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On the Morphology of the PollenGrains in ‘Armeria Sibirica’ inSpecimens from between Longitude30° W and 60° EJ. Praglowski a , G. Erdtman a & Yrjö Vasari ba Palynological Laboratory , Nybodagatan 5, 171 42, Solna,Swedenb Botanical Institute , Oulu University , Oulu, FinlandPublished online: 01 Sep 2009.

To cite this article: J. Praglowski , G. Erdtman & Yrjö Vasari (1969) On the Morphology of thePollen Grains in ‘Armeria Sibirica’ in Specimens from between Longitude 30° W and 60° E,Grana Palynologica, 9:1-3, 72-91, DOI: 10.1080/00173136909436426

To link to this article: http://dx.doi.org/10.1080/00173136909436426

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GIIASA ~ ’ . ~ I . T S O I . O G I C . ~ 9: 1-3, 1069

ON THE IIORPI-IOLOGI’ OF THE POLLEN GRAINS IN cilRMERIA SIBIRICA’ IN SPECIAIENS FROM BEI’WEEN LONGITUDE 30° \IT AND GOo E

\VITH AS XDDEXDU.\I BY Y. \‘XSAIII

.\IS received 2 3 . S I I . 6 8

‘I’lie present study of driiierin pollcii was evolied by tlie senior author’s desire to find out u-hctlier it was within thc hounds of possibility to determine if a fossil pollen grain cncountcred 1 ) ~ Dr. Y. Yasari in northern Finland (I’asari, 1OG8, p. SO) lint1 heen produced by drnicritr miritinin, or if it emanated from a nicmlxr of the niorc nor t 11 cr 11 ‘ ‘A rnierirr sib irictr corn 11 I cs ” .

The pollen grains of arctic speciiiiens of drniericr, quoted as A. labrtitlorica, have heen snit1 to he monomorphic (Irersen, 1040). l‘lic mononiorpliisni in question is licltl to lie a secondary feature, an effect OF a retrogressive transition from a cliniorpliic condition. Speci- inens of A. ldrnrloriccc (“A. sibirica”) from east Greenland, nortli- erriiiiost Scandinaria and from tlie arctic areas of tlie European part of the Soviet Union (map, Fig. 1 ) liavc more or less variable pollen grains (Figs. 2-5, Gtr-ti, 7 and 8) . ‘l’lic variations arc not confined to particular plant intliritluals as in species with typically dimorphic pollen, c.g. Arriieriti maritimti, but coesist in the inflorescences (’l’able l) , single flo\vers, and even indiriclual anthers (‘l’able 2) inresti-

It seems more liliely that the variations can he connected with nieiotic disturbances than with different stages of derelopnient. Thus in pollen grains from Alount Piiltsa, the hyperborean outpost of Swedish Lapland, the muri of the retiicrilak pollen surface are either siniplibaciilate (proritlcd with a single row of infratcctnl bac- ules; Fig. 3a) or duplibaculate (with a double row of rods; Fig. 3c).

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MOIIIWOLOGT O F TIIE POLLEX IS ‘AIINERIA SIIIIIIICA’ 73

Fig. 1. Map showing the places wlicrc the spccimcns itivcstigatctl w r c eollcctetl. The numbers corrcspontl to those in Tables 1 aIitl 2 and to the list of collcctions investigated (p. S i ) . Specirrien no. 2 (east Greenlotid), is not shown on thc map.

I’arts of pollen grains from northernmost Swxlcn arc also cs1iil)itcd in Figs. 4 f i - i . I n Figs. 4 c and f head, winding iiiriri tlcroid of spinulcs can he seen, \\-hereas in Figs. 49-i the muri are less broad, straight (or almost so) antl beset lritli spinulcs. In Figs. 4r( and b, neither Iriinina nor spinulcs can he seen. ?’lie pollen grains arc thus not mononiorpliic in the real sense of the word. \\‘it11 regard to the topography of their scsine they are instead vary variahlc. ‘Hie factual rariatioiis arc, Iio\\-erer, often so slight that they can only he rcrcalcd witli tlie aid of a microscope of high po\ver.

U’itli regard to the topography of tlie sesine sis more or less clearly delimited pollen types hare lxxn distinguislictl. Two of them (Type 1 antl Type 2) correspond more or less to the A-type in drmc- ria ~ ~ ~ ( i r i t i m n (Iwrsen, Z.C.; Ertltmnn S Dunbar, 1 %X), and one (Type 5) has ~norphological features more or less in coninion with pollen grains of the U-type in the same species. The six pollen- morphological types may briefly he described as follows.

