Liolaeminae (Squamata: Iguania:
Tropiduridae)
1
Department of Biology
2 Ecology, Evolution, and Conservation Biology
San Diego State University
University of Nevada, Reno, NV
89557 USA
San Diego, CA
92182-4614 USA
Voice:
(775) 784-1703 - FAX: (775) 784-1369
Voice: (619) 594-5394 FAX: (619) 594-5676 Email:
espin_r@unr.edu
Email: rether@sunstroke.sdsu.edu
(author
for correspondence)
The Liolaeminae (Iguania: Tropiduridae) is a large
clade of small to moderate-sized lizards that live in primarily arid
and semiarid habitats throughout most of austral South America.
They are a conspicuous component of the terrestrial vertebrate
fauna wherever they occur, and many localities are occupied by
several species. As this bibliography clearly indicates, lizards
of the Liolaeminae have been the focus of intense interest by
biologists for more than a century.
In 1989, Frost and Etheridge proposed formal
recognition of Liolaeminae as a subfamily of the iguanian family
Tropiduridae. Although
species of Liolaeminae have been coupled with a large number of
generic names (see Taxonomy), Etheridge (1995) recognized only three
as valid: Ctenoblepharys (1 species), Phymaturus (10 species) and Liolaemus (~ 150 species). Ctenoblepharys
adspersa is an
arenicolous insectivore confined to the Pacific beaches of central
Perú, where it occurs in isolation from other Liolaeminae.
Phymaturus are saxicolous,
herbivorous lizards that live at high elevations in the Andes of
central Chile and Argentina, and along the volcanic mesetas in
central and southern Argentina.
Throughout its range Phymaturus occurs
sympatrically with one or more species of Liolaemus. The number of valid species
of Liolaemus as of this writing
stands at approximately 150the uncertainty resulting from
differing opinions on the validity of some forms.
Eleven species have from two to five subspecies. The number of
Liolaemus has been growing at a rate of about
four new species each year for the past decade (Fig. 1), and at least
20 manuscript names known to us are now in preparation.
Fig. 1. Number
of Liolaemus taxa described over
time. Only taxa that are currently
considered valid are listed (see list at end of the bibliography). Incredibly, the rate of
new-species descriptions has averaged 2.7 per year since 1970 and has
risen to 3.5 per year within the last decade.
Liolaemus ranges from northern central
Perú southward through Bolivia, Chile, and Argentina to
northern Tierra del Fuego, and up the Atlantic coast of Uruguay and
southeastern Brazil. Members
of this genus occur at elevations from sea level to over 5000 m. In body size they range from
45 to just over 100 mm snout-vent length, with body masses from three
to nearly 200 g. Liolaemus
may be insectivorous or herbivorous, but the majority of species
appear to be omnivores. The
reproductive mode of Ctenoblepharys is unknown, but
it is probably oviparous. Phymaturus are viviparous, as
are about half the species of Liolaemus,
whereas the remaining species deposit eggs in varying stages of
embryonic development. The
majority of liolaemine lizards live in arid or semiarid habitats, but
the distributions of some Liolaemus extend into the humid
forest of southern Chile, and along the Atlantic coast including
several Brazilian islands. Liolaeminae
are almost entirely terrestrial, with most being microhabitat
generalists, but some are highly specialized morphologically: Ctenoblepharys
adspersa and some Liolaemus for life on aeolian
sand, and Phymaturus and some Liolaemus for life among the
crevices in outcrops of boulders.
Only one species of Liolaemus
is commonly found on trees.
Clearly the large number of species, the great range in
latitude and elevation, the occurrence of multiple syntopic species
in many regions, and the great diversity in characteristics such as
body size, morphology, diet, reproductive mode, and microhabitat
selection insures that these lizards will be the subjects of
biological investigations for many years to come. Our purpose in preparing
this bibliography and lists of valid names and synonymies is to help
facilitate those studies.
Content & How to Use
This Bibliography
The bibliography contains references written in
English, French, German, Italian, Latin, Portuguese, and Spanish, and
spans about three centuries of research.
We made every effort to locate the original work for each
citation in an attempt to avoid the errors associated with copying
bibliographies from secondary sources. We also attempted to include
the full citations for the references rather than to use
abbreviations; however, in a few cases we were unable to locate the
primary reference and thus had to rely on citations from other
resources. Originally,
this bibliography included 43 secondary references in which only
earlier works were cited, and 114 citations of published abstracts of
papers given at meetings. Because
of page limitations, these citations have been removed. Despite our attempts to make this bibliography as
error-free and comprehensive as possible, undoubtedly we have made
mistakes and missed some citations.
With the help of our colleagues we hope that we can remedy
this shortfall and keep the list up to date by receiving notices or
reprints of work relating to this fascinating group of
lizards.
The references are ordered alphabetically, by the last
name of the first author, and next by the year the paper was
published. In cases of
multiple authorship, the same principle above follows, with the
citations listed by the second authors last name and so forth. Because publication dates
are sometimes not the same as print dates, some citations are
followed by a year in brackets (e.g., [1991]).
In such instances, the year for the reference should be cited
(or searched for) using the publication date not the bracketed print
date.
A t.d. before a species name in an
annotation indicates that the reference is the source of the original
type description for a given taxon. The spelling of taxa in
annotations follows the orthography of the publications cited and
no assumptions
were made
regarding
the current identity of these taxa. Users
should refer to the Taxonomy section at the end of the bibliography
for synonomys and current useage.
Annotations are lacking for
many shorter
publications for which the contents are explicitly stated
in the title.
We are aware that the utility of this bibliography
would be greatly enhanced by the addition of one or more indexes.
We hope that future editions may include indexes, and
encourage those who are interested in particular aspects of
liolaemine biology (e.g., behavior, life history, taxonomy, etc.) to
prepare indexes for their topics of interest and share them with our
fellow researchers. We
encourage the users of this bibliography and taxonomy to send
additions and corrections to the senior author at
rether@sunstroke.sdsu.edu.
Acknowledgments
Many of our colleagues and friends assisted by sending
their references or reprints and, were it not for their efforts,
assembling this bibliography would have been much more difficult.
Among them, we would like to thank Luciano Avila, Dan
Blackburn, Félix Cruz, Kevin de Queiroz, Lee Fitzgerald,
Lígia Krause, Fernando Lobo, Ricardo Montero, Jimmy McGuire,
Scott Moody, Herman Núñez, Martha Ramírez
Pinilla, Fred Rocha, Miguel Rodriguez, Kurt Schwenk, Norm Scott,
Laura Vega, Van Wallach, and Jorge Williams.
We would like to extend a special thanks to Jorge Williams who
allowed us to incorporate numerous citations from a similar
bibliography that he was preparing. We thank Heather Powell and
the librarians directing the interlibrary loan services at San Diego
State University and at the University of Nevada, Reno for helping us
to procure some of the more difficult-to-obtain references.
Carrie Carreño, Fernando Lobo, and Heather Powell
helped to check earlier versions of the bibliography for errors.
During the preparation of this bibliography, the junior author
was supported by a Porter Fellowship from the American Society of
Physiologists and the Biological Resources Research Center at the
University of Nevada, Reno.