The Mexican Axolotl: Some Misconceptions and Problems

Author(s): Hobart M. Smith

Source: BioScience, Vol. 19, No. 7 (Jul., 1969), pp. 593-597+615
Published by: Oxford University Press on behalf of the American Institute of Biological
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The Mexican Axolotl:

Some Misconceptions and Problems
Hobart M. Smith

No popular experimental animal is

from atl, meaning "water" and having a

with the axolotl is evidenced in (1) the
combining form of merely the letter a,
derisive connotation of ugliness as apmore diversely treated taxonomically
and xolotl, meaning "dog." According to
plied, now even as in early times, to
than the axolotl, a frequently neotenic
Dr. Dibble (pers. comm.), no one meanhumans ("parece un ajolote"), and (2)
Mexican salamander properly known at
ing can be considered correct to the
the well-known myth wherein the god
all stages of its life as Ambystoma mexiexclusion of all others, any one of which
Xolotl, in attempting to escape banishcanum (Shaw, 1789). A recent survey of
is tenable and may have been an acment from the earth (death), assumed
the literature on the axolotl (Smith and
numerous forms, all monstrous (ugly)
cepted
connotation in living n.huatl, in
Smith, 1969) has revealed the existence
given
contexts.
or paired (e.g., corn or agave plants with
of well over 2750 different articles or
All meanings cited above related to
double stalks), finally assuming the form
books referring to the axolotl-most of Xolotl (Fig. 1), an Aztec god of excep- of the axolotl (whereupon Xolotl was
them original sources but many of them
tionally broad surveillance over (1) the captured, killed, and relegated to nourtextbook or reference sources. The misdead and resurrected (in which role he ishment of the sun and moon). The
conceptions made evident by this survey
took the form of a dog), (2) games (hence axolotl may be regarded as fitting the
include application of the common name,
water-sprite, water-player), (3) monpattern of the other ugly or paired disthe taxonomic status of the population to
strous or grotesque beings (e.g., conguises adopted by the god as he atwhich the name axolotl is properly ap-genital and other deformities, any repul- tempted to escape, either through the
plied, the proper scientific and genericsive-appearing objects) and (4) twins.
supposed "ugly" features of the axolotl,
names for the axolotl, the proper name
The close relation of Xolotl as a dietyor through its twin relationship, in its
more misunderstood and none has been

for the larval as opposed to the adult

stage, the degree of neoteny occurring,

and the cause for transformation. Some

of these have been briefly noted by

Maldonado-Koerdell (1964).

Application of the Common Name

The name "axolotl" (rendered

"ajolote" in Spanish) is of Aztec origin.

It has been variously interpreted in

meaning: water-slave, water-servant,

water-sprite, water-player, water-monstrosity, water-twin (Seler, 1900-01, p.

110, 111; Seler, 1902-03, p. 182, 183;

Berridge, 1935, p. 242; Deuchar, 1957,

p. 102). Dr. Charles E. Dibble, of world-

wide fame
as an
expert
n.huatl,
language
of the
Aztecs,
has on
kindly
guidedthe
a search of the extensive pertinent literature, and expresses a preference, as I do,
for still another meaning, water-dog,

matching a very popular common name

in English for the closely related larval
tiger salamander Ambystoma tigrinum
(Green). Dibble (1950) derives the name
The author is a professor in the Department of
University of Colorado, Boulder.

aquatic form, to the terrestrial (trans-

formed) salamander; the two forms

(aquatic and terrestrial) were seemingly

recognized even then as stages in the

life cycle of a single species. The mythology of Xolotl as related to the axolotl is
reviewed in numerous readily accessible

sources (Martin del Campo, 1946;

Sejourne, 1956; Caso, 1958; Soustelle,
1967).

No doubt the name axolotl was origi-

nally applied, as is true even now in

Mexico, to larvae of any of the several

species of the family Ambystomatidae
occurring within the area familiar to the
Aztecs. Presumably all are edible, and

especially about Mexico City and PItcuaro they have been long recorded as
edible (as for example in Hernindez
and Sahagun; see Dibble and Anderson,
1963). They remain today a dietary delicacy or staple in these areas.

