was never adequate until Brooke (1874, p. 36; 1878, p. 926) pointed to the fused 
cuboideonavicular and outer and middle cuneiform bones of the tarsus. 

Pudua, introduced by Garrod (1877, p. 18), is merely an invalid emendation or 
Latinization of the original barbaric name. 

Pudella, erected by Thomas (1913, p. 588) for generic distinction of Pudu 
mephistophiles de Winton from P. puda Molina, was predicated on absence of 
preorbital glands, shallow lacrimal bone, contact between premaxillary and 
nasal bones, crown length of first indsor three to four times that of ia, narrower, 
more pointed hoofs, and rhinarium larger and projecting back in midline. Some 
of the foregoing characters are equivocal, and the others are usually evaluated as 
of specific grade in related genera. For example, in P. mephistophiles, the preorbi- 
tal glands with facial opening are present, not absent as claimed, but are little 
developed, the lacrimal fossa shallow. In P. puda, the glands are large, the 
external opening conspicuous, the lacrimal fossa correspondingly deep. These 
characters parallel those distinguishing most brockets from Mazama rufina, or 
Odocoileus virginianus from O. hemionus. The premaxilla does indeed contact the 
nasals in P. mephistophiles, but may or may not in P. puda. Middle lower indsors 
are not significantly different between these spedes; hoofs are nearly as long and 
narrow, but average deeper in P. puda, and therefore appear to be more blunt. 

Lydekker (1915, p. 217), dting only the fadal glands, lacrimal bones, and 
lower middle indsor crown length as diagnostic characters, reduced Pudella to 
subgeneric rank. Pocock (1923, p. 207), however, regarded Pudella and Pudu as 
distinct genera of his subfamily Pudinae. Flerov (1952, p. 200) referred only to 
the Mazama skull mismatched with the type skin of Pudu mephistophiles for sink- 
ing Pudella in the synonymy of Mazama. Evidently, he overlooked the fact that P. 
mephistophiles was typified by the skin only with assodated leg bones, and 
Pudella was based on the holotype skin plus two topotype skins and skulls of P. 
mephistophiles. In any event, Pudu and Mazama, with P. mephistophiles included, 
were recognized as distinct genera, but other South American genera are listed 
by Flerov as subgenera of Odocoileus. Haltenorth (1963, p. 48) compromised by 
relegating Pudella to the synonymy oiPudu and redudng the latter to a subgenus 
of Mazama. Otherwise, his arrangement of remaining groups of South American 
deer agrees with that of Flerov. 



Generic Characters 

External. — Smallest of Cervidae (figs. 5, 6), legs short, body low slung, rump 
little or not raised above withers, ears more or less rounded; combined head and 
body length less than 85 cm; height at shoulder less than 50 cm; tail obsolescent, 
less than 8% of combined head and body length and concealed in rump; weight 
less than 15 kg. General body color dominantly agouti with crown and ex- 
tremities more or less saturate and contrasting with trunk; ears without white 
edging; lateral narial area and chin rufous to blackish, never marked with white. 




Odocoileus virginianus Blastocerus dichotomus 




Mazama americana 







Pudu mephistopiles Pudu puda Hippocamelus antisensis 

Fig. 5. Size relationship and body form of six South American deer, from smallest 
(Pudu) to largest (Blastocerus); the numbers refer to approximate average shoulder height 
(cm). 



16 



HERSHKOVITZ: NEOTROPICAL DEER 



17 




H ippocamelus antlscusis 



Mazama americana 



Fig. 6. Body form of four South American deer, not to scale; compare dorsal contour, 
rump height, limb proportions, neck, muzzle, ears. From top, left to right, northern pudu 
(photograph by F. Kleinschmidt, courtesy Berlin zoo); southern pudu (photograph by 
H. Fradrich, courtesy Berlin zoo); northern huemul (photograph courtesy Berlin zoo); red 
brocket (photograph courtesy San Diego zoo). 



the pale, lateral rostral band absent (figs. 7, 8); buttocks without sharply con- 
trasting patch or band. Pelage of back harsh, basal portions of hairs pithy and 
brittle; hairs of crown whorled; paired frontal tufts present and unspotted in 
females; nuchal hairs always directed back. Frontal glands not evident in 
females; external opening of preorbital (lacrimal) gland small or large (figs. 8, 9); 
forefoot and hind foot pedal glands contained in shallow interdigital trough or 
small pouch (fig. 10); specialized tarsal and metatarsal tufts weakly developed or 
absent, corresponding glands not defined in dry skin. Fore and hind hoofs 
slender, length twice or more greatest width; fore and hind dew claws longer 
than wide; soles between outer and middle digits thickly clothed with hair 
(fig. 10). 

Cranial (figs. 11-17). — Rostrum short, the diastema shorter than alveolar 
length of upper cheek teeth (pm^-m^); length of nasals less than length of fron- 
tals at midline; superior border of orbit depressed below frontal suture at same 
plane; greatest supraorbital breadth of frontals less than width of brain case; 
upper portion of supraocdpital moderately inflated, its median protuberance 



18 



FIELDIANA: ZOOLOGY 




Fig. 7. Fadal color pattern of Odocoileinae, a composite of all spedes. 

posterior auricular border 
inner auricular surface 
outer auricular surface 
mental patch 
mandibular band 
buccal patch 
gular patch 

opening of preorbital pouch 
frontal spot or tuft (marks 
frontal gland in female) 



a. 
b. 


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dark lateral rostral band 


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k. 


c. 


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d. 


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e. 


inferior orbital band 


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f. 

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h. 
i. 


superior orbital band or 
superdliary band 
anterobasal auricular patch 
posterobasal auricular patch 
anterior auricular border 


o. 
P- 

q- 

r. 



projecting beyond lambdoidal crest in lateral view; ascending ramus of pre- 
maxillary extending to, or widely separated from, lateral border of nasals; malar 
articulating with one-half or less inferior suture of fadal plate of lacrimal and 
hardly or not at all entering into composition of inferior surface of lacrimal fossa; 
transverse diameter of orbit greater than inferior margin of fadal plate of lacri- 
mal; auditory bullae little or not inflated. 



HERSHKOVITZ: NEOTROPICAL DEER 



19 





Pudu mephistophiles 



Pudu puda 




Mazama chunyi 



Fig. 8. Stylized heads of smallest South American deer showing individual color pat- 
tern, ear form, preorbital pouch opening, and dorsal outline of rhinarium. 



Dental (figs. 18-23). — Greatest mesiodistal crown length of permanent ii equal 
to or less than combined crown length of permanent i2-3 (fig. 22), all permanent 
indsors not in place before eruption of last molar; lower deciduous front teeth 
like replacements, but dii relatively broader, dia-s and dc narrower, more 
pointed, the entire series of front teeth approaching those of adult Mazama 
(fig. 22); deciduous upper canine usually present in Pudu mephistophiles, never 
persistent, but often succeeded by the permanent canine; dc absent in P. puda 
or, if present, shed after eruption of m^ and rarely, if ever, replaced (figs. 11, 13, 
14, 16). 




Mazama gouazoubira 



Hippocamelus antisensis 

Fig. 9. Heads of five South American deer; compare fadal markings, muzzle length, 
preorbital gland, ear shape; antlers of P. mqjhistophiles partially rubbed, of M. rufina and H. 
antisensis in velvet. (Photograph of Pudu puda courtesy Cologne zoo; remaining photo- 
graphs courtesy Berlin zoo.) 



20 



FRONT 



HIND 



FRONT HIND 




Pudu mepbistophiles 





Pudu puda 




f i 




Mazama rufina 






Odocoileus virginianxis 



Fig. 10. Front and hind feet (left lateral and ventral surface) of pudus, brocket, and 
Virginia deer. Pouch of interdigital gland is indicated by dashed line. 



21 



22 FIELDIANA: ZOOLOGY 

On the basis of upper molars of a specimen of Mazama americana ("M. tema") 
and one of Odocoileus virginianus, Frechkop (1965) concluded that all 
Odocoileinae (his Capreolinae) were characterized by a short crista or crochet, 
springing from the inner distal enamel border of the hypocone, labeled "x" in 
his illustrations (Frechkop, 1%5, p. 4, figs. 1, 2, 3). In a spot check of Field 
Museum specimens, this crista (fig. 23, XI) appears to be absent or incipient in 
Pudu mephistophiles and Blastoceros bezoarticus, and rudimentary to moderately 
developed in unworn molars of P. puda, usually present in Mazama americana and 
M. rufina, present or absent in Mazama gouazoubira, Blastocerus dichotomus, 
Odocoileus hemionus, and O. virginianus. In two specimens at hand of Mazama 
chunyi, the character is present in m^'^ of the type, a juvenal with unerupted m^, 
and undeterminable in the considerably worn molars of an old female (fig. 18). 

Crest XI size is variable in all spedes. Where it appears to be absent in moder- 
ately worn teeth of a given spedes, it may be found in one or more unerupted 
molars, mainly m*, of some individuals of the same spedes. In some cases, a 
crest equivalent to crista XI springs from the medial border (V) of the hypocone. 
The comprehensive account of cervid dental morphology is reserved for a future 
artide of this series. 

Dental succession. — Sequence of eruption of permanent teeth determined from 
inadequate material in the Field Museum and extrapolations therefrom, is: 

ml-m2-m3-pm4-pm3-pm2-c (P. mephistophiles) 
ml-m2-(il or m3)-pm4-pm3-pm2-i2, (i3-c). 

Antlers. — Spike-like, usually less than 10 cm measured from burr and directed 
straight back in line with dorsal plane of frontal bone. Pedides develop during 
eruption of m^, burr and antlers form during eruption of m^, well-developed 
antlers present in all 17 males examined with complete dentition. 

Antler casting is not evident in present material. Published and secondary 
data indicate, however, that new antlers grow during spring to early summer, 
mature during summer to early fall, drop in winter. Antler cyde of southern 
hemisphere pudus acdimatized in northern hemisphere zoos adjusts to north- 
em seasons. 

Vertebral formula. — Seven cervicals, 12-13 thoradc, 6-7 lumbar, 4-5 sacral, 8 
caudal; total, 38-39 vertebrae. Generally, form and relative size are the same as 
those of other cervids. 

Limb bones and proportions (fig. 24, table 2). — Metapodials extremely short and 
broad; cuboid, navicular and external and middle cuneiform bones fused into a 
single tarsal element; length of metacarpal splints II and V from approximately 
one-half to two-thirds length of metacarpals III and FV; central portions of 
metatarsal bones II and V frequently persistent as splints, each ankylosed to the 
adjacent metatarsal bone. 

Male genitalia. — In reference to Pudu puda, Brambell & Mathews (1977, p. 27) 
note, "The testes are subcutaneous, but not scrotal. The penis is not long but lies 
on the abdominal wall." Presumably, the same condition obtains in P. mephis- 
tophiles. 

Generic Comparisons 

Pudus, like other strictly New World deer, are characterized by retention of 
the distal portion of the obsolescent metacarpals II and V (telemetacarpalid) and 



HERSHKOVITZ: NEOTROPICAL DEER 23 

by division of the posterior nares into two chambers by the vertical plate of the 
vomer (figs. 2, 3, 24). In exclusively Old World Cervinae, vestiges of the lateral 
metacarpals are proximal (plesiometacarpalid) and the posterior nares are undi- 
vided.^ 

Pudus are most readily distinguished from all other deer by their small size 
(fig. 5). Only the dwarf brocket, Mazama chunyi Hershkovitz, is similarly small. 
Pudus are further distinguished from other cervids by the low-slung body with- 
out highly raised rump; short, thick legs, comparatively small, more or less 
rounded ears, small eyes, thick rhinarium with rounded nostrils, muzzle with- 
out white narial and mental patches or spots (figs. 6, 8, 9), tail completely hidden 
by rump hairs; mandibular symphysis usually more angular in horizontal plane 
than in related deer, alignment of incisors and canines of each side more nearly 
oblique than transverse. Caudal vertebrae number only 8 in Pudu, 11 in Mazama, 
and 10 or more in other cervids. The shallow interdigital depression or pocket 
containing the pedal glands in Pudu is primitive compared with the well- 
developed interdigital pouch of the hind feet, at least, of other New World deer 
(fig. 10). 

The key osteological character separating Pudu from other true deer, i.e., 
Cervinae and Odocoileinae, is the consistent union of the cuboideonavicular and 
external and middle cuneiform tarsal bones into a single bone (figs. 2, 4, 24). 
Other living cervids with similarly fused tarsal bones are the Asiatic plesio- 
metacarpalid tufted deer (Elaphodus) and muntjac {Cenmlus Blainville, the valid 
name currently replaced by the erstwhile nomen nudum Muntiacus Rafinesque 
[Int. Comm. Zool. Nomencl. 1957, Opinion 460]). It is a curious coincidence, as 
shown by Pocock (1911, p. 952, figs. 134b-c), that the interdigital integumentary 
folds containing fore and hind pedal glands are about as simple in muntjacs and 
tufted deer as in pudus. The extremely short, broad metapodials of Pudu, with 
metacarpals proportionately wider than metatarsals, are a bovine-like character 
unmatched in other cervids (table 2). 

Dental peculiarities are absent in Pudu (figs. 18-23). The form of their lower 
incisors accords with the less specialized deer type characterized by the rectan- 
gular or moderately spatulate crown of ip The more specialized crown of ii in 
Mazama and a number of other New and Old World deer is broadly spatulate or 
cuneate. In the deciduous teeth of most pudus, however, a comparatively ex- 
panded crown of ii, and attenuate iz-s, Ci adumbrate the more specialized front 
teeth of brockets (figs. 20, 22). The upper canine persists in many adult P. 
mqjhistophiles, whereas only the deciduous upper canine appears occasionally in 
immature P. puda. 



Fawnal Spotting Pattern 

The Juvenal coat of Pudu puda is spotted, whereas that of P. mephistophiles is 
not. Inasmuch as adult P. mephistophiles is more primitive than P. puda in every 
respect, including coat color, it is logical to infer that the unspotted juvenal coat 

^Among circumboreal cervids, the American moose or Palearctic elk (Alces) and caribou 
or reindeer (Rangifer) are telemetacarpalid, with the nares undivided in Alces and divided 
in Rangifer. The wapiti or deer (Cervus) is plesiometacarpalid, the nares undivided. Old 
World roebuck (Capreolus), Chinese water deer (Hydropotes) , and musk deer (Moschiis) also 
are telemetacarpalid. 



24 FIELDIANA: ZOOLOGY 

color is also more primitive. Fawns of the Andean taruga, Hippocamelus antisen- 
sis, and the marsh deer, Blastocerus dichotomus, are also unspotted. Those of 
remaining neotropical deer are spotted (cf., figs. 25-27). Judged on its own 
merits, an unspotted, unmarked, or unpatterned coat characterized by agouti 
pelage is most primitive. Conversely, any coat color pattern that departs from 
the primitive agouti as a result of pigmentary dilution or loss is derived. In the 
case of white or whitish fawnal spotting, the pigmentary loss appears in the fetal 
pelage or lanugo. The juvenal character adumbrates the adult spotting that 
appears on some axis deer {Axis), fallow deer (Dama), and some Sika deer {Cer- 
vus). In these forms, the overall trend is toward paler coloration approaching 
albinism. 

The whitish spotting pattern similar to that of fawns, or a more advanced 
whitish spot and stripe, or entirely striped pattern appears in many mammalian 
lines. This characterizes the juvenal coat of swine and tapirs, and the adult coat 
of many rodents, e.g., ground squirrels {Spermophilus) , paca (Agouti), and the 
white stripe pattern of zebras. A blackish spotting coat pattern, noted in most 
felids and many other groups, is also common, whether in the juvenal or adult 
stages, or both. In these, however, the blackish spots may be reminders of a 
more primitive blackish coat. On the other hand, whitish spotting, which entails 
loss of melanin, is derived. 

Ancestral Characters and Relationships 

Pudus sometimes are referred to as dwarf or pygmy deer, and it has been 
suggested (von Lehmann, 1960, p. 50) that these and all other spike-antlered 
deer are literally dwarfed. The ancestral pudu is unknown and it cannot be 
ascertained whether it was larger or smaller than its descendants. Nevertheless, 
of the two living kinds, the smaller Pudu mephistophiles is evidently more primi- 
tive than the larger P. puda. Brockets, genus Mazama, are also small, ranging in 
size from Mazama chunyi, which is equal to the larger spedes of Pudu, and M. 
americana, which grades into the smaller forms of the otherwise much larger, 
white-tailed deer, Odocoileus virginianus. Thus, it can be asserted that the overall 
trend is toward greater body size and increasing antler size and complication. 
No doubt, dwarfed populations and races of deer develop from time to time, but 
probably most, if not all, are ecological rather than genetic demes or races. 

Small size and rudimentary antlers restricted to the male are primitive deer 
characters. Also, the smaller the deer, the smaller and more simple its antlers, 
irrespective of the age of the individual or grade of specialization of any or all 
other of its parts. Conversely, the larger the individual, the larger the antlers 
tend to grow, and the larger the race or spedes within a given phyletic line, the 
more complex the antlers tend to become. Thus, the extremely small body and 
simple antlers of Pudu, although primitive, do not in themselves imply an an- 
cestral condition or denote a close relationship to the small-bodied, spike- 
antlered Mazama. 

