A r c h . B i o l . M e d . E x p e r .

12:179-190, 1979

L i z a r d s and rodents: an explanation for their

relative species diversity in C h i l e

Lagartijas y roedores: una explicación para su

diversidad relativa en Chile

E D U A R D O R. F U E N T E S , F A B I A N M . J A K S I C

Instituto de Ciencias Biológicas, Universidad Católica de Chile.

Casilla 114-D, Santiago de Chile

(Recibido para su publicación el 3 de diciembre de 1977)

F U E N T E S , E . R . , J A K S I Ó , F . M . L i z a r d s and rodents: an explanation for their relative species diversity

in C h i l e . ( L a g a r t i j a s y roedores: una explicación para su diversidad relativa en C h i l e ) . A r c h . B i o l .
M e d . Exper. 72.179-190, 1979.

It is discussed that the relative richness o f l i z a r d fauna in C h i l e might be related to ecological

attributes of this g r o u p ( m a r k e d altitudinal segregation, coarse-grained habitat selection, and l o w
vagility associated w i t h e c t o t h e r m y ) , which w o u l d favor a rapid speciation pattern concomitantly
w i t h Q u a t e r n a r y Glaciations.
L i z a r d s w o u l d d i v e r g e on mountaintops during interglacial periods and w o u l d m e r g e into fewer
forms o r further d i v e r g e on the l o w l a n d s during glacial periods' intermingling. It is shown that
number of subspecies per species in mountaintops is greater than on l o w l a n d s , which is consistent
with the above hypothesis hereafter referred to as " m o u n t a i n speciation m o d e " . H o w e v e r , it is also
documented that lizards might exhibit a " v a l l e y speciation m o d e " , in which case they w o u l d d i v e r g e
in non-icy valleys during glacial periods, intermingling w i t h lizards in adjacent valleys during
interglacials. A s in the previous case, merging or speciating of divergent forms w o u l d depend on
the degree of reproductive isolation attained.
A s a contrast, it is discussed the situation o f the relatively poor Chilean rodent fauna. It is shown
that although rodents exhibit some altitudinal segregation they do not form m o r e subspecies on
mountaintops than on l o w l a n d s , which is possibly related to their fine-grained habitat selection
and greater vagility associated w i t h endothermy. O n the other hand, number of subspecies per
species is positively correlated w i t h the extension of its geographical range, which strongly suggests
a " v a l l e y speciation m o d e " for rodents in C h i l e .

SPECIATION BIOGREOGRAPHY QUATERNARY GLACIATIONS VERTEBRATES CHILE

THE PROBLEM depauperated when compared with the fauna

of an area of similar climate and resources, like
C h i l e has a geographically isolated position in C a l i f o r n i a for e x a m p l e .
South A m e r i c a . T h e high A n d e s on the east, the M a m m a l s , ants and bees are about half as
barren A t a c a m a desert on the north, and the diverse in C h i l e than in C a l i f o r n i a ( 2 ) . Birds
Pacific O c e a n on the west, are all strong barriers and frogs are about the same, but lizard species
to the dispersal of most o r g a n i s m s ( 1 ) . C o n c o ­ are about one and a half times m o r e numerous
mitantly, the Chilean fauna tends to be relatively in C h i l e than in California ( 2 ) . In fact, speciation
179
180 F U E N T E S and J A K S I C