However, the ambystomatids of

Mexico represent three genera and about

a dozen species. The species historically
Fig. 1. A depiction of the Aztec god Xolotl inmost commonly encountered and probBiology, its doglike manifestation. From the Borgia ably most commonly used as food is the
codex, as rendered by Nicholson (1959).
dark-colored one apparently restricted

July

1969

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593

to Lake Xochimilco, a permanent, clearwater lake of considerable depth. Also

common in the usually temporary, shallow, muddy lakes about Mexico City are
the light-colored larvae of the tiger
salamander (Ambystoma tigrinum), a
species distributed over all except the

than in pre-Cortezian times, being limited to the single kind that occurs at Lake
Xochimilco. E. R. Dunn (1931) has been
particularly influential in this restriction.
Some (Brunst and his associates) restrict

used without ambiguity is in reference to

the particular taxon that inhabits Lake
Xochimilco, whether in the larval or in
the transformed state. This is the sense
in which the name has most often been

the name even further to the neotenic

used. A concerted effort by all authors

larvae of this single species (Brunst,
should be made to avoid usage of the
extreme western United States, and in
1955). Restriction to this particular name
spe- "axolotl" in any other sense, exsouthern Canada as well as throughout cies is appropriate not only because cept
thisof course with distinctive and approthe plateau region of Mexico south as
priate modifiers.
is the kind, presumably, most commonly
In brief, the name axolotl should be
far as the environs of Mexico City and referred to by the Aztecs but also because
Puebla. These too are used as food, and
it is the kind that has been so widely
used only in reference to the species of
they are today referred to as axolotls studied in laboratories all over the world.
ambystomatid salamander occurring in
just as is the Xochimilco type. Lake
None of the other kinds occurring so
Lake Xochimilco, but it should apply
Lerma of the Valley of Toluca is rather
widely in central Mexico have received equally to examples at any stage of
like Lake Xochimilco, being permanent,
more than superficial study. The ship- development.
deep and clear, and it has another darkment of 33 animals received in Paris in
species (Ambystoma lermaensis Taylor)1864 clearly represented the Xochimilco The Proper Generic Name

very similar to the Lake Xochimilco salapopulation, as indicated by many illustramander. The shallow lakes and ponds in
tions of specimens in that collection
However subjective the fixation of the
(Dumeril, 1866). Descendents of this
this area also contain larvae of the tiger
common name axolotl may be, the gesalamander. All are known as axolotls.
group and another received in Paris inneric name is now incontrovertibly fixed
1868
Still farther west, in the great lake
of were shipped to laboratories all
as Ambystoma Tschudi, 1838. The emenover the world, with the result that 99%dation Amblystoma, originated by
Paitzcuaro, larger and deeper than and
as clear as Lake Lerma and Lake Xochiof the experimental studies published on Agassiz in 1846, was rejected by the
the "axolotl" have been based upon
milco, is yet another specialized ambystoInternational Commission on Zoological
matid salamander (Bathysiredon dumeprogeny originating from the Paris imNomenclature (1963a), giving credit to
Tschudi for deliberate choice of the
rili Dug'es), generally known as "achoque
ports of 1864 and 1868. Even today most
work is restricted to this stock-one of
de aqua," important as food. The mounspelling he used rather than assuming
tain streams of the southern edge of the
thepurest lines of experimental animals
as was commonly done, that he meant
derive the name from the Greek words
plateau harbor another group of species
-although in recent years a few new
of ambystomatid salamanders (Rhyacosiimports have been utilized, no doubt in amblys (blunt, dull, or wide) and stoma
redon spp.) not plentiful enough to
be cases diluting or contaminating
some
(mouth). Stejneger (1907) pointed out
commonly used as food, but still known
that the name was a contraction for
the original, pure stock.
as axolotls.
In summary, basically three applica- Anabystoma, meaning "to cram into th
tions of the term "axolotl" have comVelasco (1880) notes that the name
mouth," and that Tschudi's use was calaxolotl is not limited in application by monly occurred in the past: (1) restriction
culated and deliberate. Continued use
native Mexicans to the larval stages but to the species occurring at Lake Xochiof the rejected emendation Amblystoma
is also used, with appropriate modifiers milco, whatever the stage of developis indefensible in a scientific context.
(ajolotes pelones, ajolotes mochos, ajol-ment; (2) extension to include all larval
However, many other generic names
otes sin aretes, ajolotes sordos, ajolotesambystomatids of Mexico, or at least
haveof
been used for the axolotl in the past;
de cerro, etc.), in reference to the trans- central Mexico where they are used
for
the
most popular among these is Siredon
formed individuals of any species with food; and (3) extension to all members
of 1830, a name still used occaWagler,
free-living larval stages.
the family Ambystomatidae (or even
of
sionally.
In fact, some authors (e.g.,
Application of the name axolotl has
the order Caudata) in the free-living,
Brunst and his collaborators; see Brunst,
been extended in post-Cortezian timesgilled, larval stage. The latter extreme
is consistently refer to larvae as
1955)
to larvae even of North American popu-highly undesirable, as the name axolotl
Siredon mexicanum, adults as Ambystoma
lations of the tiger salamander; refer-thereby loses any useful preciseness
and
mexicanum.
There is no scientifically
ences to the "Colorado axolotl" or "New
becomes virtually a synonym of "larva."
acceptable defense for use of more than
Mexican axolotl," etc., occasionally oc-Reference to such a stage of life history
one name for examples of a single specur. In fact the word has come to mean,in any salamander as "axolotl-like" is
cies, irrespective-of stage of life history
in some circles, simply the larval as perfectly acceptable, as are the adjectival they may represent, nor is there a need
opposed to the transformed stage of anyderivatives "axolotline" or "axolotloid."
for distinct names linked with given
ambystomatid salamander of any genusBut to refer to a larval Gyrinophilus, for
stages. Furthermore, the generic name
or species. It has even been used occaexample, or a larval Triturus, as an
Siredon was officially rejected by the
sionally in reference to the larval stage "axolotl" is an unnecessary distortion International
of
Commission on Zoological
of any type of salamander in which thereusage of the term.
Nomenclature (1963a), and accordingly
may be a transformed as well as larval
In like fashion the name axolotl loses
its use can no longer be condoned.