The affinities of Pudu, morphologically as well as historically, are with other 
neotropical cervids. However, the comparatively short muzzle, obsolescent tail, 
fused cuboideonavicular and external and middle cuneiform bones, and short, 
broad metapodials remove Pudu from all other deer and from consideration as an 
ancestral model. At the same time, primitive cervid characters of Pudu mephis- 
tophiles from which those of Mazama and other New World deer can be derived 



HERSHKOVrrZ: NEOTROPICAL DEER 25 

include the relatively large and well-developed lateral metacarpals, vertically 
inclined antorbital vacuity, shape of lower indsors, the more than sporadic 
persistence of upper canines, the comparatively simple interdigital tegumentary 
fold for containment of pedal glands, absence of specialized tarsal and metatar- 
sal tufts and glands, comparatively short, rounded ears and small eyes, slightly 
raised rump, dominantly agouti pelage, and unspotted fawn. 

The remote, possibly Miocene-Pliocene, nearest common ancestor postulated 
here must have been sylvan, smaller than living pudus, body low slung, rump 
unraised, antlers simple or rudimentary and directed back parallel to the frontal 
plane. The tail must have been comparatively well developed, middle digits, 
including hoofs, long and mobile, facial or lacrimal glands small but not 
rudimentary, tarsal, metatarsal, and frontal glands and tufts rudimentary or 
absent, ears small, rounded, nuchal hairs directed back, coronal crests or whorls 
absent, adult pelage coarse but not pithy, general coloration agouti without 
special markings, fawn unspotted, metapodials broad, lateral metacarpals nearly 
or virtually complete, the corresponding digits or dew claws well developed, 
cuboideonavicular and cuneiform bones discrete, premaxillary and nasal bones 
in contact, antorbital vacuity small, its long axis vertical, crowns of lower indsors 
more nearly square than spatulate, upper canines small and persistent, vertebral 
formula 7C, 13T, 7L, 5S, IIC. More characters can be added, but with those 
mentioned, the others fall into place. 

Origin, Dispersal, Speciation 

The origin of Pudu in terms of time and place is unknown. The ancestral stock 
of the genus, and the Odocoileinae in general, is likewise unknown. As noted 
earlier (p. 8), the Palaeomerydnae, including Blastomeryx, the imputed ancestral 
root, is a spedalized traguline-like or moschine-like group that disappeared 
without known issue in the New World. In any event, it seems likely that Pudu 
and Mazama represent divergent lines of the tiny ancestral odocoileine, which 
must have lived during the early Pliocene or late Miocene in boreal America, 
Middle America, or South America. The genus Pudu itself must be regarded as 
South American, likely since Pliocene. The distribution and interrelationship of 
its two living spedes are clear, and their evolutionary history can be recon- 
structed from available morphologic and geographic data as well as from what is 
known of Pleistocene climates (Van der Hammen, 1974). 

It is believed that the immediate ancestor of modem pudus differed little from 
Pudu mephistophiles. During what possibly was a last gladal period, the former 
must have occupied the temperate zone forest fringe at the eastern base of the 
Cordillera Oriental, in what now is Colombia, Ecuador, Peru, and, perhaps, 
northern Bolivia. When the Andean gladers retreated, the temperate zone forest 
with its fauna followed in its wake. In the case of the pudu, some populations 
moved into higher altitudes at the same latitudes, clinging to the forest fringes 
forming on the shores of large rivers, such as the Putumayo, Pastaza, Huallaga, 
and Ucayali. Other populations moved southward along the base of the eastern 
Andes into higher latitudes as far as the Strait of Magellan and westward 
through forested passes into Chile. Evidently, no relic pudu populations per- 
sisted where the climate of the original temperate lowland habitat became tropi- 
cal or subtropical (fig. 1). 

Complete geographic isolation of northern and southern pudus is reflected in 




3 
< 



E uu 



26 




Fig. 12. Adult female pudu and brocket skulls, dorsal surface. 
27 




Fig. 13. Adult female pudu and brocket skulls, palatal surface. 



28 




•T3 

C 
«5 

D 
3 



29 




Fig. 15. Adult male pudu and brocket skulls, dorsal surface. 
30 




Fig. 16. Adult male pudu and brocket skulls, palatal surface; inset, enlargement of 
premolars 2-4 with dedduous-like pm^. 



31 




Fig. 17. Mandibles of pudus and brockets, lateral surface of left rami. 

32 






Pudu mephistophiles 868A6 



^S^t&s^ 



Pudu puda 2A363 



;^#<^"^ 







Mazama rufma 


70563 




"^15^ 


^%^^ 








Mazama gouazoubira 


51091 














Mazama americana 


867A1 






n&% 








Wazama chunyi (tvne) 


73098 



Fig. 18. Upper cheek teeth (pm^-m^) of adult pudus and brockets, and dpm^"^ m'"^ of 
Juvenal Mazama chunyi (type, AMNH). 

33 



Fig. 19. Lower cheek teeth (pirij-ms) of adult pudus and brockets, and dpm2.4, m,-2 of 
Juvenal Mazama chunyi (type). 



34 




3 
3 



35 




36 




Pudu mephistophiles 
juv, 88595 ^^ ' 88496 




^^- '-' Mazama chunyi 

juv. 73098 ad. 79912 

Fig. 22. Permanent and deciduous incisor-canine series (i^a, c) in pudus and brockets; 
Mazama chunyi juv. 73098, type, AMNH. 



37 




Pudu mephistophiles 




Pudu mephistophiles 

Fig. 23. Upper right and lower left second and third molars of Pudu, the top m^ un- 
erupted of type, AMNH 73098. For explanation of symbols, see fadng page; system of 
symbols and homologies based on Hershkovitz (1977, pp. 298-301). 

38 



Explanation of Symbols Used in Figure 23. 



I 


eocene (paracone) 


1 


eoconid (protoconid) 


2 


protocone 


2 


metaconid 






3 


[paraconid, absent] 


4 


metacone 


4 


hypoconid 


5 


hypocone 


5 


entoconid 


a 


mesostyle (parastyle) 


a 


mesiostylid (parastylid) 


b 


distostyle (metastyle) 


b 


distostylid (hypoconulid) 


d 


plagioconule (metaconule) 


d 


plagioconulid 


e 


postentoconule 


e 


postentoconulid 






h 


entostylid-Zi 


i 


ectostyle-/ (stylocone) 






k 


ectostyIe-/f 


k 


ectostylid-/c* 


1 


ectostyle-/ (mesostyle) 


I 


ectostylid-/ (mesostylid) 


m 


ectostyle-m 


m 


ectostylid-m 


I 


eocrista 


I 


ecocristid 


II 


epi crista (crest labeled "+" 
by Frechkop, 1965, figs. 1,2) 


II 


epicristid 


III 


protoloph (protocrista) 


III 


protolophid 


IV 


plagiocrista (metaloph) 
IV' medial 
IV" median 


IV 


plagiocristid 


V 


entocrista 






VI 


distocrista (from postero- 
lingual cingulum C) 


VI 


distocristid 






VIII 


postentocristid 


XI 


crista-XI (individually 
variable in cervids) 






XII 


crista-XIl (individually 
variable in cervids; charac- 
teristic of notoungulates) 


' 


' 


e 


pretrigon basin or fossa 


e 


"trigonid" basin or fossa 


a 


trigon basin or fossa 


a 


talonid basin or fossa 


P 


talon basin or fossa 


/3 

en entoflexus 
ex ectoflexus 
nex neoectoflexus 


talonid basin or fossid 
S posttalonid basin 
or fossid 



* Present in lower first molars only of individual figured. 



39 



FRONT LIMB 




^ 1 



VENTRAL 



iii-h ia-iv 



sesamoid- 



\l 



Pudu mephistophiles 

86846 



Pudu puda 




HIND LIMB 

Fig. 24. Left forefeet and hind feet of pudus with vestiges of metacarpal bones II and V 
distal in position (telemetacarpalia). Digits are missing in Pudu mqjhistophiles. 



40 




Fig. 25. One-week-old southern pudu fawn (photograph by H. Fradrich, courtesy 
Berlin zoo). 




Mazama gouazoubira 



Fig. 26. One-week-old gray brocket fawn (photograph by J. Fradrich, courtesy Berlin 
zoo). 



41 



42 



FIELDIANA: ZOOLOGY 




Mazama amer icana 

Fig. 27. Red brocket doe with fawn (photograph courtesy Gladys Porter Zoo, 
Brownsville, Texas). 



the wide morphologic differences between them. The southern spedes, with its 
much greater distance from the hypothetical ancestral home, departs most from 
the postulated ancestral form. Hippocamelus, with its two spedes, the northern 
H. antisensis and the southern H. bisulcus, appears to have had a parallel history 
of dispersal and spedation. 

Most populations of the northern P. mephistophiles are still confined to the 
heads of the same Amazonian streams that defined their presumed habitats 
lower down. A few have spread across divides into watersheds drairung into the 
Caribbean or Padfic Oceans. Apparent absence of significant differences be- 
tween extant populations of northern Colombia, Ecuador, and Peru suggests 
that gene flow has been maintained between them throughout the major climatic 
event that shifted without notably altering the pudu habitat. Alternatively, the 
habitat shift was accomplished so swiftly or recently, with the actual linear 
distances covered so short, that no significant differentiation occurred in any 
population, geographically isolated as it may appear to be now or may have been 
then. 



HERSHKOVITZ: NEOTROPICAL DEER 



43 



III. THE SPECIES OF PUDUS, GENUS Pudu GRAY 
Key to the Species 

Size smaller; face, outer surface of ears, chin, forefeet and hind feet dark brown or black- 
ish; preorbital gland small, external opening inconspicuous; lacrimal fossa shallow or 
dish-shaped; long axis of antorbital vacuity nearly or quite vertical; fawns unspotted; 
(Colombia, Ecuador, Peru) Pudu mqyhistophiles (figs. 5, 6, 8, 9, 28, 29; p. 44). 

Size larger; face reddish buffy or brown; ears, narial patch, chin, forefeet and hind feet 
reddish; preorbital gland large, external opening large, conspicuous; lacrimal fossa deep 
or bowl-shaped; long axis of antorbital vacuity diagonal to nearly horizontal to basal 

plane of skull; fawns spotted; (Chile, Argentina) 

Pudu puda (figs. 5, 6, 8, 9, 36, 37; p. 60). 




Fig. 28. Northern pudus, adult male and female (photograph by H. Fradrich, courtesy 
Berlin zoo). 




Puau mephi stopni 1 es 



Fig. 29. Northern pudu, adult female, same individual shown in Figure 28. Note frontal 
tuft (photograph by H. Fradrich, courtesy Berlin zoo). 



44 FIELDIANA: ZOOLOGY 

IV. Pudu mephistophiles de W7NTON 

NORTHERN PUDU (figs. 28, 29) 

Synonymic History 

Pudua mephistophiles de Winton, 18%, Proc. Zool. Soc. London, 1896(2); 508, pL 19 
(animal) — part, not skulls shown in figs. 2, 4 [=Ma2ama]. Lydekker, 1898, Deer of all 
lands. Rowland Ward, London, p. 308, pi. 24, fig. 1 (animal) — characters. Lydekker, 
1901, Great and small game of Europe, western Asia and America. Rowland Ward, London, 
p. 374 — characters. 

[Pudella] mephistophiles, Thomas, 1913, Ann. Mag. Nat. Hist., ser. 8, 11: 588 — classification. 

Pudu (Pudella) mephistophiles, Lydekker, 1915, Cat. Ungulate Mammals Brit. Mus., 4: 217; 
ECUADOR: Pichincha (Paramo de Papallacta); characters; type history. G. M. Allen, 
1942, Extinct and vanishing mammals of the western hemisphere, p. 412 — ECUADOR; char- 
acters; type history; endangered species. Avila Pires, 1974, Publ. Biol. Inst. Invest. 
Gent., U.A.N.L., Mexico, 1:161 — distribution; conservation. 

P[udu]. mephistophiles, Hoffstetter, 1952, Mem. Soc. Geol. France, 31(66): 369— 
ECUADOR: Mazama fusca Spillmann, a synonym. 

Pudu mephistophiles, Hershkovitz, 1958, Mammalia, 22(4): 545 — metatarsal glands and tufts 
not defined. Hershkovitz, 1959, Proc. Biol. Soc. Washington, 72: 48-49, pis. 3C, 4D 
(skull), pi. 5B (front teeth) — comparisons. Cabrera, 1%1, Rev. Mus. Argentine Cienc. 
Nat. "Bernardino Rivadavia," 4(2): 344 — distribution; P. m. wetmorei Lehmann, a 
synonym. Crandall, 1964, Management of wild animals in captivity. U. of Chicago Press, 
pp. 557, 560, 594-5 — characters; no record of captivity. Grimwood, 1%9, Spec. Publ. 
no. 21, American Comm. Int. Wild Life Protection and New York Zool. Soc., p. 78 
(map), 79 — PERU: Hudnuco (Pamacucho [sic]; Carpish summit on Huanuco-Tingo Maria 
road;- upper Rio Huallaga valley, above Huanuco; near ChagUa; Yanacocha, Rio Quero, 
25 km NW Huanuco; between Huanuco and Ambo, 2700-3300 m; Quivilla road, 40 km 
from Huanuco); Junin (above San Ramon, 200-300 m; Rio Huasahuasi valley; upper Rio 
PaUcartambo, 10°50' S, 70°00' W); conservation; local names sac/ia cabra, antagllo. Dansie 
and Wince, 1971, Deer of the world. Hitchingsand Mason, Ltd., Plymouth, England, pp. 
21, 23, fig. (map, distribution)— ECUADOR; PERU. Schauenberg, 1973, Mus. Geneve, 
14(134): 12— figs. pp. 13, 14 (animal), fig. p. 15 (habitat), fig. p. 16 (head)— ECUADOR; 
COLOMBIA; history; conservation; distribution; occurrence in Peru questioned. 

P[udu]. mephistophiles mephistophiles, Mochi and Carter, 1953, Hoofed mammals of the loorld, 
Scribner's and Sons, Ltd., London, pi. VIII, text only. 

Pudu m[ephistophiles]. mephistophiles, Lehmann, 1960, Nov. Colombianas, Univ. Cauca, 
Popayan, 1(5): 282— ECUADOR; antler form. 

Pudu mephistophiles mephistophiles. Whitehead, 1972, Deer of the world. Viking Press, New 
York, pp. 29, 64, 165, map 9 (distribution) — distribution; classification. 

Pudua mephistopheles [sic], Thomas, 1908, Ann. Mag. Nat. Hist., ser. 8, 1: 350 — cranial 
characters: skull originally figured as corresponding to type skin =Mazama. Lonnberg, 
1913, Ark. Zool., Stockholm, 8 (16): 33— ECUADOR: Pichincha (Papallacta, 12,000 ft). 

Pudu mephistopheles [sic], Soukup, 1%1, Biota, 3 (28): 331— PERU. Fradrich, 1975, Zool. 
Gart., 45(1): 67, 72, figs. 5, 6, 7 (animal), fig. 9 (skuU)— ECUADOR: Cotopaxi (W of 
Salcedo); maintenance in Berlin zoo. Brambell and Mathews, 1977. Proc. Symposium 1, 
Ass. Br. Wild Animal Keepers, p. 25 — characters. 

P[udu]. mephistopheles [sic], Brokx, 1972, J. Mammal., 52(2): 360 — upper canine present in 
specimen examined. 

Pudu (Pudella) mephistopheles [sic], Lehmann, 1945, Rev. Univ. Cauca, Popayan (Colombia), 
Zool., no. 6: 76 — comparisons. 

Pudella mephistophelis [sic], Cabrera and Yepes, 1940, Mamiferos Sud- Americanos. Hist. Nat. 
Ediar (Buenos Aires), p. 278— ECUADOR; PERU. 

M[azama]. mephistopheles [sic], Flerov, 1952, Fauna U.S.S.R., n.s. no. 56, 1(2): 218 (in text), 
220 (in text) — probably a juvenal of M. rufina Pucheran. Haltenorth, 1963, Handb. Zool., 
8(32): 48— ECUADOR; PERU; characters. 

M{azama\. m[ephistopheles\. mephistopheles [sic], Haltenorth, 1963, Handb. Zool., 8(32): 
48 — classification. 

Mazama fusca Spillmann, 1931, Die Saugethiere Ecuadors im Wandel der Zeit. Univ. Cen- 
tral, Quito, 1: 37, 41 (measurements) — ECUADOR: Pichincha (type locality, Mt. Antisana 
near snow line); holotype, adult male, skull only, perhaps preserved in the Escuela 



HERSHKOVITZ: NEOTROPICAL DEER 45 

Politecnica, Quito. Hoffstetter, 1952, Mem. Soc. Geol., France, 31(66): 369, regarded as a 
synonym of P. mqjhistophiles. 

Pudu (Pudella) mq)histophiles wetmorei Lehmann, 1945, Rev. Univ. Cauca, Popayan, (Co- 
lombia), Zool. no. 6: 76, figs. p. 78, 80 (animal; skull)— COLOMBIA: Cauca (type locality. 
Paramo de San Rafael, Purace, ESE Popayan, western slope of the Cordillera Central, 
3400 m); holotype, adult male, skin and skull, Museo de Historia Natural, Universidad 
del Cauca, Popayan, no. 3801, collected 1 February 1945 by F. C. Lehmann. Cabrera, 
1%1, Rev. Mus. Argentino Cienc. Nat. "Bernardino Rivadavia," 4(2): 344 — a synonym 
of P. mephistophiles de Winton. 

