A New Synonym in Senna, Series Aphyllae (Leguminosae, Caesalpinioideae)

Author(s): Federico O. Robbiati, Ana M. Anton, and Renée H. Fortunato

Source: Systematic Botany, 39(4):1120-1126. 2014.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364414X683840

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Systematic Botany (2014), 39(4): pp. 1120–1126
© Copyright 2014 by the American Society of Plant Taxonomists
DOI 10.1600/036364414X683840
Date of publication September 3, 2014

A New Synonym in Senna, Series Aphyllae (Leguminosae, Caesalpinioideae)

Federico O. Robbiati,1,3 Ana M. Anton,1,3 and Renée H. Fortunato2,3,4,5
1
Instituto Multidisciplinario de Biologı́a Vegetal (IMBIV), Av. Vélez Sarsfield 299, 2
piso. 5000 Córdoba, Argentina.
2
Instituto de Recursos Biológicos, INTA, Hurlingham 1686, Buenos Aires, Argentina.
3
Consejo Nacional de Investigaciones Cientı́ficas y Técnicas (CONICET).
4
Universidad de Morón, Morón 1708, Buenos Aires, Argentina.
5
Author for correspondence (rhfortunato2002@yahoo.com.ar; fortunato.renee@inta.gob.ar)

Communicating Editor: Chrissen E. C. Gemmill

Abstract—Multivariate and univariate analyses based on morphological characters were carried out in order to understand the taxonomic
status of Senna spiniflora and S. chacoënsis. Fifteen morphological characters from herbarium specimens were recorded and analyzed by the
Kruskal-Wallis test and principal components analysis. The type specimens and all available herbarium material were examined for the
taxonomic study. The results showed that the taxa differed only by the character of stem pubescence and this feature displayed high
variability. In parallel, the principal components analysis demonstrated that no clearly separated groups were recognized. Based upon these
results, the synonymy of S. chacoënsis to S. spiniflora is proposed.
Keywords—Aphyllae, Fabaceae, morphometric analyses, quantitative characters, taxonomic position.

