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Grimmia dissimulata E.Maier SPECIMENS EXAMINED. — Tunisia. Mogods, Bizerte Governorate, Delegation of Tinja: Ichkeul National Park, with sporophytes, 37°08’21.00”N, 09°40’23.35”E, 20 m a.s.l. (site 2019-96), 11.IV.2019, Ben Osman & Hugonnot (TUN[TUN2019-251]); High Tell, Beja Governorate, Delegation of Thibar: Djebba, with sporophytes, 36°28’15.20”N, 09°05’57.03”E, 620 m a.s.l. (site 2019-104), 13.IV.2019, Ben Osman & Hugonnot (TUN[TUN2019-252]); Tunisian Dorsal, Kef Governorate, Delegation of Kalaat Senan, Table de Jughurta, N side, with sporophytes, 35°44’50.28”N, 08°22’44.33”E, 1090 m a.s.l. (site 2020-34), 13.III.2020, Ben Osman & Hugonnot (TUN[TUN2020-10]). REMARKS Grimmia dissimulata was found growing on calcareous rocks and screes in deep semi-wooded valleys dominated by woods composed of olive and pistachio mastic trees (Jbel Ichkeul), on a cultivated slope (fig trees, pomegranate, etc.) and xerophytic scrubland (Djebba), and in a rupicolous calcicolous community (Table de Jughurta). In Tunisia, this strongly calcicolous species was observed mainly on scree in natural habitats, which corresponds to its known ecology in Europe (Maier 2002). However, it has also been reported from such artificial substrates as limestone walls and tombstones (Smith 2004; Lüth 2012). The species bears a superficial resemblance to Grimmia meridionalis (Müll.Hal.) E.Maier, G. lisae De Not. and G. trichophylla Grev., but it can be easily distinguished by examining a transverse nerve section that shows 4 guide cells in a single layer at insertion and at the leaf base arranged (G. lisae has 6 guide cells at insertion, and G. trichophylla has additional guide cells that form an incomplete second band). From G. meridionalis, it is harder to set apart: G. meridionalis has two guide cells in the upper part of the leaf, as G. dissimulata, but these are elliptical and oblique, whereas they have a large more or less circular lumen that is not oblique in G. dissumulata. Grimmia dissimulata is a European southern-temperate species widely distributed in the Mediterranean area (Ros et al. 2013). Scattered elsewhere in Europe (Hodgetts & Lockhart 2020), it also occurs in Western Asia (Kürschner & Frey 2020). Because it is a relatively newly described species (Maier 2002), its distribution is not yet completely known (Porley 2014)., Published as part of Osman, Imen Ben, Hugonnot, Vincent, Muller, Serge D. & Daoud-Bouattour, Amina, 2022, New bryophytes for Tunisia (North Africa). Part 2: other families, pp. 173-185 in Cryptogamie, Bryologie 20 (11) on page 179, DOI: 10.5252/cryptogamie-bryologie2022v43a11, {"references":["MAIER E. 2002. - Grimmia dissimulata E. Maier sp. nova, and the taxonomic position of Grimmia trichophylla var. meridionalis Mull. Hal. (Musci, Grimmiaceae). Candollea 56: 281 - 300.","SMITH A. J. E. 2004. - The Moss Flora of Britain and Ireland. Ed. 2. Cambridge University Press, Cambridge, 1026 p.","LUTH M. 2012. - Grimmia dissimulata new to Scandinavia. Lindbergia 35: 86 - 89.","ROS R. M., MAZIMPAKA V., ABOU- SALAMA U., ALEFFI M., BLOC- KEEL T. L., BRUGUES M., CROS R. M., DIA M., DIRKSE G. M., DRAPER I., EL- SAADAWI W., ERDAG A., GANEVA A., GABRIEL R., GONZALEZ- MANCEBO J. M., GRANGER C., HERRNSTADT I., HUGONNOT V., KHALIL K., KURSCHNER H., LOSADA- LIMA A., LUIS L., MIFSUD S., PRIVITERA M., PUGLISI M., SABOVLJEVIC M., SERGIO C., SHABBARA H. M., SIM- SIM M., SOTIAUX A., TAC- CHI R., VANDERPOORTEN A. & WERNER O. 2013. - Mosses of the Mediterranean, an annotated checklist. Cryptogamie, Bryologie 34: 99 - 283. https: // doi. org / 10.7872 / cryb. v 34. iss 2.2013.99","HODGETTS N. & LOCKHART N. 2020. - Checklist and country status of European bryophytes. Update 2020. Irish Wildlife Manuals 123: 1 - 214. http: // hdl. handle. net / 2262 / 93026","KURSCHNER H. & FREY W. 2020. - Liverworts, mosses and hornworts of Southwest Asia (Marchantiophyta, Bryophyta, Anthocerotophyta): A systematic treatise with keys to genera and species occuring in Afghanistan, Bahrain, Iraq, Iran, Israel, Jordan, Kuwait, Lebanon, Oman, Qatar, Saudi Arabia, Sinai Peninsula, Syria, Turkey, United Arab Emirates and Yemen (inc. Socotra Island). Nova Hedwigia, Beihefte 149: 1 - 267.","PORLEY R. D. 2014. - Grimmia dissimulata E. Maier, in BLOCK- EEL T. L., BOSANQUET S. D. S., HILL M. O. & PRESTON C. D. (eds), Atlas of British and Irish Bryophytes. The Distribution and Habitat of Mosses and Liverworts in Britain and Ireland. Vol. 1. Pisces Publications, Newbury: 497."]}

In the Republic of Bashkortostan, 13 epilithic moss species belonging to the genus Grimmia Hedw. have been found. An annotated list of species with information about the number of findings in different vegetation zones and a brief description of habitat conditions, as well as distribution map of species are provided. In study area, Grimmia species are more common in areas with volcanic rocks outcrops, i. e. the central part and the eastern slope of the Southern Urals, as well as in the Bashkir Trans-Urals. Habitat conditions of G. laevigata, G. longirostris and G. muehlenbeckii differ significantly by temperature and humidity. The species G. plagiopodia G. teretinervis, G. alpestris, G. anomala and G. tergestina are rare in Bashkortostan and recommended to be included in the appendix to the Red Data Book of the republic.

The genus Grimmia Hedw. has been considered taxonomically difficult because of its great morphological variability, and its treatments by different specialists have led to incongruent results. One of the debates in the genus is the species status of Grimmia limprichtii Kern, an Asian-European disjunct moss species that has been considered identical to Grimmia anodon Bruch & Schimp. or Grimmia tergestina Tomm ex Bruch & Schimp. It has also been regarded as the muticous-leaved male plants of G. tergestina. Based on a detailed analysis of the type and many non-type specimens combining the molecular and morphological data, the reinstatement of species rank for G. limprichtii is proposed. The diagnostic characteristics of G. limprichtii and its distinction from some closely related species, with which it may be confused, are discussed. Grimmia obtusifolia C. Gao & T. Cao is considered a synonym of G. limprichtii based on molecular and morphological data.