in Chilean Liolaemus species has been strong tend to exhibit strong altitudinal partitioning.
enough to produce a mosaic of alternative c o m ­ Of the 36 Liolaemus species known in C h i l e ,
munities in central C h i l e ( 3 ) . O n the other hand, 20 are predominantly found above 1000 meters
endemism in m a m m a l s , ants, and bees also tends above sea level ( m a s l ) , and only 3 of them are
to be high ( 1 , 2, 4 ) . known to live above and below this altitude.
The question w e will attempt to answer is However, at different latitudes altitudinal
w h y C h i l e seems to be comparable to an isolated segregation varies. Between aproximately 18°
but complex archipelago with copious speciation and 27° latitude south by the A t a c a m a desert,
for lizards, and m o r e like an island for other there are 8 Liolaemus species above 1000 masl
groups. and only 1 below it. Further south, as the valley
Possible explanations for differences in the gets m o r e mesic, the proportion of species found
number of species between C h i l e and C a l i f o r n i a below 1000 masl increases monotonically.
could be related to the amount of time the dif­ Between 27° and 37° latitude, the ratio of species
ferent groups have had to speciate in C h i l e , o r above 1000 masl to species below is about 1.6;
to different resource availabilities in the two between 37° and 42° latitude, the ratio is 1.0,
regions. H o w e v e r , recent studies ( 2 , 5, 6 ) have and further south it is 0.0. T h i s trend reflects
shown resource supply for several groups o f the l o w productivity in the northern valley, the
organisms is quite comparable in Chile and m o r e mesic climate in the central part, and the
California. O n the other hand, paleontological low average height o f the Andes south of 42°
evidence is weak or inexistent, and therefore latitude (see 7 ) .
does not aid to explain the patterns. It is T h u s , there is strong altitudinal partitioning
conceivable though that both of these factors only in the central part of the country, roughly
could be shown to be important. around Santiago. F i g . 1 shows altitudinal species
In this contribution w e have taken an alterna­ sequences on the mesic Coastal Ranges (El
tive route which w e hope will at least offer a Roble, L a Campana) and on the drier A n d e s
partial explanation for the relative species slopes ( L a g u n i l l a s ) . T h e figure starts only at
diversity phenomenon. W e will attempt to show about 1000 masl and does not contain the species
that ecological differences between lizards and b e l o w the level (see 8 ) . Still, it can be seen that
other groups, manifest themselves in different there is species turnover on the three mountains,
speciation patterns. Because available ecological although it is not strictly parallel in all o f them.
information about most taxa is scarce, w e w i l l It is important to emphasize at this point
restrict ourselves mostly to the comparison o f that in other transects covering a w i d e r altitudinal
lizards with rodents. T h e r e is no doubt that range, some species would probably be added,
these are " p a r t i c u l a r " groups, but as will be seen and perhaps other be dropped, depending on
later in this paper, their speciation modes could slope and exposure. T h e important aspects in
offer clues for future w o r k on other taxa. Fig. 1 are the existence of species turnover and

First w e w i l l document, in a fairly detailed the lack of parallelism in the distribution of

manner, the ecological attributes of lizards in species on nearby mountains.

relation to their proposed pattern of speciation. The biological reasons for the altitudinal
We will then c o m p a r e these attributes with the species replacement are currently being investigat­
available ecological information on rodents and ed, but there are already some e m e r g i n g patterns.
their postulated speciation mode. S o m e species in F i g . 1, like L. nigroviridis and
L. leopardinus are typically high-mountain
ALTITUDINAL PARTITIONING lizards, whereas other species like L. tenuis,
L. schroederi and L. nitidus are low altitude
C h i l e is basically a mountainous country. The species further south or north ( 8 , and personal
high A n d e s border its east side, and on the west observations). Species w o u l d thus be " t r a c k i n g "
coast it is limited by the Coastal R a n g e s , with in altitude and latitude conditions w h e r e they
only a n a r r o w central valley in between. T h i s can survive and reproduce. H o w e v e r , what are
physiography is strongly reflected in the distri­ the restrictions which limit the use o f the w h o l e
bution of the organisms. T h u s , Chilean lizards altitudinal gradient by the species?
 LIZARDS A N D RODENTS 181

Fuentes and Jaksic (in preparation) show lemniscatus ( F i g . 1). L. lemniscatus is a grass-
that density of grasses can impose a strong species which uses ground preferentially ( 5 , 11).
selective pressure on lizards, such that grass At El R o b l e , as grass density decreases with
and open ground analogue species can be altitude, L. lemniscatus drops out of the c o m ­
distinguished. O p t i m a l morphologies for survival munity, being replaced by the j u v e n i l e s o f L.
in each habitat are different, and thus presence nigrouiridis (see 5 ) . In this case L. leopardinus
or absence of grass acts as a sieve separating is absent from the Coastal R a n g e s .
true alternative communities (9, 10) in each O t h e r species seem to be strongly associated
environment. with the presence or absence of a v e r y particular
We found this mechanism to explain some environmental feature. Thus, L. monticola is
latitudinal as well as altitudinal habitat partition­ a rock dweller strongly associated with the
ing. T h u s , L. nigrouiridis is preferentially a rock presence of boulders and rocks ( 8 , 11, 12). T h e s e
dweller, but can frequently be found on the features tend to be more frequent high in the
ground if grass density is l o w . A t L a g u n i l l a s mountains, close to tops and ridges, and concomi­
(east of Santiago) w e found that L. leopardinus tantly L. monticola drops out of the community
replaces L. nigrouiridis as grass density increases. at l o w altitudes. Similarly, L. tenuis is a tree
Both species are preferentially rock dwellers, dweller ( 5 , 8 ) and its presence is strongly as­
but have the option of hunting on the ground, sociated with the distribution of trees.
and L. leopardinus is more of a grass-species than Besides these particular elements, some
L. nigrouiridis. general metabolic difficulties seem to add more
The same grass-non grass phenomenon could constraints on high altitude species and to en­
be involved in setting an altitudinal limit on L. hance zonation. Shortening of the growing

