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Significance of variation and evaluation of potential characters in three morphologically similar species of Barleria sect. Barleria (Acanthaceae) in southern Africa

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Abstract

The variation exhibited within three species of Barleria (B. bechuanensis, B. irritans and B. jubata) was studied to establish whether it was discrete or continuous. Morphological characters were examined and recorded in matrices. Cluster analysis was employed to impose a hierarchical non-overlapping association among operational taxonomic units (OTUs) while ordination was used to establish whether the variation was discrete or continuous. Discrete characters were determined from quantitative morphological data using box and whisker plots. Locality information for the OTUs was obtained from herbarium labels and used to generate maps to illustrate geographic distribution of taxa. Cluster analysis and ordination demonstrated that there was discrete variation within Barleria bechuanensis, B. irritans and B. jubata, which each split into two distinct clusters, although box and whisker plots illustrated that many quantitative characters overlapped within and between species. Since clear morphological gaps between clusters are assumed to be indicators of breaks in gene flow, the distinct clusters were recognised at species level.

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Acknowledgements

The curators of B, BOL, C, GRA, J, K, M, MO, NBG, NU, PRE, STEU and WIND are gratefully acknowledged for the loan of specimens. We thank the National Research Foundation of South Africa (Grant Number 85692) and the University of the Witwatersrand, for research funds and financial support, which facilitated the field work and visits to the National Herbarium in Pretoria during the course of the study and the Management at The University of Zambia for granting study leave. We are extremely grateful to Donald McCallum for providing many of the figures and the referees for their helpful and constructive comments.

Conflict of interest

The authors declare that they have no conflict of interest.

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Authors and Affiliations

  1. School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Johannesburg, South Africa

    Florence C. Nyirenda & Kevin Balkwill

  2. Department of Biological Sciences, School of Natural Sciences, University of Zambia, Lusaka, Zambia

    Florence C. Nyirenda

Authors
  1. Florence C. Nyirenda
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  2. Kevin Balkwill
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Corresponding author

Correspondence to Kevin Balkwill.

Additional information

Handling editor: Ricarda Riina.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Online Resource 1

Matrix of qualitative, quantitative and ratios data for five Barleria species complexes (XLSX 100 kb)

Online Resource 2

Matrix of qualitative data for three species complexes in Barleria (XLSX 15 kb)

Online Resource 3

Matrix of quantitative data for three species complexes in Barleria including widths, but not lengths (XLSX 21 kb)

Online Resource 4

Cluster analysis (UPGMA) from a matrix of 26 qualitative morphological characters with eigenvalues >0.6. Correlation coefficient r = 0.9814. A phenon line is drawn at 0.67 coefficient of dissimilarity. Numbers 1–7 denote the clusters that may be distinguished at species level. X is the standard taxon. BB, BE = Putative taxa from Barleria bechuanensis; BI, BU = Putative taxa from B. irritans; EL = B. elegans subsp. orientalis (standard); JU = B. jubata (PDF 188 kb)

Online Resource 5

Two-dimensional scatter plot from Principal coordinates analysis (PcoA) showing the distribution of the OTUs in multivariate space. OTUs that are not clustering with members of the same group are indicated by arrows. BB (open circle), BE (filled triangle) = Putative taxa from Barleria bechuanensis; BI (filled circle), BU (open inverted triangle) = Putative taxa from B. irritans; EL (filled square) = B. elegans subsp. orientalis (standard); JU (open diamond) = B. jubata (PDF 31 kb)

Online Resource 6

Three-dimensional scatter plot from Principal components analysis (PCA) showing the distribution of OTUs in multivariate space. BB (open circle), BE (filled triangle) = Putative taxa from Barleria bechuanensis; BI (filled circle), BU (open inverted triangle) = Putative taxa from B. irritans; EL (filled square) = B. elegans subsp. orientalis (standard); JU (open diamond) = B. jubata (PDF 33 kb)

Online Resource 7

Variation in selected quantitative characters of selected taxa of Barleria section Barleria Group 1. Box = 25 to 75%, line = non-outlier range. BB, BE = Putative taxa from Barleria bechuanensis; BI, BU = Putative taxa from B. irritans; EL = B. elegans subsp. orientalis (standard); JU = B. jubata. a Leaf length. b Length of leaf apex spine. c Length of leaf marginal spine. d Ratio of leaf length to leaf width. e Bracteole width. f Length of bracteole apex spine. g Ratio of bracteole length to the total number of teeth on bracteole margin. h Number of teeth on the distal half of the anticous calyx lobe margin (PDF 28 kb)

Online Resource 8

Variation in selected quantitative characters of selected taxa of Barleria section Barleria Group 1. Box = 25 to 75%, line = non-outlier range. BB (open circle), BE (filled triangle) = Putative taxa from Barleria bechuanensis; BI (filled circle), BU (open inverted triangle) = Putative taxa from B. irritans; EL (filled square) = B. elegans subsp. orientalis (standard); JU (open inverted triangle) = B. jubata. a Ratio of anticous calyx lobe length to calyx lobe width. b Ratio of length of anticous calyx lobe to total number of teeth on calyx lobe margin. c Length of apex spine on posticous calyx lobe. d Number of teeth on the distal half of the posticous calyx lobe margin. e Ratio of length of posticous calyx lobe to the total number of teeth on the calyx lobe margin. f Ratio of posticous calyx lobe length to calyx lobe width (PDF 37 kb)

Appendices

Appendix 1

List of specimens examined [numbers in square brackets denote specimen codes allocated during analysis]. The specimens are arranged according to the quarter-degree grid reference system (Edwards and Leistner 1971).