Series Aphyllae Benth. belongs to the genus Senna Mill. from adjacent areas of Paraguay and Argentina. In addition
(Leguminosae) and is comprised of xeromorphic shrubs Robbiati et al. (2011) found the same situation.
and subshrubs, with deeply woody roots; the leaves of the The aim of this study is to clarify the systematics of these
adult branch are represented by minute triangular or two species that have been subjected to conflicting taxonomic
sublobate scales; the stem is junciform, green, and photosyn- opinions in the past.
thetic (Bravo 1978a; Irwin and Barneby 1982; Robbiati et al.
2011). This series was previously considered under the
Materials and Methods
genus Cassia L. by Bentham (1871), Burkart (1946, 1952),
and Bravo (1978a, 1978b, 1982). Based on vegetative and Plant Material—Field studies were conducted between 2010 and 2013.
Information regarding habitat and intra-population variation was
reproductive characters, Irwin and Barneby (1982) divided
recorded. The type collection and all available material were examined.
Cassia into three genera: Senna, Chamaecrista Moench, and Morphological Characters—Twenty-six herbarium specimens (eight of
Cassia s. s. Within ser. Aphyllae, S. spiniflora (Burkart) H. S. S. spiniflora and 18 of S. chacoënsis) were considered as operational taxo-
Irwin & Barneby and S. chacoënsis (L. Bravo) H. S. Irwin & nomic units (OTUs). We covered the whole distribution range with the
Barneby form a complex characterized by having rigid and selected specimens. Twenty-five characters were examined of which only
15 were informative (Table 1). For each specimen, floral features were
spine-tipped branches, mostly spreading at more or less 90
scored from one fully expanded and rehydrated flower, and size mea-
from the parent axis (Bravo 1978a; Irwin and Barneby 1982). surements were made with an ocular micrometer using a binocular
These species co-occur in the provinces of central and north- microscope (Carl Zeiss 475003–9902). The dimensions of the internodes
ern Argentina: Chaco, Formosa, and Santiago del Estero were taken from young branches, and the width was measured in the
median part.
(Robbiati et al. 2011).
Morphometric Analyses—The statistical analyses were performed
Burkart (1946) described Cassia spiniflora Burkart for the using INFOSTAT statistical program version 2012 (Group InfoStat,
Paraguayan Chaco region (middle Pilcomayo) as having Córdoba, Argentina). For the univariate analysis the average (M), standard
stems finely velvety-puberulent, sepals finely pilosulous, and deviation (Std), coefficient of variation (CV), minimum and maximum
racemes crowded towards the branchlet tips; and C. aphylla were recorded for each feature. To represent the variability of each char-
acter, box plots containing medians and quartiles were prepared.
Cav. var. robusta Burkart for the Argentinean Chaco region, The Kruskal-Wallis test (K) with a significance value of p > 0.05 was
which differs by glabrous stems and sepals and racemes borne used to test differences between pairs of means. Prior to carrying out
along the length of the flowering branchlets. Subsequently, multivariate analysis, the data matrix itself was examined. A Pearson’s
Burkart (1952) reported new morphological characters (i.e. Correlation was performed to identify pairs of characters with a high
degree of correlation (r > 0.6). The principal components analysis (PCA)
stiff and broad (3–4 mm) branches) and treated C. aphylla var.
was based on a correlation matrix and the results were plotted in two-
robusta as C. rigida (Hieron.) Burkart var. robusta Burkart. dimensional scatter plots. The cophenetic correlation coefficient was
Bravo (1978a), based on the same features analysed by Burkart calculated to determine goodness of fit between the Euclidian distance
(1952), recognized the taxon as a distinct species, C. chacoënsis among OTUs in the two dimensional plot.
L. Bravo; and Irwin and Barneby (1982) accepted the delimita- Geographic distribution—The geographic distribution was analysed
with software DIVA-GIS (Hijmans et al. 2005); the coordinates were
tion of the taxon but treated it as Senna chacoënsis. mapped from herbarium specimens, and when this information was
The status of Senna spiniflora and S. chacoënsis as distinct lacking, the localities were georeferenced using gazetteers or following
species from each other has also been controversial. Fortunato the georeferencing procedures of Chapman and Wieczorek (2006).
(1984) has reported S. spiniflora from northern Argentina and
pointed out variability in the pubescence of the calyx: northern
Results
Argentinean specimens slightly differ from those of Paraguay
by the calyx being externally glabrous instead of pubescent. Univariate Analysis—The average, standard deviation,
Marazzi et al. (2006) also noted that herbarium materials are CV, minimum, and maximum values of quantitative morpho-
often misidentified due to the presence of specimens morpho- logical characters are summarized (Table 2) and their variation
logically intermediate between the two species, especially is presented in box plots (Figs. 1A, 1B). The character of calyx
1120
2014] ROBBIATI, ANTON AND FORTUNATO: A NEW SYNONYM IN SENNA 1121

