Sisyrinchium humahuacense: New Plant Species from Argentina

See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/331498646
Sisyrinchium humahuacense of sect. Segetia a new species from Argentina
Article in Plant Biosystems · March 2019
DOI: 10.1080/11263504.2019.1580226
CITATIONS
READS
0
53
2 authors:
Christian A Zanotti
Agostina B. Sassone
Instituto de Botánica Darwinion
Leibniz Institute of Plant Genetics and Crop Plant Research
64 PUBLICATIONS 88 CITATIONS
24 PUBLICATIONS 60 CITATIONS
SEE PROFILE
SEE PROFILE
Some of the authors of this publication are also working on these related projects:
Monocots: Breeding system in Poa (Pooideae); Evolution in Jarava (Stipeae); Leucocoryneae (Amaryllidaceae); Tillandsia (Bromeliaceae); Orchids; Endophytes and
Dune vegetation View project
Evolutionary and Systematic studies in the tribe Leucocoryneae (AMARYLLIDACEAE, ALLIOIDEAE) View project
All content following this page was uploaded by Christian A Zanotti on 19 December 2020.
The user has requested enhancement of the downloaded file.
Plant Biosystems - An International Journal Dealing with
all Aspects of Plant Biology
Official Journal of the Societa Botanica Italiana
ISSN: 1126-3504 (Print) 1724-5575 (Online) Journal homepage: https://www.tandfonline.com/loi/tplb20
Sisyrinchium humahuacense of sect. Segetia a new
species from Argentina
Christian A. Zanotti & Agostina B. Sassone
To cite this article: Christian A. Zanotti & Agostina B. Sassone (2019): Sisyrinchium
humahuacense of sect. Segetia a new species from Argentina, Plant Biosystems - An International
Journal Dealing with all Aspects of Plant Biology, DOI: 10.1080/11263504.2019.1580226
To link to this article: https://doi.org/10.1080/11263504.2019.1580226
Published online: 04 Mar 2019.
Submit your article to this journal
View Crossmark data
Full Terms & Conditions of access and use can be found at
https://www.tandfonline.com/action/journalInformation?journalCode=tplb20
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY
https://doi.org/10.1080/11263504.2019.1580226
Sisyrinchium humahuacense of sect. Segetia a new species from Argentina
Christian A. Zanotti
and Agostina B. Sassone
Instituto de Botanica Darwinion, San Isidro, Argentina
ABSTRACT
ARTICLE HISTORY
A new species from the Andean region of northwestern Argentina is described and illustrated:
Sisyrinchium humahuacense. This species can be distinguish from its closest related species by three
main characters: the presence of a conspicuous horizontal rhizome, the colour of the flowers and the
presence of rudimentary leaves not splitting in fibers. A phylogenetic analysis based on DNA sequences supported the inclusion of S. humahuacense within the genus and as part of the section Segetia.
Also, a key to species to identify S. humahuacense is presented herein.
Received 23 May 2018
Revised 11 January 2019
Accepted 29 January 2019
Introduction
The New World tribe Sisyrinchieae (Iridaceae) encompassed
six genera: Libertia Spreng. (ca. 12 spp.), Olsynium Raf. (ca. 12
spp.), Orthrosanthus Sweet (10 spp.), Sisyrinchium L. (ca. 140
spp.), Solenomelus Miers (2 spp.) and Tapeinia Juss. (1 spp.).
The tribe is mainly distributed in temperate or tropical
regions of America (Goldblatt and Manning 2008; Karst and
Wilson 2012; Inacio et al. 2017) and South America was proposed to be the place of origin (Cocucci and Vogel 2001;
Chauveau et al. 2012). In Argentina, 52 taxa are cited, some
of them are endemic (Roitman et al. 2008) and can be found
in various habitats and growing from wet to dry grasslands,
bedrocks, prairies, swamps and also in anthropic modified
environments or in forest edges (Inacio et al. 2017).