Understanding the effects of anthropogenic disturbance on species’ behaviour is crucial for conservation planning, considering the extent of habitat loss. We investigated the influence of anthropogenic disturbances including agriculture, urbanisation, protected areas, and the presence of novel predators, on the temporal and spatial behaviour of sympatric forest antelope (Tragelaphus scriptus, Philantomba monticola, Sylvicapra grimmia, and Cephalophus natalensis) in an anthropogenic matrix containing forest fragments in the Indian Ocean Coastal Belt of South Africa. We integrated land-use types with camera-trap data and compared activity patterns using circular statistics and occupancy modelling. Antelope species overlapped in temporal and spatial distribution and exhibited diurnal activity for 50% of the time. All species exhibited nocturnal activity for ∼25–33% of all observations, except for C. natalensis. Nocturnal activity varied between species and land-use types. The predators Canis familiaris, C. mesomelas and Caracal caracal negatively influenced occupancy of P. monticola, S. grimmia and C. natalensis. Humans negatively influenced temporal activity of P. monticola, and spatial distribution of T. scriptus and S. grimmia. C. familiaris had an overall negative influence on S. grimmia. We found spatial, and to a lesser extent temporal, segregation between species. However, plasticity in activity patterns existed, which varied in response to anthropogenic effects and novel predators.

Introduction. We found an unusual specimen collected in 1975 from Xinjiang Province, China which may represent Grimmia exquisita J.Munoz. However, this species is known only from the type localitie...

International audience; Assessment of human exposure to atmospheric metals is a challenge, and mosses seem to be good biomonitors to help this purpose. Lacking roots, they are easy to collect and analyze. However, to our knowledge, no formal comparison was made between cadmium (Cd) measurements in Grimmia mosses and alternative forecasts of atmospheric Cd pollution as those produced by the CHIMERE chemistry transport model. This work aims at studying this link to improve further biomonitoring. We compare 128 Cd measurements in the cemetery mosses of Paris and Lyon metropolitan areas (France) to CHIMERE Cd atmospheric forecasts. The area to consider around the cemetery for the CHIMERE forecasts has been defined by Kendall rank correlations between both information sources—Cd in mosses and CHIMERE Cd forecasts—from different area sizes. Then, we fit linear models to those two data sets including step-by-step different sources of uncertainty. Finally, we calculate moss predictions to compare predictions and measurements in the two cities. The results show an apparent link between the Cd concentrations in mosses and CHIMERE Cd forecasts including in addition the same unique covariate, the moss support (grave or wall), in the two cities. However, this model cannot be directly transposed from region to region because the strength of the link appears to be regional.

Feng, C., Kou, J., Zhang, G.-L. & Wang, Z. 2020. Range extension of two xerophytic mosses in China based on molecular and morphological data. – Herzogia 33: 300 –308.An abnormal morphotype of Grimmia crinitoleucophaea Cardot, a species widely known as G. poecilostoma Cardot & Sebille, has been found in Inner Mongolian typical steppe, and its identification confirmed based on molecular and morphological data. Its morphological variation and taxonomic status are also discussed. Didymodon canoae C.Feng, J.Kou & L.Feng, a species previously known only from its type location in Inner Mongolia, was discovered in Tibet which extends its distribution range to a high elevation region of southwest China. Diagnostic characters, differences compared to similar species, distributional remarks and photographs of the two species are provided.

Background The phylum Tardigrada consists of over 1,300 species that inhabit terrestrial, freshwater and marine environments throughout the world. In terrestrial habitats they live primarily in mosses, lichens, leaf litter and soil, whereas tardigrades in freshwater and marine environments are mainly found in sediments and on aquatic plants. More than 65 species have been previously reported in the state of Tennessee, USA. Methods Tardigrades present in moss cushions (Grimmia sp.) collected from a xerothermic habitat on the East Tennessee State University campus, Johnson City, TN, USA, were extracted, mounted on slides, identified, and counted. Additional samples of fresh dried moss were used for integrative analyses, including morphological analysis with phase contrast (PCM) and scanning electron microscopy (SEM), as well as molecular analyses of COI, 18S rRNA, 28S rRNA, and ITS-2 of the Macrobiotus and Milnesium species. Results Five species were found, including two species new to science: Viridiscus miraviridis sp. nov. and Macrobiotus basiatus sp. nov. Viridiscus miraviridis sp. nov. differs from other members of the genus mainly by having a different type of dorsal cuticle and some other, more subtle, morphometric characters. In addition to the two new species, Viridiscus perviridis and Viridiscus viridissimus were present, and males of Vir. viridissimus were found for the first time, the first record of males in the genus Viridiscus. Macrobiotus basiatus sp. nov. is most similar to Macrobiotus nelsonae, but it differs from Mac. nelsonae mainly by the stylet supports being situated in a more anterior position, shorter and narrower egg processes, and a smaller number of areoles around the egg processes. Moreover, the identification of Milnesium inceptum was confirmed as the first record for the USA by analysis of COI.

Eight major serpentine sites in the French Massif Central have been bryofloristically surveyed and their specific assemblages studied by performing systematic releves. Seventy bryophyte taxa are reported from these serpentine areas. Archidium alternifolium, Bryum gemmilucens, Cephaloziella stellulifera, Grimmia dissimulata, Racomitrium fasciculare, R. lanuginosum and Riccia subbifurca are dominant and characteristic species. Soil and rock communities harbour typical combinations of specialized and ubiquitous taxa. The occurrence of saxicolous species (Grimmia) on the ground may be a typical feature of serpentine communities. The scarcity of liverworts is underlined, as is the complexity of the controlling factors. Lack of competition and periodic submergence are probably ecological factors of prime importance. Successional pathways are certainly insufficiently known, especially as far as bryophytes are concerned. The causes of negligible succession on skeletal soils would benefit from renewed stud...

Background and Aims Widespread and diverse in modern ecosystems, mosses are rare in the fossil record, especially in pre-Cenozoic rocks. Furthermore, most pre-Cenozoic mosses are known from compression fossils, which lack detailed anatomical information. When preserved, anatomy significantly improves resolution in the systematic placement of fossils. Lower Cretaceous (Valanginian) deposits on Vancouver Island (British Columbia, Canada) contain a diverse anatomically preserved flora including numerous bryophytes, many of which have yet to be characterized. Among them is the grimmiaceous moss described here. Methods One fossil moss gametophyte preserved in a carbonate concretion was studied in serial sections prepared using the cellulose acetate peel technique. Key Results Tricarinella crassiphylla gen. et sp. nov. is a moss with tristichous phyllotaxis and strongly keeled leaves. The combination of an acrocarpous condition (inferred based on a series of morphological features), a central conducting strand, a homogeneous leaf costa and a lamina with bistratose portions and sinuous cells, and multicellular gemmae, supports placement of Tricarinella in family Grimmiaceae. Tricarinella is similar to Grimmia, a genus that exhibits broad morphological variability. However, tristichous phyllotaxis and especially the lamina, bistratose at the base but not in distal portions of the leaf, set Tricarinella apart as a distinct genus. Conclusions Tricarinella crassiphylla marks the oldest record for both family Grimmiaceae and sub-class Dicranidae, providing a hard minimum age (136 million years) for these groups. The fact that this fossil could be placed in an extant family, despite a diminutive size, emphasizes the considerable resolving power of anatomically preserved bryophyte fossils, even when recovered from allochthonous assemblages of marine sediments, such as the Apple Bay flora. Discovery of Tricarinella re-emphasizes the importance of paleobotanical studies as the only approach allowing access to a significant segment of biodiversity, the extinct biodiversity, which is unattainable by other means of investigation. Fil: Savoretti, María Adolfina. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina Fil: Bippus, Alexander C.. Humboldt State University; Estados Unidos Fil: Stockey, Ruth A.. State University of Oregon; Estados Unidos Fil: Rothwell, Gar W.. State University of Oregon; Estados Unidos Fil: Tomescu, Alexandru M. F.. State University of Oregon; Estados Unidos