Table 1. Quantitative characters used for the phenetic analysis. Discussion

A taxonomic study and phenetic analyses were conducted to
1. Anther length of abaxial stamens (mm). 2. Anther length of median explore the taxonomic position of S. spiniflora and S. chacoënsis.
stamens (mm). 3. Asymmetric and lower petal length (mm). 4. Calyx
Among the characters analyzed, only stem pubescence showed
pubescence (hair density cm2). 5. Divergence angle of branches (degree).
6. Gynoecium length (mm). 7. Internode length (mm). 8. Internode significant differences. This feature was used by Burkart
width (mm). 9. Leaf length (mm). 10. Long sepal length (mm). (1946, 1952), Bravo (1978a), and Irwin and Barneby (1982) in
11. Pedicel length (mm). 12. Peduncle length (mm). 13. Plant height (m). the taxonomic delineation of S. spiniflora and S. chacoënsis.
14. Staminode length (mm). 15. Stem pubescence This character displayed high variability (CV = 424.26) in
(hair density cm2).
S. chacoënsis: while glabrous stems have been reported by
previous authors (Burkart 1946; Bravo 1978a; Irwin and
Barneby 1982), during this study we found several specimens
pubescence showed the highest CV for both taxa (S. spiniflora: with pubescent branches, especially in populations growing in
145.67; S. chacoënsis: 255.17), and stem pubescence displayed the central-western parts of Argentina. This result is in agree-
higher CV (424.26) for S. chacoënsis. Of the 15 features, only the ment with the morphologically intermediate individuals
character of stem pubescence showed a significant difference pointed out by Robbiati et al. (2011). On the other hand, the
between the means (H = 16 p < 0.0001). This character was character of calyx pubescence which was assigned taxonomic
used by Burkart (1946, 1952) and Bravo (1978a) to distinguish value to segregate S. spiniflora and S. chacoënsis (Burkart 1946,
S. spiniflora from S. chacoënsis. 1952; Bravo 1978a; Irwin and Barneby 1982) did not show
Principal Component Analysis—The characters of asym- significant differences and presented high CV (145.67 and
metric petal length and peduncle length showed strong cor- 255.17, respectively) for the two species, indicating that this
relation ( r > 0.63), so the former was excluded from the character is unsuitable for specific delimitation. This conclu-
principal components analysis. The first three principal com- sion concurs with the variability in the calyx pubescence in
ponents together account for 46.2% (20%, 13.8%, and 12.4%, S. spiniflora cited by Fortunato (1984). The PCA analysis
respectively) of the variance within the data. The cophenetic showed no clear differentiation between S. spiniflora and
correlation is r = 0.782, indicating a goodness of fit between S. chacoënsis, resulting in an overlap of specimens from the
the Euclidian distance among OTUs in the two dimensional central regions of distribution. This result suggests that no
plot. Loading on the first component was contributed mainly characters were useful in the segregation of the two taxa; this
by leaf length, internode length, divergence angle of branches, finding agrees with the similarities between S. spiniflora and
long sepal length, plant height, and calyx pubescence; load- S. chacoënsis reported by Fortunato (1984) and Marazzi et al.
ing on the second component was contributed mainly by (2006). Historically S. spiniflora and S. chacoënsis have been
staminode length, gynoecium length, calyx pubescence, treated as seperate species based on pubescence and inflores-
peduncle length, and long sepal length; the highest loadings cence. Results from this study show that these characters are
on the third component correspond to anther length of abax- too variable to be used for species delimitations. Therefore,
ial stamens, peduncle length, and staminode length (Table 3). synonymy is indicated.
The scatterplot of components one, two, and three resulted
in no clear differentiation between the OTUs of the taxa;
Taxonomic Treatment
moreover, they showed an overlap (Figs. 2, 3).
Geographic distribution—The northern region had more SENNA SPINIFLORA (Burkart) H. S. Irwin & Barneby emend.
specimens with pubescent calyx and stems, while the southern Robbiati & Fortunato, Iconography: Bravo, 1978a: 362,
specimens were glabrous; plants from the central area were Figure 4; 363, Figure 5. Cassia spiniflora Burkart,
very variable (Fig. 4). Darwiniana 7: 235. 1946.—TYPE: PARAGUAY. Dpto. de

Table 2. Mean ± standard deviation, coefficient of variation, and range (minimum– maximum) for the 15 characters used for phenetic analysis.
Characters numbered according to Table 1.

