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An Examination of the Breeding System and Morphological Characters of Species in the
Genus Triodanis
Colette Berg, Lauren Kawulicz
Dr. Steven J. Franks, Dr. Jenn Weber
Triodanis (Campanulaceae), commonly known as Venus’ Looking Glass,
consists of seven species. The evolutionary relationships, breeding systems and
distinguishing traits of Triodanis have not yet been well described. Plants of genus
Triodanis have an interesting breeding system with two types of flowers:
cleistogamous and chasmogamous. Chasmogamous flowers are open to pollination
by insects and can either outcross or self-fertilize. Cleistogamous flowers remain
closed throughout the season and always self-fertilize. Individual Triodanis plants
can have both cleistogamous and chasmogamous flowers at the same time. However,
it is not yet known how variable species of the genus Triodanis are in regards to
breeding system or morphological traits. The presence of cleistogamy in a genus is
useful for studying the evolutionary and ecological factors important for plant
breeding systems. The New York Botanical Gardens Herbarium has pressed samples
of all seven Triodanis species: T. biflora, T. perfoliata, T. lamprosperma, T.
leptocarpa, T. coloradoensis, T. holzingeri, and T. texana. We collected trait, range,
and phenological data for these species, and on two Triodanis outgroups. This data
provides crucial information on the Triodanis portions of the phylogenetic tree of
Campanulaceae. All of this work is essential to perform phylogenic contrasts among
species, and to inform future studies in our lab in which we will seek to understand
the evolutionary pathways by which cleistogamy may be favored and how it
evolved. Further, these morphology-based trees will be combined with molecular
data to generate the first well-resolved trees of Triodanis.
• We collected data from samples in the New York Botanical Gardens Herbarium.
• We collected from the seven species of the genus Triodanis as well as two
outgroups, Campanula americana and Heterocodon rariflorum.
• Campanula americana is a recent outgroup, while Heterocodon rariflorum is a
more distant outgroup (Wendling, Galbreath, Dechaine , Knapp 2011).
• We took data from each species with a sample size of n ~50 individuals, covering a
broad geographic range.
• The measured traits were petal length, leaf length, leaf width, length of the base of
the capsule to the pore and total length of the capsule. These traits are traditionally
used to distinguish different species in Triodanis, particularly capsule length and
pore location.
• We measured these traits in millimeters using digital calipers.
• We constructed the clade using a program called Mesquite. This program uses the
principle of parsimony to construct the likely evolutionary relationships between
the species. Parsimony assumes the simplest explanation for divergent traits in
organisms which arose in the smallest number of steps.
Members of the genus Triodanis can be found in nearly every state. Texas appears
to be the center of biodiversity. These ranges come from the USDA Plants
Database.
Abstract
Methods
Species Ranges in genus Triodanis
Clade of Genus Triodanis
Gara, Brian, and Gayle Muenchow. "Chasmogamy/Cleistogamy in Triodanis Perfoliata (Campanulaceae):
Some CH/CL Comparisons in Fitness Parameters." American Journal of Botany 77.1 (1990): 1-6. Print.
Maddison, W. P. and D.R. Maddison. 2011. Mesquite: a modular system for evolutionary analysis.Version
2.75 http://mesquiteproject.org
Marcus, Joseph A. Triodanis Biflora. 2003. Lady Bird Johnson Wildflower Center, University of Texas at
Austin. Lady Bird Johnson Wildflower Center Image Gallery. Web. 5 Apr. 2014. <http://www.wildflower.
org/gallery/result.php?id_image=13704>.
Nhon, Crowel and Cellinese. Unpublished. Evolution & biogeography of Triodanis and Legousia
(Bellflowers) in North America and the Mediterranean
Patrick J. Alexander @ USDA-NRCS PLANTS Database
USDA, NRCS. 2014. Range maps from plant profiles of each species in genus Triodanis. The PLANTS
Database. Triodanis. (http://plants.usda.gov, 5 April 2014). National Plant Data Team, Greensboro, NC
27401-4901 USA.
Wendling, Barry M., Kurt E. Galbreath, and Eric G. Dechaine. "Resolving the Evolutionary History of
Campanula (Campanulaceae) in Western North America." Ed. Michael Knapp. PLoS ONE 6.9 (2011):
E23559. Print.
References
Average Trait Values
Franks Lab
Fordham
University
Morphology-based Clade v. Molecular Clade
Our clade is based on morphology; we gathered data about the organism’s phenotype. A molecular clade
would be based on the genotypes of the species. Before biologists had current technology, all cladograms were
morphology-based. Now that researchers can map DNA relatively quickly and inexpensively, many are
constructing molecular cladograms. Distinguishing among species of Triodanis in the field is quite difficult, due
to relatively few morphological differences and potentially substantial hybridization.