Mamtimin, S., Bai, X.-L. & Zhao, D.-P. 2017. Grimmia orbicularis from China ‒ a species new to East Asia. ‒ Herzogia 30: 300–303. Grimmia orbicularis (Grimmiaceae) is reported for East Asia from China for the first time. A brief description and digital photographs of the species are provided, and the characters that distinguish it from the similar G. pulvinata are discussed.

A revision of Grimmia Hedw. (Grimmiaceae) from South Africa and Lesotho based on specimens housed in BM, BOL, G, PRE, and STU, as well as type material from B, BM, BOL, FH, G, H-BR, H-SOL, NY, and PRE, is presented. Thirteen species are recognized for South Africa and Lesotho : Grimmia consobrina Müll. Hal., Grimmia donniana Sm., Grimmia elongata Kaulf., Grimmia fuscolutea Hook., Grimmia kidderi James, Grimmia laevigata (Brid.) Brid., Grimmia longirostris Hook., Grimmia montana Bruch & Schimp., Grimmia orbicularis Wilson, Grimmia pulvinata (Hedw.) Sm., Grimmia pygmaea Müll. Hal., Grimmia sessitana De Not. and Grimmia tortuosa Hook. f. & Wilson. Keys based on morphological features, with an emphasis on costal characters, are provided. Each species is described and illustrated using key morphological and anatomical characters, such as transverse sections of leaves. Methods of specimen preparation are also explained and a glossary is supplied. Information on the geographical as well as altitudinal distribution of each species is given, with species distribution maps provided for the area covered by this study.

In analytical chemistry, biomonitoring is known as the methodology, which consider the use of living organisms to monitor and assess the impact of different contaminants in a known area. This type of monitoring is a relatively inexpensive method and easy to implement, being a viable alternative to be developed in sites where there is no infrastructure/instruments for a convenctional air quality monitoring. These organisms, having the capability to monitor the pollution, are also known as passive biomonitors (PBs), since they are able to identify possible contamination sources without the need of any additional tool. In this work, a multianalytical methodology was applied to verify the usefulness of naturally growing Grimmia genus mosses as PBs of atmospheric heavy metals pollution. Once mosses were identified according to their morphology and taxonomy, thei ability to accumulate particulate matter (PM) was determined by SEM. EDS coupled to SEM also allowed to identify the main metallic particles deposited and finally, an acid digestion of the mosses and a subsequent ICP-MS study define more precisely the levels of metals accumulated on each collected moss. The study was focused on six sampling locations from the Bilbao Metropolitan area (Biscay, Basque Country, north of Spain). The experimental evidences obtained allowed to propose naturally growing Grimmia genus as PB of atmospheric heavy metals pollution and to identify the anthropogenic sources that contribute to the emission of the airborne particulate matter rich in metals, evaluating in this sense the atmospheric heavy metals pollution of the selected locations.

Grimmia curviseta Bouman is a moss species endemic to the Canary Islands in the North Atlantic Ocean, which has a very restricted distribution across the summit areas of Tenerife and La Palma islands. Using massive sequencing, we developed ten polymorphic microsatellite markers for this species. The pattern of microsatellite alleles per locus provides preliminary evidence that G. curviseta is allodiploid. The average number of alleles per locus ranged from 2 to 7, and observed heterozygosities varied from 0.022 to 1.000. A significant genetic differentiation was observed between the Tenerife and La Palma populations, suggesting that there are some limitations to dispersal. This set of microsatellites constitutes a valuable tool to investigate patterns of genetic diversity across the distribution range of G. curviseta, information that may eventually be used to establish conservation strategies for the species. Owing to the cross-amplification with the closely related species Grimmia montana Bruch & Schimp., our study also demonstrates the utility of these markers for population-level genetic analyses in the genus Grimmia., This paper is part of the projects entitled ‘‘Lista Roja de los Briófitos de las Islas Canarias’’ funded by the Canary Islands Government (P1042004-028) and ‘‘Biogeografía y procesos evolutivos en briófitos. Estudios en las Islas Macaronésicas” funded by the Spanish Ministerio de Educación y Ciencia (CGL2005-00028). We

During investigation on the bryophytes of the Govind Wildlife Sanctuary, Grimmia nepalensis has been identified for the first time from this region. It is a new record for India and has been collected after about 150 years since its original collection from Nepal. The species is endemic to the Himalaya.

The aim of this study was to identify and determine the genetic diversity of Fasciola species in cattle from Zimbabwe, the KwaZulu-Natal and Mpumalanga provinces of South Africa and selected wildlife hosts from Zimbabwe. This was based on analysis of DNA sequences of the nuclear ribosomal internal transcribed spacer (ITS1 and 2) and mitochondrial cytochrome oxidase 1 (CO1) regions. The sample of 120 flukes was collected from livers of 57 cattle at 4 abattoirs in Zimbabwe and 47 cattle at 6 abattoirs in South Africa; it also included three alcohol-preserved duiker, antelope and eland samples from Zimbabwe. Aligned sequences (ITS 506 base pairs and CO1 381 base pairs) were analyzed by neighbour-joining, maximum parsimony and Bayesian inference methods. Phylogenetic trees revealed the presence of Fasciola gigantica in cattle from Zimbabwe and F. gigantica and Fasciola hepatica in the samples from South Africa. F. hepatica was more prevalent (64%) in South Africa than F. gigantica. In Zimbabwe, F. gigantica was present in 99% of the samples; F. hepatica was found in only one cattle sample, an antelope (Hippotragus niger) and a duiker (Sylvicapra grimmia). This is the first molecular confirmation of the identity Fasciola species in Zimbabwe and South Africa. Knowledge on the identity and distribution of these liver flukes at molecular level will allow disease surveillance and control in the studied areas.