Sisyrinchium is characterized by being perennial rhizomatous
herbs with plane or terete leaves; the basic inflorescence is a
terminal or axillary rhipidium; flowers have subequal tepals,
usually patent, in white, blue, purple, pink or yellow colour;
the androecium has filaments basally or totally connate as a
staminal column with the presence or not of trichomal elaiophores (see Cocucci and Vogel 2001; Chauveau et al. 2012);
the style of the gynoecium is entire or divided into three
branches, and seeds are globose or subglobose presenting a
dark (brown to blackish) seed coat (Goldblatt and Manning
2008; Karst and Wilson 2012; Inacio et al. 2017). Sisyrinchium
showed to be monophyletic according to morphological
(Goldblatt et al. 1990) and molecular data (Chauveau et al.
2011; Karst & Wilson 2012; Inacio et al. 2017) but there are
still problems in the circumscription of taxa by the lack of
precise species descriptions (Fachinetto et al. 2018). In addition, diagnostic characters for the recognition of species are
usually lost during the herborization process and this fact
has contributed to the great disparity in the number of recognized taxa (Ceja-Romero et al. 2009).
CONTACT Christian A. Zanotti
B1642HYD, Argentina.
ß 2019 Societa Botanica Italiana
Published online 04 Mar 2019
[email protected]
KEYWORDS
Iridaceae; ITS; psbA-trnH;
Southern Cone; Taxonomy
Over the last two centuries, infrageneric classifications of
Sisyrinchium were traditional based on morphological data.
However, recently Inacio et al. (2017) showed that these classifications based only on morphological data did not reflect
phylogenetic relationships, especially when recent phylogenetic divergences, reticulate evolution, ongoing speciation and
high phenotypic plasticity are detected within Sisyrinchium
(cfr. Henderson 1976; Cholewa and Henderson 1984; Tacuatia
et al. 2012a, 2012b, 2017). Moreover, the authors divided the
genus into 10 monophyletic sections: Hydastylus (Dryand. ex
Salisb.) Ravenna, Segetia Ravenna, Echthronema (Herb.) Benth.
& Hook.f., Spathirhachis Klotzsch ex Klatt, Viperella Ravenna,
Cephalanthum Baker, Trichoparcus C.D.Inacio, Chauveau &
L.Eggers, Morphanthus C.D.Inacio, Chauveau & L.Eggers,
Rhizilineum C.D.Inacio, Chauveau & L.Eggers, and Sisyrinchium
Lem. ex Klatt. However, the most derived infrageneric subdivisions (e.g. Sect. Morphanthus, Sect. Rhizilineum and Sect.
Sisyrinchium) were not easy to distinguish using these data
and the phylogenetic relationships among these sections
remained unresolved.
In a recent floristic study of the Argentinean flora, we
found a new species of Sisyrinchium Sect. Segetia endemic to
the High Andes of northwestern Argentina. The aim of this
study is to describe a new species and illustrate it. Data on
its habitat, distribution, ecology and phenology are also provided. We include a key to species of Sisyrinchium sect.
Segetia Ravenna and discuss the position of this new taxon
based on morphological and molecular data.
Material and methods
Taxonomic study
The morphological study was based on collections of
Sisyrinchium stored at SI (acronyms according to Thiers 2018,
Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro, Buenos Aires
2
C. ZANOTTI AND A. SASSONE
continuously updated) and the examination of type specimens
was performed through the images available at JSTOR (http://
plants.jstor.org). Sectional treatment of Sisyrinchium followed
Inacio et al. (2017). In order to assess the taxonomic position of
the analyzed taxa, an exhaustive review of the literature was
undertaken. The terminology applied to the descriptions follows
Goldblatt and Manning (2008). A distribution map was
assembled from coordinates reported on specimen labels using
the package “raster” (Hijmans 2017) available in R 3.2.2 (R Core
Team 2016), and then edited with Inkscape v. 0.92 [free opensource SVG graphics editor (Bah 2009)].
DNA isolation, amplification, sequencing and
phylogenetic analyses
Genomic DNA was isolated from silica-dried leaf tissue following a modified CTAB protocol (Doyle and Doyle 1987).
Based on the resolution obtained by previous results for genetic markers in Iridaceae (Chauveau et al. 2011; Inacio et al.