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74

Fig. 2. Specimen from east Greenland, Alakenzie Bay. x 1500.--n-b, type 5; c-d, types 1-2, with short spinules; e-11, type 1 (two different grains e, f and 9, h), spinules distinct, comparatively long.

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Fig. 3. Specimen from Sweden, 3Iount Paltsa. x 1500.--a-b, type 5, muri winding, lumina small; c-d, type 6, niuri duplibaculate, spinules missing; e-f, type 1; g-h, type 2. Long spinules present in e-h.

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7G

Fig. 4. Specimen from Forwag, Birtavarre. x 1500.--n-b, type 4; c-f, two grains of type 5 (c , d and e, f ) ; g-i, type 1.

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JIOIIPIIOLOGS O F TILE POLI.I;S IS ‘ A 1 I ~ I I ~ I I l . i SII3IIIIC. i ’ 77

Fig. 5. Spccinien from Sorway, Mount Parras. x l5OO.-Two grains (u, b and c, d) of type 5, e and cl exhibit thick niuri supported by bacules with swollen distal end; (e, f and g, h) of typc 1.

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7s J. PRAGI.O\\’SI<I AXD G . ERDTSIAS

Fig. G. a-d, Specimen from Kolguyev. x 1500.--a-b, type 5; c-d, type 1.-e-g, x 1200; stigmas in spccirnen from Xorway, Jloant Parras (e); east Grcenland, Alakenzie Bay (f); USSR, Waygach (6).

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l l O l ~ l ’ l l O I ~ O G Y O F TIIE P O L L E S IN ‘ARJII:I<IA SIIHIIIC.1’ i 9

Fig. 7. Specimen from USSR, \\’aygach. x 1500.-a-d, typc 1: a-c, part of a pollen surface in three successive foci; note the spinriles in a, b, d.-e-f, tgpc 5.

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so

Fig. 8. SpeciInen from USSR, Sovafa Zenilya. x 1500.--n-b, type 4, grain intcctate, baculate, bacules with distinct capita; c-tl, transitional grain betwen types 4 and 5, partially subtectate, baeules with swollen ends.-e-f, type 3, with long luminal bac- u1cs.-g-h, type 2, spinules n o t very distinct.

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~ I O R P I I O L O G T O F TIIE POI,LEX IX ‘ARMEIIIA SIDIRICI’ 81

Fig. 9. Specimen from USSR, \Yaygacli. One-anther-slidcs. x 1250.--n-c, “anther-l”, with a, type 4; 6, typc 5; c, typc 1. ti-f, “anther-2”, with d, types 4, 5; e-f, type 1.

6 - 699762 Gram Pal. vol. 9: 1-3

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52

Fig. 10. Specimen from USSR, Kovaya Zemlya. One-anther-slides. x 1250.-u-b, “anther-1”, with u, type 5; b, type 1; c-f, “antherd”, with c, type 4; d, type 5; e, type 1; f, type 6.

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Fig. 11. drrneria rnorifirna, recent grains. x 1200.-a-b, specimen from JIunso ~ i e m Stockholm, A-line; c-d, specimen from Bjorkii, B-line.

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S4 J. PHAGLO\\’SliI A S D G . ERDTJIIS

Fig. 12. Fossil pollen grain of Arrneria froni northern Finland; (cf. Y. \lasari, 1967). x 1300.-Tliis grain is more or less similar to pollen grains of type 1 (“A-Type”); e.g.

Figs. 2e, fi 3e-h and 4g-i.

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SIOI~PIIOLOGT OF TIIE POLLEN IN ‘ARSIERIA SInInIch’ 85

Table 1 . Rela fiiw freqrrencks (percentnges) of pollen groin types in inflorescences of “drmerict sibirica”. I00 grains = 100 per cent except in specimen 2, iuhere the percentages are based irpori 80 grains only. The figures 2-8 in the uppermost roiu ofthe table refer lo the map, Fig. I , p. 73.

2 3 4 5 6 7 8

Pollen types

’. (“A-type”) 68 91 90 96 87 51 41 2 3 24 26 4 2 26 5 (“B-type”) 32 4 6 4 13 21 7 6 5 2 4

Table 2 . Pollen types in 16 “one-anther” pollen slides (nurnber ofgrains). The figures 2-8 in the uppermost roiu of the table refer to the map, Fig. I , p . 73.

