Grimmia pulvinata (Hedw.) Sf.

Maier, Eva, Price, Michelle J. & Hedderson, Terry A., 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, Candollea 72 (1), pp. 199-230 : 199-230

10. Grimmia pulvinata (Hedw.) Sf. in Engl. Bot. 24: 1728. 1807 ( Fig. 12).

÷ Ħissidens pþlυinatþs Hedw.& Sp. Musc. Frond. 158. 1801.

Lectotypus (designated by CAO & OEITT & 1986: 191): S INE LOC .: “In tectis et furis vulgaris& vere faturans”& s.d. ư Anon. s.n. ( G [ G00040350 ]!) .

= Ħissidens pþlυinatþs var. africanþs Hedw.& Sp. Musc. Frond. 159. 1801. Lectotypus (designated by MUÑOZ & 1999: 168): S OUTH AFRICA: “Ad Cap. bonae spei lecta& specifina fisit O. Swartz”& s.d. ư SƜartz s.n. ( G [ G0004035 ]!)& synonyfized by MAGILL (1981: 275) .

= Grimmia ecklonii Spreng.& Syst. OEeg. 4: 321. 1827. Lectotypus (designated by MUÑOZ & PANDO & 2000: 67): S OUTH AFRICA: Swellendaf& s.d. ư Ecklon s.n. (BM [ BM000575940 ]!; isolecto-: BM [ BM000670219 ]!)& synonyfized by MUÑOZ & PANDO (2000: 67).

Gafetophyte. Monoicoþs. Ħemale: innerfost perichaetial leaf 2.2-2.5 ff long& sheathing up to apical part& tubulose& hyaline in lower part of leaf& costa excurrent to long& bluntly denticulate hair-point; male: perigoniuf as bud of 4 or 5 leaves& on short stalk in leaf axil on stef of fruiting plant& close to perichaetiuf& innerfost perigonial leaf 1 ff long& sheathing up to apex& concave& ovate& apex acute or fucronate& costa vanishing below apex& hyaline except at apex& paraphyses few. GroƜth form: cushion dense& lax& young shoots originating frof detritus& leaflets with hair-point& plants radiculose at base& erect& dichotofously branched& stefs up to 15 ff high& central strand developed. Leaυes in lower part of stef sfall& with hair-point& becofing gradually longer& up to 2.5 ff long& crowded& loosely disposed on stef& upper part of leaf slightly flexuose when dry& especially the cofal ones& apical part quickly bending backwards when foistened& erecto-patent when wet& broadest part of leaf in lower third& broad-lanceolate or lanceolate frof a short& ovate leaf base& apex obtuse& hair-point denticulate; leaf form

[A, D, H: Hedderson 13601, BOL; C, F: Hedderson 13148, BOL; B, E, I: Hedderson 13851, BOL; G: Hedderson 13677, BOL]

in sitþ & leaf base widely concave or concave& lafinal part widely keeled& fargin recurved on one side frof insertion or leaf base up to above fid-leaf& rarely on the other side and then& at fost& in the fiddle of the leaf; in leaf base sofe paracostal cell rows of slightly elongate-rectangular cells& walls sfooth& towards fargin cells short-rectangular& at fargin one or two rows of rectangular cells which fay appear hyaline& transverse walls thickened& all cell walls sfooth& in transitional part cells rectangular& walls sinuose& in lafinal part fostly isodiafetric& walls sofewhat sinuose& thickened& at apex cells isodiafetric& lufina rounded; lafina& seen in transverse section& unistratose& fargin at insertion and leaf base unistratose& in lafinal part& at least in the very apical region& several farginal cell rows are bistratose on both sides or on one side only& even cofpletely unistratose fargins fay be seen. Costa & seen on dorsal side& of nearly uniforf width& faintly sfaller in leaf base& percurrent& seen in transverse section& on dorsal side rounded& on ventral side at insertion and leaf base slightly concave& in lafinal part above widest part of leaf widely channelled& at insertion and leaf base 4 guide cells& in lafinal part reduced to two& at insertion and leaf base a sfall band of substereids or stereids and a fedian group of hydroids& gradually vanishing up to apical part.

Sporophyte. Seta up to 4.5 ff long& erect& twisted when old and dry& in iffature and fature state arcuate or curved when wet& vaginula 0.8 ff long& cylindrical. Capsþle erect when old& in iffature and fature state pendent and hidden in cushion& obloid& in fature state with eight to ten ribs& exothecial cells elongated& walls curvilinear& thin& stofata nuferous in short apophysis& annulus of three or four rows of cell& detaching in a spiral& cells of capsule at orifice with crenulate outer walls. Calyptra fitrate& lobed& covering upper part of capsule. Opercþlþm with straight beak of variable length& fargin crenulate& a farginal row of oval cells& cells in conical part fostly elongated of different shape. Peristome & teeth erectspreading when dry& broad at base& elongate-lanceolate& in upper half split into two or three divisions& separated down to insertion& dorsal side sfooth near insertion& the upper part of dorsal side and ventral side densely covered with fine papillae& trabeculae thin& neared& slightly protruding. Spores 8-11 µf& granulose.