P{udu\. mq)histophiles ivetmoorei [sic], Hoffstetter, 1952, Mem Soc. Geol. France, 31(66): 
369— COLOMBIA. 

P[udu]. m[ephistophiles]. wetmorei, Mochi and Carter, 1953, Hoofed mammals of the world, 
Scribner's and Sons, Ltd., London, pi. VIII, text only. 

Pudu mephistophiles wetmorei, Lehmann, 1959, Nov. Colombianas, 1(4): 202, fig. (animal), 
fig. 3 (antlered skull) — COLOMBIA: Cauca (Paramo de Guanacas, partial albino; Paramo 
de las Delicias, albinos); Valle (Paramos de Barragan, east of Tulua); description; skull; 
hunting; conservation. Whitehead, 1972, Deer of the world. Viking Press, New York, pp. 
29, 64, 169, map 9 (distribution) — classification; distribution. 

P[udu]. mephistophelis [sic] wetmoorei [sic] Hoffstetter, 1952, Mem Soc. Geol. France, 
31(66): 369— COLOMBIA. 

Pudu mephistophelis [sic] wetmori [sic], Keirans, 1973, J. Med. Entemol., 10(3): 254 — 
COLOMBIA: Cauca (Pusna, 2800 m); tick parasite {Ixodes montoyanus). 

M[azama\. m[ephistopheles]. [sic] wetmorei, Haltenorth, 1963, Handb. Zool., 8(32): 48 — 
COLOMBIA. 

Pudu sp. Keirans, 1973, J. Med. Entemol., 10(3): 254— COLOMBIA: Cauca (Malvasa, 3500 
m, 3000 m); tick parasite (Ixodes montoyanus). 

Type. — Originally described from the skin and attached leg bones of an imma- 
ture female, and a mismatched skull of Mazama rufina, both presented in 1896 to 
the British Museum (Natural History), by Ludovic Soderstrom. The mismatch 
was discovered by Thomas (1908, p. 350) and the type was restricted to the skin 
BM 96.1.28.5, by Lydekker (1915, p. 217). 

Type locality. — Said to be the Paramo de Papallacta, but probably taken in 
temperate zone forests below paramo zone, southeast of Quito, eastern slope of 
the Cordillera Oriental, Pichincha, Ecuador. The altitude is said to be between 
3,6(X)-3,700 m above sea level, but probably should be estimated at 200 or 300 m 
lower. 



Distribution 
(fig. 30) 

Restricted to the temperate zone forests and fringing grasslands or paramos 
above treeline in the Cordillera Central of southern Colombia, from the depart- 
ments of Valle and Tolima southward through the Cordillera Oriental of Ecuador 
and Peru into the department of Junin. The parts of the Andean chains actually 
occupied are poorly known, and distributional gaps between locality records in 
northern Ecuador and central Peru are unresolved. 

According to Lehmann (1945, p. 76), the Colombian pudu is known to occur in 
the Cordillera Central, from the Paramo and Valle de las Papas and probably the 
Paramo del Lebrero, northeast of the Valle de las Papas; northward through the 
paramos of Sotara, Purace, Las Delidas, Guanacas, and Moras, in the depart- 
ment of Cauca, to those of Las Hermosas in the department of Valle. The pudu 
has also been reported by Lehmann (1959, p. 203) to the north in the region of 
the Paramos de Barragan east of Tulua, department of Valle. In Lehmann's 
opinion, the range never extended farther north because of a deep depression in 




Fig. 30. Distribution of Pudu mepbistophiles de Winton, based on museum specimens 
and published 19th and 20th century locality records (dots). For locality names, see corre- 
sponding numbers in "Gazetteer" (p. 80). 



46 



HERSHKOVITZ: NEOTROPICAL DEER 47 

the Andes that acts as a climatic barrier. Altitudinal range is given as between 
3,000 and 4,000 m. 

Dr. Hernando Chirivi Gallego, head of the Laboratorio de Fauna of INDE- 
RENA (Institute de Desarrollo de los Recursos Naturales Renovables), Minis- 
terio de Agricultura, Bogota, informs me (/>i litt., 23 April 1979) that the range of 
Pudu mephistophiles may extend as far north as Mt. Quindio (4°42'N, 75°25'W). 
He adds that a skull of a pudu from the Paramo de Moras is in the INDERENA 
collection, and that pudus are well known by the name venado chonta in the 
paramos of Azufral, Cumbal, and Chiles in the department of Narifio. 

The parts of paramos presumably inhabited by pudus are actually sub- 
paramos, mostly — if not entirely — secondary in origin and characterized by low, 
thin, fire-resistant forest growths or matorrales, usually dominated by Polylepis. 
The steep, eastern slope of the Cordillera Central in the upper Rio Magdalena 
valley, below the Paramo de las Papas in the department of Huila, where 1 found 
signs and remains of pudus, ranges from 2,700 to 4,000 m above sea level and is 
thickly forested. 

The seeming absence of pudus from the Colombian Cordillera Oriental, which 
joins with the Cordillera Central at the Paramo de las Papas, is perplexing. The 
head of the Rio Magdalena, which marks the dichotomy between the eastern 
and western chains, is narrow and shallow enough at altitudes above 2,000 m for 
a pudu to jump, walk, or step across, and no doubt pudus do as occasions 
require. Farther down and eastward, the crest of the Cordillera Oriental loses 
altitude, and, passing the point (Picos de Fragua) where it turns sharply north- 
eastward, it forms a deep saddle at the divide between the Rio Suaza (Mag- 
dalena) and Rio Orteguaza (Caqueta). Here the climate is comparatively dry, 
upper tropical to subtropical, and acts as a barrier to northern spread of many 
temperate zone mammals, including pudus, but serves (or served) as a pass for 
tropical zone Amazonian mammals. The Colombian Cordillera Occidental, ex- 
cept for a few isolated peaks and subparamos, is too low to provide suitable 
habitat for pudus. 

The pudu has not been recorded from Carchi and Imbabura, the two Andean 
provinces of northern Ecuador, but is known there by the name sacha cabra. One 
was captured with hunting dogs in the virgin forests northeast of San Gabriel, 
Carchi, during my explorations there in 1934. 

Dr. Fernando Ortiz Crespo of the Pontifica Universidad Catolica del Ecuador 
in Quito, informs me (in litt., 6 March 1979) that he heard rumors of pudus being 
sighted around Volcan Chiles in the western Andes at the Ecuadorian- 
Colombian border. Dr. Ortiz mentions that the scarcity of the species may be 
due to competition with Mazama in forests between 2,500-3,200 m altitude, and 
to the absence of suitable habitat higher up. He observes, however, that the 
animal is rarely encountered because of its nocturnal or, at least, crepuscular 
habits and the difficulty of access to its extremely accidented habitat where it can 
be hunted only with dogs. He particularly notes that pudus are not confined to 
the isolated patches of Polylepis forest found in paramo and that these animals, 
including the holotype recorded from the Paramo de Papallacta, may have origi- 
nated in forests well below the paramo zone. 

The occurrence of pudus in the southern half of Ecuador, south of the Rio 
Pastaza, and northern Peru, north of Huanuco department, is not on record. 
However, Dr. Antonio Brack Egg, a distinguished naturalist, formerly of the 
Direccion General Forestal y de Fauna, Lima, believes {in litt.) that pudus may be 
present in the paramos of Huancabamba at the headwaters of the Rios Quiroz, 



48 FIELDIANA: ZOOLOGY 

Chinchipe, and Huancabamba, in the departments of Piura and Amazonas on 
the Peruvian-Ecuadorian border. Grimwood (1%9, p. 79) provides the following 
details on the occurrence of pudus in central Peru: 

The occurrence of this genus within Peruvian limits does not appear to have beeri 
previously recorded. There is, however, a misidentified mounted specimen of Pudu 
mephistophiles (De Winton) in the museum of the Colegio "Leoncio Prado" at 
Huanuco, which was shot some years ago at Pamacucho, 4 kms. from that town; and 
in 1965 I obtained the skin and skull of a male of that species which had been killed 
near Carpish summit on the Huanuco-Tingo Maria Road. Subsequently, I have seen 
the animal alive in that region and have been shown skins or photographs of other 
animals killed there and in a number of localities in the upper Huallaga valley ab>ove 
the town of Huanuco. 

The distribution of this spedes appears to be very limited. At Carpish it is quite 
common in the forests on both sides of the divide, from the summit at approximately 
3,000m. down to about 2,300m. on the eastern side and down to where forest gives 
way to cultivation at about 2,000m. on the western side. It is also known from similar 
country near Chaglla, 30 kms. further south. 

In the upper Huallaga basin the species occurs in a much drier habitat, and has been 
shot at Yanacocha, on the Rio Quere [sic=Quero], 25 kms. northwest [sic=SW] of 
Huanuco; at Pamacucho; and in a number of small patches of woodland between 
Huanuco and Ambo which lie at 2,700 to 3,300m., are no more than a few square 
kilometers in size, and are separated from each other by many miles of arid hillside. It 
is also reported to occur in an isolated patch of forest at 3,300m., 40 kn\s. from 
Huanuco on the Quivilla road. 

I believe Pudu mqjhistophiles also to occur in parts of the Department of Junin, 
although I have seen no specimen from that region. Local hunters, however, have 
unhesitatingly identified my Carpish specimen as the same animal as one known 
locally as "Antagllo" which is reported to be very plentiful at elevations of from 2,000 
to 3,d00m. on the forested heights above San Ramon and also in the valley of the Rio 
Hausanhausi [sic], to the north. The antagllo is only doubtfully known in the 
Oxapampa area, but is reliably reported from the upper Rio Paucartambo at approxi- 
mately 76.00'W, 10.50'S. 

The Peruvian population of Pudu mephistophiles appears to be entirely isolated from 
the nearest known population in Ecuador, since no small deer of any kind is known in 
any area north of Huanuco where 1 have made enquiries. 

The specimen from Carpish mentioned above was shipped to the Field 
Museum but never arrived (Grimwood 1969, p. 80). To date, the only preser\'ed 
specimen of a Peruvian pudu known to me is the pelt from Bosque Zapatogocha, 
Huanuco, presented to the Field Museum by Peter Hocking. 

Early reports of the occurrence of pudus in southern Peru and Bolivia prove to 
have been based on pelts or bones of the dwarf brocket, M. chunyi (Hershkovitz, 
1959a). 



Conservation 

The late Professor Lehmann warned in 1960 (p. 282) that the pudu was being 
hunted almost daily in the Colombian highlands by campesinos and often by 
villagers who sometimes killed as many as six or seven deer in a single organized 
hunt. At this rate of attrition, he argued, the pudu's extinction was foreseeable 
in the near future. Lehmann added that account must be taken of the limited 
range of the species (as known then) and of the increasingly large portions that 
man was converting to his own use. This alone, he declared, is enough to doom 
the pudu unless the government provides the vigilance and control needed to 



HERSHKOVITZ: NEOTROPICAL DEER 49 

ensure survival of the natural fauna and flora by establishment of appropriate 
nature reserves. 

The Colombian government has since (1968) founded the Purace National 
Park and others. Pudus of the Purace region, the first known from Colombia, 
served Lehmann for the original description of his Pudu mephistophiles wetmorei. 
Oddly, Hunsaker (1972, p. 445) mentions the mountain tapir {Tapirus punchaque) 
but overlooks the pudu among the protected species of this park. 

Present status of P. mqthistophiles in Colombian National Parks has been 
summarized by Dr. Hernando Chirivi Gallego (see above, p. 47). The informa- 
tion is abstracted below with some comments. For location of parks listed, see 
"Gazetteer" (p. 80) and map (fig. 30, p. 46). Additional information on Colom- 
bian national parks is provided by Hunsaker (1972) and Harroy (1972). 

LOS NEVADOS, Cordillera Central, astride boundaries between departments of Caldas, 
Resaralda, Quindio, and Tolima; 4°45'N, 75°20'W; 50,000 ha (200 sq miles); established 
1977. Not plotted on map. Figure 30. 

"Local informants distinguish a small reddish deer whose description agrees with that of 
Mazama rufina, and a smaller dark one like Pudu mephistophiles" (Chirivi"). 
This park lies within the range of Mazama rufina, but well north of the northernmost 
authenticated locality recorded for the northern pudu. 

LAS HERMOSAS, Cordillera Central, departments of Valle and Tolima, established 1977. 
"A mounted specimen of Pudu mephistophiles is in the Hunters Club of Ibague, Tolima. H 
was taken in the pdramo above Roncesvalles, Tolima" (ChirivQ. 

Lehmann (1945, p. 76) also recorded the pudu from the Valle side of the Paramo de las 
Hermosas. 

NEVADO DEL HUILA, departments of Huila, Tolima, and Cauca. 

"There is no information but in all probability the pudu is present" (ChirivO- 

PURACfi, Cordillera Central, Cauca; 80,000 ha (293 sq miles), 2,500-4,700 m; established 
1968. 

"A specimen (skin and skull) from Laguna de Cusiyaco near the headwaters of Rio 
Magdalena, is preserved in the Instituto de Ciendas Naturales, Universidad Nadonal de 
Colombia, Bogota" (ChirivO. 

MUNCHIQUE, Cordillera Central, Cauca. 

"Dr. Heliodoro Sanchez, Chief, Division of National Parks, INDERENA, observed a 
pudu from a distance of a few meters on the western slope of Cerro de Munchique, 
between 2,500-2,600 m" (Chirivi). 

CUEVA DE LOS GUACHAROS, Cordillera Oriental, Huila, 15,000 ha (61 sq miles), 
1,700-4,000 m. 

"There are vague rumors of the existence of pudus in the highest mountains surround- 
ing the Park" (ChirivQ. 

Pudu hunting in Colombia, Dr. Chirivi informs me, has been prohibited since 
1969. However, without the necessary vigilance in the mostly unsupervised 
parks, survival of the species, where presently known to occur, is dubious. 

In Ecuador, the Parque Nacional de Cotopaxi harbors pudus. The pair cap- 
tured west of Salcedo and exhibited in the Berlin zoo (fig. 28) must have origi- 
nated in this park. There may be more havens. Dr. Fernando Ortiz Crespo 
informs me (see above, p. 47) that the Ecuadorian Ministerio de Agricultura and 
Ganaderia plans to establish nature reserves that include pudu habitats. Among 
areas under consideration are the slopes of Mt. Cotacachi, Imbabura Province; 
the eastern flanks of Mt. Tungurahua, Tungurahua Province; Mts. Altar and 
Sangay in Morona-Santiago; and the eastern slope of the Cordillera Oriental at 



50 FIELDIANA: ZOOLOGY 

the latitude of Mt. Cayambe, Napo, which would incorporate the type locality of 
Pudu miyhistophiles. Dr. Ortiz adds that conservationists can speed up the estab- 
lishment of such reserves by exerting pressure on the highest governmental 
levels. 

I.R. Grimwood, Technical Adviser on Wild Life for the British Ministry of 
Overseas Development, on assignment to the Peruvian Servido Forestal y de 
Caza, is equally urgent in his assessment of the situation in Peru. He (1969, 
p. 80) gave this observation: 

It is improbable that this species will survive for much longer in the isolated forest 
patches in which it occurs in the upper Huallaga basin, where it is relentlessly hunted 
with dogs and guns. It has, however, outlasted the white-tailed deer in many of them 
because it does not break cover, as does that species. . . . The species is also hunted in 
the Carpish, Chaglla and San Ramon areas, but in these regions destruction of habitat 
for settlement is likely to present a greater threat. Control of hunting is, however, 
necessary. 

Pudu mephistophiles does not occur in any existing or projected national park or 
reserve [cf., Dourojeanni, 1968]. The creation of a special sanctuary, for which the 
high ground above the San Ramon gorge is suggested, is therefore desirable. 

In a personal communication dated 1 April 1979, Dr. Antonio Brack expresses 
doubt that the pudu ever occurred in the San Ramon region. In any event, he 
adds, the original environment has been destroyed and the region now is un- 
suited for a reserve. Brack further states that the only area where pudus are 
known to occur in abundance is in the Cordillera de Carpish, department of 
Huanuco, where the government plans to establish a reserve when funds and 
national conditions permit. In Brack's opinion, Pudu mephistophiles in Peru is not 
seriously threatened, as long as good habitat over an extensive zone of the 
eastern slope of the Andes remains closed to highways. 

A national park worth mention because it may shelter pudus in an area where 
never recorded is that of San Andres de Cutervo (6°13'S, 78°42'W), Cajamarca, 
altitude, 2,200-3,000 m. It is the smallest of Peru's national parks (2,500 ha). 
Harroy (1972, p. 77) describes it as ". . . totally protected: no occupation, no 
exploitation; lumbering is now forbidden almost everywhere. Staff headquarters 
at the town of San Andres de Cutervo, 3 km from the Park." 

Vernacular Names 

The Colombian pudu, according to Lehmann (1945), is called venado conejo 
(rabbit deer). Those with antlers in velvet may be distinguished as caciques 
(chiefs). Melanic individuals of the Valle de las Papas, Cauca, are recognized as a 
distinct variety called venado chonta (chonta is a species of palm with dark trunk). 
On the eastern slope of the Paramo de las Papas, Huila, where I worked, venado 
chonta is the name applied to the small, red mountain brocket, Mazama rufina; 
whereas the sympatric pudu is called venado churucu (or churuco, also used for 
the blackish, woolly monkey, Lagothrix lagothricha lugens). 