stage.

On the other hand, in some circles,

particularly among taxonomists and
naturalists, the name axolotl has acquired
a more instead of less precise application

594

preciseness when applied indiscrimi-

nately even to Mexican ambystomatids,

whether in reference to the larval stage
alone or the transformed adults as well.
The only sense in which the name can be

BioScience

Vol.

The number of generic names used for

the axolotl is astonishing, but even more
astonishing is the fact that at least seven
of these (Axolot, Axolotus, Philhydrus,
Phyllhydrus, Siredon, Sirenodon, Stego-

19

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No.

adults vs. light above and below, in

transformed adults, dark above with large
light spots, light below). The axolotl presumably freely interbreeds under artificial conditions with the surrounding populations clearly referable to Ambystoma
tigrinum, as several authors (e.g., Geyer
and Freytag, 1949; Humphrey, 1967)

have reported fertile hybrids between

laboratory stocks. Yet we have no evidence that in nature there is any hybridization whatever between the two

Fig. 2. A larval Ambystoma mexicanum, total length 211 mm., taken August, 1967, at
Xochimilco, D. F., Mexico, by Thomas H. Fritts. (Photo courtesy Edwin D. Pentecost,
taken about three months after capture.)

porus) antedate the generic name that

now is officially designated for it! They
were long overlooked, however, or were
superseded in popularity by Ambystoma,
so that unnecessary confusion would result from their resurrection. The 1963(a)
Opinion of the ICZN officially suppressed
all these names. Still other generic names
have been used for the axolotl, but are
either junior synonyms of Ambystoma
Tschudi, 1838, or were used incorrectly;

examples are Gyrinus, Proteus, Lusus,

Axolotes, Piscis, Siren, and Hypochthon.