EL ROBLE CAMPANA LAGUNILLAS

1000 1200 1400 1600 1800 2000 1000 1200 1400 1600 I8D0 20CO I0CO 1400 1830 2200
T 1 1 1 1 1 1 1 1 1 1 1
L_ nigroviridis

L leopardinus

L_ schroederi

L_ tenuis

L. monticola

L nitidus

L. lemniscatus

Lgravenhorsti

L. fuscus

C maculatus

Fig. 1. Altitudinal species replacement of lizard species.

Altitudinal species sequences at the closeby mountains El Roble, Campana, and Lagunillas are compared. Data for El
Roble were obtained from Pinto el al. (43). Data for Campana from Jerez and Ortiz (44). Notice that species sequences
differ on the three mountains. See the text for further explanation and discussion.
182 F U E N T E S and J A K S I C

season with altitude imposes strong restrictions 1. Species which live on mountains and do not
on some species. Fuentes and D i Castri ( 1 3 ) form subspecies. H o w e v e r , this is strictly not
noted that high altitude Liolaemus species are an exception but sustaining evidence, since
forced to be o m n i v o r o u s , incorporating vegetable these species are known from only one locality
tissues into their diet. They interpreted this and therefore give strong evidence for genetic
plant consumption as compensation for l o w and isolation on mountaintops. Species in this
unpredictable insect densities in those e n v i r o n ­ category are L. signifer, L. ornatus, L. moc-
ments. Furthermore, Fuentes ( 5 ) found that quardi, L. pantherinus, L. fitzgeraldi, L.
large Callopistes maculatus eats vegetation at lorenzmulleri, L. buergeri, L. knegi, L.
its limit of altitudinal distribution, but not paulinae, L. constanzae, and L. darwxni
further d o w n . A g a i n , it w a s suggested that as (see 8 ) .
favorable season shortens with altitude, lizards 2. M o u n t a i n species at high latitudes do not
have to be less insectivorous and have to c o m ­ form subspecies. H o w e v e r , at high latitudes
pensate the l o w consumption of insects by eating mountains are very l o w , and distinction bet­
plant material. T h e p r o b l e m is that only large ween mountain and valley becomes arbitrary
lizards can use vegetation efficiently, and not (see 7 ) . A g a i n , this is expected from the h y ­
all plants are edible (13, 14). Besides the pothesis and does not contradict it. Species
physiological difficulties for most lizards of in this category are L. magellanicus, L. kingi,
eating l o w caloric plant material ( 1 5 , 1 6 ) , high L. lineomaculatus, L. d'orbignyi, and L.
altitude Liolaemus also tend to be viviparous fitzingeri (see 8 ) .
(8, 17). T h i s indicates that shorter favorable
T h u s , the pattern of divergence on mountains
seasons at higher altitudes could impose alti­
but homogeneity at l o w altitudes, confirms the
tudinal restrictions to egg development of some
suggestion based on ecological data regarding
Liolaemus species. O n the other side, l o w alti­
isolation of lizard populations living at high
tudinal limits are likely to be associated with
altitudes. A s w e saw before, habitat selection
particular habitat features ( i . e . rocks and trees),
(most likely in relation with interspecific c o m ­
and diffuse competition ( 9 ) in these m o r e stable
petition) can set barriers to movement of lizards
environments.
on an altitudinal gradient. It is likely then that
In sum, in central C h i l e there is altitudinal
these barriers have a l l o w e d for intraspecific dif­
species turnover of lizards and this pattern seems
ferentiation in mountain species (see Fig. 2 ) .
to be associated with alternative morphologies
and behavior of the species. In general, species Interestingly, the degree of differentiation
with optimal fitness at one point in the altitudi­ between mountain populations has been c o m ­
nal gradient are not equally adapted to other parable t o , or greater than differences observed
habitats, suggesting the presence of ecological between mainland and island populations. T h e r e
barriers to lizard movement between mountains. are no island species absent from the mainland,
and the existing island forms are o n l y subspecies
of mainland species ( 8 ) .

