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Pollen morphological study of some species of the genus Rubus L. (Rosaceae) of the flora of Armenia Cover

Pollen morphological study of some species of the genus Rubus L. (Rosaceae) of the flora of Armenia

Hacquetia
Volume 24 (2025): Issue 1 (June 2025)
By: Alla Hayrapetyan,  Marieta Asatryan,  Hasmik Sonyan,  Karine Balayan and  Dmitri Beketovski  
Open Access
|Feb 2025

Figures & Tables

Figure 1:

Average sizes of pollen grains (P × E) in some species of the genus Rubus L.Our data: blue – polar axis (P), green – equatorial diameter (E). Data from literature: orange – polar axis (P), yellow – equatorial diameter (E). (1) – Li et al. (2001), (2) – Lechnovich et al. (2020), (3) – Tomlik-Wyremblewska (1995), (4) – Monasterio-Huelin & Pardo (1995).Slika 1: Povprečna velikost pelodnih zrn (P × E) nekaterih vrst rodu Rubus L.Naši podatki: modra – polarna (vertikalna) os (P), zelena – ekvatorialna (horizontalna) os (E). Podatki iz literature: oranžna – polarna (vertikalna) os (P), rumena – ekvatorialna (horizontalna) os (E). (1) – Li et al. (2001), (2) – Lechnovich et al. (2020), (3) – Tomlik-Wyremblewska (1995), (4) – Monasterio-Huelin & Pardo (1995).
Average sizes of pollen grains (P × E) in some species of the genus Rubus L.Our data: blue – polar axis (P), green – equatorial diameter (E). Data from literature: orange – polar axis (P), yellow – equatorial diameter (E). (1) – Li et al. (2001), (2) – Lechnovich et al. (2020), (3) – Tomlik-Wyremblewska (1995), (4) – Monasterio-Huelin & Pardo (1995).Slika 1: Povprečna velikost pelodnih zrn (P × E) nekaterih vrst rodu Rubus L.Naši podatki: modra – polarna (vertikalna) os (P), zelena – ekvatorialna (horizontalna) os (E). Podatki iz literature: oranžna – polarna (vertikalna) os (P), rumena – ekvatorialna (horizontalna) os (E). (1) – Li et al. (2001), (2) – Lechnovich et al. (2020), (3) – Tomlik-Wyremblewska (1995), (4) – Monasterio-Huelin & Pardo (1995).

Figure 2:

Comparison of the average values of pollen polar axis (P) of investigated Rubus species using Simpson and Roe test.Slika 2: Primerjava povprečnih vrednosti polarne (vertikalne) osi (P) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.
Comparison of the average values of pollen polar axis (P) of investigated Rubus species using Simpson and Roe test.Slika 2: Primerjava povprečnih vrednosti polarne (vertikalne) osi (P) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.

Figure 3:

Comparison of the average values of pollen equatorial diameter (E) of investigated Rubus species using Simpson and Roe test.Slika 3: Primerjava povprečnih vrednosti premera ekvatorialnega premera pelodnega zrna (E) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.
Comparison of the average values of pollen equatorial diameter (E) of investigated Rubus species using Simpson and Roe test.Slika 3: Primerjava povprečnih vrednosti premera ekvatorialnega premera pelodnega zrna (E) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.

Figure 4:

Comparison of the average values of pollen apocolpium diameter of investigated Rubus species using Simpson and Roe test.Slika 4: Primerjava povprečnih vrednosti premera apokolpija (brazde) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.
Comparison of the average values of pollen apocolpium diameter of investigated Rubus species using Simpson and Roe test.Slika 4: Primerjava povprečnih vrednosti premera apokolpija (brazde) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.

Figure 5:

Comparison of the average values of pollen mesocolpium width of investigated Rubus species using Simpson and Roe test.Slika 5: Primerjava povprečnih vrednosti premera mezokolpija (brazde) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.
Comparison of the average values of pollen mesocolpium width of investigated Rubus species using Simpson and Roe test.Slika 5: Primerjava povprečnih vrednosti premera mezokolpija (brazde) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.