The proposition that an artificial generic distinction be maintained between

populations. They may possibly intergrade at points of actual range contact,

and thus constitute conspecific subspecies, but in the absence of such evidence
it is preferable to assume that their gene

pools are essentially separate and that

even in Xochimilco stocks. Morphological

the two taxa constitute species in nature.
gaps, if they exist, are quite subjective.
Thorough field studies will be essential
for definitive conclusions; an educated
The small number of gill rakers, long
thought to be a useful distinction from
guess of specific relationship is the best
A. tigrinum with more numerous gill
that can now be provided. Salthe (1967)
rakers, is excellent only in comparison
suggests that the courtship patterns of
with northern tiger salamanders! Thethe two species appear to be importantly
number is the same in comparison with
different; if differences exist they may
act as a reproductive barrier in nature:
tiger salamanders from nearby localities.
The taxonomic status of all these popula"What is suggested . . . is a possible close
tions is open to question and can be
relationship between A. mexicanum and
established definitively only by intensive the opacum group as well as a possibly

more distant relationship between A.

restudy.
Information now available indicates

mexicanum and A. tigrinum than are

that the Xochimilco population consistindicated by Tihen." Courtship records

the axolotl and its relatives in Ambystoma

ently differs from the surrounding are,
popuhowever, sorely deficient.
(Smith, 1939) was effectively demolished
lations in habitat (deep, permanent, clear
Regardless of conspecificity or bispeciby Tihen (1958), who demonstrated that
water vs. shallow, temporary, turbid
ficity of the axolotl and tiger salamander,
the relationship was so close that to accept
water), development (commonly neotenic
it is an operational certainty that at least
generic distinction would make a mockery
two taxa are involved; they are taxovs. seldom neotenic), and color/pattern
of taxonomy. The problem of replacement
(dark with black flecks, above and below,
nomically distinct at least at the subspeciof Ambystoma 1838 by the earlier Siredon
with light flecks above in transformed
fic level. The fairly recent habit of some

1830, which the 1939 solution was proposed to obviate, was eliminated by action of the ICZN (1963a).
Populational Relationships

The information on populational relationships of the axolotl that is mandatory

for a definitive determination of taxo-

nomic status is, most regrettably, lacking.

At one time morphological, chromatic,

and developmental data gave adequate

justification, on the basis of the criterion
of gapsize, for regarding the Xochimilcolike population as a species distinct from
all others. The greater diversity of samples now available reveals that these distinctions are actually not clearcut. Ambystoma lacustris of Lake Zumpango is virtually identical in color and pattern and

so is Ambystoma lermaensis of Lake

Lerma. Neoteny, long a treasured characFig. 3. A fully transformed Ambystoma mexicanum, total length 891 mm. (same data and
ter, is now known to be widely variable
credit as Fig. 2).

July

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595

workers (even Mexican, e.g., Tamayo,

Among them are Siren pisciformis Shaw, mation. Brunst (1955) remarks that indi-

1962: 86,320) to equate the axolotl and 1802; Siredon axolotl Wagner, 1830; Axol- viduals that do metamorphose seldom
otes guttata Owen, 1844; Siredon hum- live long thereafter. Herpetologists as
the tiger salamander and to combine
indiscriminately all work on either taxonboldtii Dumeril, Bibron and Dumeril, well as experimental zoologists have emunder the name Ambystoma tigrinum is 1854; Gyrinus edulis, idem; Lusus aqua- braced the view of obligatory neoteny;