M O V E M E N T S BETWEEN MOUNTAINS H e r e w e refer only to islands connected with

the mainland during the last glaciation, that is
H e l l m i c h ( 1 2 , 18) noted that high altitude species separated from the continent by at most 100
tend to diverge more than species inhabiting the meters deep channel (19, 20). T h i s restriction
valley. In fact, using the data by H e l l m i c h ( 1 2 ) excludes mainly the Juan Fernández archipelago
and D o n o s o - B a r r o s ( 8 ) , w e have been able to and some large islands on the southern channels,
show that species living under 800 masl tend to both of which are not known to h a v e lizards. It
have fewer subspecies than species above that also excludes Easter Island, since it is oceanic.
level (Fischer's exact probability test, P < . 0 0 1 ) . Typical island subspecies are L. cyanogaster
Furthermore, species living above and below braltstroemi, L. pictus chiloensis, L. pictus
8 0 0 masl (L. nitidus, L. fuscus, and L. lemnisca- major on the southern C h i l o é archipelago (43°
tus) do not form subspecies. T h e r e are t w o types latitude south), L. nigromaculatus ater, and L.
of exceptions to this rule: nigromaculatus sieversi at about 30° latitude.
 LIZARDS A N D RODENTS 183

Fig. 2. Altitudinal islands in central Chile.

Distribution of heights above 1000 masl (black) or above 600 masl (dashed) are comparable to an altitudinal mainland on
the east, with an archipelago on the west. Glacial snows could have been as low as 900 masl (see 21) and therefore the
density of altitudinal islands is a measure of alternative recolonization opportunities in central Chile during inter-
glacial times. See the text for discussion and more explanations

In sum, adaptations of lizards to their habitats would have favored resource partitioning and
seems to be fine enough to entail reproductive hence n e w forms capable o f coexistence.
isolation between populations on nearby moun­ By the above mechanism, races and perhaps
tains. species formation would follow interglacial
isolation on separate mountains. During
A T W O W A Y SPECIATION MODEL glaciations, populations formerly on different
ranges w o u l d intermingle and further diverge,
Based on research in the European Alps, Hell- or if differences achieved are not strong enough,
mich ( 1 2 ) suggested the Andean races o f Liolae- they w o u l d m e r g e into one species. A t any rate,
mus would develop on different mountains this is a speciation model which in repeated
during interglacial periods and eventually come glaciation events could generate great diversity.
into contact with each other during glacial In central C h i l e there have occurred three to
periods. T h a t is, as snow level approaches l o w e r four glaciations (21) and there are separate
altitudes, high altitude lizard species and sub­ ranges along the A n d e s , as well as in the Coastal
species w o u l d shift their distributions closer to R a n g e s ( 7 ) , thus a l l o w i n g for multiple species
the valley, with many of them c o m i n g in contact formation within even a small latitudinal range.
with each other. If reproductive difference achiev­ The essential factors in this speciation mode is
ed during the isolation in the interglacial period that species partition the habitat altitudinally
were pronounced enough, natural selection and that during interglacial periods, movements
184 F U E N T E S and JAKSIC