Figure 6:

Pollen grains of Rubus armeniacus Focke. a–h – LM micrographs: a – 3-zonocolp-orate pollen grain, b – 4-zonocolp-orate pollen grain with additional small slit-like apertures at the pole (marked with arrows), c – spheroidal os (marked with arrow), d, e – thickened edges of colpi (marked with arrows), f – geniculum (marked with arrow), g – exine, columellae layer (marked with arrow), h – finely reticulate exine ornamentation; i, j – SEM micrographs: i – pollen grain in equatorial view, colpus, j – striate-perforate exine ornamentation in combination with finely striate-microreticulate one; scale bars: a–i – 10 μm, j – 1 μm.Slika 6: Pelodna zrna vrste Rubus armeniacus Focke. a–h – LM mikrografi: a – 3-zonokolporatna pelodna zrna, b – 4 zonokolporatna pelodna zrna z dodatno odprtino na polih (označeno s puščico), c – sferoidna os (označeno s puščico), d, e – odebeljen del brazde (označeno s puščico), f – genikulum (označeno s puščico), g – eksina, kolumelna plast (označeno s puščico), h – fino mrežasta ornamentacija eksine, i, j – SEM mikrograf: i – pelodno zrno v ekvatorialnem pogledu, brazda, j – črtasto-luknjičasta ornamentacija eksine v kombinaciji s fino črtastomikroretikuliranim okrasjem; merilce: a–i – 10 μm, j – 1 μm.
Pollen grains of Rubus armeniacus Focke. a–h – LM micrographs: a – 3-zonocolp-orate pollen grain, b – 4-zonocolp-orate pollen grain with additional small slit-like apertures at the pole (marked with arrows), c – spheroidal os (marked with arrow), d, e – thickened edges of colpi (marked with arrows), f – geniculum (marked with arrow), g – exine, columellae layer (marked with arrow), h – finely reticulate exine ornamentation; i, j – SEM micrographs: i – pollen grain in equatorial view, colpus, j – striate-perforate exine ornamentation in combination with finely striate-microreticulate one; scale bars: a–i – 10 μm, j – 1 μm.Slika 6: Pelodna zrna vrste Rubus armeniacus Focke. a–h – LM mikrografi: a – 3-zonokolporatna pelodna zrna, b – 4 zonokolporatna pelodna zrna z dodatno odprtino na polih (označeno s puščico), c – sferoidna os (označeno s puščico), d, e – odebeljen del brazde (označeno s puščico), f – genikulum (označeno s puščico), g – eksina, kolumelna plast (označeno s puščico), h – fino mrežasta ornamentacija eksine, i, j – SEM mikrograf: i – pelodno zrno v ekvatorialnem pogledu, brazda, j – črtasto-luknjičasta ornamentacija eksine v kombinaciji s fino črtastomikroretikuliranim okrasjem; merilce: a–i – 10 μm, j – 1 μm.

Figure 7:

Pollen grains of Rubus caesius L. a–f – LM micrographs: a–b – pollen grains in polar view (a – colpi, b – exine, columellae layer), c – spheroidal os (marked with arrow), d–f – pollen grains in equatorial view (e – os area, marked with arrows), f – granulate exine ornamentation; g–h – SEM micrographs: g – pollen grains in polar and equatorial view, h – striate-perforate exine ornamentation in combination with finely striate-microreticulate one; scale bars: a–g – 10 μm, h – 1 μm.Slika 7: Pelodna zrna vrste Rubus caesius L. a–f – LM mikrografi: a–b – pelodno zrno v polarnem pogledu (a – brazda, b – eksina, kolumelna plast), c – sferoidna os (označena s puščico), d–f – pelodno zrno v ekvatorijalnem pogledu (e – osno območje, označeno s puščico), f – zrnata ornamentacija eksine, g–h – SEM mikrograf: g – pelodno zrno v polarnem in ekvatorialnem pogledu, h – črtasto-luknjičasta ornamentacija eksine v kombinaciji s fino črtasto-mikroretikulirano ornamentacijo; merilce: a–g – 10 μm, h – 1 μm.
Pollen grains of Rubus caesius L. a–f – LM micrographs: a–b – pollen grains in polar view (a – colpi, b – exine, columellae layer), c – spheroidal os (marked with arrow), d–f – pollen grains in equatorial view (e – os area, marked with arrows), f – granulate exine ornamentation; g–h – SEM micrographs: g – pollen grains in polar and equatorial view, h – striate-perforate exine ornamentation in combination with finely striate-microreticulate one; scale bars: a–g – 10 μm, h – 1 μm.Slika 7: Pelodna zrna vrste Rubus caesius L. a–f – LM mikrografi: a–b – pelodno zrno v polarnem pogledu (a – brazda, b – eksina, kolumelna plast), c – sferoidna os (označena s puščico), d–f – pelodno zrno v ekvatorijalnem pogledu (e – osno območje, označeno s puščico), f – zrnata ornamentacija eksine, g–h – SEM mikrograf: g – pelodno zrno v polarnem in ekvatorialnem pogledu, h – črtasto-luknjičasta ornamentacija eksine v kombinaciji s fino črtasto-mikroretikulirano ornamentacijo; merilce: a–g – 10 μm, h – 1 μm.