rum, idem; Piscis lubricus, idem; Siredon my own proposal (Smith, 1939) for arbialba Dumeril, 1870; Ambylstoma Weis- trary retention of Siredon was based upon
ably distinct and failure to distinguish
that premise, in turn acquired by permanni Wiedersheim, 1879; and Siredon
them conceals other differences that may
sonal communication with other herpetolexist and courts discrepancies in experiedule Dughs, 1884. Several of these
specific epithets have additionally been ogists. Both Dunn (1931) and Schmidt
mental results readily explicable by
(1960), highly influential herpetologists
used with other generic names.
proper segregation of experimental
animals.
Although the name mexicanus was first of their era, adhered to the "obligatory
proposed in conjunction with the mascu- neotene" school of thought.
The species as a whole, however, is not
Specific and/or Subspecific Names line name Gyrinus, its ending must (in
conformance with the International Code
so limited in ability to transform. Threefourths of a shipment of 22 larvae reon Zoological Nomenclature) agree with
One of the curious oversights of
ceived in 1967 at the University of Illinois
that of whatever generic name it is assoworkers who would regard the axolotl
through the efforts of Mr. Tom Fritts
ciated with. Ambystoma is neuter, and
and tiger salamander as conspecific is the
promptly transformed (although no effort
therefore the specific epithet takes the
fact that, up until 1963, the proper name
was made to produce metamorphosis),
neuter form, mexicanum.
for the composite "species" would be
the rest remaining in the larval state
Ambystoma mexicanum, not A. tigrinum,
The Family Name
about a year longer despite being kept
since Gyrinus mexicanus Shaw, 1789,
under identical conditions. Brunst (1955)
long antedates Salamandra tigrina Green,
likewise observed that specimens in some
1825. In fact, the earliest published epi- Even the proper orthography of the
of the new shipments received in this
thet (specific or subspecific name) now name of the family to which the axolotl
country directly from Mexico transform
referable to Ambystoma is Shaw's mexi- belongs has been subject to differing
canum. Even the type of the genus,
opinions of acceptability of the variants very often. A shipment of several speciLacerta subviolacea Barton, 1804 - Ambystomidae, Amblystomidae, Amby- mens to the University of Auckland, New
Lacerta maculata Shaw, 1802, now Amby-stomatidae and Amblystomatidae. With Zealand, included some which later
stoma maculatum (Shaw, 1802), is more the rejection of Amblystoma, both family transformed (James D. Fawcett, pers.
comm.). Numerous records exist of sponrecent than mexicanum (a generic type names derived from it are, of course,
is, however, not required to be the ear-eliminated. The ICZN (1963a) officially taneous transformation, beginning with
the epochal announcement of Marie von
designated the orthography Ambystomaliest in the genus-it needs only to be
Chauvin (1876), which precipitated an
tidae as correct by placing that name on
explicitly designated).
incredibly prolific and agitated discourse
In 1963, however, the ICZN first placed the Official List of Family Group Names
mexicanus on the Official List of Specific in Zoology. It must thus be accepted in a scientific community already "sold"
on the obligatory nature of neoteny in the
Names (ICZN, 1963a) and later the same despite etymological and grammatical
year placed tigrina on the same list
arguments to the contrary (Stejneger, axolotl. Her observations were not widely
repeated, and thus the impression still
(ICZN, 1963b).. Ironically, these actions 1907).
remained that transformation is an excepprecipitated a unique nomenclatural
tional thing. Weismann's discourse (1882)
Fixation of Neoteny
problem, since any " . . . specific name
is one of the most elaborate documentaonce stabilized in this way is to be used
The
scientific
literature
is
saturated
tions of the puzzlement of scientists of
in preference to any other name for the
time about larval reproduction and
species concerned .. ." (Hemming and
with statements that the axolotl is a the
fully
transformation
in the axolotl, and its
Noakes, 1958). The problem is: if mexi- or virtually obligatory neotene which
can
significance.
canum and tigrinum are considered con-be forced to transform only by resort
to
One of the conventional defenses for
specific, which name is valid? The Code artificial means, for example, by introrequires that each one be valid, since
the
"obligatory neotene" school was, and
ducing iodine into the water, or injecting
is, the supposition that in nature the
each is on the Official List, yet obviously thyroxine, thyroid extract, thyroid
only one can be used. A proposal is now glands, or pituitary extract into the larvae. axolotl never spontaneously transforms.
before the ICZN to validate tigrinum for This impression stems largely from the Even Brunst (1955) regarded natural
the species, and it may safely be assumed fact that, as received by Brunst (1955), transformation as nonexistent or very
that it will be approved. Therefore, if the the most popular stock for experimentsrare. This supposition is in error, as
axolotl is regarded as a subspecies con- carried on in laboratories all over EuropeVelasco (1880) states at length that metaspecific with the tigrinum, it should be is one in which metamorphosis rarely morphosed individuals are well known at
designated Ambystoma tigrinum mexi- occurs, if ever. The stock used by Dr. Xochimilco as well as elsewhere in
canum. If held as separate species, the R. R. Humphrey (pers. comm.) for severalMexico.
names mexicanum and tigrinum are in- decades in this country also never yielded Accordingly, it is well established that
violable for the axolotl and tiger sala- transforming individuals among the sev-both in the fresh laboratory imports and
mander respectively.
eral thousands reared to reproductive
in nature spontaneous transformation of
Other specific epithets have been ap- stages. In this stock, neoteny is indeed the axolotl does occur frequently and
does not require heroic measures such
plied to the axolotl; all are junior syn- apparently obligatory and artificial
means must be utilized to force transforas iodine or hormone application.
onyms of Shaw's Gyrinus mexicanus.
most unfortunate. The taxa are remark-

596

BioScience

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No.