between mountains are restricted and comparable On the other side, the same design operating
to movements between mainland and islands by valley speciation would at most double the
on the continental shelf. In the remaining part number of species per cycle. T h e point is that
of this contribution we shall refer to this speciation rate depends directly on number o f
speciation mode as " m o u n t a i n speciation". isolates and amount of intermingling. C l e a r l y ,
A second mode of speciation, not exclusive speciation rate of groups using both speciation
of the former, is also likely to have operated in modes would be considerably higher than
C h i l e . D u r i n g glaciations, pockets of non snowed speciation rate of groups using only one mode.
habitats or at least with short g r o w i n g seasons, For all the reasons given above, lizards seem
have been inferred to occur in the southern to fall into the mountain speciation mode. T h e
A n d e s ( 2 1 ) . T h e s e pockets being more or less evidence given on their altitudinal stratification
isolated from each other, can lead to reproductive and on the isolation between mountains, with
isolation between populations. During inter- presence of subspecies and even species on some
glacial periods, the formerly isolated populations mountains during the present interglacial, point
w o u l d intermingle and further differentiate, or in this direction. H o w e v e r , in addition lizards
m e r g e into fewer forms. F o r analogous reasons could have also used the valley speciation mode.
as before, w e will refer to this speciation mode Unfortunately, since w e are living on interglacial
as " v a l l e y speciation". In valley speciation, the period at present and therefore during an inter­
essential features are restricted movements mingling phase for valley speciation, it is not
between valleys or pockets during glacial possible to clearly determine its importance.
periods, and m i x i n g during interglacial periods. Possible examples of valley speciation in
Fig. 3 shows the t w o speciation modes in a lizards could be the geographical subspecies
diagram. Note that for three mountains and patterns of L. platei and L. nigromaculatus.
t w o valleys, as shown in F i g . 3, each cycle o f L. platei platei is roughly distributed between
mountain speciation would at most triple the 26° and 32° latitude south, whereas its only
number of species (species 1 and 2 in Fig. 3 ) . known subspecies L. platei curicensis is restrict-

BEGINNING END OF GLACIAL INTERGLACIAL

INTERGLACIAL INTERGLACIAL

MOUNTAIN / 1
SPECIATION,

DIFFERENTIATION OF POPULATIONS DIFFERENTIATIONS OF

POPULATIONS

VALLEY
SPECIATION

DIFFERENTIATION OF POPULATIONS

Fig. 3. Speciation modes.

Mountain and valley speciation are asynchronous with regard to the phases of differentiation and of mixing of differentiat­
ed populations. In mountain speciation, differentiation occurs during interglacial periods and mixing in valleys
during glaciations. T h e opposite is true of valley speciation, where differentiation occurs in isolated habitat pockets or
val leys during gl acial periods. See th e text for further discussion.
 LIZARDS A N D RODENTS 185

ed to a few valleys of C u r i c o , at 35° latitude C o d y et at. ( 2 ) used beta-diversity, that is