Taxon S. spiniflora S. chacoënsis

Character/value M Std CV Min–Max M Std CV Min–Max

1 4.34 ±0.66 15.24 3.00–5.10 4.38 ±0.49 11.13 3.75–5.00

2 2.90 ±0.60 20.53 2.25–4.00 2.90 ±0.82 28.22 2.50–4.00
3 14.7 ±2.20 14.95 11.0–17.0 13.85 ±1.97 14.24 9.00–17.0
4 1.75 ±2.55 145.67 0.00–6.00 0.78 ±1.99 255.17 0.00–7.00
5 85.3 ±4.75 5.56 75.0–90.0 80.39 ±11.3 14.13 50.0–90.0
6 10.03 ±2.47 24.68 7.20–14.0 11.11 ±1.28 12.64 7.00–13.0
7 15.5 ±3.46 22.35 10.0–20.0 13.17 ±5.19 39.44 6.00–24.0
8 3.79 ±0.68 17.93 2.50–4.50 2.85 ±0.97 34.15 1.20–4.50
9 0.72 ±0.24 34.15 0.50–1.00 0.85 ±0.17 19.45 0.50–1.00
10 5.56 ±1.05 18.88 4.00–7.50 5.11 ±1.01 19.72 3.00–7.00
11 5.78 ±2.91 50.40 0.75–10.0 6.75 ±1.77 26.19 4.00–10.0
12 4.91 ±1.86 37.93 2.50–8.00 3.81 ±2.50 65.6 1.50–11.0
13 1.80 ±0.21 11.52 1.50–2.00 1.83 ±0.18 9.74 1.50–2.00
14 1.10 ±0.23 20.9 0.75–1.50 1.19 ±0.27 22.41 0.96–1.75
15 8.25 ±3.15 38.2 4.00–12.0 0.11 ±0.47 424.26 0.00–2.00
1122 SYSTEMATIC BOTANY [Volume 39

Fig. 1A. Box plots representing the variability of the quantitative characters in S. spiniflora and S. chacoënsis. The box represents the interquartile
range; upper horizontal line (bar) is the uppermost value; lower horizontal line is the lowermost value; circle within the box and the bar in the middle
of the box represent the mean and the median, respectively. Points represent outliers.
2014] ROBBIATI, ANTON AND FORTUNATO: A NEW SYNONYM IN SENNA 1123

Fig. 1B. Continuation of Fig. 1A.

1124 SYSTEMATIC BOTANY [Volume 39

Fig. 2. Principal components analysis (PCA) scatter plots of the first two components. The squares represent S. spiniflora samples and the triangles
represent S. chacoënsis samples. The morphological characters used in this analysis are listed in Table 1.

Boquerón: Entre Toba Quemado y Magariños, Sector New York Bot. Gard. 35: 569. 1982. —TYPE: ARGENTINA.
Pilcomayo, Sept. 1938, Rojas 8269 (holotype: SI!). Prov. Santiago del Estero: Gral. Taboada Añatuya, 27 Jan
Cassia aphylla Cav. var. robusta Burkart, Darwiniana 7: 237. 1944, Soriano 571 (holotype: SI!).
1946. syn. nov. Cassia rigida (Hieron.) Burkart var. robusta
Shrubs 1.5–2 m tall; branches 1.2–4.5 mm wide, glabrous
Burkart, Legum. Argent. (ed. 2): 544. 1952. Cassia
to pubescent, divaricate, divergence angle 50
–90
; leaf 0.5–
chacoënsis L. Bravo, Darwiniana 21: 359. 1978. Senna
1 mm long, ± 0.5 mm wide. Racemes borne along the length
chacoënsis (L. Bravo) H. S. Irwin & Barneby, Mem.
of the flowering branchlets to crowded toward tip of branch-
lets. Bracts 1–1.5 mm long, 0.25–0.75 mm wide; peduncle
1.5–11 mm long, 0.25–0.5 mm wide; pedicel 0.75–10 mm long,
Table 3. Contributions of individual characters to the first three 0.25–0.5 mm wide, both the peduncle and pedicel glabrous to
multivariate axes of the principal components analysis (PCA). Characters pubescent. Calyx glabrous to pubescent, the 2 smaller sepals
numbered according to Table 1. 3–4 mm long, 2–2.25 mm wide and the 3 larger sepals 3–
7.5 mm long, 3–4 mm wide. Petals 9–17 mm long, 4–8 mm
Character PC1 PC2 PC3
wide. Androecium: 3 anthers of long abaxial stamens 3–
1 –0.09 –0.19 0.59 5.1 mm long, and filament 1–8 mm long; 4 anthers of median
2 0.27 0.23 –0.02
stamens 2.5–4 mm long, and filament 1–1.5 mm long and
4 0.29 –0.37 0.27
5 0.37 –0.02 –0.09 3 staminodes 0.75–1.75 mm long, and filament 1–2 mm long.
6 0.03 –0.45 0.19 Gynoecium 7–14 mm long, 0.75–1 mm wide, glabrous. Pods
7 –0.40 0.12 –0.26 7–9 cm long, 4–5 mm wide, glabrous. Seeds 3.5–5 mm long,
8 –0.23 0.15 0.11 2.5–3.5 mm wide.
9 0.41 –0.01 –0.17
10 –0.35 –0.30 0.17 Distribution and Habitat—Senna spiniflora occurs in
11 –0.25 –0.19 –0.13 central and northern Argentina (Córdoba, Chaco, Formosa,
12 –0.10 0.31 0.47 Salta, Santa Fe, and Santiago del Estero provinces) and south-
13 –0.29 0.11 –0.08 ern Paraguay (Boquerón department) (Fig. 4). This species
14 0.02 0.51 0.33
grows in Chaco Biogeographic regions usually on clay and
15 0.19 0.19 0.21
salty soils (Cabrera 1971).
2014] ROBBIATI, ANTON AND FORTUNATO: A NEW SYNONYM IN SENNA 1125