Cause of Transformation

quickly eliminated, producing a pure

least 14 other generic and 10 other specineotenic line. The native stock, however, fic names have been applied to the
The occurrence of spontaneous transstill retains a high frequency of indi- axolotl.
formation in the axolotl and in the re2) The same scientific name must be
viduals with the genotype facilitating
transformation.
lated tiger salamander has been genused for all stages of the single species.
3) The common name axolotl has been
It seems apparent, therefore, that the
erally regarded as subject primarily to
an environmental influence. Weismann
role of the environment to determine
very broadly applied, but should be rewhether a given individual will transform
(1882) sought endlessly for some environstricted to the single dark species occurin Lake Xochimilco. The name is of
or not is subordinate to the geneticring
commental factor to account for the neoteny,
and many authors have concluded that
Aztec origin and means "waterdog."
position of that individual. Meaningful
experiments upon transformation will
in family name to which the axo"conditions" were primarily responsible
4) The
for the maintenance of neoteny, or
the
for
future require careful attention
lotlto
is assigned has been officially fixed
resort to transformation. The mere introthe particular genome represented
as the
byAmbystomatidae.
duction of an axolotl (or tiger salamander
the experimental animals. That two 5)
real
The axolotl is not an obligatory
larva) into a "stress" situation (e.g., shalstrains or cryptic species may exist in
the
neotene,
but frequently transforms in
nature.
axolotl as well as in the neotenic races of
low water, different pH, inanition, excessive warmth) might trigger metathe tiger salamander is a possibility
6) Transformation is genetically limmorphosis, and indeed many cases of
worthy of concerted attention.
ited, some individuals being incapable of
metamorphosis were associated with
Another aspect in need of investigation spontaneous transformation even under
stress phenomena of one sort or another.
is the statement by Velasco (1880) that stress, others readily transforming in
What has been less frequently observed
the color of both the axolotl and the tiger presence of stress. Whether populais that not all individuals react the same
salamander is correlated with turbidity, tional strains or heterogeneous variants
to these stresses; some transform readily,
light specimens (presumably tigrinum) are involved is unknown. Likewise unwhereas others do not.
becoming as dark as the axolotl when kept known are the effect of turbidity upon
It seems quite evident, on the basisinof
clear water (he did not mention recip- color of larvae, the degree of correlation
data now available, that the capacityrocal
to experiments). What bearing this
of development of the 6th aortic arch
transform is a genetic factor, and that
in
observation
has upon the status of the with neoteny, and the degree of sympatry
the axolotl some individuals cannot transaxolotl remains to be determined, but
or parapatry, if any, of the axolotl with
form no matter what stress is imposed,
since dark color is supposedly diagnosticthe adjacent race of the tiger salamander.
whereas other individuals do transform
of the axolotl, it may have some significance.
if they encounter almost any sort of stress.

References

Few if any, apparently, bear a genetic

Likewise intriguing is the observation
composition that requires transformation
by Brunst (1955) that neoteny is linked Berridge, W. S. 1935. All about Reptiles and
Batrachians. Harrap, London. 271 p.
irrespective of presence of stress, whereas
with absence of the ventral part of the
Brunst, Victor V. 1955. The axolotl (Siredon
most salamanders do, including most
6th aortic arch in tigrinum, and that only mexicanum). Lab. Invest., 4: 45-64, 429-449.
geographic races of the tiger salamander. in those individuals with a complete 6th
Caso, Alfonso, 1958. The Aztecs: People of

The survival value of neoteny is quite aortic arch is transformation possible.

apparent in regions of a hostile environ- Applicability of this correlation to the

ment outside of the water. It is also

axolotl merits examination.

the Sun. University of Oklahoma Press,

Norman. 125 p.