The most widely used name for the pudu in Ecuador and Peru is sacha cobra 
(wdld goat). The notation "local name ollique" appears on the collector's tag of a 
specimen from Cayambe, Ecuador, preserved in the British Museum (Natural 
History). Grimwood (1969) includes fl«teg//o as the name used in Junin, Peru. 

The Chilean name "pudu" now is universal for both recognized spedes of the 
genus Pudu. I follow Haltenorth (1963, p. 48) in distinguishing the two as 
"northern pudu" and "southern pudu." 



HERSHKOVITZ: NEOTROPICAL DEER 51 

Taxonomy 

The original description of Pudu mephistophiles de Winton is based on a skin 
with foot bones intact, and the skull of a subadult brocket, apparently Mazama 
rufina Pucheran. The skin and mismatched skull were received from Mr. Ludovic 
Soderstrom, British consul in Quito, Ecuador. De Winton (1896, p. 511) exam- 
ined the foot bones still in the skin and observed that "Osteologically they agree 
with P. humulis [=Pudu puda], the ectocuneiform and navicular-cuboid bones 
being all in one." 

The erroneous association of the type skin and foot bones with a Mazama skull 
was discovered by Thomas (1908) when he received two additional specimens 
from Mr. Soderstrom, both with properly matched skins and skulls. 

The three known skins of P. mephistophiles from the type region in northern 
Ecuador average slightiy more saturate in general body color than southern 
Colombian representatives of wetmorei, and one of them agrees with the 
holotype of wetmorei as described. A skin from the Cordillera de los Llanganates, 
south of the type region of mephistophiles, with a buffy or olivaceous-appearing 
coat is absolutely indistinguishable from the majority of Colombian specimens I 
examined, including four topotypes of wetmorei. An adult male and female, also 
from Llanganates, reported by Fradrich (1975, p. 75) are said to "differ insignifi- 
cantly from the more northern wetmorei. " In the absence of any notable difference 
in color or any other character including antier shape, which varies even more 
widely, there appear to be no grounds for recognition of wetmorei as a race of 
P. mephistophiles. 

Characters 

Size. — Smallest of living deer; measurements are given in Table 3. 

Coloration and pelage (figs. 31-34). — General body color buffy agouti (oliva- 
ceous) to reddish brown agouti, the middorsum dominantiy dark brown, pelage 
stiff, adpressed, the hairs pithy; cover hairs of middorsum about 35 mm long, 
guard hairs like cover hairs, but about 5 mm longer, basal half of both types pale 
brown or grayish brown, terminal half dark brown (eumelanin) with buffy to 
reddish (pheomelanin) subterminal band 0.5 mm wide; fine brown curly wool 
hairs thinly spread and completely hidden to view by cover hairs. Head, in- 
cluding muzzle, chin, outer surface of ears, contrastingly colored dark brown or 
blackish (eumelanin), the hairs generally unhanded, specialized markings ab- 
sent; inner surface of ears sharply defined whitish, grayish, or pale buff; rostral 
hairs banded, usually with a single, narrow pheomelanic ring; orbital ring or 
stripe absent; neck, including nape, dominantiy buffy, the individual hairs with 
broad pheomelanic bands. Belly, inner sides of thighs and arms buffy to rufous, 
the hairs unhanded; forefeet and hind feet dark brown contrasting with buffy to 
reddish agouti of flanks; hoofs extremely long and narrow. 

Fawns unspotted, pelage coarse, less hispid, more relaxed, color pattern more 
uniform, without markedly contrasting dark head and legs. 

Color variation. — General body color of the holotype is reddish brown; back of 
head, nape, and shoulders more or less like back, rump more saturate than 
middle of back. Of two juvenals in the British Museum (Natural History), one, a 
topotype from Papallacta, is slightiy more saturate throughout than the 
holotype; the other, from Mt. Cayambe in type region, is slightiy paler. 



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HERSHKOVITZ: NEOTROPICAL DEER 

PELAGE 



53 




pelage 
upper mld-dorsum 



wool 



guard cover 
Fig. 31. Pelage and hair types plucked from middorsum of Pudu mqjhistophiles. 



The 23 southern Colombian specimens examined, all from type region of Pudu 
mqjhistophiles wetmorei Lehmann, vary from reddish brown to olivaceous buff or 
ochraceous in general body color; coats of individuals taken in May, June, Sep- 
tember through February appear new or slightly worn. 

In his description of P. mqjhistophiles wetmorei from southern Colombia, 
Lehmann (1945, p. 78) noted that bright reddish, as well as nearly black, indi- 
viduals had been observed in the savannas of Paletara, and that he had seen a 
half dozen or so pudus as dark as the holotype. A female from the Paramo de 
Guanacas, described and figured by Lehmann, is a partial albino with whitish 
head, throat, nape, and shoulder bands, and a sprinkling of whitish hairs over 
the remainder of the otherwise uniformly brownish body. Lehmann (1959) also 
mentioned that completely albino pudus and albino Andean brockets, Mazama 
rufina, had been taken in the same region and in contiguous Paramo de las 
Delidas to the north. 



54 



FIELDIANA: ZOOLOGY 




Fig. 32. Longitudinal section of hair from Pudii mqjhistophilcs , n\idback; scanning elec- 
tron microscopic views at magnifications given. Hair structure is not discussed in Part I. A, 
cuticle showing irregular crenate pattern of overlapping scales; x 1,CXX). B, same as A, 
X 2,000. C cuticle and air pockets of cortex, x 700. D, cortex, x 700. 



The single Peruvian specimen at hand, from 10,000 ft in Bosque Zapatogocha, 
Carpish Divide, Huanuco, is juvenal or subadult. Head and body length of 790 
mm (tail 50, hind foot 260, ear 60), recorded on the field label indicates an adult, 
but the same measured on the tanned skin is not more than 700 mm. The 
comparatively long and lax pelage agrees most nearly with that of a juvenal from 
Malvasa, Colombia, with head and body length about 640 mm measured on the 
tanned skin. However, the Peruvian specimen differs from the Malvasa sample 
and all other FMNH specimens from Colombia, plus one ANSP skin from 
Ecuador, by its generally saturate brown color and lack of contrast between its 



HERSHKOVITZ: NEOTROPICAL DEER 

SATURATION PROCESS 



55 



•PHEOMELANIN PATHWAY 
red-yellowH 



EUMELANIN PATHWAY - 
^^■black-brown 




saturate saturate 
monochromic dichromic 
pheomelani n 



modified AGOUTI modified 

agouti (with agouti 

eume1 anin 

base] 



saturate 

monochromic 

eumelanln 



Fig. 33. Saturation, the evolutionary pathway by which mammals have diverged in 
coloration from a hypothetically common, dominantly agouti -colored ancestor. This pro- 
cess terminates in either uniformly eumelanin (blackish) hairs, as in Pudu mephistop'hiles, or 
pheomelanin (reddish) hairs, as in Pudu puda. For source of figures and evolutionary 
account of mammalian tegumentary color patterns, see Hershkovitz, 1977, p. 92. 



dark browm back, rump, and outer sides of forelegs and hind legs. This recently 
accessioned individual has not been compared with the dark pudus from 
Ecuador now in the British Museum, or with those recorded by Lehmann from 
southern Colombia. The Peruvian specimen may represent a dark race separable 
from the typical Ecuadorian form. 

Rhinarium. — Somewhat bulbous, with posterodorsal border projecting 
backward along midline. 

Tegumentary glands (figs. 10, 11, 14). — Preorbital gland in preserved or tanned 
skin is represented by an inconspicuous pouch opening about Viz cm in diameter. 
Forefeet and hind feet pedal glands are contained within shallow, interdigital, 
trough-like skin fold; tarsal and metatarsal glands, if present, are not evident on 
dry or tanned skin. 

Hoofs and toes (fig. 10). — Hoofs are comparatively long and narrow, greatest 
length being about three to five times the width measured on plantar surface. 
Front toes, including hoofs, are comparatively widespread, as seen in dry skin 
and noted in the living animal (Fradrich, 1975, p. 72) and presumably are en- 
dowed with unusual power of abduction. , ' 



56 



FIELDIANA: ZOOLOGY 
PUDU PUDA PUDU MEPHISTOPHILES 



CROWN 



EAR 



MUZZLE 



MID-BACK 



UPPER ARM 



CHIN 



CHEST 



BELLY 



HIP 



LEG 



FOOT 



ir:.>tfA.i.-!V.Uk J U. ...'....iT^ 



■^^:.':.;-W'i.v,'"r"j;,4^»jjr 



n~] 



pheotnelanin 



slightly or not pigmented 



Fig. 34. Randomly selected hairs plucked from 11 chromogenetic fields of preserved 
skins of Pudu piida and P. mqjhistophiles. 



Skull (figs. 11-17). — Lacrimal fossa shallow or dish-shaped without fenes- 
trations, its ventral border hardly or not at all encroaching on malar and maxil- 
lary bones; superior border of ascending ramus of premaxillary expanded, con- 
tacting lateral margin of nasals; anterodorsal border of lacrimal flat; long axis of 
antorbital vacuity nearly or quite vertical; plane area of vacuity generally much 
less than one-half, rarely slightly more than plane area of facial plate of lacrimal; 
perforations on upper surface of zygomatic process of squamosal, if present, 
usually less than 1 mm in diameter each; angle of mandible produced well 
beyond posterior plane of condyle as an expanded subrectangular process; lunar 
notch between angle and condyle, deeply excised, concavity between base of 
angle and ramus well defined. 



HERSHKOVITZ: NEOTROPICAL DEER 57 

Cranial measurements are given in Table 3. 

Dentition. — Upper permanent canine persistent in eight of 15 skulls with com- 
plete permanent dentition; deciduous canine present in two of three young with 
only first molars erupted, and in two of three with second molars erupted; 
metaloph and metaconule present, postentoconule absent in fully erupted mo- 
lars (fig. 23). 

The form of left pm^ in FMNH 24362 (fig. 16) is that of a deciduous tooth. In 
size and wear, however, it agrees with the upper third premolar of the right side 
and of other pudus. There is no sign of a replacement tooth. Ostensibly, the 
deciduous -like premolar is the permanent tooth. 

Antlers. — Adult male skulls examined were collected October through Feb- 
ruary and one in July in the department of Cauca, Colombia. All support 
polished antlers, the longest of which exceeds 90 mm (table 3). Antlers of the 
holotype of Pudu mephistophiles wetmorei, collected February 1945, are also 
polished (Lehmann, 1960, p. 283). The adult male of a pair of Ecuadorean pudus 
that arrived in the Berlin zoo during April rubbed the velvet from its antlers in 
June (Fradrich 1975, p. 76). 

The foregoing suggest that antlers of adult male pudus in southern Colombia 
and northern Ecuador may be shed from February into March, are in velvet from 
March into June and possibly July, and are polished from June or July until cast. 

I find no information on antlers of Peruvian representatives of the spedes. 

A skull of a three-horned pudu from Paramo de Guanacas, Cauca, Colombia, 
without date of collection, is figured by Lehmann (1959, p. 204). The antlers, still 
in velvet, are said to be fully developed and completely separated. Two antlers 
occupy normal positions and in the photograph appear to be more or less sym- 
metrical. The third antler, slightly shorter than the others, arises directly to the 
right of the normal right antler. 

Vertebral formula. — In a male (FMNH 86846) from Malvasa, Cauca, Colombia: 7 
cervical, 13 thoracic, 6 lumbar, 4 sacral, 8 caudal. 



Comparisons 

Pudu mephistophiles is distinguis|ied from P. puda, its only congener, by char- 
acters already mentioned under generic headings (p. 15), key to the spedes (p. 
43), and in the description of the spedes (above and pp. 65, 70). The only other 
deer with which the northern pudu need be compared is the smallest known 
brocket, Mazama chunyi. 

Using greatest skull length as most reliable indicator of overall body size in 
present material, that of one fully adult Mazama chunyi (all available) is 146 mm, 
whereas mean and extremes of 14 adult Pudu mephistophiles are 132.5 mm (127- 
142). Externally, as represented by four skins, including holotype, M. chunyi 
more nearly resembles pale Colombian representatives of P. mephistophiles than 
the saturate Peruvian individual at hand. The brocket is separable by its white 
narial (trimmed away in holotype) and mental patches, tail distinct from rump, 
the underside sharply defined white, ears probably similarly short and rounded 
but edges white, their outer surface brownish like muzzle and crown, not black- 
ish, coronal hairs not whorled, body color dominantly pheomelanin, not 
eumelanin or agouti, lower portion of legs not markedly different in color from 
upper legs or flanks, and fawn spotted. 



58 FIELDIANA: ZOOLCX^Y 

In Mazama chunyi, as in all brockets, external cuneiform bone is present as 
a discrete tarsal bone, and crown of first permanent incisor is broadly spatulate, 
its greatest length more than combined length of iz-a- For additional characters, 
see comparison between Pudu and Mazama (p. 22). 

Specimens Examined 

Total 29. COLOMBIA. Cauca: Paramo de Purace, 4 (USNM, 1; FMNH, 1; 
AMNH, 2); Malvasa, 15 (FMNH); Quintana, 1 (FMNH); La Vega, 1 (FMNH); 
Paletara, 2 (AMNH); Coconuco, 1 (AMNH). ECUADOR. Pichincha: Paramo de 
Papallacta, 2 including holotype (BM); Mt. Cayambe, 1 (BM); Tungurahua: Cor- 
dillera de los Llanganates, 1 (ANSP). PERU. Hudnuco: Bosque de Zapatogocha 
above Acomayo, 1 (FMNH). 

V. BIOLOGY OF Pudu mephistophiles 
Habitat 

In tracking the northern pudu, I found the animal confined to the central 
Andean forests in southern Colombia and the eastern Andean forests in north- 
ern Ecuador, at altitudes between 2,700 and 4,000 m. The severely acddented 
terrain where the animal still may be found probably is its last retreat rather than 
the predilect abode of this persistently persecuted little animal. 

The southern Colombian habitat is described by Lehmann (1945, p. 78) as the 
dense brush or matorrales of the paramos and the sedge and frailejdn (Espeletia) 
covered swamps of the Andean summits. I have never seen signs of pudus 
where grasses or frailejdn dominate. However, Lehmann points out that pudus 
sometimes attempt to escape pursuing dogs by climbing trunks of trees inclined 
over streams or bluffs, which occur only in the temperate zone forests below true 
paramo. In any event, pudus may invade borders of interdigitating grasslands or 
paramo to eat the succulent herbs and shoots that appear with the first rains 
after the dry season burnings. 

The foUov^ng is Grimwood's comparison (1969, pp. 77) of the habitat oiPudu 
mephistophiles in central Peru with that of the very similar-appearing pygmy 
brocket, Mazama chunyi of southern Peru: 

The two spades occupy the same type of habitat, avoiding the cold and damp of the 
highest woods and of the thickets above tree line, and [are] found on the eastern 
[Andean] slopes only from about 3,200 m downwards, in regions where [the alder] 
Alnus jorulensis is often the dominant tree. Both spedes are also known from a rather 
different, drier habitat, in isolated patches of forest or woodland in the high valleys 
immediately to the west of the main eastern watershed. 

Habits 

Little is known of the behavior of the northern pudu in the wild beyond what 
can be inferred from its habitat (see preceding section) and gross morphology. 
Grimwood (1%9, p. 77) compared the pudu with the very similar Mazama chunyi 
and noted that, "Both are very small nocturnal deer, found only singly or in 
pairs, which emerge in clearings or fields at dusk. Both are much given to 
whistling and both are said often to be dazed if disturbed by day, so that they 
can sometimes be caught by hand." 

Lehmann (1945, p. 78) observed that, when pursued by dogs, the pudu runs 
in a weaving pattern. Grimwood also noticed that, in flight, the animal does not 



HERSHKOVITZ: NEOTROPICAL DEER 59 

break cover, but circles in the thickets and among the cliffs. Its long, fine, 
widespread hoofs facilitate climbing, and, as previously noted (Lehmann, 1945, 
p. 76), the pudu sometimes escapes dogs by running or climbing up inclined 
trees. Grimwood believes, on the basis of information received from hunters, 
that the little deer is as nimble on rocks as the African klipspringer (Oreotragus). 

Maintenance in Captivity 

Northern pudus are frequently captured alive, but, according to Lehmann 
(1945, p. 78), they do not survive long in captivity. One pudu in Lehmann's 
possession in Cali, Colombia, became very tame and particularly fond of milk, 
but died in two months. Lehmann adds that a friend, Guillermo Valencia, con- 
fessed his inability to keep captured pudus alive for more than a few weeks 
despite all efforts, including maintenance in a natural environment. 

The first northern pudus received in a European zoo were two females from 
Ecuador that arrived dead or dying 23 November 1968, at the Berlin zoological 
garden. This report, by Fradrich (1975, p. 73), includes a notice without particu- 
lars of the later accession in 1970 of a northern pudu by the same zoo. Sub- 
sequently, Fradrich continues, a pair captured in Ecuador, west of Salcedo be- 
tween Mt. Cotopaxi and the Cordillera de los Llanganates, arrived at th e Berlin 
zoo in good condition 12 April 1972. After the usual quarantine, the pair was 
placed on public view and fed a standard brocket diet (Fradrich, 1974). The male 
died 6 August 1972. The autopsy showed pasteurellosis, tracheobronchitis, and 
fibrinous pneumonia. The female died of food poisoning 11 October 1972. 