Fig. 3. Scatter plots of the first and third components. The squares represent S. spiniflora samples and the triangles represent S. chacoënsis samples.
The morphological characters used in this analysis are listed in Table 1.

Acknowledgements. We are grateful to the curators and directors of Fortunato, R. H. 1984. Dos Leguminosas nuevas para la Flora Argentina.
the BAB, CORD, CTES, LIL and SI herbaria for permission to study and/or Darwiniana 25: 267–268.
loans of specimens. We would like to express our gratitude to Dr. Michael Hijmans, R. J., L. Guarino, A. Jarvis, R. O’Brien, P. Mathur, C. Bussink,
Nee and Dr. Jorge Chiapella for their critical reading of the manuscript M. Cruz, I. Barrantes, and E. Rojas. 2005. DIVA-GIS 7.1.7. Available
and to Mr. Marcelo Gritti (CONICET) to help with the preparation of in http://www.diva-gis.org.
the illustration. The authors also thank the Consejo Nacional de Irwin, H. S. and R. C. Barneby. 1982. The American Cassinae: A synop-
Investigaciones Cientı́ficas y Técnicas (CONICET) for the fellowship tical revision of Leguminosae, tribe Cassiae subtribe Cassiinae
given to Federico O. Robbiati and the Universidad Nacional de Córdoba in the New World. Memoirs of the New York Botanical Garden 35:
(UNC) for providing facilities and workplace. This research was supported 567–570.
by grants CONICET (PIP 112-200801-00323 and PIP 11220080101557) and Marazzi, B., R. H. Fortunato, P. K. Endress, and R. Spichiger.
Instituto Nacional de Tecnologı́a Agropecuaria (INTA). Fortunato and 2006. Senna (Cassiinae, Leguminosae) in Paraguay: synop-
Anton are members of CIC (CONICET). sis, occurrence, ecological role and ethnobotany. Candollea 61:
315– 329.
Robbiati, F. O., L. Ariza Espinar, A. M. Anton, and R. H. Fortunato. 2011.
Cambios nomenclaturales en el género Senna Ser. Aphyllae
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550 S
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430 S 63
070 W, 17 Mar 2013, Fortunato 10221 (BAB); 70 km al E de 61
520 W, 15 Mar 2013, Fortunato 10204 (BAB); 10 km al O de Ing. Juárez,
Monte Quemado por ruta nacional 16 en dirección a Taco Pozo, 26
040 S por ruta nacional 81 en dirección a la provincia de Salta, 23
510 S 61
570
62
110 W, 13 Nov 2001, Fortunato 7237 (BAB). Dpto. Gral. Taboada, W, 15 Mar 2013, Fortunato 10211 (BAB). Prov. Salta: Dpto. Anta,
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560 S 62
300
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