A large amount of data for inconspicuous taxa is stored in natural history collections; however, this information is often neglected for biodiversity patterns studies. Here, we evaluate the performance of direct interpolation of museum collections data, equivalent to the traditional approach used in bryophyte conservation planning, and stacked species distribution models (S‐SDMs) to produce reliable reconstructions of species richness patterns, given that differences between these methods have been insufficiently evaluated for inconspicuous taxa. Our objective was to contrast if species distribution models produce better inferences of diversity richness than simply selecting areas with the higher species numbers. As model species, we selected Iberian species of the genus Grimmia (Bryophyta), and we used four well‐collected areas to compare and validate the following models: 1) four Maxent richness models, each generated without the data from one of the four areas, and a reference model created using all of the data and 2) four richness models obtained through direct spatial interpolation, each generated without the data from one area, and a reference model created with all of the data. The correlations between the partial and reference Maxent models were higher in all cases (0.45 to 0.99), whereas the correlations between the spatial interpolation models were negative and weak (−0.3 to −0.06). Our results demonstrate for the first time that S‐SDMs offer a useful tool for identifying detailed richness patterns for inconspicuous taxa such as bryophytes and improving incomplete distributions by assessing the potential richness of under‐surveyed areas, filling major gaps in the available data. In addition, the proposed strategy would enhance the value of the vast number of specimens housed in biological collections.

Two species, Tortula brevissima Schiffn. and Grimmia indica (Dixon & P. de la Varde) Goffinet & Greven are added to the Chinese moss flora. The former is newly reported for China and East Asia. The latter has been found in Tibet for the first time. Brief morphological descriptions, distribution details, and digital photographs of the two species are provided.

10. Grimmia pulvinata(Hedw.) Sm. in Engl. Bot. 24: 1728. 1807 (Fig. 12). [Fissidens pulvinatus Hedw., Sp. Musc. Frond. 158. 1801. Lectotypus (designated by Cao & Vitt, 1986: 191): Sine loC .: ���In tectis et muris vulgaris, vere maturans���, s.d., Anon. s.n. (G [G00040350]!). 5 Fissidens pulvinatus var. africanus Hedw., Sp. Musc. Frond. 159. 1801. Lectotypus (designated by Mu��oz, 1999: 168): South AfriCA : ���Ad Cap. bonae spei lecta, specimina misit O. Swartz���, s.d., Swartz s.n. (G [G0004035]!), synonymized by Magill (1981: 275). 5 Grimmia ecklonii Spreng., Syst. Veg. 4: 321. 1827. Lectotypus (designated by Mu��oz & Pando, 2000: 67): South AfriCA : Swellendam, s.d., Ecklon s.n. (BM [BM000575940]!; isolecto-: BM [BM000670219]!), synonymized by Mu��oz & Pando (2000: 67). Gametophyte. Monoicous. Female: innermost perichaetial leaf 2.2-2.5 mm long, sheathing up to apical part, tubulose, hyaline in lower part of leaf, costa excurrent to long, bluntly denticulate hair-point; male: perigonium as bud of 4 or 5 leaves, on short stalk in leaf axil on stem of fruiting plant, close to perichaetium, innermost perigonial leaf 1 mm long, sheathing up to apex, concave, ovate, apex acute or mucronate, costa vanishing below apex, hyaline except at apex, paraphyses few. Growth form: cushion dense, lax, young shoots originating from detritus, leaflets with hair-point, plants radiculose at base, erect, dichotomously branched, stems up to 15 mm high, central strand developed. Leaves in lower part of stem small, with hair-point, becoming gradually longer, up to 2.5 mm long, crowded, loosely disposed on stem, upper part of leaf slightly flexuose when dry, especially the comal ones, apical part quickly bending backwards when moistened, erecto-patent when wet, broadest part of leaf in lower third, broad-lanceolate or lanceolate from a short, ovate leaf base, apex obtuse, hair-point denticulate; leaf form in situ, leaf base widely concave or concave, laminal part widely keeled, margin recurved on one side from insertion or leaf base up to above mid-leaf, rarely on the other side and then, at most, in the middle of the leaf; in leaf base some paracostal cell rows of slightly elongate-rectangular cells, walls smooth, towards margin cells short-rectangular, at margin one or two rows of rectangular cells which may appear hyaline, transverse walls thickened, all cell walls smooth, in transitional part cells rectangular, walls sinuose, in laminal part mostly isodiametric, walls somewhat sinuose, thickened, at apex cells isodiametric, lumina rounded; lamina, seen in transverse section, unistratose, margin at insertion and leaf base unistratose, in laminal part, at least in the very apical region, several marginal cell rows are bistratose on both sides or on one side only, even completely unistratose margins may be seen. Costa, seen on dorsal side, of nearly uniform width, faintly smaller in leaf base, percurrent, seen in transverse section, on dorsal side rounded, on ventral side at insertion and leaf base slightly concave, in laminal part above widest part of leaf widely channelled, at insertion and leaf base 4 guide cells, in laminal part reduced to two, at insertion and leaf base a small band of substereids or stereids and a median group of hydroids, gradually vanishing up to apical part. Sporophyte. Seta up to 4.5 mm long, erect, twisted when old and dry, in immature and mature state arcuate or curved when wet, vaginula 0.8 mm long, cylindrical. Capsule erect when old, in immature and mature state pendent and hidden in cushion, obloid, in mature state with eight to ten ribs, exothecial cells elongated, walls curvilinear, thin, stomata numerous 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 mitrate, lobed, covering upper part of capsule. Operculum with straight beak of variable length, margin crenulate, a marginal row of oval cells, cells in conical part mostly 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 smooth near insertion, the upper part of dorsal side and ventral side densely covered with fine papillae, trabeculae thin, neared, slightly protruding. Spores 8-11 ��m, granulose. Diagnostic characters. ��� Gametophyte. Leaves broadest in the lower third; margin recurved on one side from insertion or leaf base up to above mid-leaf, rarely so on the other side, and if so only in the middle of the leaf; leaf with one or two basal paracostal cell rows rectangular, towards margin cells short-rectangular, at margin one or two rows rectangular with thickened transverse walls which may appear hyaline, all cell walls smooth. Distribution, habitat and ecology. ��� Grimmia pulvinata is a very widespread species that occurs on every continent except Antarctica. It is often common in temperate areas of the northern hemisphere, with some extensions northward into boreal zones, and (rarely) southward to subtropical areas. In the southern hemisphere this species is recorded from Australia, New Zealand, southern South America (Chile and Uruguay), and from southern Africa. Although frequently considered cosmopolitan, it is actually largely absent from the tropics and subtropics. In sub-Saharan Africa, for example, it appears to be largely restricted to the study area and adjacent Namibia, and is not recorded for any other country on the continent except Ethiopia. It is thus probably better considered a temperate species with some poleward extensions. In South Africa and Lesotho (Fig. 2 D), G. pulvinata is one of the two most common species of Grimmia encountered. It occurs over a broad altitudinal range (50 to 1,910 m), and in most of the major biomes, except for the various savannah and tropical thicket and forest types. Most collections are from quartzitic sandstones, but it also occurs on most other rock types (e.g. shales, granites, basalt) present in the region. Notes. ��� A total of 167 specimens were seen of which 67 had sporophytes, although only 6 had capsules that were in a suitable state for examination whilst 9 were immature, and the remaining 52 were decomposed. The leaf shape in Grimmia pulvinata is variable. Leaves can vary from short ovatelanceolate to elongated ovate-lanceolate, and small lanceolate forms also appear. The hair-points can be longer than the lamina, whilst at the other extreme some plants have leaves with only a few hyaline cells at the apex or even lack the hair-point altogether. A specimen from the summit of the Zuurberge Range (Alexander s.n., BOL) where all leaves are muticous was annotated by Dixon as ��� G. pulvinata var. submutica ���, a name that was never published. The stratosity of the leaf margins is variable. In different leaves, even on the same plant, unistratose or bistratose margins may be observed. For the correct identification of G. pulvinata, and for its differentiation from G. orbicularis, the stratosity of the leaf margins is thus not very useful. A stable distinguishing character is provided by the paracostal cell rows in the leaf base. In G. pulvinata these (Fig. 12 F, G) are slightly elongate-rectangular, becoming short-rectangular towards the margins and all cell walls are smooth. In G. orbicularis on the other hand (Fig. 11 E, F), these are elongate-rectangular, with nodulose walls towards the costa, but rectangular to quadrate with smooth walls at the margins. Other species have been subject to frequent misidentification as G. pulvinata, and amongst the material at BOL and PRE specimens of G. consobrina, G. fuscolutea, G. laevigata, and G. pygmaea were found to have been erroneously attributed to G. pulvinata. Grimmia pulvinata grows from 250 to 2,000 m in Europe and North America (records from above 2,000 m should be checked), from 1,000 to 1,800 m in Asia, and from 50 to 1,910 m in South Africa. Grimmia pulvinata thus seems to be restricted to altitudes from sea level to 2,000 m across its range. Selected specimens examined. ��� South AfriCA . Prov. Western Cape: Cape Peninsula, Silvermine NR, Silvermine Valley, just above ���Sunbird Environmental Recreation Centre���, 34��06���28���S 18��24���37���E, c. 50 m, 6.IV.2000, Hedderson 13148 (BOL); Ceres District, Oliphants River Mountains. Beaverlac area, Ratel���s River, c. 400 m, 32��54���27���S 19��04���01���E, 8.I.2001, Hedderson 13601 (BOL); Citrusdal Region, Cederberg, Cederberg State Forest, Welbedacht Kloof, 900- 1300 m, 32��24���30���S 19��10���24���E, 17.II.2001, Hedderson 13677 (BOL); c. 4 km along R355 from intersection with R46 between Ceres and Touwsrivier Steep, c. 850 m, 33��11���10���S 19��47���05���E, 4.VI.2001, Hedderson 13851 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 220-223, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Cao, T. & D. H. Vitt (1986). A taxonomic revision and phylogenetic analysis of Grimmia and Schistidium (Bryopsida, Grimmiaceae) in China. J. Hattori Bot. Lab. 61: 123 - 247.","Munoz, J. (1999). A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Missouri Bot. Gard. 86: 118 - 191.","Magill, R. E. (1981). Bryophyta. I (1). In: Leistner, O. A. (ed.), Fl. Southern Africa.","Munoz, J. & F. Pando (2000). A world synopsis of the genus Grimmia (Musci, Grimmiaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 83."]}