Figure 8:

Pollen grains of Rubus candicans Weihe ex Rchb. a–f – LM micrographs: a–c – pollen grains in polar view (a – colpus membrane ornamentation, b – os area, c – exine, columellae layer, marked with arrows), d–f – pollen grains in equatorial view (d – colpus without genuculum, e – colpus with genuculum, marked with arrows, f – thickened edges of colpi); g–h – SEM micrographs: g – pollen grains in polar and equatorial view (colpus with genuculum, marked with arrow), h – finely striate-perforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.Slika 8: Pelodna zrna vrste Rubus candicans Weihe ex Rchb. a–f – LM mikrografi: a–c – pelodna zrna v polarnem pogledu (a – ornamentacija membranebrazde, b – območje osi, c – eksina, kolumelna plast, označeno s puščico), d–f – pelodno zrno v ekvatorialnem pogledu (d – brazda brez genikuluma, e – brazda z genikulumom, označeno s puščico, f – odebeljen del brazde); g–h – SEM mikrograf: g – pelodna zrna v polarnem in ekvatorialnem pogledu (brazda z genikulumom, označeno s puščico), h – fino črtasto-luknjasta ornamentacija eksine (SEM); merilce: a–g – 10μm, h – 1 μm.
Pollen grains of Rubus candicans Weihe ex Rchb. a–f – LM micrographs: a–c – pollen grains in polar view (a – colpus membrane ornamentation, b – os area, c – exine, columellae layer, marked with arrows), d–f – pollen grains in equatorial view (d – colpus without genuculum, e – colpus with genuculum, marked with arrows, f – thickened edges of colpi); g–h – SEM micrographs: g – pollen grains in polar and equatorial view (colpus with genuculum, marked with arrow), h – finely striate-perforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.Slika 8: Pelodna zrna vrste Rubus candicans Weihe ex Rchb. a–f – LM mikrografi: a–c – pelodna zrna v polarnem pogledu (a – ornamentacija membranebrazde, b – območje osi, c – eksina, kolumelna plast, označeno s puščico), d–f – pelodno zrno v ekvatorialnem pogledu (d – brazda brez genikuluma, e – brazda z genikulumom, označeno s puščico, f – odebeljen del brazde); g–h – SEM mikrograf: g – pelodna zrna v polarnem in ekvatorialnem pogledu (brazda z genikulumom, označeno s puščico), h – fino črtasto-luknjasta ornamentacija eksine (SEM); merilce: a–g – 10μm, h – 1 μm.

