The genus
Quercus is well-known for having low barriers to gene flow
among different species. This hybridization can lead, in
areas where multiple taxa coincide, to the production of hybrid
individuals. These individuals often show intermediary
morphological characteristics to the two parental taxa. In
areas of widespread gene exchange and introgression this can
lead to a wide diversity of plant forms across the landscape.
The prevalence of hybridization in groups such as Quercus
has led to a difficulty in delimiting and even defining what a
species is.
While it can be
relatively straightforward to identify hybrid individuals once
the characters of the parental taxa are well-known, this
morphological determination of the prevalence of hybridization
can be misleading and not indicate the full extent of the hybrid
occurrence. Genetic data are thus key to determine the
relative genomic contribution from various parental forms.
The Four Corners area
of Arizona, Colorado, New Mexico, and Utah is principally
dominated by two species of Quercus, Q. gambelii
and Q. turbinella.
Across the region various hybrid populations and
individuals are known which have been commonly referred to as
Q. undulata. Quercus undulata is a variable complex of
hybrid derived forms extending across the southwestern United
States and influenced by numerous parental taxa (Tucker, 1961;
Tucker et al., 1961). The Four Corners area also shows a few
more geographically restricted forms of
Quercus. Plants from the Navajo Basin of Utah and
Arizona have normally been treated as a distinct variety of
Q. havardii a species better known from NW Texas and
adjacent New Mexico and Oklahoma. This taxon, Q.
havardii
var. tuckerii, has been hypothesized to represent a
variable hybrid form and may not be allied to Q. havardii.
Following this rational it was recently transferred to a
descretely named taxon, Q. welshii. An additional
discrete variety of Q. gambelii has also recently been
described from SE Utah, Q. gambelii var. bonina.
Range maps ©
Flora of North America
Seminar Goals
The goal of this seminar is to lean how natural
hybridization can act as a means of speciation and evaluate
patterns of hybridization in a group in which hybridization is
known to be an important factor in evolution. To do this
we will be studying Quercus across the Four Corners
region to evaluate the extent and patterns of hybridization
using a variety of organismic biology tools. While we will
all be studying the same system differing groups of students
will focus on the completion of different parts of the study.
1. Mapping of distribution of parental species
using herbarium specimen databases and GIS.
2. Morphometric analysis of leaf form among
various taxa.
3. Genetic analysis using Inter-simple Sequence
Repeat markers (ISSR)
Calendar for
Student Presentations
6 October - Rae Landers -
Taxonomic identity of
Quercus coccifera L. in the
Iberian Peninsula is maintained in spite of widespread
hybridisation, as revealed by morphological, ISSR and ITS
sequence data.
20 October - Matthew Schendel -
Morphometric variation in oaks of the Apostle Islands in
Wisconsin: Evidence for hybridization between Quercus rubra
and Quercus ellipsoidalis (Fagaceae).
27 October - Eric Ireland -
Assessing the threat from hybridization to the rare endemic
Physaria bellii Mulligan (Brassicaceae).
3 November - Brenna Christie -
Gene flow and hybridisation in a mixed oak forest (Quercus
pyrenaica Willd. and Quercus petraea
(Matts.) Liebl.) in central Spain.
10 November - Haley Nichols -
Natural
hybridization and hybrid zones between Quercus crassifolia
and Quercus crassipes (Fagaceae) in Mexico:
Morphological and molecular evidence.
Papers for
Student Presentations
Each title links to a PDF of
the article.
Borazan,
A. and M. T. Babaç. 2003.
Morphometric
leaf variation in oaks (Quercus) of Bolu, Turkey.
Ann. Bot. Fennici 40: 233-242.
González-Rodríguez, A, D. M. Arias, S.
Valencia, and K. Oyama. 2004.
Morphological and
RAPD analysis of hybridization between Quercus affinis
and Q. laurina (Fagaceae), Two Mexican red oaks.
American Journal of Botany 91: 401-409.
González-Rodríguez, A, J. F. Bain, J.L.
Golden, and K. Oyama. 2004.
Chloroplast DNA
variation in the Quercus affinis–Q. laurina complex in
Mexico: geographical structure and associations with nuclear and
morphological variation. Molecular Ecology 13: 3467-3476.
Hokanson,
S.C., J.G. Isebrands, R.J. Jensen, and J.F. Hancock, 1993.
Isozyme
variation in oaks of the Apostle Islands in Wisconsin: genetic
structure and levels of inbreeding in Quercus rubra and
Q. ellipsoidalis (Fagaceae). American Journal of
Botany 80: 1349-1357.
Howard,
D. J. R. W. Preszler, J. Williams, S. Fenchel, and W. J.
Boecklen. 1997.
How discrete
are oak species? Insights from a hybrid zone between Quercus
grisea and Quercus gambelli. Evolution 51:
747-755.