(see 2 2 ) . Both D o n o s o - B a r r o s ( 8 ) and M u l l e r species turnover over comparable habitat tran­
and H e l l m i c h ( 2 2 ) insist on the subspecies level sitions, to describe species turnover of m a m m a l s
of differentiation, although this is not crucial and lizards in California and C h i l e . T h e y found
for the argument. It is reasonable to assume that beta-diversity on altitudinal transects is
L. platei once had a wider distribution, covering quite comparable for rodents and lizards in
the gap between 32° and 35° latitude south. central C h i l e . T h i s means not only that lizards
During the last glaciation, the previously and mammals change with altitude, but that
continuous distribution of L. platei could have in central C h i l e — at least around the latitude of
been fragmented. L a t e r , during the interglacial, Santiago — they seem to change at about the
L. platei w o u l d have persisted only in its known same rate.
distribution, with extinctions between 32° and T h e biological compromises involved in the
35° latitude. Here, competition with other altitudinal segregation by rodents are not as
Lwlaemus species could have been very important, clear as in lizards. In N o r t h A m e r i c a , structural
since they reach high species densities in that habitat ( 2 7 , 28, 2 9 ) and productivity ( 3 0 , 3 1 )
latitudinal range ( 8 ) . Fuentes and Jaksic (in have been invoked in relation to habitat par­
preparation) offer evidence suggesting that titioning, but about the Chilean situation there
L. platei — a non grassy species ae is in fact is no published information.
replaced by L. lemniscatus — a grassy species —
W h a t e v e r the reasons are which explain the
at about 32° latitude south. O n the other hand,
altitudinal partitioning phenomenon in rodents,
the permanence of L. platei curicensis since the
o n e requirement of mountain speciation is not
last glaciation is not clearly understood, but
met by these vertebrates. W h e r e a s lizards living
could be related to the local physiography.
on different mountaintops tend to be relatively
The second evidence for valley speciation isolated, the same is not true for rodents. Our
in lizards is the kreiss of L. nigromaculatus. evidence for isolation of lizards is the greater
L. nigromaculatus and its numerous subspecies, race formation between mountaintops than
form a latitudinal cline ( 2 , 12), suggested to be between localities in the valley. For rodents,
a manifestation of secondary contact (12). we used data by O s g o o d ( 1 ) and Miller and
Interestingly, the discussed L. platei — L. Rottmann ( 3 2 ) , and found this difference not
lemniscatus transition occurs at about this same to be statistically significant (Fischer's exact
latitude ( 3 2 ° ) and shows clear signs of h y b r i d i z ­ probability test, P > . 3 4 ) . T h a t is, species restric­
ation and secondary contact (Fuentes and Jaksic, ted to l o w altitudes tend to form as many sub­
in p r e p a r a t i o n ) . ' It is very likely therefore, that species ( o r to be as isolated) as species living
there w a s a glacial barrier to dispersal at about on mountaintops.
that latitude. N o w a d a y s w e w o u l d be o b s e r v i n g
The above result is particularly important
the merging and differentiation of formerly
for our speciation model, since the three m a m -
isolated populations. M o r e evidence on areas o f
malogically better known islands off the coast
secondary contact, and k n o w l e d g e o f which valleys
of C h i l e ( T i e r r a del Fuego, C h i l o e and M o c h a )
w e r e glaciated (see 21) are needed before w e
w e r e all connected to the mainland during the
can postulate a set of barriers to dispersal in the
last glaciation ( 2 1 ) , and n o w a d a y s they exhibit
past.
subspecies of mainland forms. T i e r r a del Fuego
Rodents present a clearly contrasting situation has three subspecies, C h i l o e four, and Mocha
in comparison to lizards. Rodents exhibit alti- three (see 1, 3 2 ) .
tudinal habitat partitioning (see 2 3 , 24, 25, 2 6 ) T h e evidence suggests then, there has been
but, as w i l l be seen, also m o r e movement between time for rodent subspeciation on mountains, as
mountains. F i g . 4 shows the patterns of altitu- well as intermingling of populations during the
dinal segregation at about 18° latitude south, present interglacial, and hence the lack of dif­
and M a l l e c o ( 3 9 ° latitude). It can be seen that ferentiation.
in both situations there are species restricted This difference in mobility between lizards
to mountain tops, and species living further and rodents is likely to be related to their mode
down. of thermoregulation, and energy requirements.
186 F U E N T E S and J A K S I C

T A R A F » C A ( I 8 * - 2 0 " L A T SUR) M A L L E C 0 ( 3 8 ' - 3 9 ° L A T SUR )

1000 2000 3000 4030 5000 1500 2000

~r T
A cirwco
0. glifoidas

P oaqoodi
A. botiviaww
A sublimit
E typu»
L- viicqcio
Ch. tahomqe
A. «don
Ch. brevicouda
N. ebriwus
C. lepidus

Fig. 4. Altitudinal species replacement of rodents .'pedes.

Altitudinal species sequences in central Chile (Malleco) and northern Chile (Tarapacâ). Data for Malleco were obtained
from Greer (24). Data for Tarapacâ were taken from Miller and Rottmann (25), Spotorno (26), and records of Museo
Nacional de Historia Natural (Chile). See the text for discussion.

Rodents are endotherms and do not have the of geographical distribution measured in degrees
capability to hybernate, at least in C h i l e ( 1 , 2 4 ) , (r = .82, P < . 0 0 0 1 ) . T h a t is, species distributed
so they are more likely to periodically shift over long latitudinal stretches tend to have m o r e
their altitudinal range and diet. In contrast, subspecies than m o r e restricted ones. This is
lizards are temporal opportunists, have relatively evidence (certainly not conclusive) for valley
l o w e r energetic requirements, and restrict their isolation during glacial periods, and secondary
activity to the season of m a x i m u m productivity, contacts during interglacials.
not shifting in altitude. In the sense described In conclusion, our evidence points towards
by L e v i n s ( 3 3 ) , lizards w o u l d be coarsely grained a t w o - m o d e speciation for lizards, but a valley
and use one environmental type preferentially, speciation m o d e for rodents. In agreement w i t h
whereas rodents w o u l d be more finely grained this is the fact that C h i l e has m o r e lizards than
tending to use more environmental types, related California, whereas the opposite is true for
to the p r o p o r t i o n o f their occurrence. Rodents, rodents. T h e former w o u l d speciate faster than
being less specialized than lizards, w o u l d have the latter, and w o u l d thus have been able to
exhibited more between-year i n t e r m i n g l i n g and compensate for the more diverse centers of
greater transition probabilities between moun­ origin of t h e ' C a l i f o r n i a fauna. R o d e n t s , on the
tains, and hence less subspeciation than moun­ other hand, depending on more stringent
tain lizards. speciation conditions, w o u l d have speciated at