Figure 9:

Pollen grains of Rubus canescens DC. a–f – LM micrographs: a – pollen grains in semipolar view, b – synaperturate pollen grain in polar view (synaperture marked with arrow), c–f – pollen grains in equatorial view (c – os, d – genuculum, e – thickened edges of colpi, marked with arrows, f – striate-reticulate exine ornamentation); g–h – SEM micrographs: g – pollen grains in semipolar and equatorial view, h – finely striateperforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.Slika 9: Pelodna zrna vrste Rubus canescens DC. a–f – LM mikrografi: a – pelodno zrno v semipolarnem pogledu, b – po daljšnici anastamozna odprtina (označeno s puščico), c–f – pelodno zrno v ekvatorialnem pogledu (c – os, d – genikulum, e – odebeljeni rob brazde, označen s puščico, f – črtasto-mrežasta ornamentacija eksine; g–h SEM mikrograf: g – pelodno zrno v semipolarnem in ekvatorialnem pogledu, h – fino črtasto-luknjasta ornamentacija eksine (SEM); scale bars: a–g – 10 μm, h – 1 μm.
Pollen grains of Rubus canescens DC. a–f – LM micrographs: a – pollen grains in semipolar view, b – synaperturate pollen grain in polar view (synaperture marked with arrow), c–f – pollen grains in equatorial view (c – os, d – genuculum, e – thickened edges of colpi, marked with arrows, f – striate-reticulate exine ornamentation); g–h – SEM micrographs: g – pollen grains in semipolar and equatorial view, h – finely striateperforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.Slika 9: Pelodna zrna vrste Rubus canescens DC. a–f – LM mikrografi: a – pelodno zrno v semipolarnem pogledu, b – po daljšnici anastamozna odprtina (označeno s puščico), c–f – pelodno zrno v ekvatorialnem pogledu (c – os, d – genikulum, e – odebeljeni rob brazde, označen s puščico, f – črtasto-mrežasta ornamentacija eksine; g–h SEM mikrograf: g – pelodno zrno v semipolarnem in ekvatorialnem pogledu, h – fino črtasto-luknjasta ornamentacija eksine (SEM); scale bars: a–g – 10 μm, h – 1 μm.

Figure 10:

Pollen grains of Rubus cartalinicus Juz. a–e – LM micrographs: a–c – pollen grains in polar view (a – pollen and colpus membrane ornamentation, b–c – synaperturate pollen grains and os area, marked with arrows), d–e – pollen grains in equatorial view (d – os, e – thickened edges of colpi); f–g – SEM micrographs: f – pollen grains in equatorial and semipolar view, g – finely striate-perforate exine ornamentation; scale bars: a–f – 10 μm, g – 1 μm.Slika 10: Pelodna zrna vrste Rubus cartalinicus Juz. a–e – LM mikrografi: a–c – pelodno zrno v polarnem pogledu (a – pelod in ornamentacija membrane brazde, b–c pelodno zrno z anastamozno odprtino, označeno s puščico), d–e – pelodno zrno v ekvatorialnem pogledu (d – os, e – odebeljen del brazde); f–g – SEM mikrograf: f – pelodno zrno v ekvatorialnem in semipolarnem pogledu, g – fino črtasto-luknjasta ornamentacija eksine; merilce: a–f – 10 μm, g – 1 μm.
Pollen grains of Rubus cartalinicus Juz. a–e – LM micrographs: a–c – pollen grains in polar view (a – pollen and colpus membrane ornamentation, b–c – synaperturate pollen grains and os area, marked with arrows), d–e – pollen grains in equatorial view (d – os, e – thickened edges of colpi); f–g – SEM micrographs: f – pollen grains in equatorial and semipolar view, g – finely striate-perforate exine ornamentation; scale bars: a–f – 10 μm, g – 1 μm.Slika 10: Pelodna zrna vrste Rubus cartalinicus Juz. a–e – LM mikrografi: a–c – pelodno zrno v polarnem pogledu (a – pelod in ornamentacija membrane brazde, b–c pelodno zrno z anastamozno odprtino, označeno s puščico), d–e – pelodno zrno v ekvatorialnem pogledu (d – os, e – odebeljen del brazde); f–g – SEM mikrograf: f – pelodno zrno v ekvatorialnem in semipolarnem pogledu, g – fino črtasto-luknjasta ornamentacija eksine; merilce: a–f – 10 μm, g – 1 μm.