2017), the plastid region psbA-trnH and the ribosomal internal
transcribed spacer (ITS1-5.8S-ITS2) were selected. DNA amplification was conducted in a volume of 25 ll polymerase chain
reaction (PCR) containing 20–40 ng of DNA template and a
final concentration of 1 PCR Buffer minus Mg, 2.5 mM MgCl2,
0.025 mM of each dNTP, 0.2 lM of both forward and reverse
primers and 1.25–2.5 U of Taq Polymerase (Invitrogen Life
Technologies, S~ao Paulo, Brazil) following the indications
described in Chauveau et al. (2011). To assess the phylogenetic position of the new taxa, 92 sequences representing 38
species, were downloaded from GenBank (https://www.ncbi.
nlm.nih.gov/genbank/) and detailed in Table 1.
Editing and assembling of sequences were conducted in
Chromas Pro version 1.34 (Technelysium Pty, Ltd., Tewantin,
Australia). Quality of sequences was assessed by visual
inspection of the chromatograms. Sequences were aligned
using MUSCLE integrated to MEGA5 (Tamura et al. 2011) and
then revised and edited manually using BioEdit (Hall 1999).
The results from different phylogenetic inference methods
were compared with reconstructed trees using both parsimony and Bayesian approaches were performed with TNT
ver. 1.1 (Goloboff et al. 2008) and MrBayes v.3.2.6
(Huelsenbeck and Ronquist 2001), respectively. Models of
molecular evolution were taken from Inacio et al. (2017). For
Bayesian analysis, three-independent runs were completed to
ensure that the analyses converged on the optimal tree set.
Each analysis implemented four simultaneous chains and ran
for 1 1010 generations. Tree space was sampled every
100th generation for a total sample of 10,000 trees per analysis. Each independent run reached stationary prior to the
20,000th generation. All output searches were analyzed with
TRACER (Rambaut et al. 2014) to determine convergence,
and we discarded the first 25% of trees as burn-in. Bayesian
and modeltests analyses were performed via the CIPRES
Gateway (Miller et al. 2010). Consensus trees were visualized
and edited using FigTree 1.4.3 (Rambaut 2009). Final figures
were edited using Inkscape v. 0.92 (Bah 2009). Sequences
produced in this study were deposited in GenBank and listed
in Table 1.
Table 1. Taxon and Genbank accession number of specimens included in
this study.
Taxon
Libertia chilensis (Molina) Gunckel
Libertia sessiliflora (Poepp.) Skottsb.