2 3 4 5 6 7 8

Anther no. 1 2 3 4 5 6 7 8 9 1 0 1 1 1 2 1 3 1 4 1 5

Pollen types

(“A-type”) 29 23 21 22 13 23 23 8 19 27 20 5 4 18 9 2 3 3 6 9 4 5 (“B-type”) 2 2 3 6 1 2 5 18 9 2 6 2 3 1 6

Total 31 25 21 28 13 24 23 10 19 27 27 5 28 31 26

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D i a g n o s e s: Type i‘ (“A-?’ypc”). Figs. 2e-11, 3c-It, 494, iie-li, Gc, d

Pollen grains ciircticulate. JIuri straight, simplibaculate. Their tectal part ahout 3 ,u thick (the distinct, supratectal spinulcs-about 0.5-1 .5 / I long-not inclutlcd). The infratcctal, 1)aculatc zone of the muri is ahout 5-G p thick. Luniina angular to slightly rounded, occasionally with stray hacules (shorter than the mural bacdcs, cf. Fig. Gc).--?‘his type is more or Iess frequent in a11 specimens in- vestigated. It is similar to the pollen grains in the A-line of drmeria marifiziici, although less roliust.

Type 2 (“A-Type”). Figs. 39, 11, Sg, 11 ‘ ~ i e r e is only a vcrx slight tliflerencc bet\vecn this type and type 1 :

the muri are winding, wliercas those in type 1 are usually more straight.-Type 2 has h e n encountered in all specimciis invcs- tigatetl.

Type 3. Figs. 8 e and f Pollen grains only partially and at the same time less distinctly

reticulate (cureticulate). Jluri usually straight. ‘I‘hcir tectal part is more \\.Cali ant1 thin than in types I anti 2. Spinules indistinct or nI)scnt. Luniinal haciilcs of about the same length or occasionally sliglitly longer than the mural and usually iiiore nunicrous than the latter. This type is niorc or less intcrinctliate bet\vecn types 1 antl 2 on one hand and type 4 on the other.-Pollen grains of this type have only heen encountered in the specimens from the eastern sector of the area esaminctl, i.e. from Waygach antl Novaya Zemlya.

Type 6. Figs. 4n, b, 8a-d Pollen grains usually intcctate, occasionally partially subtectate

(Figs. 8 ( I , c). Scsine consisting of densely spaced 1)acules. These are of varying thickness and rclati\*ely shorter than those in types 1 antl 2 . Their distal end is sonictiiiies distinctly swollen, pro\~itlcd \villi a “captit” (I7&. 8n-d). Kcsine conipnratively thicker than in types 1 and 2. - I’olleii grains of type 4 havc heen seen in slides made from specinlens froni iforthcrn Scandinavia (Fig. 4, locality Uirta- varre) and Kovaya Zemlyn (Fig. 8).

Type 5 (“B-Type”). Figs. 3 a , b, 4c-f, Gn, b, ’ i e , f. Pollen grains cureticulate with a faint tendency toumds a rcti-

haculatc condition. Jluri usually relatively robust, winding, occa- sio nnlly fr a g ni en t etl and provitl ed with stray in t crcala t cd is01 a t ed

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AIOIIPIIOLOGY O F TIIE POLLES IS ‘A I ~ J I E R I A S I ~ I R I C A ’ 87

bacules n.itli su-ollen apices. Their tectal part smooth or occasionally provided with indistinct spinulcs. Luniina (if present) comparatively small, usually without 1,aculcs. Infratcctal bacules clumsy, shorter than in types 1 ant1 2; their distal part sometimes relatirely large (diameter about 2-4 p; cf. Figs. 5 c , d). - ‘I‘his type occ~irs in all spccinicns investigated. I t is some\rhat reminiscent of the pollen grains in the B-Type of drmeria maritima. Type G. Fig. 3c.

Pollen grains with more or less deriating or irregular sesinc. Graint from Mount PSltsa with duplibaculate muri (Fig. 3 c), ant1 various others difficult to classify (c.g. “pilate” grains with ab- nornially large capita encountered in Russian collections) have been referred to this type.

S p cc i ni c n s i n y es t i ga t e (1 The specimens investigated come from the lierbarium (S) of

i\’aturhistorislta riltsmuseet, Stocliliolm. They arc as follows (the numbers correspond to those of the localities indicated in the map, Fig. I). ‘I‘hey also correspond to the numbers of the illustrations. X.U.: Nunnbcr 2 (cast Greenland) is not slio\rn on the map.

2. “ilrnicriri uiilgmis Willd. var. sibirica (Turcs.)”, Fig. 2-east Greenland, Alaltcnzic Bay, 31 July 1900; I<. A. G. Gredin s.n. - SPL slide 5122.

3. “drniericc sibirica Turcz.”, Fig. 3-S\vcdcn, AIount PKltsa, 5 August 1939; H. Smith s.11. - SPL slide 5070.