In sum, rode do not comply with the a slower rate and w o u l d therefore not have been

requirements for mountain speciation because able to compensate for the diversity of centers

of their high mobility during interglacials. On of origin in the Californian rodent fauna.

the other hand, rodents seem to have speciated T h e by n o w well k n o w n radiation of altiplano
in the valley mode. Number of subspecies of rodents ( 2 3 , 26, 3 4 ) w o u l d thus be linked to the
rodents is significantly correlated with breadth existence of numerous v a l l e y s and surrounding
 LIZARDS A N D RODENTS 187

mountains, which w o u l d h a v e provided during south and southwest (Sonoran and Mexican
glacial times the reproductive isolation needed deserts), the Chilean immigration has been
for speciation. T h e geographical and ecological relatively low. For several faunistical groups,
proximity of these valleys during the inter- speciation in the country has been copious.
glacials, their isolation from each other by the A m o n g these, lizards (Liolaemus) seem to be
high mountains, and the vaster areas associated an excelent e x a m p l e o f adaptive radiation. T h e i r
with the altiplano, would have provided the speciation has been explained as a result of the
conditions for greater speciation of rodents in greater speciation rate as compared to other
the northern region than in the more extended groups, particularly rodents. T h e basic difference
southern A n d e s . between these t w o types of vertebrates seems to
Other groups besides rodents and lizards are reside in their different mobilities, ultimately
more difficult to explain, and w e do not have related to their endothermy or ectothermy. A s
the required information to examine tlieir a consequence of this difference, lizards w o u l d
speciation patterns. H o w e v e r , it is possible that speciate during and between glaciations, whereas
both birds and frogs also have valley speciation rodents would do so only during glacial periods.
patterns. Birds have high mobilities and are It is possible the speciation of birds and frogs
not expected to show the large phenotypic dif- could be explained by the same scheme.
ferences between mountains as lizards d o . C o d y
( 3 5 , 3 6 ) has shown Chilean birds tend to segre-
gate more by regions than by habitat types. RESUMEN
Speciation in these animals would probably
operate not only between valleys, but between C h i l e es un país relativamente rico en lagartijas,
distant and isolated ones. Vuilleumier (37) pero pobre en roedores, en comparación con
postulated that the Nothofagus bird fauna as- lugares que exhiben características similares de
sociated with forest speciated phyletically, which clima y recursos. En este trabajo se intenta res-
strongly suggests that long distances are requir- ponder por qué nuestro país ha sido análogo a
ed for genetic isolation in these animals. un archipiélago para las lagartijas (alta diferen-

Birds would thus be expected to exhibit ciación específica e intraespecífica), pero más

relatively lower endemism than lizards or rodents. bien como una isla para los roedores (menor

The similarity of C a l i f o r n i a and Chile bird diversidad y diferenciación intraespecífica).