Neomarica candida Sprague
Olsynium biflorum (Thunb.) Goldblatt
Olsynium filifolium (Gaudich.) Goldblatt
Olsynium frigidum (Poepp.) Goldblatt
Olsynium philippii (Klatt) Goldblatt
Orthrosanthus multiflorus Sweet
S. alatum Hook.,
S. angustifolium Mill.
S. californicum (Ker Gawl.) Dryand.
S. chiricanum Woodson
S. convolutum Nocca
S. cuspidatum Poepp.
S. decumbens Ravenna
S. demissum Greene
S. foliosum I.M.Johnst.
S. graminifolium Lindl.
S. humahuacese (Voucher: Zuloaga et al. 14236 )
S. humahuacese (Voucher: Zuloaga et al.16351)
S. humahuacese (Voucher: Zanotti, C. et al. 1023)
S. hoehnei I.M.Johnst.
S. jamesonii Baker
S. jamesonii
S. jamesonii
S. jamesonii
S. laxinervium Ravenna
S. laxinervium
S. laxinervium
S. laxinervium
S. limarinum Ravenna
S. littorale Greene
S. marginatum Klatt
S. micranthum Cav.
S. minus Engelm. & A.Gray
S. minutiflorum Klatt
S. palmifolium L.
S. palustre Diels.,
S. platense I.M.Johnst.
S. praealtum Kraenzl.
S. praealtum
S. pusillum Kunth
S. pusillum
S. rosengurttii I.M.Johnst.
S. sellowianum Klatt
S. tinctorium Kunth
S. trinerve Baker
S. uliginosum Ravenna
S. vaginatum Spreng.
ITS
KX432914
KX432915
KX432916
KF577162
KF577192
KF577191
JN389212
KX432917
KF577161
MG215718
KF577179
KF577206
KF577195
KF577136
KX432925
KF577182
KF577152
KF577193
MH340538
MH340539
MH340540
KF577208
KF577131
KF577130
KF577127
KF577125
KX432934
KX432933
KF577132
KF577129
KF577194
JN389247
KX432939
KF577196
KF577150
KF577151
HQ607073
KF577124
HQ607075
KX432948
KF577126
KX432949
KF577121
KX432955
KF577157
KF577198
KX432961
KF577169
KX432962
psbA-TRNH
KX432555
KX432556
KX432557
KF577323
KF577353
KF577352
KF577365
KX432558
KF577322
KC704323
KF577341
KF577367
KF577298
KF577297
KX432566
KF577343
KF577313
KF577354
MH998362
MH998363
MH998364
KF577333
KF577289
KF577287
KF577293
KF577292
KX432575
KF577291
KX432574
KF577294
KF577355
KF577340
KX432579
KF577307
KF577311
KF577312
KF577315
KF577286
KF577316
KF577288
KX432589
KX432590
KF577283
KX432594
KF577318
KF577358
KX432601
KF577329
KF577350
Taxonomic treatment
Sisyrinchium humahuacense C.A. Zanotti, sp. nov.
(Figures 1 and 2)
Type
Argentina. Jujuy. Dep. Humahuaca. Ruta Provincial 13, Abra
camino a Iruya, 22 530 1600 S; 65 140 5800 W, 4000 m a.s.l, 12
March 2013, Zuloaga, F.O, Aagesen, L, N. Deginani, S.
Nomdedeu & C.A. Zanotti 14236 (holotype, SI [barcode
092977]!; isotype, BA!).
Description
Perennial herb 3–13 cm high with conspicuous horizontaloblique rhizomes up to 3 cm long and with coarse roots.
Leaves basal mixed with old castaneous and copious rudimentary leaves of the previous cycles not splitting into fine
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY
3
Figure 1. Sisyrinchium humahuacense. A. Plant. B. Detail of the adaxial surface of the blade. C. Cross section of the blade. D. Flower with four stamens and stigmatic
branches. E. Flower with three stamens and stigmatic branches. F. Fruit (All based on Zuloaga, F.O et al. 14236, holotype: SI, except F, from Zanotti & Panizza 1023).
fibers; basal leaves 6–9 0.2–0.4 cm, erect to slightly curved,
folded, linear-ensiform, glabrous, with 6–8 prominent veins,
apex acute, with basally scarious margin ca. 0.5 mm long.
and with wax plates in the rest of the margin. Scape
2–5 0.1–0.15 cm, simple, ancipitate, narrowly winged, with
a terminal bract. Terminal bract 2–2.5 0.2–0.3 cm, folded
with the dorsal face costate and with basally scarious margin.
Inflorescence of one rhipidia sessile, enclosed by spathes,
with 1–2(–3) flowers, opening one at a time. Spathes
1–2 0.1–0.4 cm acute, with the dorsal face costate and with
the margin scarious from the base to the apex, ca. 0. 1 cm
wide. Pedicel 12–15 mm long., glabrous, erect or curved,
sometimes longer than the spathe. Tepals 1–1.2 cm 1.5–4 mm, blue, subequal, spreading, oblanceolate, rounded.
Flowers with 3(4) stamens and stigmatic branches.
Filaments white, glabrous, basally connate for 1–2 mm, then
free for 2.5–3 mm, ascending to patent. Anthers 2.5–3 mm
long., yellow, dorsifixed, versatile, incurved at dehiscence.
Ovary glabrous, 2–2.5 1–1.5 mm. Style ca. 4 mm long,
white, glabrous, style branches 2.5–3 mm long., alternate to
the stamens, ascending to patent, stigma capitate, papillate.
Capsules ellipsoid, 0.4–0.5 0.3–0.35 cm, dark.
Phenology
Flowering and fruiting from February to March.
Ecology and distribution
Sisyrinchium humahuacence was collected in the High Andes
of north-western Argentina, in the province of Jujuy, between
3950 and 4200 m a.s.l, on a sandy or rocky and bare ground.
Among the most frequent species of the surrounding vegetation are: Adesmia minor (Hook. & Arn.) Burkart var. caespitosa
(Phil.) Ulibarri & Burkart, Astragalus crypticus I.M. Johnst.
Gutierrezia mandonii (Sch. Bip.) Solbrig, Pseudognaphalium tarapacanum (Phil.) Anderb., Spergula villosa Pers., Stevia tarijensis
Hieron. and Werneria villosa A. Gray. (Figure 3).
Etymology
The species name is a reference to the holotype was found
in the department of Humahuaca (Jujuy, Argentina).