4. “Arnieria clongatci (Hoflin.) I<och vnr. sibiriccc (Turcz.)”, Fig. 4 --Nor\ray, Lyngcn, I3irtavarre, .July 1899; E. Hagliintl & J. Iiiillstriini s.n. - SI’L 5072.

5 . “drnieria sibiriccc Turcz.”, Fig. 5 - Norway, Lyngcn, Xlount I’arras, G August 1939; H. Smith s.11. - SPL 5051.

G. “drnieria sibirica Turcz.”, Fig. G - USSl\, I<olguycr, 10 J d y 1925; A. Tolniatchew s.n. - SI’L 5124.

7. “drnicritr sibirica Turcz.”, Fig. 7 - USSR, l\Taygach, 12 Aiigust 1907; 0. Ekstam s.11. - SPL 5125.

8. “drmeria sibirica Turcz.”, Fig. 8 - USSR, Xoraya Zemlyn, Xlatochltin Schar, 12 August 1895; 0. Elistall1 s.11. - SI’L 5118.

D is c u s s i o n Table 1 illustrates the occiirrence of the sis pollen types in seven

specimens from east Greenland in the west to Novaya Zcnilya in the

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88 J. PRhGLO\\’SKI AXD G . ERDTMAK

east. Only one inflorescence from each specimen was investigated. As can be seen in Tables 1 and 2, grains of types 1 and 2 (“A-Type”) and type 5 (“I3-Type”), are much more frequent than grains be- longing to the remaining intermediate types. In all the Scandinavian samples high percentages (up to 96%) of types 1 + 2 (“A-Type”) prevailed. In east Greenland and in Soviet Russia, however, types 1 and 2 are less dominating. This may possibly point to a certain pollenmorphological “type-stability”, more pronounced in the Scan- dinavian area. As seen from Table 1, the pollen-frequency of types 1 and 2 amounts to about 50% in a specimen from The Isle of Waigatsch, whereas the remaining pollen grains belong to types 3, 5 and, subordinately, to type G (aberrant grains). In the Isle of Kol- gujev the proportion betwxn types 1 + 2 and type 5 is about G : 1 , whereas in the specimen from east Greenland it is about 2: 1 . In the specimen from Noraya Zemlya types 3 and 4 constitute each one fourth of the total of the grains. It should be noted that this specimen was robust but smaller than the other specimens. In addition the stamens and stigmas were distinctly shorter.

The examination of one-anther-slides (Table 2) exhibited similar trends as previously observed in “inflorescence-analyses” of “Ar- nieria sibirica”. It s h o w that one to four of the six more or less inter- grading pollen types can be produced within the confines of a single anther. Each anther produces about 20-30 pollen grains. This rnay be contrasted with Trifolium pratense (about 200 grains), Calluna uolgaris (about 500 tetrads) etc.

Fifteen anthers from seven localities investigated were examined. There are four types among 28 pollen grains in one anther from a specimen from the Isle of Waygach. Eighteen grains belong to type 5 (“B-Type” Fig. Db). In an “inflorescensces analysis” from the same specimen type 5 constitutes about 20 %. r‘One-anther-slides” made from material from the specimen from Xovaya Zemlya show trends (Table 2) Fig. 10, analogous to those observed in the “in- florescence analysis” of the same specimen. Table 2 also shows that pollen grain types (all from the same inflorescence) can vary from one anther to another. Among the anthers investigated from the speci- men from the Isle of Kolgugev, two contained “monomorphic” pollen, whereas three pollen types were found in a third (unopened) anther from the same inflorescence. An analogous state of things prevailed in the specimen from Isle of M’aygach.

Herbarium material is not wry adequate for a study of small

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details of stigmas. The photomicrographs (Fig. 6 e-9) seem to indicate that the stigmas, in conformity with the observation of Iversen, are more or less “monomorphic” (“B-Type”). Fig. 11 exhibits the pollen of recent Armeria marifinin, A and B line. Fig. 12 s h o w the fossil Arnieria pollen grain (slide made by Dr. 1’. Yasari). With due consideration of the morphological features illustrated and described above n-e feel inclined to suggest that Vasari’s grain (Fig. 12) has been produced by “Armeria sibirica”, not by Ammeria marifima s.str. (Fig. 11).

A c k n owl e d g e ni en t s \Ye are indebted to the Director of the Botanical Section, Museum

of Natural History, Stockholm, Prof. T. Norlindh for kind permission to take polliniferous material of “Armeria sibirica” from the her- baria in his charge. \Ve also thank Prof. Vasari for kindly letting us avail ourselves of his original slide with the solitary fossil pollen grain of Arnieria and for his concluding relllarlis on this find.