faunas remains to be explained, but it is probably Se muestra que las lagartijas exhiben una
a compound phenomenon with local speciation fuerte segregación altitudinal, que es particular-
and relatively more immigration from distant mente patente en la zona central de C h i l e (véase
localities. F i g . 1). L a razón para este r e e m p l a z o altitudi-
Frogs might have speciated locally ( 3 8 ) and nal de especies estaría relacionado con la selec-
could exhibit valley and perhaps some mountain ción de ciertas características del habitat por
speciation. F r o g s seem to c o m p l y with the l o w parte de las lagartijas, que les proporcionarían
mobility requirement, but to our k n o w l e d g e it condiciones óptimas para su sobrevivencia y re-
is not k n o w n h o w much altitudinal stratification producción. En el texto se analizan numerosos
they exhibit. It is suggestive though that total casos en que esta situación es evidente.
species counts for C a l i f o r n i a and C h i l e frogs Intimamente ligados a la selección del habitat
are about the same (see 39, 4 0 ) . M o r e research se encontrarían las morfologías y el comporta-
on this point is necessary. miento de las lagartijas, por lo que fenotipos
óptimos en un cierto lugar no lo serían en o t r o ,
CONCLUSIONS en donde podrían ser reemplazados por fenotipos
(especies) diferentes. L a estrecha relación que
It has been assumed — and there are good existiría entre fenotipo y ambiente seleccionado,
reasons for it — that C h i l e and C a l i f o r n i a offer determinaría la partición altitudinal del gradien-
resources for comparable faunas ( 4 1 , 4 2 ) . H o w - te de habitats en las montañas, restringiendo
ever, whereas C a l i f o r n i a has received immigration el flujo de individuos desde ciertas porciones del
of species from centers in the north ( G r e a t B a s i n ) , gradiente (en las cuales constituyen el fenotipo
188 F U E N T E S and JAKSIC

óptimo) a estratos similares en montañas cer- citan posibles ejemplos de especiación "de
canas. v a l l e " . L a suma de estos dos m o d o s de especia-

Este r a z o n a m i e n t o basado en datos ecológi- ción, actuando durante sucesivos ciclos glacia-

cos, se ajusta al hecho que las lagartijas que están les, explicaría entonces la gran diversidad de

restringidas a las montañas muestran mayor lagartijas presentes en C h i l e .

divergencia (mayor cantidad de subespecies) C o m o contraste se presenta el caso de los roe-

q u e aquellas que viven en el valle. El m a y o r dores chilenos, que son bastante menos diversos

aislamiento de las cumbres habría entonces per- q u e las lagartijas. Se discute que, aun cuando

mitido la mayor diferenciación intraespecífica los roedores exhiben partición altitudinal en

que se observa en las montañas. En este sentido, montañas (véase Fig. 4), también presentan

las regiones montañosas se asemejarían a un m a y o r m o v i l i d a d entre ellas. A m o d o de a p o y o

conjunto de islas en las cuales se puede producir a esta proposición, se muestra que los roedores

divergencia intraespecífica, debido a la restric- de montaña no forman significativamente más

ción del flujo de individuos entre aislados conti- subespecies que los de valle. Esto, junto a otras

nentales (véase F i g . 2 ) . evidencias, sugiere que los roedores especiarían

al modo " d e v a l l e " .
A n t e r i o r m e n t e se ha sugerido que las razas En conclusión, los autores proponen que la
de Lwlaemus se desarrollarían en montañas diferencia en diversidad de lagartijas y roedores
durante los períodos interglaciales, entrando en podría deberse a sus distintos modos de especia-
contacto con otras razas similares cuando el ción: " d e m o n t a ñ a " y " d e v a l l e " para el primer
descenso de la nieve (durante los períodos gla- g r u p o , pero sólo " d e v a l l e " para el segundo
ciales) las empujara a bajar al valle. En esta
situación, o se refuerzan los mecanismos de ais-
lamiento reproductivo, o las especies incipientes ACKNOWLEDGMENTS
se refunden en una sola. En C h i l e han habido
tres o cuatro glaciaciones, de manera que el m e - Johanna Filp and Patricio Sánchez made useful suggestions
canismo de especiación propuesto podría generar on the manuscript. W e are also grateful to Angel Spotorno
gran diversidad en sucesivas recolonizaciones for his advice on altitudinal distribution of Chilean rodents,
de las montañas. L o s factores esenciales en este and to José Yáñez, curator at the Museo Nacional de Histo-
ria Natural (Chile), for allowing us to use localities infor-
modelo d e especiación requieren d e la partición
mation on mammals under his care. Research reported in
altitudinal del habitat en las montañas, y la res-
this contribution has been generously supported by L'niver-
tricción del m o v i m i e n t o de poblaciones que v i - sidad Católica de Chile through grant DIUC 113/76 to
ven en distintas montañas durante los inter- Eduardo R. Fuentes.
glaciales. A este modo de especiación, los autores
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