Grimmia Hedw., Sp. Musc. Frond.: 75. 1801. Note. ��� A description of the genus Grimmia can be found in Maier (2010: 14)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on page 205, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Maier, E. (2010). The genus Grimmia Hedw. (Musci, Grimmiaceae) - A morphological-anatomical study. Boissiera 63."]}

5. Grimmia kidderi James in Bull. Torrey Bot. Club 6: 54. 1875 (Fig. 7). Lectotypus (designated by Mu��oz, 1999: 143): frAnCe . French Southern and Antarctic Territories: Kerguelen Island, 1874, Kidder s.n. (FH!; isolecto-: FH!). Gametophyte. Dioicous. Female: innermost perichaetial leaf up to 4 mm long, slightly sheathing up to shoulder at midleaf, costa stout, excurrent to scarcely denticulate hair-point; male: perigonia not seen. Growth form: cushion dense, adherent to substrate with grayish, in wet state hyaline, rhizoids, interwoven with young shoots, originating from older stem parts, leaflets scale-like, concave, muticous or with a hyaline end cell, in dry state firmly appressed to stem, apices spreading, producing bristly aspect, plants erect, sparsely branched, stems up to 10 mm high, central strand small. Leaves from lower part of stem 0.3-0.5 mm long, concave, muticous, becoming progressively longer, up to 1.7 mm long, loosely arranged to stem, erect when dry, moving weakly when moistened, erect or erecto-patent when wet, from ovate leaf base narrowly lanceolate, tapering to obtuse apex, muticous or with hairpoint of different length, bluntly denticulate; leaf form in situ, from insertion up to apical part concave or above broadest part of leaf widely keeled, in apical part keeled, margins plane throughout; basal paracostal cells elongate-rectangular, thickwalled, more or less nodulose, towards margin rectangular, becoming short-rectangular to quadrate in transitional part, walls smooth, thickened, especially the transverse walls, cells in laminal part isodiametric, lumina rounded; seen in transverse section, leaf base unistratose, in lower laminal part uni- to bistratose, apical part bistratose, occasionally with tristratose marginal cell rows. Costa, seen on dorsal side, of even width throughout, not reaching apex, seen in transverse section, costa rounded, on ventral side widely channelled, in upper part of leaf channelled or narrowly channelled, at insertion and leaf base 4 guide cells, in laminal part 2, with hydroids. Sporophyte. Seta slightly curved, 1.2 mm long, vaginula 0.6 mm long, with ochrea. Capsule hidden in the leaves, ovoid, smooth, after spore release enlarged at orifice, exothecial cells elongated, walls thick, in surface view strongly thickened, stomata rare in capsule base, annulus disintegrating in fragments. Calyptra not seen. Operculum conical, beak long, oblique, smooth at margin, all cells rounded, thick-walled. Peristome teeth reflexed in dry state, entire, on inner and upper outer side densely ornamented with sharp papillae, the lowest outer plates finely and sparingly papillose, trabeculae close together, small, scarcely protruding. Spores 10-13 ��m, finely granulose. Diagnostic characters. ��� Gametophyte. Perichaetial leaves markedly longer than vegetative leaves; leaves narrowly lanceolate, concave nearly throughout, margins plane. Sporophyte. Capsule immersed. Distribution, habitat and ecology. ��� Grimmia kidderi is a southern-temperate species recorded from Argentina, Chile, the Kerguelen Islands, Ascension Island, Tristan da Cunha and South Africa. In South Africa and Lesotho G. kidderi is rare and currently known only from two localities on the high summits of the northern and middle Cederberg from 1,150-1,870m, as well as from a single locality on the Great Escarpment of the Eastern Cape Province (Fig. 1 E). This species is found in fynbos with outcropping sandstone, and on sandstone cliffs and seasonally wet rock slabs. Whilst apparently genuinely rare in the west, it may be under-collected in the eastern part of its range. Notes. ��� The four specimens seen were sterile. The description of the sporophyte herein is based on Skottsberg 371 (H-BR) from the Patagonian region of Argentina (Maier, 2010: 180). The stratosity of the lamina in this species is variable. Some collections of G. kidderi from the study area have bulging cells with hyaline outer walls on the dorsal surface of the leaves. This phenomenon is not restricted to this species and seems to be due to harsh conditions under which the plants are growing. Selected specimens examined. ��� South AfriCA . Prov. Western Cape: Cederberg. Zuurvlakte, 1150 m, 32��36���32���S 19��12���12���E���, 27.II.2000, Hedderson 13081 (BOL); Citrusdal Region, Cederberg State Forest, E side of Langberg, and slopes of Shadow Peak, 1550-1870 m, 32��23���20���S 19��10���25���E, 17.II.2001, Hedderson 13724 (BOL); ibid. loc., Hedderson 13744 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 212-213, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. (1999). A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Missouri Bot. Gard. 86: 118 - 191.","Maier, E. (2010). The genus Grimmia Hedw. (Musci, Grimmiaceae) - A morphological-anatomical study. Boissiera 63."]}