Additional material examined
Argentina. Jujuy. Dpto. Humahuaca: Ruta Provincial 13, de
ndor, 4000 m a.s.l, 17 Feb 1997,
Iturbe a Iruya, Abra del Co
4
C. ZANOTTI AND A. SASSONE
Figure 2. Sisyrinchium humahuacense. A. In natural habitat on rocky soil. B. Idem, on sandy soil. C. D. Flower. E. Flower, fruit and terminal bract. F. Rhizome. A and
D from Zuloaga et al 14236. B, C, E and F, from Zanotti & Panizza 1023. Photos by Zanotti, C.A).
Figure 3. Distribution map indicating the four known localities of Sisyrinchium humahuacense in Jujuy, Argentina.
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY
5
Figure 4. Bayesian inference tree of the genus Sisyrinchium (Iridaceae) based on the combined data matrix of ITS and psbA-TRNH. Posterior probability values
>0.75 and bootstrap values >75% are indicated above branches.
Zuloaga et al. 5972 (SI); ıdem. loc., 4000 m. a.s.l., 15 Mar
2018, Zuloaga et al. 16351 (SI); ıdem loc., near the border
with the province of Salta towards Iruya, 3800 m. a.s.l., 15
Mar 2018, Zuloaga et al. 16345 (SI); ıdem loc., Abra Colorada,
3960 m a.s.l, 11 Feb 1998, Morrone et al. 2439 (SI). Dpto.
Tumbaya: Cuesta de Lipan, ladera norte del Cerro Lipan Sur,
23 430 1700 S, 65 390 4800 4197 m a.s.l, 25 Feb 2018, C. A. Zanotti
& A.M. Panizza 1023 (SI, BA).
Phylogenetic relationships
The aligned ITS sequence data matrix consists of 612 characters of which 167 were parsimony informative, whilst psbAtrnH matrix consists of 607 characters of which 57 were parsimony informative. The analyses of individual regions produced congruent results and then, the two matrices were
analyzed together. The combined matrix of ITS and psbAtrnH regions resulted in 1219 characters of which 224 were
parsimony informative. The analysis yielded four most parsimonious trees of length 851 steps (CI ¼ 0.55; RI ¼ 0.81). The
analyzed specimens of Sisyrinchium humahuacense were
placed into the monophyletic of S. sect. Segetia (PP ¼ 0.99;
BS MP ¼ 99; Figure 4). The three specimens included,
resulted monophyletic in all the parsimony trees explored,
and moderate support (PP ¼ 0.8, BS MP ¼ 90), but never
resulted nested with the other species of the section.
Key to species of Sisyrinchium Sect. Segetia based on Inacio
et al. (2017).
1. Plants with conspicuous horizontal-oblique rhizome, with
rudimentary leaves of previous cycles, never split into fine
fibers; flowers blue. Fruit ellipsoid . . . . . . . . . . .S. humahuacense
10 . Plants without horizontal-oblique rhizome, with rudimentary leaves of previous cycles splittied into fine fibers; flowers
yellow or white. Fruit obloid . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Terminal bracts longer (2 times) than the longest spathe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..3
20 . Terminal bracts equal or shorter than the largest spathe. 5
3. Leaves 3–5 mm wide . . . . . . . . . . . . . . . . . . . . S. praealtum
6
C. ZANOTTI AND A. SASSONE
30 . Leaves 1–2 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . ..4
Explorer project. Finally, our thanks to the reviewers and editor for
improving this manuscript.
4. Leaves longer or equal than the scape . . . . . . . S. jamesonii
40 . Leaves always shorter than the scape . . . . . . . . S. trinerve
5. Plants 25 cm long . . . . . . . . . . . . . . . . . S. unispathaceum
50 . Plants 5–15 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
ORCID
Christian A. Zanotti
http://orcid.org/0000-0002-4804-7655
6. Plants no more than 7 cm high . . . . . . . . . . . . . . S. pusillum
60 . Plants over than 7 cm high . . . . . . . . . . . . . . . . ..S. brevipes
Discussion
The circumscription of the genus Sisyrinchium has been problematic, and it was demonstrated that the use of traditional
morphology is not enough to delimit taxa, suggesting that a
multidisciplinary approach is necessary to describe infrageneric taxa (Inacio et al. 2017 and references therein). Until the
appearance of molecular phylogenetic reconstructions
(Chauveau et al., 2011; Karst and Wilson, 2012; Alves et al.,
2014), the number of available names, the lack of the diagnostic characters, and the problem with herborization process made the circumscription of taxa within and among
genera of Iridaceae, a true challenge. Inacio et al. (2017) concluded that molecular and morphological dataset is relevant
for the circumscription of the different sections within
Sisyrinchium. The inclusion of S. humahuacense in the Sect.