Figure 11:

Pollen grains of Rubus sanctus Schreb. a–c – LM micrographs: a – pollen grain in polar view, b–c – pollen grains in equatorial view (c– colpus with genuculum, marked with arrow); d–e – SEM micrographs: d – pollen grain in equatorial view, e – finely striate-perforate exine ornamentation; scale bars: a–d – 10 μm, e – 1 μm.Slika 11: Pelodna zrna vrste Rubus sanctus Schreb. a–c – LM mikrografi: a – pelodno zrno v polarnem pogledu, b–c – pelodno zrno v ekvatorialnem pogledu (c– brazda z genikulumom, označeno s puščico); d–e – SEM mikrograf: d – pelodno zrno v ekvatorialnem pogledu, e – fino črtastoluknjasta ornamentacija eksine; merilce: a–d – 10 μm, e – 1 μm.
Pollen grains of Rubus sanctus Schreb. a–c – LM micrographs: a – pollen grain in polar view, b–c – pollen grains in equatorial view (c– colpus with genuculum, marked with arrow); d–e – SEM micrographs: d – pollen grain in equatorial view, e – finely striate-perforate exine ornamentation; scale bars: a–d – 10 μm, e – 1 μm.Slika 11: Pelodna zrna vrste Rubus sanctus Schreb. a–c – LM mikrografi: a – pelodno zrno v polarnem pogledu, b–c – pelodno zrno v ekvatorialnem pogledu (c– brazda z genikulumom, označeno s puščico); d–e – SEM mikrograf: d – pelodno zrno v ekvatorialnem pogledu, e – fino črtastoluknjasta ornamentacija eksine; merilce: a–d – 10 μm, e – 1 μm.

List of investigated species and specimens of the genus Rubus (ERE – Herbarium of the Institute of Botany after A_ Takhtajyan, National Academy of Sciences, Yerevan, Armenia)_Tabela 1: Seznam preučevanih vrst in primerkov rodu Rubus (ERE – Herbarij Botaničnega inštituta A_ Takhtajyan, Nacionalna akademija znanosti, Erevan, Armenija)_

SpeciesHerbarium sheet information
R. armeniacus FockeERE, 80977; ERE, 93906
R. caesius L.ERE, 56383; ERE, 63234; ERE, 63238
R. candicans Weihe ex Rchb.ERE, 58307; ERE, 63240
R. canescens DC.ERE, 58846; ERE, 98402
R. cartalinicus Juz.ERE, 63221
R. caucasicus FockeERE, 58313
R. ibericus Juz.ERE, 135970
R. idaeus L.ERE, 98394
R. sanctus Schreb.ERE, 56378; ERE, 56381; ERE, 63978; ERE, 98366
R. takhtadjanii Mulk.ERE, 64509

Some palynological and karyological features of the representatives of the genus RubusL_ of the flora of Armenia (± SD – standard deviation, CV% – coefficient of variation, LM – light microscopy, SEM – scanning electron microscopy)_Tabela 2: Nekatere palinološke in kariološke značilnosti predstavnikov rodu RubusL_ v flori Armenije (± SD – standardni odklon, CV% – koeficient variacije, LM – svetlobni mikroskop, SEM – vrstični elektronski mikroskop)_