13. Grimmia tortuosa Hook. f. & Wilson in London J. Bot. 3: 540. 1844 (Fig. 15). Lectoypus (designated by Mu��oz, 1999: 174): United Kingdom . Falkland Islands Dependencies: ��� Antarct. Exp. 1938-1843���, s.d., Hooker s.n. (BM [BM001007061]!; isolecto-: NY [NY01162773]!). Gametophyte. Monoicous (fide Cao & Churchill, 1995 and Mu��oz, 1999). Neither sporophytes nor perigonia seen. Growth form: cushions dense, compact, adherent to substrate by smooth rhizoids, plants sap green, radiculose at base, ascending, branched, stems up to 10 mm high, central strand well-developed or not at all, with intermediate states on one and the same stem. Leaves on lower part of stem muticous, 0.3- 1.0 mm long, suddenly becomimg markedly longer, up to 2-3 mm long, crowded, contorted, forming comal tuft when dry, scarcely moving when moistened, erecto-patent, stiff when wet, from elongate leaf base lanceolate, tapering to acute or obtuse apex, hair-point of different lengths, bluntly denticulate; leaf form in situ, at insertion and in leaf base concave, in transitional part widely keeled, in lower laminal part keeled, in upper laminal part narrowly or very narrowly so, margins plane throughout; basal cells hyaline, elongate-rectangular, longitudinal and transverse walls of even thickness, thin and smooth, scarcely nodulose, towards margin the hyaline cell rows gradually vanishing, outermost row reaching up to the broadest part of leaf, thus forming between hyaline basal cells and thicker walled chlorophyllose lamina cells a more or less well expressed delimitation, running obliquely from costa to margin, lamina cells isodiametric, lumina rounded; lamina, seen in transverse section, unistratose throughout, margin unistratose at leaf base, in upper half of leaf some cell rows bistratose or tristratose. Costa, seen on dorsal side, small at leaf base, markedly stout in laminal part, indistinct at apex, percurrent or excurrent to hair-point, seen in transverse section, at insertion and leaf base rounded, from there up to apex prominent, on ventral side at insertion plane, in transitional zone widely channelled, in upper half of leaf narrowly channelled, from insertion up to below apical part 4 guide cells, in apical part 2 guide cells, at insertion and leaf base costa composed of nearly uniform substereids, transformed in laminal part to a large band of stereids, dorsal cells with large lumina, no hydroids seen throughout. Sporophyte. Not seen. Diagnostic characters. ��� Gametophyte. Leaves with margins plane on both sides throughout. Costa in laminal part prominent, with a large band of stereids, and hydroids lacking. Distribution, habitat and ecology. ��� Grimmia tortuosa is currently known only from the type locality in the Falkland Islands, and the locality newly reported here from the study area. In South Africa and Lesotho (Fig. 2 F), Grimmia tortuosa is currently known from a single locality at 1,940 m on the Waiihoekberge, at the southern end of the Hex River Mountains, where it occurs on damp, hard, quartzitic sandstone slabs in a shallow ravine where snow occasionally lies in late winter. Other species associated with it (e.g. Blindia magellanica M��ll. Hal., Platyneurum praealtum (Mitt.) Ochyra & Bednarek- Ochyra) also exhibit similar disjunctive distributions. Notes. ��� The description here is based on a unique, sterile specimen (Hedderson 14459, BOL). The descriptions given by Cao & Churchill (1995) and Mu��oz (1999) as well as the protologue in Hooker & Wilson (1844) have also been taken into consideration for the description. Although geographically incongruous, the plants from South Africa match with both the description and the types of G. tortuosa, including the distinctive characters of a costa that is exceptionally rich in stereids below the middle of the leaf (see Fig. 15 E). Specimen examined. ��� South AfriCA . Prov. Western Cape: Ceres area, Waaihoek Berge, Waaihoek Peak, c. 1940 m, 33��28���15���S 19��18���54���E, 18.III.2002, Hedderson 14459 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on page 227, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. (1999). A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Missouri Bot. Gard. 86: 118 - 191.","Cao, T. & S. P. Churchill (1995). An assessment of Grimmia tortuosa Hook. f. & Wils. (Bryopsida: Grimmiaceae). Nova Hedwigia 60: 515 - 518.","Hooker, W. J. & W. Wilson (1844). Musci antarctici. London J. Bot. 3: 533 - 556."]}