Segetia is well supported (PP ¼ 0.99; BS MP ¼ 99; Figure 4)
by the molecular analyses. The monophyly is recovered with
three specimens from different localities (Table 1) in all the
analysis performed and the sibling species are not clear yet,
being necessary more exhaustive molecular analysis.
Regarding morphological characters, S. humahuacense
approaches to the description of this section except for its
synapomorphies: the colour of the tepals (blue vs. yellow/
white) and by the shape of the fruit (ellipsoid vs. obloid)
(Inacio et al. 2017), and mainly for the presence of a conspicuous horizontal rhizome with copious rudimentary leaves
of the previous cycles not splitting into fine fibers.
It should be taking into account that Sisyrinchium laxinervium Ravenna included in the Sect. Hydastylus (Ravenna,
2003) and then relocated to Sect. Segetia (Inacio et al. 2017)
is not included in the taxonomic key since the name was not
validly published according to Art. 40.6 of the ICBN (McNeill
et al. 2012). Furthermore, the diagnostic characters described
by Ravenna (2003) clearly discriminate S. laxinervium from S.
humahuacense: the colour of the flowers are yellowish (vs.
blue), all parts of the plant are larger, the presence of previous cycles of the leaves are splitting into fine fibers and, also
its distribution (S. laxinervium is endemic to Peru).
Acknowledgements
The authors thank the Director of Instituto Darwinion (IBODA-CONICET)
for photographs and kind support and to Francisco Rojas for the illustration of the species. We also thank L. Giussani and S. Arroyo-Leuenberger
for useful comments on earlier drafts of the manuscript. This study was
supported by CONICET (Argentina), grants from the National Geographic
References
Alves TLS, Chauveau O, Eggers L, Souza-Chies TT. 2014. Species discrimination in Sisyrinchium (Iridaceae): assessment of DNA barcodes in a
taxonomically challenging genus. Mol Ecol Resour. 14:324–335.
Bah T. 2009. Inkscape: Guide to a Vector Drawing Program (Digital Short
Cut).
pez-Ferrari AR. 2009. Una nueva espeCeja-Romero J, Espejo-Serna A, Lo
cie de Sisyrinchium (Iridaceae) del estado de Guanajuato, Mexico.
Acta Bot Mex. 87: 83–90.
Chauveau O, Eggers L, Raquin C, Silverio A, Brown S, Couloux A, Cruaud
C, Kaltchuk-Santos E, Yockteng R, Souza-Chies TT, et al. 2011.
Evolution of oil-producing trichomes in Sisyrinchium (Iridaceae):
insights from the first comprehensive phylogenetic analysis of the
genus. Ann Bot. 107(8):1287–1312.
Chauveau O, Eggers L, Souza-Chies TT, Nadot S, Annals S, August N,
Chauveau O, Eggers L, Souza-Chies TT, Sophie N. 2012. Oil-producing
flowers within the Iridoideae (Iridaceae): evolutionary trends in the
flowers of the New World genera. Ann Bot. 110(3):713–729.
Cholewa AF, Henderson DM. 1984. Biosystematics of Sisyrinchium Section
Bermudiana (Iridaceae) of the Rocky Mountains. Brittonia. 36:342–363.
doi.org/10.2307/2806596
Cocucci AA, Vogel S. 2001. Oil-producing flowers of Sisyrinchium species
(Iridaceae) and their pollinators in southern South America. Flora.
196(1):26–46.
Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure from small
quantities of fresh leaf tissues. Phytochem Bull. 19:11–15.
Goldblatt P, Rudall P, Henrich JE. 1990. The genera of the Sisyrinchium
alliance (Iridaceae: Iridoideae): Phylogeny and relationships. Syst. Bot.
15: 497–510.
Fachinetto J, Kaltchuk-Santos E, Inacio CD, Eggers L, de Souza-Chies TT.