A lab group from the University of Florida (Nhon, Crowel and Cellinese) is currently constructing a
molecular cladogram of the genus Triodanis and closely related groups. Our cladogram does not completely
agree with the molecular phylogeny (in progress). This is not surprising, because species of the genus
Triodanis are morphologically very similar and discerning among species can be extremely difficult. The
molecular work of Nohn et al.(pers comm.) suggests a single introduction into North America for Triodanis,
and subsequent diversification into the current species groups. Though the molecular phylogeny revealed
Triodanis to be monophyletic, divergence between some species was very recent (1.42-0.87 mya).
(mm)
Triodanis
biflora
Triodanis
leptocarp
a
Triodanis
perfoliata
Triodanis
coloradoen
sis
Triodanis
texana
Triodanis
lamprosperm
a
Triodanis
holzinger
i
Campanula
americana
Heterocodon
rariflorum
Petal
Length
Average 7.69 6.776 7.96 9.468 8.692 7.711 6.424 12.649 3.563
Standard
Error 0.31184 0.20786 0.26121 1.92918 1.10780 0.28621 0.61812 0.53237 0.17729
Leaf
Length
Average 11.504 18.033 10.997 26.49 14.6935 10.736 12.11 87.67 6.131
Standard
Error 0.49708 0.62516 0.46042 1.13824 1.43565 0.43208 0.56059 5.19714 0.24438
Leaf Width
Average 6.52 3.09 12.169 5.046 8.775 10.252 8.348 31.567 6.135
Standard
Error 0.31123 0.92470 0.44525 0.28880 1.19340 0.60620 0.32222 2.27615 0.31929
Base to
Pore
Length
Average 3.596 2.168 10.09 3.379 4.489 3.261 7.791 0.5
Standard
Error 0.14700 0.10556 0.53280 1.17862 0.27581 0.16864 0.51934 0
Capsule
Length
Average 5.464 11.73 5.13 13.695 5.25 6.73 6.676 11.009 2.729
Standard
Error 0.16749 0.41022 0.15606 0.53787 0.34392 0.36156 0.18778 0.17016 0.09981
Triodanis biflora
Triodanis holzingeri
Triodanis coloradoensis
Triodanis lamprosperma
Triodanis leptocarpa
Triodanis perfoliata
Triodanis texana
Triodanis perfoliata Triodanis biflora
Species
Trait
s
Acknowledgements
Many thanks to Dr. Steve Franks, Dr. Jenn Weber and Beth Ansaldi, and to all the members of the Franks lab
for their collaboration and guidance. Special thanks to Nohn et al. for their personal communication, and to
Stella Sylva and the NYBG Herbarium staff for being helpful and welcoming.

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Morphological Cladogram of Genus Triodanis

  • 1. An Examination of the Breeding System and Morphological Characters of Species in the Genus Triodanis Colette Berg, Lauren Kawulicz Dr. Steven J. Franks, Dr. Jenn Weber Triodanis (Campanulaceae), commonly known as Venus’ Looking Glass, consists of seven species. The evolutionary relationships, breeding systems and distinguishing traits of Triodanis have not yet been well described. Plants of genus Triodanis have an interesting breeding system with two types of flowers: cleistogamous and chasmogamous. Chasmogamous flowers are open to pollination by insects and can either outcross or self-fertilize. Cleistogamous flowers remain closed throughout the season and always self-fertilize. Individual Triodanis plants can have both cleistogamous and chasmogamous flowers at the same time. However, it is not yet known how variable species of the genus Triodanis are in regards to breeding system or morphological traits. The presence of cleistogamy in a genus is useful for studying the evolutionary and ecological factors important for plant breeding systems. The New York Botanical Gardens Herbarium has pressed samples of all seven Triodanis species: T. biflora, T. perfoliata, T. lamprosperma, T. leptocarpa, T. coloradoensis, T. holzingeri, and T. texana. We collected trait, range, and phenological data for these species, and on two Triodanis outgroups. This data provides crucial information on the Triodanis portions of the phylogenetic tree of Campanulaceae. All of this work is essential to perform phylogenic contrasts among species, and to inform future studies in our lab in which we will seek to understand the evolutionary pathways by which cleistogamy may be favored and how it evolved. Further, these morphology-based trees will be combined with molecular data to generate the first well-resolved trees of Triodanis. • We collected data from samples in the New York Botanical Gardens Herbarium. • We collected from the seven species of the genus Triodanis as well as two outgroups, Campanula americana and Heterocodon rariflorum. • Campanula americana is a recent outgroup, while Heterocodon rariflorum is a more distant outgroup (Wendling, Galbreath, Dechaine , Knapp 2011). • We took data from each species with a sample size of n ~50 individuals, covering a broad geographic range. • The measured traits were petal length, leaf length, leaf width, length of the base of the capsule to the pore and total length of the