Palynological Laboratory Kybodagatan 5 171 42 Solna. Sweden

Addendum: Phytogeographical considerations on the finding of Armeria scabra Pall.’ as subfossil in Kuusamo,

North-East Finland I ’ R J 6 VASARI

The single pollen grain of the Armeria type that was found by Vasari (1967) in the Late-glacial (probably Pre-boreal) deposits of AIaanselHnsuo in Iiuusanio, Korth-East Finland, has now been identified by Praglowski & Ertttnian as being “producetl by ‘Armeria sibirica’, not by Armeria maritinia s.slr.” (see the present paper, p. 89). This calls for some considerations on the phytogeographical significance of this interesting find.

The Armeria species are entomophilous with sticky pollen (cf.

In this report, the name Armeria scabra Pall. has been used following the example of Iversen (1940) and the nomenclature of Hylander (1955). Evidently, in this case there is a question of the subspecies known as A. scabra ssp. labraiforica (Wallr.) nov. comb. (syn. A. sibirica Turcz.) (Iversen, 1940).

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90 J. PIIAGLOn’SI<I A S D G . ERDTAIAS

Hcgi, 1927, p. lSSG and Ilaltcr, 19-15, p. 771) . This, together with the lo\\., hcrbaceoiis habit of tlie plants, makes it seem highly 1111-

likely that tlie pollen grain in tlie deposits at Jlaansel5nsuo \\.ere a result of long-(listance transportation-it can he regarded as reason- ably certain that A. scabrrr had indeed heen then growing on the shorcs of the little tarn on tlie spot, outside the liinits of its present distribution (cf. Hu l th , 1950, map 130s).

At present, drrrierin scabui is, in Fennoscandia, a typical repre- sentative of a floral element 1;nown as tundra and fjcltl plants with a “centric” distribution pattern, in other \vords being confined to oiie or more clearly defined “centres” (cf. e.g. Xordliagen, 193G; Xaiinfcldt, 1947, p. 5G and Kalela, 19G1, pp. 310-313). They are in general calcicole plants with a clear preference for continental cli- mate. ?’hey arc, as a wliole, IJiotically \veal; plants, able to grow only in places where the vegetation is not closed ant1 wlicrc there is also plenty of bare inincral soil, such as on sand and gravel slopes, shores etc.

Decause of tlic lack of subfossil finds of plants of this group, one lias heen compelled to rely almost entirely upon their present distribution patterns wlicn considering their immigration history in Fcnnoscandia. Laying particular stress upon their pcctiliar eco- logical dernantls, the most continental phases of tlie Late-glacial, the Oldest and Older Dryas (Zones I a and Ic) have generally been regarclet1 as tlie periods most favourable for tlic spreading and thriving of the group of centric tundra antl fjeltl plants (Kalela, 19G1). \\’hen t li e climate b eca me warmer and ni or c liu in id t 11 eir ex p a nsi on lvas clieckctl. The present range lias been regarded as more or less concordant with their former distribution pattern.

The finding of sub-fossil rirmcria sccibra in I’re-boreal deposits outside the present range of tlie species is of quite considerable in- terest as it Jirstly proves that tlic area of this species (antl probably of many others, of this group, too) lias l>c.en witler and iiiore con- tinuous than is evident from the present distribution pattern.

Secondly, it sliou-s that the expansion of this species (and tlic eleinent represented by it) is not liltely to be limited only to the Older Dryas period (Zone I) but continued all through the period when Late-glacial conditions prevailed, even until the Pre-boreal. This may also be of some relevance to tlie question of the climatic cliar- acter of the Pre-boreal in Eastern Fcnnosltandia for which both a continental and a maritime climate have been attributed (cf. the

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JIORPIIOLOGY O F TIIE POLLEK IX ‘ A I I I I E I ~ I A SIUIRICA’ 91

discussion in, for example, Yasari, 19G2, pp. 81-88 S 123 and Tolonen, 19G7, p. 355).

Thirdly, a n e w member thus becomes ntltletl to tlie rich element of old t u n d r a a n d fjeltl plants within t h e Iiuusa111o-Salla district (cf. e.6. Iialcla, 19G1, pp. 314-315). Evidently, tlie said element w a s originally iiiucli richer than it is at present. A11 the species that grew i n that area i n Late-glacial times, however, were not able to survive there lip to the present time, noth\vitlistaiitling the coiiinioiiness of ra ther “exceptional” hahitats there (cf. also t h e literature cited i n Vasari, 1962, pp. 82-83).

Uotanical Institute Oulu University Oulu, Finland

REFERENCES

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