Chauvin, Marie von. 1876. Uber die Ver-

apparent that in the axolotl, as in The

some
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Amblystoma. Z. Wiss. Zool., 27(4): 522-535.
high-altitude races of the tiger salamanhe has seen two types, only one of which
Deuchar, Elizabeth M. 1957. Famous animals.
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neoteny has not been completed, contraryXochimilco, requires clarification. The
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to the case of the related Paitzcuaro speimplication is that in southern parts of
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actual strains exist-maintained by extencolored type (tigrinum?) occurs, whereas
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(1955), or whether there is a heterosumably mexicanum) occurs. Existence
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of both species in one lake, even with
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ecological segregation (assuming that the
Dum6ril, Auguste Henri Andre. 1866. ObserThe fact that the descendents of the
southern parts are shallower and/or more vations faites a la menagerie des reptiles du Museum d'Histoire Naturelle sur
Paris imports very soon established a
turbid), is highly questionable.
population of universally neotenic axolreproduction des axolotls, batraciens urod6les ' branchies ext6rieures et sur les
otls (as is documented by the experience
Summary
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thousands of animals) is readily explicable 1) The axolotl appears to be specifiDunn, Emmett Reid. 1931. Present status of
on the basis of the inadvertent but strong cally and certainly is subspecifically disthe axolotl problem. Anat. Rec., 49 (1)
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Uber Kreuzungen zwischen Tigersalaare very difficult to breed successfully in designated Ambystoma tigrinum meximander (Ambystoma tigrinum) and Axolotl
the laboratory. Those Paris individuals canum if held as conspecific with tigri(Ambystoma mexicanum) und ihre F2and descendents which possessed the num, or Ambystoma mexicanum if reGeneration. Mitt. Mus. Nat. Vorg. Naturgene or genes for transformation were garded as allospecific with tigrinum. At
wiss. Arb. Magedeburg, 2 (2): 9-23.
Continued on p. 616

July 1969

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597

the models show the effect of the rain.

.1963b. Opinion. 663. Salamandra tigrina

Green, 1825 (Amphibia): validation under

The constants were 2400 cpm, 4820 cpm,
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3, respectively. While about 16% more

Maldonado-Koerdell, Manuel. 1964. Algunos
activity was applied to Tree I than the
problemas de sistemitica y distribuci6n
other trees, Tree 2 had more activity
de la familia Ambystomidae (Urodela, Am-

translocated to the fruit. The difference

phibia) en la repdblica mexicana. Mem.

Acad. Nac. Cienc. 'Ant. Alzate', 59(3/4):
in the equilibrium constants was the effect
379-388.
of the loss of activity by the washing of*
Martin del Campo, Rafael. 1946. Xolotl y los
the rain. As in the first experiment,
gemelos y monstruos. An. Inst. Biol. (Mex.),

washed-off activity was confined to the 17: 343-345.

peat on the plastic. Because of the large
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changed after 10 days and fresh peat and Protombdico e historiador del Rey de
Espaia, Don Felipe II, en las Indias Occiplastic spread for the next 16 days.
dentales, islas y tierra firma del mar oceano.

Obras completas. Tomo I. Vida y obra de

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y Nueva Espaha en la tpoca de Felipe II.

Summary

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experiments. For a given amount of effort,

Nicholson, H. B. 1959. Los principales dioses

mesoamericanos. Pp. 161-178, figs. 1-47,
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one has the alternatives of either (1)

greenhouse experiments with a highly

ciones Antropol6gicas de Mdxico. M6xico,

D. F. Vol. 1. 478 p.

artificial environment, but with more

statistical control, or (2) field experiments
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When you need a good

MICROSCOPE ILLUMINATOR

by Raul Noriega et al., Centro de Investiga-

look at

Salthe, Stanley N. 1967. Courtship patterns and

phylogeny of the urodeles. Copeia, 1967 (1):
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merit. The concentration of mixed radio-

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sample activities that were considerably Thames and Hudson, London, 192 p.
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TIYODA

TIYODA OPTICAL COMPANY

348 Sixth Street

San Francisco, California 94103

Watson & Viney, London. 146 p.

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Further conclusions on uptake and

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be drawn after complete analysis
of

Smith, Hobart M., and Rozella B. Smith. 1969.

An Analytical Bibliography of the Axolotl.
(In preparation.)

the results.

Continued from p. 597

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July

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