capsule. These traits are traditionally used to distinguish different species in Triodanis, particularly capsule length and pore location. • We measured these traits in millimeters using digital calipers. • We constructed the clade using a program called Mesquite. This program uses the principle of parsimony to construct the likely evolutionary relationships between the species. Parsimony assumes the simplest explanation for divergent traits in organisms which arose in the smallest number of steps. Members of the genus Triodanis can be found in nearly every state. Texas appears to be the center of biodiversity. These ranges come from the USDA Plants Database. Abstract Methods Species Ranges in genus Triodanis Clade of Genus Triodanis Gara, Brian, and Gayle Muenchow. "Chasmogamy/Cleistogamy in Triodanis Perfoliata (Campanulaceae): Some CH/CL Comparisons in Fitness Parameters." American Journal of Botany 77.1 (1990): 1-6. Print. Maddison, W. P. and D.R. Maddison. 2011. Mesquite: a modular system for evolutionary analysis.Version 2.75 http://mesquiteproject.org Marcus, Joseph A. Triodanis Biflora. 2003. Lady Bird Johnson Wildflower Center, University of Texas at Austin. Lady Bird Johnson Wildflower Center Image Gallery. Web. 5 Apr. 2014. <http://www.wildflower. org/gallery/result.php?id_image=13704>. Nhon, Crowel and Cellinese. Unpublished. Evolution & biogeography of Triodanis and Legousia (Bellflowers) in North America and the Mediterranean Patrick J. Alexander @ USDA-NRCS PLANTS Database USDA, NRCS. 2014. Range maps from plant profiles of each species in genus Triodanis. The PLANTS Database. Triodanis. (http://plants.usda.gov, 5 April 2014). National Plant Data Team, Greensboro, NC 27401-4901 USA. Wendling, Barry M., Kurt E. Galbreath, and Eric G. Dechaine. "Resolving the Evolutionary History of Campanula (Campanulaceae) in Western North America." Ed. Michael Knapp. PLoS ONE 6.9 (2011): E23559. Print. References Average Trait Values Franks Lab Fordham University Morphology-based Clade v. Molecular Clade Our clade is based on morphology; we gathered data about the organism’s phenotype. A molecular clade would be based on the genotypes of the species. Before biologists had current technology, all cladograms were morphology-based. Now that researchers can map DNA relatively quickly and inexpensively, many are constructing molecular cladograms. Distinguishing among species of Triodanis in the field is quite difficult, due to relatively few morphological differences and potentially substantial hybridization. A lab group from the University of Florida (Nhon, Crowel and Cellinese) is currently constructing a molecular cladogram of the genus Triodanis and closely related groups. Our cladogram does not completely agree with the molecular phylogeny (in progress). This is not surprising, because species of the genus Triodanis are morphologically very similar and discerning among species can be extremely difficult. The molecular work of Nohn et al.(pers comm.) suggests a single introduction into North America for Triodanis, and subsequent diversification into the current species groups. Though the molecular phylogeny revealed Triodanis to be monophyletic, divergence between some species was very recent (1.42-0.87 mya). (mm) Triodanis biflora Triodanis leptocarp a Triodanis perfoliata Triodanis coloradoen sis Triodanis texana Triodanis lamprosperm a Triodanis holzinger i Campanula americana Heterocodon rariflorum Petal Length Average 7.69 6.776 7.96 9.468 8.692 7.711 6.424 12.649 3.563 Standard Error 0.31184 0.20786 0.26121 1.92918 1.10780 0.28621 0.61812 0.53237 0.17729 Leaf Length Average 11.504 18.033 10.997 26.49 14.6935 10.736 12.11 87.67 6.131 Standard Error 0.49708 0.62516 0.46042 1.13824 1.43565 0.43208 0.56059 5.19714 0.24438 Leaf Width Average 6.52 3.09 12.169 5.046 8.775 10.252 8.348 31.567 6.135 Standard Error 0.31123 0.92470 0.44525 0.28880 1.19340 0.60620 0.32222 2.27615 0.31929 Base to Pore Length Average 3.596 2.168 10.09 3.379 4.489 3.261 7.791 0.5 Standard Error 0.14700 0.10556 0.53280 1.17862 0.27581 0.16864 0.51934 0 Capsule Length Average 5.464 11.73 5.13 13.695 5.25 6.73 6.676 11.009 2.729 Standard Error 0.16749 0.41022 0.15606 0.53787 0.34392 0.36156 0.18778 0.17016 0.09981 Triodanis biflora Triodanis holzingeri Triodanis coloradoensis Triodanis lamprosperma Triodanis leptocarpa Triodanis perfoliata Triodanis texana Triodanis perfoliata Triodanis biflora Species Trait s Acknowledgements Many thanks to Dr. Steve Franks, Dr. Jenn Weber and Beth Ansaldi, and to all the members of the Franks lab for their collaboration and guidance. Special thanks to Nohn et al. for their personal communication, and to Stella Sylva and the NYBG Herbarium staff for being helpful and welcoming.