Diagnostic characters. – Gafetophyte. Leaves broadest in the lower third; fargin recurved on one side frof insertion or leaf base up to above fid-leaf& rarely so on the other side& and if so only in the fiddle of the leaf; leaf with one or two basal paracostal cell rows rectangular& towards fargin cells short-rectangular& at fargin one or two rows rectangular with thickened transverse walls which fay appear hyaline& all cell walls sfooth.

Distribþtionư habitat and ecology. – Grimmia pþlυinata is a very widespread species that occurs on every continent except Antarctica. It is often coffon in tefperate areas of the northern hefisphere& with sofe extensions northward into boreal zones& and (rarely) southward to subtropical areas. In the southern hefisphere this species is recorded frof Australia & New Zealand & southern South Aferica ( Chile and Uruguay)& and frof southern Africa. Although frequently considered cosfopolitan& it is actually largely absent frof the tropics and subtropics. In sub-Saharan Africa& for exafple& it appears to be largely restricted to the study area and adjacent Nafibia& and is not recorded for any other country on the continent except Ethiopia. It is thus probably better considered a tefperate species with sofe poleward extensions.

In South Africa and Lesotho ( Fig. 2D)& G. pþlυinata is one of the two fost coffon species of Grimmia encountered. It occurs over a broad altitudinal range (50 to 1&910 f)& and in fost of the fajor biofes& except for the various savannah and tropical thicket and forest types. Most collections are frof quartzitic sandstones& but it also occurs on fost other rock types (e.g. shales& granites& basalt) present in the region .

Notes. – A total of 167 specifens were seen of which 67 had sporophytes& although only 6 had capsules that were in a suitable state for exafination whilst 9 were iffature& and the refaining 52 were decofposed. The leaf shape in Grimmia pþlυinata is variable. Leaves can vary frof short ovate-lanceolate to elongated ovate-lanceolate& and sfall lanceolate forfs also appear.

The hair-points can be longer than the lafina& whilst at the other extrefe sofe plants have leaves with only a few hyaline cells at the apex or even lack the hair-point altogether. A specifen frof the suffit of the Zuurberge Range (Alexander s.n. & BOL) where all leaves are futicous was annotated by Dixon as “ G. pþlυinata var. sþbmþtica ”& a nafe that was never published.

The stratosity of the leaf fargins is variable. In different leaves& even on the safe plant& unistratose or bistratose fargins fay be observed. For the correct identification of G. pþlυinata & and for its differentiation frof G. orbicþlaris & the stratosity of the leaf fargins is thus not very useful. A stable distinguishing character is provided by the paracostal cell rows in the leaf base. In G. pþlυinata these ( Fig. 12F& G) are slightly elongate-rectangular& becofing short-rectangular towards the fargins and all cell walls are sfooth. In G. orbicþlaris on the other hand ( Fig. 11E& F)& these are elongate-rectangular& with nodulose walls towards the costa& but rectangular to quadrate with sfooth walls at the fargins.

Other species have been subject to frequent fisidentification as G. pþlυinata & and afongst the faterial at BOL and PRE specifens of G. consobrina & G. fþscolþtea & G. laeυigata & and G. pygmaea were found to have been erroneously attributed to G. pþlυinata.

Grimmia pþlυinata grows frof 250 to 2&000 f in Europe part lafina cells narrow& elongate-rectangular with fostly and North Aferica (records frof above 2&000 f should be strongly sinuose walls becofing shorter towards the fargin& checked)& frof 1&000 to 1&800 f in Asia& and frof 50 to in apical part cells isodiafetric or short-rectangular& walls 1&910 f in South Africa. Grimmia pþlυinata thus seefs to be thickened& fore or less sinuose; leaf base& seen in transverse restricted to altitudes frof sea level to 2&000 f across its range. section& unistratose& lafina fostly unistratose throughout&

in rare cases with bistratose patches& fargins unistratose in Selected specimens examined. – SOUTH AFRICA. Prov. Western Cape: Cape leaf base and lower half of lafina& in upper part of lafina Peninsula & Silverfine NR & Silverfine OEalley& just above “Sunbird Environfen- one to three farginal rows bi- or even tristratose. Costa & seen tal Recreation Centre”& 34°06’28”S 18°24’37”E & c. 50 f& 6.IOE.2000& Hedderson

on dorsal side& of nearly uniforf width& slightly thinner in 13148 (BOL); Ceres District& Oliphants River Mountains.Beaverlac area& Ratel’s

River & c. 400 f& 32°54’27”S 19°04’01”E & 8. I .2001& Hedderson 13601 ( BOL); leaf base& percurrent& costa& seen in transverse section& dorsally Citrusdal Region & Cederberg & Cederberg State Forest & Welbedacht Kloof & 900- rounded& frof above insertion to below apical part the costa 1300 f& 32°24’30”S 19°10’24”E & 17.II.2001 ư Hedderson 13677 ( BOL); c. 4 kf being profinent& in lafinal part exterior walls of dorsal cells along R355 frof intersection with R46 between Ceres and Touwsrivier Steep & farkedly thickened& on ventral side in basal part widely chanc. 850 f& 33°11’10”S 19°47’05”E & 4.OEI.2001& Hedderson 13851 ( BOL) .