Barleria jubata S.Moore [JU]

Namibia: –2016 (Otjiwarongo): Okosongomingo (–BC), 11 Jun 1939, Volk 2249 (M) [JU108]; 13.9 miles S. of Otjiwarongo on road to Okahanja, Farm Tweekoppies (–DA) de Winter 2739 (K) [JU117]; Okanjanda, Erongo Mts (–DA), Jun–Jul 1916, Pearson 9895 (BOL, NBG) [JU007]; Farm Graslaagte, on Grey Mountain (also found on the Ozondjaheberg) (–DB), 28 Nov 1952, Walter 283 (B, M) [JU106]. –2017 (Waterberg): 65 km E of Otjiwarongo, Waterberg plantation, on the slope down from the plantation to the plain (–AC), 3 Mar 1968, Wanntorp and Wanntorp 641 (K) [JU116]; Upper slopes at S.W. point of the Omuverume Plateau (–AC), Rutherford 352 (WIND) [JU107]; Okarakowisa, north of Farm Bergtuine OTJ 455, on Waterberg Plateau on red sandstone cliffs (–AD), Geiss 12375 (MO; PRE) [JU009]; Waterberg–Abhang (–AD) Volk 2472 (M) [JU109]; Waterberg Plateau (–CA), Boss 34948 (PRE) [JU008]; Klein Waterberg above Okosongomingo (–CA), 15 July 1954, Schelpe 201 (J) [JU144].

Barleria damarensis T.Anderson

Namibia: –1714 (RuacanaFalls): 20 miles southern Ruacana, Bergveld (–AD), Giess and Leippert 7588 (NBG) [JU141].

Barleria bechuanensis C.B.Clarke [BB]

South Africa: –2723 (Kuruman): Bophuthatswana, Nyara Farm, NW-facing on slight slope of plain (–AD), 27 Feb 1982, Gubb 258/39 (PRE) [BB086]; Newstead Farm, on level plain at top of hill (–CB), 4 Mar 1981, Gubb 184-41 (PRE) [BB088]. –2724 (Taung): Vryburg, Kormutsetla Farm, on gently sloping plain to the W (–AA), 12 May 1982, Gubb 323-95 (PRE) [BB099]; Dwaalvlakte (–AB), 1912, Sharpe 8148 (PRE) [BB010]; Barkly West, Boetsap, rocky slope (–CD), 17 Feb 1945, Brueckner 136 (PRE) [BB080]. –2725 (Bloemhof): Cawoods’ Hope (–DA), Mar 1912, Burt-Davy 12930 (BOL) [BB002]. –2824 (Kimberly): Barkly West, Koopmansfontein, Agric. Res Station (–AA), 14 Apr 1964, Nursey 133 (PRE) [BB005]; Newlands (–AD), 17 Mar 1939, Lewis s.n. (NBG) [BB142]; Newlands (–AD), Mar 1939, Esterhuysen 1008 (BOL) [BB006]; 3 mls east of Kimberly (–DA), 11 May 1961, Schlieben 8715 (PRE) [BB101]. –2924 (Hopetown): Riet Pan, Mar 1920 (–CA), Wilman 16418 (BOL) [BB003]. S.loc., s.d., s.col. (curator Pretoria 198), (PRE) [BB100].

Barleria sp. aff. B. bechuanensis C.B.Clarke [BE]

South Africa: –2723 (Kuruman): 5.3 km south southeast of Kuruman (–AD), 8 Apr 2000, Balkwill, McDade and Lundburg 11815 (J) [BE120]; 11 m N W of Kuruman, 1280 m (–AD), 19 Feb 1956, Leistner 593 (PRE) [BE119]; Dunmore Farm, S—facing on level talus slope (–DD), 7 Mar 1981, Gubb 171-82 (PRE) [BE087]; Killarney Farm, in a pan (depression) (–DD), 19 Feb 1981, Gubb 166-65 (PRE) [BE089]. –2823 (Griekwastad): east of Postmasburg, west of Owendale, facing railway to Blinkklipkop Station, on the loop, 28°14.87′S 23°17.34′E (–AB), 8 Apr 2000, Balkwill, McDade and Lundburg 11818 (J) [BE121]; about 7 m NE of Postmasburg, Blinkklipkop (–AC), 26 Mar 1959, Leistner 1396 (K, SRGH, NBG, PRE, M) [BE085]; Asbestos Hills, Daniel’s Kuil (–BA), Mar 1939, Esterhuysen s.n. (BOL) [BE122]; near Daniel’s Kuil (–BA), Rogers 10413 (BOL) [BE124]; Daniel’s Kuil on hills near Ouplaas, (–BA), Apr 1940, Esterhuysen 2046 (BOL) [BE082]; Warrenton (–BB), 11 Mar 1902, Adams 17 (BOL) [BE125]; Daniel’s Kuil (–DA), Mar 1939, Esterhuysen 1007 (BOL) [BE084]. –2824 (Kimberly): Barkly West, Koopmansfontein, Agric. Res Station, at A53 (–AA), 26 Mar 1937, Acocks 2121 (PRE) [BE118]; Riet Pan (–DB), Apr 1919, Moran 15919 (BOL) [BE004].