SpeciesPolar axis (P) (μm) ± SD (CV%)Equatorial diameter (E) (μm) ± SD (CV%)P/EColpusExine ornamentationChromosome number (2n)
apocolpium diameter(μm) ± SD (CV%)mesocolpium width (μm) ± SD (CV%)LMSEM
R. armeniacus Focke23.5–25.8 / 24.5 ± 0.7 (2.7%)16.9–22.8 / 19.4 ±1.8 (9.2%)8/7–8/6 (subprolate)4.8–5.9 / 5.3 ± 0.4 (7.1%)8.4–12.5 / 10.4 ± 1.0 (9.8%)finely reticulatefinely striate-perforate or finely striate-microreticulate28 (Krahulcová & Holub, 1997)
R. caesius L.22.2–28.9 / 25.1 ± 1.8 (7.2%)19.7–24.1 / 20.9 ± 1.6 (7.4%)8/7–8/6 (subprolate)4.5–5.3 / 4.9 ± 0.3 (5.3%)14.1–15.8 / 14.9 ± 0.4 (2.8%)finely reticulatefinely striate-perforate or finely striate-microreticulate28 (Krahulcová & Holub, 1997)
R. candicans Weihe ex Rchb.20.3–25.4 / 22.9 ± 1.9 (8.4%)17.5–22.1 / 19.0 ± 1.6 (8.7%)8/7–8/6 (subprolate)5.0–5.6 / 5.3 ± 0.1 (2.8%)14.2–15.4 / 14.8 ± 0.3 (2.0%)granulatefinely striate-perforate21 (Gustafsson, 1943)
R. canescensDC.23.3–25.8 / 24.5 ± 0.7 (3.0%)17.7–20.5 / 19 ± 0.9 (4.9%)8/7–8/6 (subprolate)4.2–5.1 / 4.7 ± 0.2 (4.8%) (sometimes synaperturate)10.5–14.3 / 12.4 ± 1.0 (7.7%)finely striate reticulatefinely striate-perforate, striae linked together by bridges14 (Krahulcová & Holub, 1997)
R. cartalinicus Juz.21.8–30.1 / 25.8 ± 3.1 (11.8%)22.1–23.5 / 22.9 ± 0.5 (2.0%)8/8–8/7 (prolate spheroidal)2.8–4.5 / 3.6 ± 0.4 (11.6%) (sometimes synaperturate)14.4–16.8 / 15.6 ± 0.6 (3.8%)finely striate reticulatefinely striate-perforate–
R. caucasicus Focke21.7–33.2 / 26.3 ± 4.1 (15.7%)16.8–22.5 / 18.0 ± 3.6 (19.8%)8/6–8/4 (prolate)4.7–5.9 / 5.24 ± 0.4 (7.5%)18.5–19.5 / 19.0 ± 0.3 (1.7%)granulatefinely striate-perforate–
R. ibericus Juz.20.5–22.5 / 21.4 ± 0.7 (3.1%)14.6–18.1 / 16.2 ± 1.2 (7.6%)8/6–8/4 (prolate)3.5–4.8 / 4.2 ± 0.3 (7.8%)11.8–14.6 / 13.2 ± 0.7 (5.3%)finely reticulate––
R. idaeus L.18.5–25.3 / 21.8 ± 2.5 (11.6%)14.1–18.9 / 16.5 ± 1.6 (9.6%)8/6–8/4 (prolate)3.8–5.7 / 4.8 ± 0.5 (10.0%)10.5–14.9 / 12.7 ± 1.1 (8.7%)finely reticulate–14, 21 (Chen et al., 1993; Ivanova, 2005)
R. sanctus Schreb.15.5–33.5 / 19.2 ± 5.3 (27.7%)12.1–17.5 / 14.4 ± 1.6 (11.3%)8/6–8/4 (prolate)4.8–6.1 / 5.4 ± 0.3 (6.0%)10.9–12.5 / 11.7 ± 0.4 (3.4%)granulatefinely striate-perforate, diameter of perforation varies14 (Runemark, 2006)
R. takhtadjanii Mulk.12.9–15.6 / 14.2 ± 0.8 (5.6%)14.1–16.8 / 15.4 ± 0.9 (6.1%)6/8–8/6 (subspheroidal)5.3–6.9 / 6.1 ± 0.4 (6.6%)9.8–11.5 / 10.7 ± 0.4 (4.0%)finely reticulate––
DOI: https://doi.org/10.3986/hacq-2025-0006 | Journal eISSN: 1854-9829 | Journal ISSN: 1581-4661
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Language: English
Page range: 67 - 81
Submitted on: Apr 26, 2024
Accepted on: Nov 18, 2024
Published on: Feb 21, 2025
Published by: Slovenian Academy of Sciences and Arts
In partnership with: Paradigm Publishing Services
Publication frequency: 2 issues per year
Keywords:
brambles,
pollen aperture,
exine ornamentation,
light microscopy,
scanning electron microscopy
Related subjects:
Life sciences,
Plant science,
Ecology,
Life sciences, other

© 2025 Alla Hayrapetyan, Marieta Asatryan, Hasmik Sonyan, Karine Balayan, Dmitri Beketovski, published by Slovenian Academy of Sciences and Arts
This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 License.

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