Jensen, R. J., S. C. Hokanson, J. G. Isebrands, and J. F.
Hancock. 1993.
Morphometric variation in oaks of the Apostle Islands in
Wisconsin: Evidence for hybridization between Quercus rubra
and Quercus ellipsoidalis (Fagaceae). American
Journal of Botany 80: 1358-1366.
Kothera, L., S. M. Ward, S. E. Carney.
2007.
Assessing the threat from hybridization to the rare endemic
Physaria bellii Mulligan (Brassicaceae). Biological
Conservation 140: 110-118.
Lau, C. P. Y., L. Ramsden, and R. M.
K. Saunders. 2005.
Hybrid
origin of "Bauhinia blakeana" (Leguminosae:
Caesalpinioideae), inferred using morphological, reproductive,
and molecular data. American Journal of Botany 92: 525-533.
Muir, G. and C. Schlötterer. 2005.
Evidence for shared
ancestral polymorphism rather than recurrent gene flow at
microsatellite loci differentiating two hybridizing oaks (Quercus
spp.). Molecular Ecology 14: 459-561.
Rubio de Casas, R., E. Cano, L.
Balaguer, E. Pérez-Corona, E. Manrique, C. García-Verdugo, P.
Vargas. 2007.
Taxonomic identity of Quercus coccifera L. in the
Iberian Peninsula is maintained in spite of widespread
hybridisation, as revealed by morphological, ISSR and ITS
sequence data. Flora 202: 488-499.
Tovar-Sánchez,
E and K. Oyama. 2004.
Natural
hybridization and hybrid zones between Quercus crassifolia
and Quercus crassipes (Fagaceae) in Mexico:
Morphological and molecular evidence. American Journal of
Botany 91: 1352-1363.
Valbuena-Carabaña, M, S. C. González-Martínez,
V.L. Sork, C. Collada, A. Soto, P.G. Goicoechea, and L. Gil.
2005.
Gene flow and hybridisation in a mixed oak forest (Quercus
pyrenaica Willd. and Quercus petraea
(Matts.) Liebl.) in central Spain. Heredity 95: 457-465.
Zalapa, J. E., J. Brunet and R. P. Guries. 2009.
Patterns of
hybridization and introgression between invasive Ulmus pumila
(Ulmaceae) and native U. rubra. American Journal of
Botany 96: 1116-1128.
Important
Literature (Useful background reading for proposal preparation)
The following literature
should serve as a base for your understanding of natural
hybridization in Quercus and the specifics of the Four
Corners species of Quercus.
Anderson, E. and G. L. Stebbins, Jr. 1954.
Hybridization as an evolutionary stimulus. Evolution 8:
378-388.
Baack, E. J. and L. H. Rieseberg. 2007.
A genomic view of introgression and hybrid speciation.
Current Opinion in Genetics & Development 2007. 17: 513–518.
Cristofolini, G. and S. Crema. 2005.
A morphometric study of the Quercus crenata species
complex (Fagaceae). Botanica Helvetica 115: 155-167.
Mallet, J. 2007.
Hybrid speciation. Nature 446: 279-283.
Petit,
R., C. Bodenes, A. Ducousso, G. Roussel, and A. Kremer. 2003.
Hybridization
as a mechanism of invasion in oaks. New Phytologist 161:
151-164.
Rieseberg, L. H. 1995.
The Role of
Hybridization in Evolution: Old Wine in New Skins. American
Journal of Botany 82: 944-953.
Soltis,
P. S. and D. E. Soltis. 2009. The role of hybridization in plant
speciation. Annual Review of Plant Biology 60: 561-588.
Tucker,
J. M. 1961.
Studies
in the Quercus undulata complex. I. A preliminary
statement. American Journal of Botany 48: 202-208.
Tucker,
J. M. 1963.
Studies in the Quercus undulata complex. III. The
contribution of Q. arizonica. American Journal of
Botany 50: 699-708.
Tucker, J. M. 1970.
Studies in the
Quercus undulata complex. IV. The contribution of Quercus
havardii. American Journal of Botany 57: 71-84.
Tucker,
J. M. 1971.
Studies in the Quercus undulata. V. The type of
Quercus undulata. American Journal of Botany 58:
329-341.
Tucker,
J. M., W. P. Cottam, and R. Drobnick. 1961.
Studies in the
Quercus undulata complex. II. The contribution of Quercus
turbinella. American Journal of Botany 48:
329-339.
Van
Valen, L. 1976.
Ecological species, multispecies, and oaks. Taxon 25: 233-239.
Welsh,
S. L. 1986.
Quercus (Fagaceae) in the Utah flora. Great Basin
Naturalist 46: 107-111.
Whittemore, A. T. and B. A. Schaal. 1991.
Interspecific gene flow in sympatric oaks. Proceedings of
the National Academy of Sciences, USA. 88: 2540-2544.
Last Updated 26 March 2010
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