I find no other published records of Pudu mephistophiles in zoological gardens. 

Ungulate Associations 

Cervids occurring within parts of the range of Pudu mephistophiles are the 
brockets, Mazama rufina and M. americana, white-tailed deer, Odocoileus vir- 
ginianus, and taruga, Hippocamelus antisensis (fig. 5). The latter, known from the 
Andes of Ecuador, Peru, Bolivia, and northern Argentina is nearly, if not al- 
ready, extinct in Ecuador and Peru. The pinchaque or woolly or mountain tapir, 
Tapirus pinchaque, associates with the pudu in Colombia and Ecuador; its num- 
bers are dwindling. 

The dwarf deer, Mazama chunyi, is common in the southern Peruvian depart- 
ments of Cuzco, Puno, and Madre de Dios in habitats appropriate for Pudu 
mephistophiles, but the latter is unknown south of the department of Junin. 
Information on the distribution of small deer in southern Peru is meager, and 
identifications often have been equivocal or based on hearsay. The pudu, com- 
mon as it appears to be in central Peru, was not reliably reported from that 
country until 1969 (Grimwood, p. 79), and the first museum-preserved speci- 
men is recorded here. M. chunyi, long confused with P. mephistophiles, was not 
recognized as a distinct spedes until 1959. The two little deer may yet prove to be 
sympatric. 

Reproduction 

A female from Ecuador that arrived at the Berlin zoo 12 April 1972 died 11 
October 1972 carrying a fully developed female fetus (Fradrich, 1975, p. 76). 
Grimwood (1969, p. 80) reports that, in central Peru, nearly full-term fetuses 
were taken from pudus killed in April and November. 



60 FIELDIANA: ZOOLOGY 

The meager data indicate that, in Pudu mephistophiles, a gestation period of 
seven months is followed by a postpartum estrus as in Pudu puda. 

VI. Pudu puda MOLINA 

SOUTHERN PUDU (figs. 36, 37) 

Synonymic History 

Capra puda Molina, 1782, Saggia storia Nat. Chile, libr., 4: 308, 343. 

Capra puda, Anonymous ("An American Gentleman"), 1808, The geographical, natural and 
civil history of Chile, 1: 217, 240 — "Translation from the original Italian of Abbe don J. 
Ignatius Molina, Saggia . . . ." Printed for J. Riley, Middletown, Conn. 

[Ovis] Pudu [sic], Gmelin, 1788, Linnaeus Syst. Nat., ed. 13, 1: 201 — classification. 

Ovis Pudu, Shaw, 1800, Gen. Zool., 2(2): 392. 

[Aries]pudu, Fischer, 1814, Zoognosia Tab. Syn., 3: xvi, 406. 

A[ntelope]. Pudu, Blainville, 1816, Bull. Soc. Philomatique, pp. 76, 80 — classification. 

M[azama]. pudu, Rafinesque, 1817, American Monthly Mag., 1: 437 — regarded as a goat 
antelope. 

Mazama pudu, Vanoli, 1967, Saugetierk. Mitteil., 15(2): 155, figs 1-10 (adults, young) — 
CHILE: Osorno-Llanquihue; habits, habitat, reproduction; food; growth; development- 
antlers; enemies; disease; conservation. 

Mazama [{Oreamnos)] puda, Rafinesque, 1817, American Monthly Mag., 2: 44 — 
classification. 

M{azama (Pudu)]. pudu, Haltenorth, 1963, Handb. Zool., 8(32): 48— classification. 

Antilocapra pudu, Lesson, 1842, Nouv. Tabl. Reg. Animal, Mamm., p. 182 — classification. 

Cer^MS pudu. Gay and Gervais, 1846, Ann. Scd. Nat., (3)5: 88 — CHILE: classification; de- 
scription; measurements; Cervus humilis Bennett, a synonym. Gay, 1847, Hist. Chile, 
Zool., 1: 158, Atlas, pi. 9 (animal), pi. 1 (skull) — CHILE: from Cauquenes toChiloe; Indian 
name, pudu or puudu. Pucheran, 1852, Arch. Mus. Hist. Nat. Paris, 6: 482 — characters; 
measurements; history. Sclater, 1866, Proc. Zool. Soc. London, 1866: 104, fig. 
(animal) — menagerie specimen. Sdater, 1871, Proc, Zool. Soc. London, 1871: 238, pi. 17 
(animal) — menagerie specimen; antler shedding. Garrod, 1877, Proc. Zool. Soc. Lon- 
don, 1877: 4, 13, 14 — alimentary tract; penis; cranial characters; classification. Huet, 
1890, Bull. Soc. Nat. Acclim. France, 1888/1889, p. 86— coloration. 

C[ervus]. pudu, Wagner, 1855, Schreber's Saugth., Suppl., 5: 390, pi. 26 (animal) — 
characters; humilis Bennett a synonym. 

Coassus [(Pudu)] Pudu Gray, 1850, Gleanings Knowlsely Menag., p. 69 — characters. Gray, 
1852, Proc. Zool. Soc. London, 1850: 242 — CHILE: Concepcion; Chiloe; classification; 
characters; part synonymy: Capra pudu Molina, Cen'ws pudu Gervais, Cervus humilis 
Bennett. 

Pudu pudu, Sdater, 1870, Proc. Zool. Soc. London, 1870: 116 — dassification. Wolffsohn 
and Porter, 1908, Rev. Chilena Hist. Nat., 12: 74 — CHILE: Valdii'ia; local names, venado, 
venadito, pudu (Araucano). Pocock, 1911, Proc. Zool. Soc. London, 1910: 967, 978 — 
interdigital gland. Lydekker, 1915, Cat. Ungulate Mammals Brit. Mus., 4: 215 — 
characters; synonymy; CHILE: Maule; Constitucion; Concepcidn (Concepdon); Cautin 
(Temuco). Synonyms, humilis Bennett, chilensis Gray. Pocock, 1923, Proc. Zool. Soc. 
London, 1923: 201, fig. 13 (foot gland), fig. 14fc (rhinarium), fig. 15 (ear) — external 
characters. Wolffsohn, 1923, Rev. Chilena Hist. Nat., 27: 160— CHILE: Concepcidn (Pedro 
Valdivia); Maule (Constitudon); measurements. Yepes, 1929, Rev. Chilena Hist. Nat., 
33: 472 — CHILE: Maule (Constitudon) to Magallanes; ARGENTINA: western Neuquen to 
western Santa Cruz. Junge, 1932, Zool. Garten, 5: 249 — sensitivity to heat. Cabrera and 
Yepes, 1940, Mamiferos Sud- Americanos. Hist. Nat. Ediar (Buenos Aires), p. 278, pl. 88 
(animal); characters; distribution. Yepes, 1943, Rev. Argentina Zoogeogr., 3: 75 — 
ARGENTINA: Neuquen (Cordillera del Trompul, San Martin de Los Andes; Brazo Co- 
rrentoso, south of Cordillera del Trompul; La Lolog, San Martin de Los Andes); local 
name, corzuelo. Osgood, 1943, Field Mus. Nat. Hist., Zool. Ser., 30: 228— CHILE: 
Magallanes (Isla Riesco; Rio Baker): Llanquihue (Cayetue [sic], Lago Todos Santos; Pe- 
trohue, Lago Todos Santos); Chiloe; Cautin (Pitrufquen); Concepcidn (San Pedro); Arauco 
(Santa Juana; Vega Blanca, Sierra Nahuelbuta); Valdivia (Valdivia); characters; variation; 



HERSHKOVITZ: NEOTROPICAL DEER * 61 

habits. Schmidt, 1944, Argentinische Sdugetiere. (Buenos Aires); pp. 170, 237, 5 figs, 
(adults and young), 1 fig. (skull) — ARGENTINA: Rio Negro (Lago MascardQ; Neuqu^ 
(Rio Quillen). Flerov, 1952, Fauna U.S.S.R., n.s., no. 56, Mammals, 1(2): 220 (in text)— 
listed. Junge, 1954, Bol. Jardin Zool., Concepdon (Chile), 1: 3, fig. (animal) — two 
females in captivity. Hershkovitz, 1958, Mammalia, 22(4): 545 — metatarsal glands and 
tuft not defined. Mochi and Carter, 1953. Hoofed mammals of the world, Scribner's and 
Sons, Ltd., London, pi. VIII, fig. 16 (animal) — characters. Hershkovitz, 1959, Proc. Biol. 
Soc. Washington, 72: 45, pis. 3D, 4B (skull) — comparisons. Walker, 1964, Mammals of the 
world, Johns Hopkins Press, Baltimore, 2:1399, fig. (animal)— CHILE; "BOLIVIA." 
Crandall, 1964, The management of wild animals in captivity. U. of Chicago Press, pp. 557, 
560, 594 — characters; exhibition in zoos. Greer, 1965, Publ. Mus. Midhigan St. U. Biol. 
Ser., 3(2): 146— CHILE: Malleco (Laguna Malleco; Galvarino, 10 km W, 61 m); Angol 
(Cordillera de Nahuelbuta); measurements. Junge, 1966, Int. Zoo Yearb. 6: 263 — CHILE: 
Angol; Malleco; Valdivia (Mariquina); habits in captivity. Windecker, 1967, Fr. Kolner Zoo, 
10(2): 58, 4 figs. incl. 2 col. (animals) — Cologne zoo; characters of adult and young; 
gestation; antler shedding; diet in captivity; hunting in wild. Pefaur, Hermosilla, Di 
Castri, Gonzalez and Salinas, 1968, Rev. Soc. Med. Vet. (Chile), 18(1-4): 4, 5, 6, 7, 8, 11, 
14 — CHILE: Curico (?); Nuble; Concepdon; Arauco; Bio-Bio; Malleco; Cautin; Valdivia; Osomo; 
Llanquihue; Chiloe; Aysen; Magallanes; distribution; disease (hoof-and-mouth); en- 
dangered spedes. Hick, 1968, Fr. Kolner Zoo, 4(10): 111, 16 photographs, 2 in color 
(female and young) — gestation, birth and chronology of growth and behavior in Co- 
logne zoo. Hick, 1969, Int. Zoo Yearb., 9: 110— CHILE: Llanquihue (Lago 
Llanquihue) — mating; antler growth and shedding; molt; birth, growth, behavior of 
young in Cologne zoo. Kohls, 1969, J. Med. Entomol., 6(3): 280— CHILE: Valdivia (Santo 
Domingo, 10 km SE Valdivia; Callumapu, 15 km W. Valdivia); ticks (Acarina): Ixodes 
taglei, I. stilesi. Hick, 1979, Fr. Kolner Zoo, 4(10): 113, fig. (week old young)— birth in 
Cologne zoo. Hick, 1971, Z. Kolner Zoo, 14(2); fig. p. 83 (mother v^ath F2 young S) — 
breeding in Cologne zoo. Dansie and Wince, 1971, Deer of the world. Hitchings and 
Mason, Ltd., Plymouth, England, pp. 21, 23, fig. (animal), fig. (map, distribution) — 
CHILE. Wetterberg, 1972, Oryx 11(5): 347, fig. (animal) fig. (map, Parque Nadonal 
Vicente Perez Rosales) — CHILE: Llanquihue (Parque Nadonal Vicente Perez Rosales); 
ARGENTINA: Neuquen (Parque Nahuel HuapO; habitat; conservation; food; habits; pre- 
dators. Koulischer, Tyskens and Mortelmans, 1972, Acta Zool. Path. Antwerp, 56: 25, 
28, fig. 2 (karyotype) — karyotype (2n=70; FM=74). Whitehead, 1972, Deer of the world. 
Viking Press, New York, pp. 29, 64, 65, 169, map 9 (distribution), figs. 24a, b (animal) — 
characters; distribution. Avila Pires, 1974, Publ. Biol. Inst. Invest. Cient. U.A.N.L., 
Mexico, 1: 61 — distribution; conservation. Spotorno and Fernandez-Donoso, 1975, 
Mamm. Chrom. Newsl., 16(1): 17, fig. 10 (karyotypes) — CHILE: Llanquihue (Llan- 
quihue); karyotype (2n=70, FM=74); weights. Fradrich, 1975, Zool. Gart., 45(1): 67, figs. 
1, 2 (animal), fig. 3 (mother with fawn), fig. 4 (fawn), fig. 8 (skull) — characters; mainte- 
nance in Berlin zoo; antiers; breeding. Czernay, 1977, Zool. Gart., 47(3/4): 226, figs. 2, 3 
(growth graph), fig. 4 (male), figs. 5, 6 (mating), fig. 7 (fawn), figs. 8, 9, 10, 11 (mother 
and young), figs. 12, 13 (family group), fig. 4 (antlers in velvet). Brambell and Mathews, 
1977, Proc. Symp. 1, Assoc. Brit. Wild Animal Keepers, p. 25 — characters, reproduction, 
management in captivity. 

P[udu]. pudu, Wolffsohn, 1927, Rev. Universitaria, Univ. Catolica Chile, p. 240— CHILE: 
hunting, domestication. Schneider, 1946, Bol. Soc. Biol. Concepdon, 21: 78 — CHILE: 
Concepdon (Nahuelbuta; Cayumanque; Manquimavida); habits. Hoffstetter, 1952, Mem. 
Soc. Geol. France, 31(66): 369— CHILE. 

Pudu puda. Flower, 1929, List of vertebrated animals Zool. Soc. London 1828-1927, p. 
325— listed. 

P[udu]. puda, Zuckerman, 1952, Proc. Zool. Soc. London, 122: 901 — births in London zoo, 
23 May 1932 to 29 May 1933. 

P[uda]. pudua [sic], Carette, 1922, Rev. Mus. la Plata, 26: 445 — characters. 

Pudu pudua, Castellanos, 1940, Rev. Chilena Hist. Nat., 44: 159— ARGENTINA: southern 
Andes near Chilean border. 

Pudua pudu, Lydekker, 1898, Deer of all lands, p. 307 — characters; distribution. Matschie, 
1899, Sitz.-Bericht Gesellsch. Naturf. Berlin, p. 130— distribution. Lydekker, 1901, Great 
and small game of Europe, western and northern Asia and America. Rowland Ward, London, 
p. 373 — characters. 



62 FIELDIANA: ZOOLOGY 

Nanelaphus Pudu, Fitzinger, 1873, Sitzb. Akad. Wiss. Wien (1873), 68(1): 361— 
classification. 

Ovis dubia Afzelius, 1815, N. Act. Reg. Soc. Sd., Upsalla, 7: 217 — new name for Copra pudu 
Molina; work not seen. 

Cewus humilis Bennett, 1831, Proc. Zool. Soc. London, 1831: 27 — CHILE: precise locality 
unknown; type female, skin originally in the collection of the London Zoological Soci- 
ety, now in the British Museum (Natural History) no. 55.12.24.248. Gay and Gervais, 
1846, Ann. Sci. Nat., Paris, ser. 3, 5: 90 — a synonym of Cen'us pudu [sic] Molina. Fraser, 
1849, Zoologica typica, pi. 14 and text — CHILE: Concepcion; Chiloe. 

C[enms]. humilis, Sundevall, 1846, K. Svenska Vet. Akad. Handl., 1844: 183— part, CHILE: 
specimen seen in Paris museum. 

Cervus [{Capreolus)] humilis, Lesson, 1842, Nouv. Tabl. Reg. Anim., p. 172 — classification. 

Pudu humilis, Gray, 1852, Cat. Mamm. Brit. Mus., Ungulata, p. 240, pi. 36, fig. 1 (skull 
captioned Pudu chilensis) — CHILE: characters; part synonymy. Gray, 1872, Cat. rumi- 
nant Mamm. Brit. Mus., p. 93. Gray, 1873, Handlist ruminant animals Brit. Mus., p. 
163— type history; CHILE. Jentink, 1887, Cat. Osteol. Mus. Pays-Bas, 9: 153— CHILE. 
Jentink, 1892, Cat. Syst. Mamm. Mus. Pays-Bas, 11: 183— CHILE. 

Pudua humilis, Garrod, 1877, Proc. Zool. Soc. London, 1877: 18 — classification. Sclater, 
1877, List Vertebrated Animals Zool. Soc. London, p. 139, fig. 18 (animal) — male ex- 
hibited live. Brooke, 1879, Proc. Zool. Soc. London, 1878: 927— CHILE. Sdater, 1883, 
List Vertebrated Animals Zool. Soc. London, p. 174, fig. 28 (animal) — 2 males, 1 female 
exhibited live. Sdater, 18%, List Vertebrated Animals Zool. Soc. London, p. 185 — 1 
male, 2 females exhibited live. De Winton, 18%, Proc. Zool. Soc. London, 18%: 507, 
figs. 1, 3 (skull); characters; comparisons. Kidd, 1900, Proc. Zool. Soc. London, 1900: 
686— hair pattern. Reiche, 1903, Anal. Mus. Nac. Chile, p. 46— CHILE: Arauco (Isla de la 
Mocha, extirpated). 

Coassus humilis, Riitimeyer, 1881, Abh. Schweiz. Palaont. Gesellsch., 8: 31, 93 — cranial 
characters; relationships; distribution. 