2018. Multidisciplinary approaches for species delimitation in
Sisyrinchium (Iridaceae). Plant Species Biol. 33(1):3–15.
Goldblatt P, Manning JC. 2008. The Iris Family: natural history and classification. Portland: Timber Press.
Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for phylogenetic analysis. Cladistics. 24(5):774–786.
Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl Acids Symp
Ser. 41:95–98.
Henderson DM. 1976. A biosystematic study of Pacific Northwestern
blue-eyed grasses (Sisyrinchium, Iridaceae). Brittonia. 28(2):149–176.
Hijmans RJ. 2017. Raster: Geographic Data Analysis and Modeling. R
package version 2.6-7. Available from https://CRAN.R-project.org/
package=raster
Huelsenbeck JP, Ronquist F. 2001. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics. 17(8):754–755.
Inacio CD, Chauveau O, Souza-Chies TT, Sauquet H, Eggers L. 2017. An
updated phylogeny and infrageneric classification of the genus
Sisyrinchium (Iridaceae): challenges of molecular and morphological
evidence. Taxon. 66(6):1317–1348.
Karst L, Wilson CA. 2012. Phylogeny of the New World Genus
Sisyrinchium (Iridaceae) based on analyses of plastid and nuclear DNA
sequence data. Syst. Bot. 37(1):87–95.
McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth DL,
Herendeen PS, Knapp S, Marhold K, Prado J, et al. 2012. International
Code of Nomenclature for algae, fungi, and plants (Melbourne Code)
adopted by the Eighteenth International Botanical Congress,
Melbourne, Australia.
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY
Miller MA, Pfeiffer W, Schwartz T. 2010. Creating the CIPRES Science
Gateway for inference of large phylogenetic trees. In: Gateway
Computing Environments Workshop GCE, New Orleans (LA), USA.
R Core Team 2016. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.
Document freely available on the internet at: http://www. r-project.
org2015.
Rambaut A. 2009. FigTree, a graphical viewer of phylogenetic trees.
Edinburgh, UK: Institute of Evolutionary Biology University of Edinburgh.
Available from: http://tree.bio.ed.ac.uk/software/figtree/
Rambaut A, Suchard MA, Xie D & Drummond AJ. 2014. Tracer v1.6,
Available from http://beast.bio.ed.ac.uk/Tracer.
Ravenna PF. 2003. Revisional studies in the genus Sisyrinchium
(Iridaceae) XI. Onira 8(13):48–54.
Roitman G, Castillo A, Maza I. 2008. Iridaceae. In: Zuloaga, FO, Morrone
O, Belgrano MJ, editors. Catalogo de las plantas vasculares del Cono
Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monogr Syst
Bot from Missouri Bot Gard. Vol. 107. Saint Louis (MO); Missouri
Botanical Garden Press, pp. 423–453.
View publication stats
7
Tacuatia LO, Eggers L, Kaltchuk-Santos E, Souza-Chies TT. 2012a.
Population genetic structure of Sisyrinchium micranthum Cav. (Iridaceae)
in Itapu~a State Park, Southern Brazil. Genet Mol Biol. 35(1):99–105.
Tacuatia LO, Souza-Chies TT, Flores AM, Eggers L, Siljak-Yakovlev S,
Kaltchuk-Santos E. 2012b. Cytogenetic and molecular characterization
of morphologically variable Sisyrinchium micranthum (Iridaceae) in
southern Brazil. Bot J Linn Soc. 169(2):350–364.
Tacuatia LO, Kaltchuk-Santos E, Souza-Chies TT, Eggers L, Forni-Martins
ER, Pustahija F, Robin O, Siljak-Yakovlev S. 2017. Physical mapping of
35S rRNA genes and genome size variation in polyploid series of
Sisyrinchium micranthum and S. rosulatum (Iridaceae: Iridoideae). Plant
Biosyst – An Int J Deal with All Asp Plant Biol. 151:403–413.
Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. 2011.
MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Mol
Biol Evol. 28(10):2731–2739.
Thiers B. 2018. Index Herbariorum: a global directory of public herbaria
and associated staff. [Bronx (NY)]: New York Botanical Garden’s Virtual
Herbarium. [accessed 2018 Jan 12]. Available from http://sweetgum.
nybg.org/ih.