6. Grimmia laevigata (Brid.) Brid. , Bryol. Univ. 1: 183. 1826 (Fig. 8). [Campylopus laevigatus Brid., Muscol. Recent. Suppl. 4: 76. 1819. Lectotypus (designated by Cao & Vitt, 1986: 181): itAly : sine loc., s.d., Anon. s.n. (B!). 5 Grimmia senilis Shaw in Cape Monthly Mag. 17: 380. 1878. Typus: South AfriCA . Prov. Cape: Dry karroo dyke on bare rock, Longhope Cookhouse, 900 m, IV.1921, Sim 9956 (holo-: PRE!), synonymized by Magill (1981: 277). Gametophyte. Dioicous. Female: innermost perichaetial leaf up to 2 mm long, sheathing up to broadest part of leaf, narrowed at leaf base, ovate to broad-ovate, lower half hyaline or only some rows of hyaline cells at margin, vanishing at mid-leaf, costa obscure in apical part, excurrent in long, denticulate hair-point; perigonia as multifoliose buds at tips of branches; male: innermost perigonial leaf up to 1 mm long, sheathing, broad-ovate, suddenly narrowed to acute apex with hyline cell, hyaline in lower part, costa vanishing below apex, paraphyses short, numerous. Growth form: cushion lax, young shoots mostly present, originating from decomposing plants, with scale-like leaflets appressed to stem, costa vanishing below acute apex, with sharply pointed hyaline cell or short hair-point; plants erect, scarcely branched, slightly radiculose at base, stems up to 20 mm high, central strand welldeveloped. Leaves in lower part of stem scale-like, becoming gradually longer to tip of stem, 1.2-1.8 mm long, rarely up to 3 mm long, imbricate, appressed to stem when dry, older leaves bending backwards when moistened, younger leaves moving slightly, erect or spreading when wet; from short (~1/5 of leaf length), rounded, half-sheathing, slightly decurrent leaf base, lingulate or broad-lanceolate, tapering to obtuse or rounded, even acute apex, hair-point roughly denticulate, occasionally nearly smooth; leaf form in situ, widely concave or concave throughout, margin plane; some rows of basal paracostal cells rectangular, walls smooth or faintly nodulose, towards margin in sheathing part cells isodiametric or transversely rectangular or oval, transverse walls thicker than longitudinal walls, in laminal part cells homogeneous, rounded, walls thick; leaf base, seen in transverse section, unistratose, in transitional part bistratose in places, in laminal part bi- to tristratose, at margin from insertion up to apical part one or more cell rows unistratose, in apical part at least one side unistratose, at apex bistratose. Costa, seen on dorsal side, at leaf base large, from above widest part of leaf up to apex indistinct, excurrent, in laminal part dorsal cells not different from lamina cells, seen in transverse section, on dorsal side at insertion and lower part of leaf flat, weakly convex, at upper part slightly rounded, on ventral side at leaf base widely channelled, in upper part narrowly so, at insertion and leaf base from 7 to 11 ventral cells, most of them guide cells, a small median band of substereids, interrupted by 3 groups or one large central group of hydroids, substereids and hydroids vanishing in apical part, in transitional part number of guide-cells reduced to 4, in laminal part with 2 guide cells, sunken into narrow channel, adaxial cell walls strongly thickened. Sporophyte. Seta up to 3 mm long, straight, vaginula 1 mm long, cylindrical. Capsule emergent, obloid, smooth, exothecial cells elongated, of variable shape, slightly curvilinear, stomata in short neck, annulus of three rows of cells, detaching spirally in groups. Calyptra mitrate, in upper part brownish, lobed, covering upper part of capsule. Operculum conical, beak straight, blunt, margin uneven or crenulate, one or two marginal rows of rounded cells, in conical part irregular, walls thickened, curvilinear, faintly nodulose. Peristome teeth erect when dry, broad at base, slit half way down to two or three branches or perforate, lower dorsal side smooth, upper dorsal and ventral sides sparingly to densely covered with rounded papillae, trabeculae broad, distant, protruding, in upper third thin. Mature IX. Spores 12-16 ��m, smooth. Diagnostic characters. ��� Gametophyte. Leaves scale-like on young shoots and from lower part of stem with short hairpoint or at least with a sharp hyaline end cell (Figs. 8 B-D); base of stem leaves short, rounded, half-sheathing, at margin from insertion up to apical part one or more unistratose cell rows. Costa with hydroids, in upper part of leaf with 2 guide cells sunken into a narrow channel, their adaxial cell walls strongly thickened (Fig. 8 H), a specific character of G. laevigata. Sporophyte. Peristome teeth slit or perforated. Distribution, habitat and ecology. ��� Grimmia laevigata is widespread across temperate areas of the globe, with extensions into the tropics, and it occurs on all continents except Antarctica. This species is very common in arid and semi-arid zones, and especially in regions with a Mediterranean-type climate. In Africa it is common across the desert and semi-desert areas, and also occurs on higher summits in tropical regions, but it is apparently absent from west- and central-tropical Africa. In South Africa and Lesotho, G. laevigata, along with G. pulvinata, is one of the most widespread species in the study area where it has been recorded from all the major biomes (Fig. 1 F). It is especially common on shale substrates throughout the various types of karoo vegetation, occurring from 80 m to 2,250 m. Notes. ��� A total of 133 specimens were seen. Of these 16 had sporophytes, 5 of which had capsules that were in good condition. The leaf form and length are very variable, the hairpoint is short or elongate, sharply and densely denticulate, rarely nearly smooth. The normally bi-stratose lamina may show partially tristratose patches. In rare cases the costa may be overlaid ventrally by supplementary cells on one side or both sides of the leaf axis, thus covering the guide cells. A specimen of G. laevigata that is cited in Dixon & Gepp (1923) is housed in PRE as ���G. campestris Burchell [ex Hook.], no. 135, coll. Rehmann, Cape Town, s.d.���. Selected specimens examined. ��� South AfriCA . Prov. Northern Cape: Namaqualand, Richtersveld Mountains, Khubus, narrow valley running east of town below Vandersterrberg, 28��25���09���S 17��02���38���E, up to c. 800 m, 23.IV.2000, Hedderson 13219 (BOL, G). Prov. Western Cape: Koue Bokkeveld Mountains, Twee Riviere, Suikerbossie Farm, c. 900 m, 32��40���30���S 19��16���02���E, 7.IV.2001, Hedderson 13782 (BOL); c. 6 km von Clanwilliam dam, c. 200 m, 7.IX.1990, L��benau SA 62 (G, STU)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 213-215, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Cao, T. & D. H. Vitt (1986). A taxonomic revision and phylogenetic analysis of Grimmia and Schistidium (Bryopsida, Grimmiaceae) in China. J. Hattori Bot. Lab. 61: 123 - 247.","Magill, R. E. (1981). Bryophyta. I (1). In: Leistner, O. A. (ed.), Fl. Southern Africa.","Dixon, H. N. & A. Gepp (1923). Rehmann's South African Mosses. Bull. Misc. Inform. Kew 6: 193 - 238."]}