Pudu chilensis Gray, 1852, Cat. Mamm. Brit. Mus., Ungulata, pi. 36, fig. 1 (skull)— CHILE: 
(no predse locality); name in caption for figured skull or same spedmen denominated 
Pudu humilis in text (p. 240). Gray, 1873, Hand-list of edentate, thick-skinned and 
ruminant mammals in the British Museum, p. 163 — adult female, skull nos. 972 and 
50.11.29.6 (same figured 1852), skin of type skull received from Zoological Sodety of 
London. 

"Chevreuil," Poeppig, 1829, Ferussac, Bull. Sd. Nat. Geol., Paris, 19: 96— CHILE: high 
altitudes; size. 

"pudu," Poeppig, 1835, Reise in Chile, Peru and Amazonstr., 1: 450 — CHILE: Bio Bio 
(Anhjco). Ulmer, 1959, America's First Zoo, 11(1): 7— exhibited 1877 in Philadelphia 
Zoological Garden. 

T}/pe. — None in existence; name based on native accounts of the animaL 
Type locality. — Southern provinces of Qiile. 

Distribution 

(fig. 35) 

Forests of southern Chile and some adjacent parts of south w^estem Argentina; 
in Chile, from the province of Maule, southward nearly or quite to the Strait of 
Magellan; in Argentina, from southwestern Neuquen, southward along the An- 
dean foothills and mountain passes into southwestern Santa Cruz province; 
altitudinal range from sea level to nearly 1,000 m above. 

Conservation 

Pudus are still hunted in Chile, among other areas, as a cheap source of 
protein or as an expensive form of sport and their habitat is rapidly being de- 
stroyed, except possibly in the following national parks: 

NAHUELBUTA, Arauco-MaUeco, 1,000-1,500 m altitude, 5,414 ha (20 sq miles). Pudus 




Fig. 35. Distribution of Pudu puda Molina, based on museum spedmens and published 
19th and 20th century records (dots). For locality names, see corresponding numbers in 
"Gazetteer" (p. 80). 



63 



64 FIELDIANA: ZOOLOGY 

have been recorded from the Cordillera de Nahuelbuta, but no information is at hand 
regarding the present status of the native mammals of this park. 

TOLHUECA, Malleco, 1,500-2,000 m, 3,000 ha (12 sq miles). Although this park is within 
the geographic range of and provides a suitable habitat for pudus, it hardly is large 
enough to support more than a small population. 

VICENTE PtREZ ROSALES, Llanquihue, 135,175 ha (522 sq miles). According to Chileafi 
authorities surveyed by Harroy (1971, p. 168), hunting is permitted in the park. How- 
ever, Wetterberg (1972, p. 348) reports pudus abundant and generally unmolested in the 
area. 

VILLARICA, Cautln, 13,780 ha (53.1 sq miles); ". . . inaccessible and has few inhabitants," 
according to Harroy (1971, p. 168), and incorporates good pudu habitat. 

TORRES DEL PAINE, Magallanes, 10,000—20,000 ha. 

Three additional parks are located within the geographic range of Pudu puda, 
namely: 

LAGO SAN RAFAEL, Aisen (590,000 ha); PUYEHU£, Valdivia (75,000 ha); and 
PIRIHUEICO, Valdivia (13,950 ha). These are not regarded as conservation areas, due to 
the high number of human habitations and excessive exploitation, such as unrestricted 
or uncontrolled hunting, fishing, lumbering, stock grazing, and rapid destruction of 
forest. 

The future of the pudu in Argentina is clouded, Dennler de la Tour (1957, p. 
159) observes, "Among the arumals most in need of protection in Argentina are 
the Andean deer 'huemul,' Hippocamelus bisulcus, and the dwarf deer 'pudu/ 
Pudua pudu. These are threatened not only by 'development' and poaching, but 
by the introduction of the European red and fallow deer in the park areas." A 
decade later, Nottebohm (1967, p. 117) warned that disappearance of the 
huemul and pudu from Argentine territory is imminent. The following is a list of 
Argentine national parks, from north to south, where pudus (and huemuls) live 
now or lived in the past: 

LOS COPAHUES, northwestern Neuquen, 80,000 ha (310 sq nules). The pudu is not 

known to occur in northern Neuquen at present, if it ever did before. 
LAGUNA BLANCA, central Neuquen, 1,000-2,000 m, 11,250 ha (40 sq miles); doubtful if 

the pudu occurred in the area surrounding the lake. 
LANlN, southwestern Neuquen, 600-3,776 m, 395,000 ha (1,500 sq miles). This area 

adjoins Nahuel Huapi National Park and has the same fauna, including pudu and 

huemul. 
NAHUEL HUAPt southwestern Neuquen, 720-3,554 m, 785,000 ha (3,000 sq miles) of 

which only 60,000 ha (260 sq miles) is nature reserve, the remainder being private 

lumbering, farming, and grazing property. According to Dennler de la Tour (1957, p. 

160), "The huemul and the pudu have been almost eradicated by poaching, by the dogs 

of sheepbreeders and by the introduction of European deer." 
LOS ALERCES and ANEXO PUELO, western Chubut, 400-2,280 m, 280,000 ha (1,100 sq 

miles). Although the pudu has not been recorded here, the park lies within its probable 

geographic range and has a suitable habitat. 
PERITO FRANCISCO P. MORENO, west central Santa Cruz, 800-2,770 m, 115,000 ha 

(400 sq miles). Dennler de la Tour (1957, p. 163) includes the huemul and guanaco in the 

local fauna, but does not mention the pudu, which may occur nearby if not inside the 

park. 
LOS GLAQERES, southwestern Santa Cruz, 300-2,441 m, 670,000 ha (2,600 sq miles). 

Dennler de la Tour (1957, p. 162) reports that the "huemul and pudu are present, though 

reduced in numbers." 

Taxonomy 

Molina (1782, p. 308-309) described the "pudu {Capra puda) as a kind of wild 
goat about the size of a six-month-old kid, brown in color, the male with small 
horns. The animal is called venado or roebuck by the Spaniards but improperly as 



HERSHKOVITZ: NEOTROPICAL DEER 65 

it has all the characters of a goat except for lack of a beard and in that its horns 
are smooth, rounded and divergent." 

Gmelin (1788, p. 201) referred the pudu to the genus Ovis, but rendered the 
original specific name "pudu," a misspelling used by nearly all subsequent au- 
thors. 

Rafinesque (1817a, p. 437) thought the pudu was akin to goat antelopes when 
he referred it to his newly described genus Mazama, together with Mazatna ovina 
{Oreamnos americanus americanus Blainville). Later he (1817b, p. 44) included 
Mazama puda, Mazama dorsata (=0. americanus americanus), and Mazama sericea 
(=0. americanus americanus) in "a particular subgenus (or perhaps genus) which I 
shall call Oreamnos, distinguished by the horns slightly curved backwards or 
outwards, often rough or annulated, and long hair, besides living in mountains: 
if the horns should prove hollow, it will have much affinity with the genus 
Rupricapa [sic]." This judgment was made earlier by Blainville (1816, p. 76). 

The true nature of Capra puda Molina remained equivocal until 1846 when Gay 
& Gervais (p. 88) demonstrated the cervid characters of two pudu skins and one 
skull that Gay brought to the Paris museum. They also showed that Cervus 
humilis Bennett, described in 1831 on the basis of a female that had lived in the 
gardens of the London Zoological Society in 1830, represented the same species. 

The name Pudu chilensis Gray appears first as the caption for the figure of a 
skull in Gray's Catalogue of the Specimens of Mammalia in the Collection of the British 
Museum (1852b, pi. 36, fig. 1). This name is valid, although it almost certainly is a 
lapsus for, and definitely a synonym of, P. humilis Bennett, the only spedes Gray 
(1852b, p. 240) actually described in the text. 

The preceding synonymy includes a complete bibliographical history of Pudu 
puda Molina. 

Characters 

Size. — Average larger than Pudu mephistophiles, smaller than Mazama chunyi. 

Coloration and pelage. — General body color buffy agouti with middorsum usu- 
ally contrastingly colored reddish brown agouti; pelage stiff, adpressed, the 
hairs pithy; dorsal cover hairs about 30 mm long, guard hairs like cover hairs 
about 5 mm longer, basal half of both types grayish, of terminal half brownish 
(eumelanin) with subterminal pheomelanin band on back, flank, rostrum, 
cheek, and neck; crown, outer surface of ears, supraorbital stripe or drcumorbi- 
tal band, narial and mandibular patches ochraceous to rufous, terminal portions 
of hairs uniformly pheomelanin, inner surface of ears, chin, throat like preced- 
ing but paler, the uniformly reddish parts contrasting with surrounding buffy or 
reddish agouti areas, including rostrum, cheeks, nape, neck, shoulders, flanks; 
chest, belly, inner sides of upper parts of front and hind limbs reddish, lower 
parts of limbs entirely reddish, hairs unhanded. 

Fawns spotted, pelage less hispid than that of adults. 

Color variation. — The dominant pheomelanin ranges in coloration from buffy 
to reddish or reddish brown. Of four adult males from Chiloe taken in January, 
one is in new buffy pelage, another is in new reddish pelage, and two are in old 
reddish pelage. The remaining adults and unspotted subadults at hand with 
known dates of capture are in unworn pelage, all from Chile, as follows: 

February — buffy 6 , Chiloe. 

April — reddish 9, Puerto Montt; rufous 9, Temuco. 



66 FIELDIANA: ZOOLOGY 

April — buffy 9, Cautin. 
July — buffy 6 , Angol. 
September — buffy 9, Angol. 
October — buffy 9, Valdivia. 
November — buffy 6 , Petrohue. 

One female (without collection date) from San Pedro, Concepdon, is in worn, 
faded, buffy pelage. The condition of the fur in this example, however, may be 
the result of exposure to sunlight and wear after the specimen was prepared as a 
round study skin. Two undated males, one from Arauco, are in unworn, buffy 
pelage. 

Worn pelage appears to be associated with the reddish color, but not certainly 
with the buffy. New pelage, however, may be buffy or reddish. There is no 
indication in present material that an animal may molt from one dominant color 
to the other. 

Molt. — Winter molt begins in fall, summer molt in spring. Starting with birth- 
date, fawns begin to shed the juvenal spotted coat during the sixth week and 
lose it completely by end of the second month (Vanoli 1%7, for Chilean pudu; 
Hick, 1%7/1968, and Fradrich, 1975, for German zoo pudus). 

Rhinarium. — Not notably swollen or bulbous, posterodorsal border slightly 
convex or forming a straight line between nostrils. 

Tegumentaiy glands. — Preorbital gland large, the opening wide, conspicuously 
displayed; interdigital glands of forefoot contained in deep trough or pocket of 
hind foot in well-defined pouch; tarsal and metatarsal glands not evident in 
available material. 

External genitalia. — According to Garrod (1877, p. 9, fig. 16), the glans penis is 
similar to that of Odocoileus and Blastoceros {Ozotoceros) . 

Skull. — Lacrimal fossa large, bowl-shaped and often fenestrated, the maxillary 
bone contributing to ventral border, malar bone excluded; anterodorsal border of 
fadal plate of lacrimal bone slightly to moderately convex or convoluted; 
superior border of ascending ramus of premaxillary bone not expanded, and 
either contacting or widely separated from lateral margin of nasal bone; long axis 
of antorbital vacuity diagonal to nearly horizontal, plane area of vacuity from 
approximately one-half to nearly as large as plane area of fadal plate of lacrimal; 
perforations present on upper surface of root of zygomatic process of squamosal 
bone, the diameter of posteriormost usually more than 1 mm; angle of mandible 
subdrcular, produced little beyond posterior plane of condyle; lunar notch be- 
tween condyle and angle little exdsed, concavity between base of angle and 
ramus more or less defined. 

Cranial measurements are given in Table 4. 

Dentition. — Upper dedduous canine, present in some fawns, shed without 
replacement after eruption of first molar; metaloph absent in fully erupted mo- 
lars, postentoconulid and corresponding crista rudimentary in unerupted, ab- 
sent in fully erupted molars. 

Antlers (table 5). — Growth completed and spikes polished during summer, 
shed in winter same day new growth begins. In high, southern hemisphere 
latitudes, mature antlers are rubbed dean of velvet in December and dropped in 
July (Vanoli, 1%7). Antlers of pudus introduced into northern European zoos 
maintain southern periodidty until cast. During the next one or two years, antler 
development and other vital processes adjust to reversed seasons. Antlers attain 




^ Pudu puda \ 

Fig. 36. Southern pudus, females, in Cologne zoo (photograph courtesy Cologne zoo). 




Fig. 37. Southern pudus in Berlin zoo. Left, breeding male; center, male in velvet; rear, 
five-month-old male; right, female. [Photograph by J. Fradrich, courtesy Berlin zoo; cap- 
tion from H. Fradrich (1975, p. 69).] 

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70 RELDIANA: ZOOl-OGY 

polished maturity from end of May to mid-July, are shed between mid- 
December and end of January (table 5). Yearling antlers are little more than 
buds. Second-year spikes grow to about 30 mm. Recorded lengths of third-to 
eighth year antlers range from 63 mm to 90 mm (Vanoli 1%7, p. 162). 

Vertebral formula. — In a male (FMNH 24363) from Quellon, Isla de Chiloe: 7 
cervical, 12 thoracic, 7 lumbar, 5 sacral, 8 caudal. In two specimens (MCZ 5022; 
6029) from Chile, precise locality unknown: 7, 7; 13, 13; 6, 7; 4, 4; 8, 5(-l-?). 

Karyotypes 

The chromosomes of a male pudu studied by Koulischer et al (1972) have a 
diploid number of 70, with fundamental number of 74. Sixty-six autosomes are 
acrocentric or telocentric and one pair is metacentric. The X chromosome, a 
metacentric, is the largest of the complement; the Y chromosome, also metacen- 
tric, is the smallest. A few satellites are seen attached to the largest autosome 
pair. 

Spotomo & Fernandez-Donoso (1975) found virtually identical karyotypes in a 
male and female pudu captured alive in Llanquihue. Satellites attached to the 
largest autosome pair are evident in the illustration of the karyotype, and the 
metacentric X chromosomes are by far the largest of all. 

The pudu karyotype is grossly similar to that of Odocoileus. The only other 
neotropical deer for which karyotypic data are available is the red brocket, 
Mazama americana, with formulae, as dted by Taylor et al (1969, p. 349), "2n=68 
and NF=74 (Taylor et al [Taylor, Hungerford, and Snyder, unpublished] 1968)." 
According to the authors dted, an extra pair of metacentric autosomes in the 
brocket accounts for the lower diploid number of chromosornes. It is curious, 
however, that Benirschke (1977, p. 52, pi. 9) found 50 chromosomes in father 
and son and 49 in the mother of a family group of Mazama americana. 

Comparisons 

All distinctive characters of Pudu puda can be derived from an ancestral form, 
such as P. mephistophiles. Average size is larger, general body form less squat, 
more brocketlike, wdth slightly longer muzzle, less-rounded, more elongate 
ears, and larger eyes; tegumentary glands, most notably lacrimal and pedal, 
more developed; general body coloration of adult less agouti, the monochromal 
hairs pheomelanin; fawn with characteristic cervid spotted pattern; cranial parts 
more advanced, the lacrimal pit large, more deeply excavated, antorbital vacuity 
larger, its long axis more nearly horizontal; permanent upppr canines almost 
entirely lost. The same relationship holds for all other spedfic characters de- 
scribed elsewhere. In virtually every case the differences are quantitative. It 
remains to be tested whether or not complete reproductive isolation between the 
spedes has been achieved. 

Specimens Examined 

Total 31. CHILE. Concepcion: San Pedro, 1 (FMNH); Arauco: Santa Juana, 1 
(FMNH); Sierra Nahuelbuta, 1 (FMNH); Isla Mocha, 1 (AMNH); Malleco: Angol, 
3 (AMNH); Cautin: Maquehua, 1 (AMNH); Pitrufquen, 2 (FMNH); Tolten, 2 
(FMNH); Valdivia: Valdivia, 1 (FMNH); Llanquihue: Petrohue, 1 (FMNH); Puerto 
Montt, 1 (AMNH); Chiloe: Isla Chiloe, 8(5 FMNH; 1 USNM; 2 MCZ); Lago 
Cucao, 1 (AMNH); Quellon, 6 (FMNH); Rio Inio, mouth, 1 (FMNH). 



HERSHKOVITZ: NEOTROPICAL DEER 71 

VII. BIOLCX5Y OF Pudu puda 

Habitat 

The Chilean habitat of Pudu puda is the humid forest or Valdivian biotic dis- 
trict, which lies south of the Santiagan. As described by Osgood (1943, p. 30), 
the Valdivian 

. . . extends along the coast and through the mountains to the vicinity of the Gulf of 
Penas, about lat. 47°S. There is some interdigitation at the northern boundary where 
humidity increases with altitude, but the change is fairly abrupt from a region [the 
Santiagan] of moderate rainfall and relatively light vegetation to one of great humidity 
and dense forest. [Footnote: ". . .rainfall at Valparaiso 20-40 inches; at Valdivia, 
80-200 inches."] In the heart of this district, vegetation is widely varied and highly 
exuberant. Heavy forest growth is combined with dense undergrowth and the 
numerous cryptograms that go wdth an atmosphere constantly charged with moisture. 