Barleria irritans Nees [BI]

South Africa: –3124 (Hanover): Noupoort, Karoo veld (–BB), Dec 1961, Repton 5725 (PRE) [BI095]. –3325 (Port Elizabeth): Kirkwood, Zuurberg National Park, 33°18′26″S, 25°23′11″E, Fynbos (–AD), 17 Apr 1999, Brand 133 (PRE) [BI093]; Uitenhage, 2.5 miles from Barkley Bridge along Alexandria railway line (–CA), 10 Febr 1953, Marais 162 (i and ii) (K, PRE) [BI145 and BI067]; Uitenhage (–CA), 13 Dec 1847, Prior s.n. (K, PRE) [BI068]; Uitenhage Cap. b. sp. (–CA), Ecklon 384 (79.2) (M, PRE) [BI104]; 11 km N of Uitenhage, Uitspan experimental plot of Department of Plant Protection Unit (–CD), 10 Mar 1979, Mauve 5238 (PRE) [BI075]; Kommands Kraal (–CD), Zeyher 1421 (NBG) [BI073]; Uitenhage (–CD), Harvey s.n. (BOL) [BI077]; Addo Elephants National Reserve, open veld (–DC), 12 Feb 1954, Brynard 330 (PRE) [BI072]; –3326 (Grahamstwn): Albany, Hell Poort entrance slightly <16 m from Grahamstown on Cradock road (–AB), 21 Jul 1975, Brink 369 (PRE) [BI097]; Double Drift, Fish River Valley, 33°05′20″S, 26°47′05″E, hillside above river (–AD), 8 Mar 1981, Phillipson 216 (GRA, PRE) [BI074]; Alexandria, limestone ridge S.E. of Sandflats (–CA), 2 Apr 1952, Archibald 4234 (GRA, K, PRE) [BI071]. –3327 (Peddie): 2000 ft. (–AA), 1897, Sim 20021 (NU, PRE) [BI079].

Barleria sp. aff. B. irritans Nees [BU]

South Africa: –3224 (Graaff Reinet): Hillside on Farm Reitvlei (–AB), 3 Mar 1930, Galpin 9972 (K, PRE) [BU070]; Spitskop (–AD), 1 Apr 1912, Burtt-Davy 13503 (PRE) [BU091]; Swart River near Graaff Reinet (–BC), 3 Dec 1950, Willems 703 (NBG) [BU143]; 10 miles south of Graaff Reinet (–BC), 3 Jan 1956, Theron 1853 (GRA, K, PRE) [BU105, BU092]; near Graaff Reinet (–BC), Mar 1867, Bolus 563 and 10441 (BOL) [BU066]; veld flats (–BC), 9 May 1954, Henrici 4958 (PRE) [BU094]; Jansenville, Jockomst, in open veld (–DC), 18 Apr 1932, Long 561 (PRE) [BU078]; Jansenville (–DC), 3 Apr 1981, Olivier 2806 (PRE) [BU069].

Information on Electronic Supplementary Material

Online Resource 1. Matrix of qualitative, quantitative and ratios data for five Barleria species complexes.

Online Resource 2. Matrix of qualitative data for three species complexes in Barleria.

Online Resource 3. Matrix of quantitative data for three species complexes in Barleria including widths, but not lengths.

Online Resource 4. Cluster analysis (UPGMA) from a matrix of 26 qualitative morphological characters with eigenvalues >0.6.

Online Resource 5. Two-dimensional scatter plot from Principal coordinates analysis (PcoA) showing the distribution of the OTUs in multivariate space.

Online Resource 6. Three-dimensional scatter plot from Principal components analysis (PCA) showing the distribution of OTUs in multivariate space.

Online Resource 7. Variation in selected quantitative characters of selected taxa of Barleria sect. Barleria Group 1.

Online Resource 8. Variation in selected quantitative characters of selected taxa of Barleria sect. Barleria Group 1.

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Nyirenda, F.C., Balkwill, K. Significance of variation and evaluation of potential characters in three morphologically similar species of Barleria sect. Barleria (Acanthaceae) in southern Africa. Plant Syst Evol 304, 77–92 (2018). https://doi.org/10.1007/s00606-017-1452-5

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  • DOI: https://doi.org/10.1007/s00606-017-1452-5

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