According to Wetterberg (1972, p. 348): 

. . . The most favorable pudu habitat in the [Vicente Perez Rosales National] Park 
is . . . classified as the Valdivian Forest Type, and the Coigue [Coihue] Forest Type. In 
the lower altitudes, pudu are often found in the understory thickets consisting of 
small trees of tiaca {Caldcluvia paniculata) , avellano {Guevina avellano), luma (Amonyrtus 
luma), arrayan (Myceugenella apiculata) and huedhued {Pernettya furiens). I never saw 
tracks at elevations above 1200 m. nor have sightings been reported by others. At 
these higher ranges, the understory is often quila {Chusquea quilla) and colihue (C. 
colue). 

Greer (1965, p. 146) notes that pudus have been seen regularly in the Laguna 
Malleco area, w^hich ". . . is heavily forested with coihue (Nothofagus dombeyi), 
rauli (N. alpina) and roble (N. obliqua) along with dense patches of cane [Chusquea 
sp.]." 
The habitat in adjacent parts of Argentina is similar to the Chilean. 

Maintenance in Captivity 

In sharp contrast to its northern congener, the southern pudu adjusts readily 
to captivity, whether as household pet in its native lands, or as resident in 
zoological gardens in Argentina, Chile, Belgium, England, Germany, Holland, 
and the United States. Doubtiess, it can survive in any temperate zone climate 
where deer are kept. The female sent to the London zoo in 1830, which served as 
type of Cervus humilis Bennett (=Pudu puda), may have been the first pudu 
shown in a European zoo. Since then, according to the International Zoo Yearbook, 
published since 1960 by the London Zoological Society, the animal has been kept 
and successfully bred in the Berlin, Cologne, Erfurt, Hellenthal, Wuppertal, 
Rotterdam, and London zoos. Birth records, for which dates are available, are 
listed in Table 6. 

A pudu breeding farm, that began with a pair captured in the wild, was 
established in 1938 in Neuquen, Argentina, by Otto Neumeyer (in Schmidt, 
1944, p. 239). A total of eight individuals, including third-generation young, was 
reported in 1944. 

In 1976, according to the International Zoo Yearbook for 1978, only 12 animals 
were living in zoos, as follows: Antwerp 5, West Berlin 1, Cologne 1, Erfurt 1, 
Wuppertal 3, and Chilian (Chile) 1. 

The official studbook keeper, as of 1978, is J. P. Guillermo Staudt, Director, 
Wildfreigehege Hellenthal, 5374 Hellenthal-Eifel, West Germany, acting under 
supervision of Carlos Junge, Director, Jardin Zoologico de Chilian Viejo, Chilian, 
Chile. 



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72 



HERSHKOVITZ: NEOTROPICAL DEER 73 

Ungulate Associations 

The huemul, Hippocamelus bisulats, is the only other native cervid in Chile, and 
the only one that might occur within the Argentine portion of the range of Pudu 
puda. Historically, the range of the grassland vicugna {Vicugna vicugna) and 
guanaco {Lama guanicoe) may have contacted that of the pudu in Chile and 
Argentina, but probably without sympatry. 

Numbers and Aggregations 

The Abbe Molina (1782, p. 308) notes that, with onset of winter in the moun- 
tains, pudus descended in flocks ("a truppe") to the warm coastal plains of the 
southern provinces. Gay (1847, p. 159) says they lived in small flocks in the 
central valley between the Cordilleras. However, accounts of modern authors 
indicate that the pudu, an inhabitant of forest thickets, is a sedentary and soli- 
tary animal except during the fall rut when it pairs and in spring when the single 
young accompanies its mother. Housse (1953, ex Wetterberg, 1972, p. 349) al- 
lows that pudus may have occurred in small herds before the last half of the 19th 
century, when the animals abounded, but "now that they have decreased, these 
groups of two or three are not a herd in formation but rather families not yet 
dispersed." Natives of the Nahuelbuta region informed Greer (1965, p. 146) that 
"pudus do not change places of habitation and are in the area in equal numbers 
in all seasons." 

Osgood (1943, p. 230) spoke of the animal living in very dense forest, and of 
only once catching a glimpse of one. Those collected on his two Chilean expedi- 
tions evidently were solitary animals driven from cover by dogs. Schneider 
(1946, p. 78) encountered the pudu singly or in pairs in the heavily forested parts 
of the Cordillera de Nahuelbuta, the Cayumanque, and Manquimavida. The 
semidomesticated pudus that Vanoli (1967) observed in gardens or small pre- 
serves were loners, pairs, or family groups consisting of parents with their 
young of the season. 

The largest known concentration of pudus lives on the privately owned Mar- 
garita Island, Lake Todos Los Santos, Llanquihue, within the Vicente Perez 
Rosales National Park. At this island, Wetterberg (1972, p. 348) reported "a herd 
of 10-15 with no natural predators or dogs to disturb them, and no interference 
apart from an occasional stick thrown at them when they wander too close to the 
vegetable garden. Some of these pudus were brought to the island by colonists 
on the mainland whose dogs had chased them into lakes or streams where they 
might have drowned." Wetterberg himself saw only two captive pudus in the 
town of Peulla, and a pair "casually walking down the road near the hotel at La 
Ensenada in the early evening." 

Migrations 

It has been said (Molina, 1782, p. 309) that herds of pudus come down from 
the Cordillera at the beginning of winter to feed on the plains of the southern 
provinces of Chile. Yepes (1943, p. 75) states that the deer entered Argentina 
from Chile through numerous passes in the humid, forested zones of the Andes. 
He records pudus from elevations as high as 900 m (Lago Lolog) above sea level, 
but most of the mountain passes to which Yepes alludes are much higher. The 
pudu does not descend the Argentine slope of the Andes to the Patagonian 
scrublands. 



74 FIELDIANA: ZOOLOGY 

Occurrence of seasonal migrations has not been verified. According to Osgood 
(1943, p. 229), "In early accounts and many later ones based on them, [the pudu] 
is usually spoken of as an animal of the cordillera, but this appears not to be the 
case since it is actually recorded only from heavily forested regions either near 
sea level or at very moderate elevations." 

Present information indicates that the pudu is a sedentary, cryptic animal, its 
movements localized and likely limited to the narrowly circumscribed territory 
of the individual or family group, or to the quest of new territory by the matured 
young of the year. 

Locomotion, Play 

Information on pudu locomotion consists of a few observations of wild or 
captive animals in flight or play. Vanoli (1967, p. 155) compares the low, 
crouching, zigzagging flight of the short-legged pudu with movements of a 
young boar. Playful, developing young bound and gambol like kids (Hick, 
1967/1968). Adults often engage in playful flight in early morning or evening, 
running and doubling until they pant with tongue hanging out (Vanoli, 1967). 
Junge (1966, p. 264) describes the female of the pair in the Chilian (Chile) zoo as 
"sometimes very playful toward evening and early in the morning. She runs 
about the enclosure like a puppy, zig-zagging, skipping and jumping on the 
spot till she is so exhausted that the foam shows on her muzzle. Sometimes the 
more sedate male joins in these games for a moment but he soon gives up or 
gives his high-spirited mate a rather firm dig in the ribs with his horns. The 
female then quickly ceases her playfulness and retires to the hut." 

Enemies 

Principal natural enemies of Pudu puda include the native foxes (Dusicyon 
spp.), puma {Felis concolor), spotted cats (F. guigna, F. colocolo), and possibly the 
eagle owl (Bubo). The introduced enemies — man and domestic dogs — have ex- 
terminated the pudu throughout most of its range and greatly reduced its num- 
bers in the remainder. 

Diseases and Parasites 

Hoof-and-mouth disease among free-living pudus was reported by Vanoli 
(1967, p. 162). He also mentioned infestation with ticks. Two kinds of parasites 
have since been identified as Ixodes taglei and /. stilesi by Kohls (1969, p. 280). 

Food and Water 

Free-living pudus in Chile, according to Vanoli (1%7, pp. 156-157) and 
others, feed on many kinds of native and introduced trees, shrubs, and vines. 
They browse mainly on foliage, but also eat bark, twigs, buds, blossoms, fruits, 
berries, acorns and other nuts, vegetables, and ferns. Branches of the Rosaceae, 
including roses, raspberries, and blackberries, are preferred above all (Junge, 
1966, p. 264). Grass is seldom eaten, but plantain {Plantagd), clover, other weeds, 
and cereals are. Salt is taken eagerly. Pudus thrive on the regular vegetarian zoo 
fare, including commercial food pellets (Czernay, 1977; Hick, 1967/68). Caged 
young also eat insects (Hick, 1969, p. 115). Perhaps many kinds of invertebrates 
are consumed in the wild by pudus of all ages. Vanoli (1967) observes that. 



HERSHKOVITZ: NEOTROPICAL DEER 75 

among the pudus in reserves and gardens of the Osorno-Llanquihue district, 
females are fastidious eaters, nibbling a leaf here and another there, seldom 
eating large quantities of any one kind, except roses, which they especially 
appreciate. In contrast, males, particularly old bucks, satisfy themselves with 
basic foods. 

The free-living, semi domesticated pudus that Vanoli (1967, p. 162) observed 
in Qiile drank water in summer and winter, even when available food was not 
dry. 

Earthquake Sensitivity 

The semidomesticated, free-living pudus of the Osorno-Llanquihue region 
observed by Vanoli (1967, p. 159) panicked with the onset pf the 1960 earthquake 
in southern Chile. Their lacrimal glands were everted as they fled from one 
hiding place to another. The subsequent eruption of volcanic ash caused them to 
cower in their retreats for long periods, with infrequent sallies for foraging. 

Unusual behavior of animals prior to earthquakes has attracted man's atten- 
tion through the ages. For detailed discussions of the subject, see Evernden 
(1976). 

Temperature Regulation 

Pudus normally live in cool, shaded forests. With regard to those in the Con- 
cepcion (Chile) zoo, Junge (1932, p. 241) noted, "Should the animal be exposed 
to the full glare of the sun for 1 or 2 hours it would die the evening of the same 
day." According to Junge (1966, pp. 263-264), pudus at the Chilian Viejo (Chile) 
zoo breathed with their mouths open when the temperature rose to 32° C (= 90° 
F). "Cooling the enclosure by spraying," he writes, "was not a success as the 
animals were more troubled by a damp heat than by ^ dry one. On very hot days 
they withdrew completely into the darkness of their huts to rest or ruminate." In 
winter, however, as shown by Vanoli (1967, p. 162, fig. 10), free-living pudus 
emerge into clearings to bask in the warmth of the high sun. 

Reproduction, Growth, Development (Tables 7-9) 

The female is sexually mature when about six months old (Vanoli, 1967, p. 
160; Hick, 1967/68, p. 111). Males may become sexually mature at about the same 
age, according to Hick (1967/68, p. Ill; 1%9, p. 110), who has the word of a 
Chilean resident that a six-month-old female pudu "was covered by a male of 
the same age" and gave birth about seven months later. The successful mating of 
a nine-month-old Erfurt zoo male was reported by Czernay (1977, p. 230). Al- 
though Vanoli (1967, p. 157) mentions a fertile mating by a one-year-old male, 
he (1967, p. 163) sets the age of male sexual maturity at \y^ years. 

The rut occurs in the fall. Competition or battles between males for possession 
of the female is unknown. Vanoli (1967) summarizes the mating behavior of a 
pair of free-living, semidomesticated Chilean animals as follows. The buck is 
drawn by pheromones to the estrous female living within or near his home 
range. On meeting, they engage in bouts of butting, the blows usually directed 
to flanks and groin. This courtship intensifies to a point at which both animals 
hurl themselves against each other. The buffeting is followed by mutual groom- 
ing, with the female carefully licking the male from top to bottom and the buck 
reciprocating less diligently and thoroughly. The rut lasts several days, and. 



Table 7. Growth and development of a male Pudu puda in the Cologne zoo, Germany.* 
Day Remarks 

1 Bom 13 November 1%7 before 8:00 A.M.; bleats; mother eats fetal membranes; 

fawn rises, walks to distant corner of cage, lies down; suckles standing up 
whilst being groomed by mother; weight "about 1000 g" shoulder height "about 
25 cm" ( = 15 cm?). [Weight at birth 370 g, shoulder height 15 cm "after a few 
hours ran about and hid in a tree in the bush grass," (V). Born London Zoo, 
1975, in usual sleeping area; father present; mother active but returns to feed 
young, licking fawn's anal region while suckling; first suckling attempt at about 
lOhr(B).] 

2 Changes resting place repeatedly; bleats to call mother; utters alarm cry when 

frightened. [Definite suckling at 27 hr; suckles for about 5 min at 2-4 hr inter- 
vals; gains about 80 g daily (B).] 

3 Rests hidden under straw; gambols; nibbles on lettuce and straw but does not 

swallow; suckles at intervals during day; preorbital glands everted when 
frightened; goes to mother for nursing. [Fed only in early morning, noon, and 
evening (V). Runs playfully (B).) 

4 Suckles regularly at 10:00, 12:00, 16:00, and 18:00 hours; eats dirt; cowers when 

frightened. [Ingests dirt and gravel (V).] 

5 Enjoys standing in water tank. 

8 Tests food eaten by mother but spits it out. 

9 [Tests solid food (B).] 
12 Eats leaves. 

14 Eats lettuce; stands in water trough for longer periods. [Height, 20 cm; eats tender 

leaves; runs along with adults (V).] 

15 Straw covering cage floor replaced by sand and cinders; prefers to sit on cinders; 

rises on cage bars. 

16 Cage movements follow regular pattern; changes resting place. 

17 No longer hides; rests in center of cage. 
19 Lowers head and butts mother. 

22 Butts mother and observer (Hick); eats bananas and commercial food pellets for 

first time. 
24 Following fright attempts to climb onto mother's back. 
26 Takes to cover more readily when frightened; groomed less often by mother; 

alternates running with butting. 
28 Eats hay; takes grasshopper from observer's hand; fawn and mother often rest 

near each other. 
30 [Last seen suckling (B).] 
33 Leaps over mother's back in play. 

37 Licks salt stone. 

38 Caught and ate cockroaches. 

39 Mother goes to resting young for nursing. 

40 Young nearly resembles mother in size, spotted coat disappearing; both use same 

place for defecation and urination. 
52 Spot pattern virtually gone. 
60 Weaned; rejected by mother when attempts to suckle. [Nearly as large as mother; 

shoulder height about 30 cm, spotted pelage gone (V). Faunal stripes disappear 

(B).] 
62 Eats adult fare. 

65 Accustomed to solid food; no longer attempts to suckle. 
90 [Faunal spots gone (B).] 

180 6 young sexually mature. [9 young sexually mature (V).] 
270 [First antlers appear in S (V), 6 sexually mature (Czemay, 1977, p. 230).] 

* Source: Hick (l%7/68; 1%9), with interpolations in brackets from Vanoli, 1%7 (V), free 
living, Chile; Brambell & Mathews, 1977 (B), London zoo; and Czemay, 1977, Erfurt Zoo. 

76 



Table 8. Chronology of weight increment, actual and proportional, in a male and 
female Pudu pmda born in the Thiiringer zoo, Erfurt, Germany, from day 1 to day 98 for 
both, to sexual maturity (182 days) and beyond (205 days) for female only. 





9* 


Daily average 


cJ** 


Daily average 


Day 


(kg) 


increment 


(kg) 


increment 


1 


0.78 


11.5% 


- 


- 


2 


0.87 


- 


.70 


- 


2-3 


- 


9.2% 


- 


14.3% 


4 


1.03 


- 


.90 


- 


4.5 


- 


3.4% 


- 


7.8% 


6 


1.10 


- 


1.04 


- 


6-7 


- 


8.2% 


- 


8.2% 


8 


1.28 


- 


1.21 


- 


8-11 


- 


2.7% 


- 


5.4% 


12 


1.42 


- 


1.47 


- 


12-15 


- 


3.9% 


- 


3.7% 


16 


1.64 


- 


1.69 


- 


16-17 


- 


3.4% 


- 


2.1% 


18 


1.75 


- 


1.76 


- 


18-20 


_ 


3.8% 


- 


3.3% 


21 


1.95 


- 


2.04 


- 


21-24 


- 


2.1% 


- 


2.3% 


25 


2.11 


- 


2.23 


- 


25-29 


- 


2.7% 


- 


2.1% 


30 


2.40 


- 


2.46 


- 


30-33 


- 


1.0% 


- 


1.3% 


34 


2.50 


- 


2.69 


- 


34-37 


- 


1.6% 


- 


0.4% 


38 


2.66 


- 


2.73 


- 


38-42 


- 


1.4% 


- 


1.5% 


43 


2.85 


- 


2.94 


- 


43-47 


- 


1.8% 


- 


1.6% 


48 


3.10 


_ 


3.17 


- 


48-55 


- 


0.8% 


- 


1.1% 


56 


3.30 


- 


3.44 


- 


56-62 


- 


1.7% 


- 


1.0% 


63 


3.70 


- 


3.67 


- 


67 


3.85 


- 


3.80 


- 


63-70 


- 


0.7% 


- 


0.8% 


71 


3.90 


- 


3.90 


- 


71-80 


- 


0.4% 


- 


0.3% 


81 


4.05 


- 


4.0 


' - 


81-98 


- 


0.6% 


- 


0.6% 


99+ 


4.60 


- 


4.40 


- 


112 


4.80 


- 


■- 


- 


118 


5.03 


- 


- 


- 


146 


5.45 


- 


- 


- 


160 


5.77 


- 


- 


- 


170 


6.05 


- 


- 


- 


182 


6.25 


- 


- 


- 


195 


6.20 


- 


- 


- 


205 


6.25 


- 


- 


- 



Source: Czernay (1977, pp. 238, 239). 

* Born 2 May 1975 in Erfurt zoo. ** Bom 3 September 1974 in Erfurt zoo. + After 99 
days, male resisted too vigorously for accurate weighing. 



77 



78 FIELDIANA: ZOOLCXiY 

Table 9. Three oldest longevity records for Pudu puda. 



irliest recorded 




Last recorded 




Zoo 




date 


Age 


date 


Interval 


locality 


Reference 


10 July 1909 


Adult 9 


1 Jan. 1917 


7 yr, 6 mo 


Berlin 


Fradrich (1975:71) 


9 Nov. 1966 


Adult 6 


10 June 1974 
(living) 


8 yr, 7 mo 


Berlin 


Fradrich (1975:71) 


14 Feb. 1866 


Newborn 


17 Feb. 1875 


9 yr, 5 days 


London 


Jones, in Crandall 
(1964:595)» 



* The information was contained in an unpublished mimeographed manuscript dated 
1958. 1 find no reference to this pudu in publications of the Zoological Society of London. 



during a three-day peak the buck mounts the doe repeatedly. He thrusts with- 
out panting or other signs of effort, but the female reacts to each thrust with a 
bleat. 

During the rut of the following year, Vanoli reports, the male served the same 
female, as well as a six-month-old, sexually mature virgin. No courtship pre- 
ceded his repeated mountings of the younger animal. On the approach of the 
buck, she arched her back, then stooped and pressed against the ground in 
response to the male's probings. She was silent throughout the act. However, 
the older doe responded wdth bleats to the thrusting of the male, as she had the 
year before. 

Newborn pudus are dropped in the spring of northern or southern hemi- 
spheres after a gestation of about seven months (202 to 223 days). Neumeyer (in 
Schmidt, 1944) reports six births in Neuquen, Argentina, from November 
through January, four occurring in November alone. Two births in the Osomo- 
Llanquihue region recorded by Vanoli (1967) occurred in November and De- 
cember. A late January birth in an unspecified Chilean locality was mentioned by 
Hick (1969, p. 110), and two pregnant females that arrived in the Berlin zoo 
produced in November of the same year according to Fradrich (1975). Eighteen 
other parturitions occurred in western European zoos during the following 
months; the details are given in Table 6. 



Jan. 


Feb. 


Mar. 


Apr. 


May June July Aug. 


Sept. 


Oct. 


Nov. 


Dec. 


1 








1 


11 1 1 1 


1 





1 






The 13 (72%) spring (April, May, June) and summer (July, August, September) 
births were produced by captive does that had borne young the preceding year 
in the same zoos. A wild, captured female that arrived pregnant in the Cologne 
zoo was responsible for the single November birth. The history of the lone 
January birth, also in the Cologne zoo, is not given. 

Fifteen births in the Berlin, Erfurt, Cologne, and Wuppertal zoos from 1967 to 
the end of 1973, compiled by Fradrich (1975, p. 70), are distributed by months as 
follows, with the January birth in the Cologne zoo excluded: 

April May June July August 
14 6 3 1 

Sex of 22 young born in the same four German zoos is 11 males and 11 
females. Newborn listed in Table 6 include 10 males, eight females, and five with 
no sex recorded. Twins have not been recorded. 

Composition of milk is shown in Table 10. 



HERSHKOVITZ: NEOTROPICAL DEER 79 

Table 10. Composition of milk produced on 32nd lactation day (4 June 1975) by wild- 
caught pudu {Pudu puda) maintained in Erfurt zoo since August 1971, compared with other 
ungulates and Homo. 



-1.5 



Component 


Pudu 


Mule deer 


Roebuck 


Goat 


Cow 


Hui 


a. Fat 


3-4 


10.5 


6.7 


4.07 


2.8-3.5 


3.5 


b. Albumin 


2.03 


9.56 


8.8 


3.76 


3.5 


1.2- 


c. Lactose 


7-8 


3.91 


3.9 


4.44 


4.7 


7 


d. Potassium 


81 


- 


- 


- 


150 


53 


e. Calcium 


37 


- 


- 


- 


130 


31 


/. Phosphorous 


46 


- 


- 


- 


97 


15 


g. Sodium 


110 


- 


- 


- 


53 


15 


h. Chlorine 


22 


- 


- 


- 


100 


36 



Values: a — c = g/100 ml; d — h = mg/100 ml after electrolysis. 
Source: Czernay (1977, p. 235). 

Longevity 

The three oldest longevity records for zoo-kept Pudu puda are listed in Table 9. 
Age at accession of each of the first two animals listed probably is more than one 
year. 

VIII. SUMMARY 

The South American deer of the genus Pudu, comprised of two of the smallest 
living species, is characterized and compared with other artiodactyls, particu- 
larly with respect to limb proportions and metapodial morphology. A hypotheti- 
cal reconstruction of pudu history is presented, and the genus is arranged sys- 
tematically as representative of tribe Pudini, subfamily Odocoileinae, family 
Cervidae. The northern Andean species, P. mephistophiles de Winton, is shown 
to be primitive, and the southern Chilean-Argentine species, P. puda Molina, 
derived. Both species are described, characterized, and compared: their geo- 
graphic ranges are plotted, and their life history notes are summarized. The 
status of wdldlife conservation of the pudu also is discussed. 

IX. ACKNOWLEDGMENTS 

Thanks are expressed to the authorities of the institutions listed in the "Ab- 
breviations" section for permission to study the material in their charge. Most of 
the pen drawings were prepared by Marion Pahl, former Field Museum staff 
artist; others were contributed by Zorica Dabich of the present staff. Most 
photographs of skulls were taken by Curtis Bean, a work-study student, later a 
volunteer, the remainder by staff photographer Ron Testa, who also processed 
the prints. Sarah Derr Bruner typed the manuscript, and technical assistant 
Barbara Brown helped with research and preparation of the manuscript for 
publication. 

I am indebted to the following persons and their institutions for providing 
photographs of living pudus and brockets used in the comparisons and descrip- 
tions of external characters: 

Dr. Don D. Farst, Director, and Violet K. Springman, Information Coor- 
dinator, Gladys Porter Zoo, Brownsville, Texas. 

Dr. Hans Fradrich, Assistant Director, Zoologischer Garten, Berlin. 

Uta Hick, Scientific Assistant, Zoologischer Garten, Koln (Cologne). 

Dr. Clyde A. Hill, San Diego Zoological Society. 



80 FIELDIANA: ZOOLOGY 

X. GAZETTEER AND EXPLANATION OF MAPS 
(figs. 30, 35) 

Localities mentioned in the text are listed from north to south by country, 
province, or department, and by nearest town, subdivision, or topographic fea- 
ture where pudus were captured, observed, or reported as probably present. 
Documentation for specimens I examined is included with their respective 
localities. Documentation for other recorded localities is contained in the litera- 
ture dted in text, including the annotated synonymies for each of the two 
species. The numbers identify the localities plotted in Figures 30 and 35. 

COLOMBIA— PHd« mephistophiles 
Valle (Valle del Cauca) 

1. Barragan,4'X)l'N, 75°52'W. 

2. Las Hermosas (Paramo), 3°48'N, 75°56'W. 

2. Las Hermosas. 
Cauca 

3. Moras (Paramo), 2°50'N, 76°12'W. 

4. Delicias (Paramo), 2°38'N, 76°14'W. 

4. Malvasa, 2°34'N, 76°04'W; 3,000-3,500 m. 

K. von Sneidern, November-December 1956; January 1957, September 1957, Oc- 
tober 1957, December 1957; January-February 1958, 15 (FMNH). 

5. Munchique (Cerro), 2°32'N, 76°57'W. 

6. Pusna. See Pusha. 

6. Pusha (Cerro), Totoro, 2°30'N, 76°24'W; 2,800 m. 
6. Guanacas (Paramo), 2°27'N, 76°15'W. 
6. Quintana, 2°24'N, 76°25'W. 

K. von Sneidern, November 1945, 1 (FMNH). 
6. San Rafael (Paramo), 3,400 m. 

Adjacent to Paramo de Purace (q.v.). 
6. Purace (Paramo), 2°21'N, 76°23'W; 2,800 m, 3,480 m, 3,500 m. 

F. Lehmann, July 1945, 1 (USNM), February, May 1953, 2 (AMNH); 1 (FMNH). 

6. Coconuco, Paletara, 2°16'N, 76°22'W; 3,200 m. 
H. Van Deusen, 1 (AMNH). 

7. Sotara (Paramo), 2°12'N,76°31'W. 

7. Paletara, 2°10'N, 76°26'W. 
H. Van Deusen, 2 (AMNH). 

8. La Vega, 2°05'N, 76°50'W; 3,500 m. 

K. von Sneidern, November 1956, 1 (FMNH). 

9. Las Papas (Paramo), 1°55'N, 76°36'W; 4,000 m. 
9. Cusiyaco (Laguna), 1°55'N, 76°36'W. 

9. Lebrero (Paramo), NE Paramo de las Papas (q.v.). ' 

9. Valle de las Papas, 1°52'N, 76°40'W. 
Tolima 

10. Los Nevados. 

11. Roncesvalles, 4°01'N, 75^2'W. 
Huila 

12. Nevado del Huila. 

13. San Antonio, upper Rio Magdalena, 1°57'N, 76°29'W; 2,700-4,000 m. 
P. Hershkovitz, August-September 1951 (observation). 

14. Cueva de Los Guacharos, ca. 1°35'N, 76°00'W. 
Narino 

15. Azufral (Paramo), 1°05'N, 77°43'W. 

16. Cumbal (Paramo), 0°57'N, 77°52'W. 

17. Chiles (Paramo), 0°48'N, 77°57'W. 
ECUADOR— Pudu mephistophiles 

Carchi 

18. Atal, Hacienda El Vinculo, 0°35'N, 77°44'W. Cordillera Pimanpiro, W slope Cor- 



HERSHKOVITZ: NEOTROPICAL DEER 81 

dillera Oriental. 
P. Hershkovitz, October 1934 (report). 
Pichincha 

19. Cayambe (Mt.), 0°2'N, 77°59'W. 
L. Soderstrom, May 1898, 1 (BM). 

20. Deleted. 

21. Papallacta, 0°22'S, 78°08'W. 

L. Soderstrom 1896, 1 (BM), May 1897, 1 (BM). 

21. Papallacta (Paramo), 3,600-3,700 m. See Papallacta. 

22. Antisana (Mt.), 0°30'S, 78°08'W. 
Cotopaxi 

23. Parque Nacional de Cotopaxi. See Salcedo. 

23. Salcedo, W of, 1'^2'S, 78°34'W; 3,500 m. 
Tungurahua 

24. Llanganates (Cordillera), 1°13'S, 78°15'W. 
Andes-Amazon Expedition 1935-1936, 1 (ANSP). 

PERU— Pudu mephistophiles 
Hudnuco 

25. Quivilla road (Quivilla=9°32'S, 76°41'W); 3,300 m. 
40 km NW Huanuco (q.v.). 

26. Carpish (summit), 9°42'S, 76WW; 2,000-3,000 m. 
On Huanuco- Tingo Maria Road. 

26. Zapatogocha (Bosque), Acomayo, 9°43'S, 76°04'W; 3,100 m. 
P. Hocking, November 1972, 1 (FMNH). 

26. Chaglla, 9°50'S, 75°54'W. 

27. Yanacocha, Rio Quero, 9°48'S, 76°25'W. 
27. Huanuco, above, 9°55'S, 76°14'W. 

27. Pamacucho, 4 km from Huanuco (Grimwood, 1969, p. 79). 

27-28. Huanuco and Ambo, between, 9°55'S, 76°14'W— 10°07'S, 76°10'W; 2,700- 
3,300 m. 

28. Ambo, lOWS, 76°10'W. See Huanuco and Ambo, between. 
Junin 

29. Paucartambo (Rio), 10°57'S, 75°16'W. 

29. San Ramon, above, 11°08'S, 75°20'W; 2,000-3,000 m. 

30. Huasahuasi (Rio), 11°15'S, 75°37'W. 

CHILE— Pwdw puda 
Maule 

1. Constitucion, 35°20'S, 72°25'W; 1 m. 

2. Cauquenes, 35°58'S, 72°21'W; 143 m. 
Concqjcion 

— Pedro Valdivia, not located 

3. Concepcion, 36°50'S, 73°03'W; 13 m. 

4. San Pedro, 36°51'S, 73°02'W; 17 m. 
CarlosS. Reed, 1 (FMNH). 

5. Manquimavida, 36°57'S, 73°01'W. 

5. Cayumanque, 36°40'S, 72°33'W. 
Bio Bio 

6. Antuco, 37°20'S, 71°41'W; 602 m. 
Arauco 

7. Santa Juana, 37°11'S, 72°52'W; 75 m. 
W. H. Osgood, 1 (FMNH). 

8. Cordillera de Nahuelbuta. See Sierra Nahuelbuta. 

8. Sierra Nahuelbuta, 37°48'S, 73°02'W; highest point, 1,440 m. 
W. H. Osgood, November 1939, 1 (FMNH). 

8. Vega Blanca. See Sierra Nahuelbuta. 

9. Isla de la Mocha. See Isla Mocha. 
9. Isla Mocha, 38°22'S, 73°56'W. 

D. S. Bullock, December 1932, 1 (AMNH). 



82 FIELDIANA: ZOOLOGY 

Malleco 

10. Angol, 37°48'S, 72°43'W; 150 m. 

D. S. Bullock, September 1929, 2 (AMNH), July 1932, 1 (AMNH). 

11. Laguna Malleco, 38°13'S, 71°49'W. 

12. Galvarino, 38°24'S, 72°47'W; 61 m. 
Cautin 

13. Temuco, 38°44'S, 72°36'W; 113 m. 
13. Maquehue, Temuco. See Maquehua. 

13. Maquehua, Temuco, 38°46'S, 72°39'W. 
D. S. Bullock, April 1910, 1 (AMNH). 

14. Pitrufquen, 38°59'S, 72°39'W; 93 m. 

Carlos S. Reed, June 1926, 1 (FMNH), November 1955, 1 (FMNH). 

15. Tolten, 39°13'S, 73°13'W. 

Carlos S. Reed, March, May 1926, 2 (FMNH). 
Valdivia 

16. Callumapu, 39°36'S, 72°13'W. 

17. Valdivia, 39°48'S, 73°14'W; 7 m. 
Carlos S. Reed, October 1936, 1 (FMNH). 

17. Santo Domingo, 39°53'S, 73°12'W. 

18. Mariquina, 40°08'S, 72°20'W. 
Llanquihue 

19. Vicente Perez Resales, Parque Nadonal, 41°00'S, 72n5'W. 
19. Petrohue, Lago Todos Santos, 41°08'S, 72°25'W. 

W. H. Osgood, November 1939, 1 (FMNH). 
19. Cayetue. See Cayutue. 

19. Cayutue, Lago Todos Santos, 41°14'S, 72°17'W; 221 m. 

20. Lago Llanquihue, 41°08'S, 72°48'W; 51 m. 

21. Llanquihue, 41°15'S, 73°01'W; 54 m. 

22. Puerto Montt, 41°28'S, 72°57'W; 1 m. 
F. M. Chapinar, April 1920, 1 (AMNH). 

Chiloe 
— Isla Chiloe. 
Carlos S. Reed, December 1925, 2 (FMNH), February 1933, 1 (FMNH); B. Dean, 1 
(USNM); Gerrard, 2 (MCZ); no data, 2 (FMNH). 

23. Lago Cucao, Isla Chiloe, 42°38'S, 74°07'W. 
Junius Bird, 1936, 1 (AMNH). 

24. Quellon, Isla Chiloe, 43°08'S, 73°40'W. 
W. H. Osgood, January 1923, 6 (FMNH). 

25. Rio Inio, mouth, Isla Chiloe, 43°21'S, 74 WW. 
W. H. Osgood, January 1923, 1 (FMNH). 

Aisen 

26. Rio Baker, 47°30'S, 73°37'W. 
Magallanes 

27. Isla Riesco, 53°00'S, 73°37'W. v 
ARGENTINA— PMdH puda 

Neuquen 

28. Quillen (Rio), 39°23'S, 70°56'W. 

29. Lago Lolog, San Martin de los Andes, 40°05'S, 71°20'W. 

30. Cordillera del Trompul, San Martin de los Andes, 40°08'S, 71°26'W. 
30. Lago Lacar, 40°11'S, 71°30'W. 

30. Brazo Correntoso, San Martin de los Andes, south of Cordillera del TrompuL 

31. Lanin Parque National, 39°30'S, 71°30'W; 600-3,776 m. 

32. Nahuel Huapi Parque National, 41°00'S, 71°30'W. 
Rio Negro 

33. Lago Mascardi, 41°21'S, 71°52'W; 798 m. 
Chubut 

34. Los Alerces Parque National, 42°50'S, 71°50'W. 
Santa Cruz 

35. Perito Frantisco P. Moreno Parque National, 47°50'S, 72°15'W; 800-2,770 m. 

36. Los Glatieres Parque National, 50°00'S, 73°30'W. 



HERSHKOVITZ: NEOTROPICAL DEER 83 

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