4. Grimmia fuscolutea Hook. , Musci. Exot. 1: 63. 1818 (Fig. 6). Lectotypus (designated by Mu��oz & Pando, 2000: 31): South AmeriCA : sine loc., s.d., Humboldt 50 [H 2673] (BM [BM000670173]!; isolecto-: BM000535435!). 5 Grimmia sanii Greven in Bryologist 99: 429. 1996. Typus: South AfriCA . Prov. Natal: Crows��� Nest. Mount aux Sources, 3048 m, VII.1947, Schelpe 2115 (holo-: PRE!; iso-: BOL! [3 packets filed under G. pulvinata (Hedw.) Sm.]), synonymized by Mu��oz (1999: 138). Gametophyte. Monoicous. Female: innermost perichaetial leaf up to 3.8 mm long, sheathing in lower part, shape and cell pattern as in stem leaves, lower third hyaline, costa excurrent to elongated, denticulate, slightly decurrent hair-point; male: perigonia below perichaetium in leaf axil as bud on short stalk, innermost perigonial leaf 0.8 mm long, slightly sheathing, ovate, apex rounded or obtuse, hyaline up to 2/3 of leaf length, costa percurrent, paraphyses short, few. Growth form: cushion compact, adherent to substrate with rhizoids, from where the young shoots originate, leaflets patent, muticous, apex acute, plants erect, branched in upper part, rhizoids in leaf axils, stems up to 15 mm high, central strand well-developed. Leaves up to 1.5 mm long, crowded, imbricate, appressed to stem and somewhat contorted when dry, slowly spreading when moistened, erecto-patent and stiff when wet, from ovate base lanceolate or broad-lanceolate, slightly asymmetric, tapering to acuminate apex, hair-point of different length, smoothly denticulate; leaf form in situ, concave at leaf base, with a more or less expressed plica near the costa on one side, and another one in the middle of the lamina on the other side, keeled in lower laminal part, narrowly keeled in upper laminal part, margin recurved or revolute on larger side from leaf base up to laminal part, on other side more or less recurved, in upper part both sides plane; all basal cells elongated, walls smooth or nodulose, thin- or thick-walled, at margin two or three rows of hyaline cells, vanishing above leaf base, in transitional part and lower half of lamina cells rectangular, walls strongly or weakly sinuose, arranged as strict, perpendicular rows parallel to costa, upper cells short-rectangular, walls sinuose; seen in transverse section, leaf base unistratose, laminal part unistratose, bistratose in places, apical part partly bistratose, in laminal part one or two marginal cell rows bistratose, dorsal and ventral exterior cell walls scarcely or distinctly bulging or occasionally with joint thickenings. Costa, seen on dorsal side, at insertion and leaf base weak, enlarged in laminal part, excurrent to hairpoint, in muticous leaves vanishing below apex, seen in transverse section, costa on dorsal side at insertion and in leaf base rounded, in laminal part prominent, mammillose, on ventral side channelled, narrowly so in upper part, at insertion and leaf base 4 ventral cells, 2 median ones are guide cells, the 2 outer ones belong partly to basal paracostal cells, above broadest part of leaf reduced to 2 narrow elliptical guide cells, obliquely arranged to leaf axis, at insertion and leaf base a median group of hydroids or stereids, transformed to substereids in upper laminal part, vanishing in apical part. Sporophyte. Seta arcuate, up to 2.5 mm long, vaginula 0.8 mm long, cylindrical, ochrea small. Capsule emergent, horizontal or pendent, ovoid, of various size, with apophysial part, plicate, in mature state nearly smooth, constricted below orifice, exothecial cells irregularly elongated, pentagonal, walls curvilinear, thin or thick (depending on focussing), stomata large, numerous at apophysis, annulus of 3-4 cell rows, detaching as spirals. Calyptra mitrate, lobed, covering operculum. Operculum conical, beak obtuse of variable length, margin smooth, some marginal rows of rounded or short-rectangular cells, in conical part cells rounded-rectangular, thinwalled. Peristome teeth spreading when dry, lanceolate, broad at base, perforated longitudinally or slit into two branches, at dehiscence joined at base, separating subsequently, lower dorsal side covered with prickly papillae, upper dorsal and ventral sides densely covered with pointed papillae, trabeculae thin, in upper part distant, scarcely protruding. Spores 10-12 ��m, granulose. Diagnostic characters. ��� Gametophyte. Leaves basally plicate on one side near the costa (Fig. 6 C), basal cells elongated (Fig. 6 D). In transverse section, costa with small, elliptical guide cells oriented obliquely to leaf axis in laminal part. Distribution, habitat and ecology. ��� Disjunctively distributed, Grimmia fuscolutea is known in the northern hemisphere from the European Alps, and from scattered localities in the Himalaya (India, Nepal) and from Japan. In the southern hemisphere this species occurs along the South American Andes as far north as Mexico and in Africa from South Africa and Reunion Island. In South Africa and Lesotho (Fig. 1 D), G. fuscolutea occurs in mid- to high-altitude sites (730-3,370 m) from the northern Cederberg, south through the Cape Fold Mountains, and eastward along the Great Escarpment (where it is known from only three localities), to the Drakensberg. It grows on hard, quartzitic sandstones, basalt or dolerite in fynbos and grassland habitats. Notes. ��� A total of 32 specimens were seen, 17 with sporophytes but only one with a capsule in a suitable state for examination. Selected specimens examined. ��� South AfriCA . Prov. Natal: Underberg, Distr. Sani Pass, 8.XI.1973, Hilliard & Burtt 7115 (BOL). Orange Free State Prov. Witzieshoek, base of Sentinel, 2740 m, Schelpe 7682 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 210-211, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. & F. Pando (2000). A world synopsis of the genus Grimmia (Musci, Grimmiaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 83.","Munoz, J. (1999). A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Missouri Bot. Gard. 86: 118 - 191."]}

1. Grimmia consobrina M��ll. Hal. , Syn. Musc. Frond. 1: 785. 1849 (Fig. 3). Lectotypus (designated by Mu��oz, 1999: 176): Chile : sine loc., s.d., P��ppig s.n. (NY [NY00322587]!; isolecto-: BM [BM000670949, BM000670950]!). Gametophyte. Dioicous. Female: innermost perichaetial leaf tubulose, to 3 mm long; male: plants in separate cushions with stem leaves mostly muticous, perigonia in multi-foliose buds, terminal and in leaf axils, several on a stem, innermost perigonial leaf concave. Growth form: dense cushions, adhering to substrate, plants radiculose at base, ascending, weakly branched, stems 0.5 to 1.5 mm high, capsule-bearing plants reaching up to 2.5 mm in height, central strand weak. Leaves becoming longer from stem base to tip, 1.5-3.0 mm long, of irregular length in capsule-bearing plants, imbricate, loosely arranged and slightly turned around stem, apices spreading when dry, older leaves quickly bending backwards when moistened, younger ones slowly so, patent to patulous when wet, in comal tuft patulous, from elongate-ovate, at insertion slightly narrowed, decurrent, auriculate leaf base tapering to acuminate apex, symmetric or falcate, muticous or with very short to elongate, bluntly denticulate hair-point, development of brood-bodies occurs on dorsal side of lamina in transitional part, destroying lamina cells only; leaf form in situ, at insertion and leaf base concave, lower laminal part keeled, upper part narrowly so, margin on one side recurved from insertion up to mid-leaf; basal paracostal cells elongate-rectangular, walls smooth or nodulose, towards margin several rows of shortrectangular or nearly quadrate, hyaline cells with smooth or nodulose walls reaching up to transitional part, at margin 2 rows of short- to elongate-rectangular hyaline cells with smooth walls, vanishing in transitional part or shortly above it, cells in transitional part and lower half of lamina rectangular with nodulose walls, in upper half nearly isodiametric, lumina irregular, oval or rounded; in transverse section leaf unistratose throughout, rarely bistratose in laminal part, at margin from above insertion up to below apical part one row of cells bistratose, in apex 2 or more rows bistratose. Costa, seen from dorsal side, strikingly small and thin from insertion up to broadest part of leaf compared to its stoutness in the laminal part, excurrent to hair-point, in transverse section costa weak on dorsal side at insertion, from leaf base up to apex rounded, channelled on ventral side to upper laminal part, narrowly so in apical part, at insertion 4 guide cells, the 2 outer belonging partly to paracostal cells, from leaf base up to apex 2 guide cells, a band of stereids, in upper laminal part a group of hydroids or substereids, in smaller plants stereids throughout. Sporophyte. Seta arcuate wet, straight when dry, to 3.5 mm long. Capsule obloid, ribbed, pendent or horizontal, apophysis well-developed, stomata numerous in 2 rows in the apophysis, annulus of 3-4 rows of cells, detaching as spirals. Calyptra conical, mitrate, lobed, covering operculum. Operculum conical, rostrate, beak straight or slightly oblique. Peristome teeth erect when dry, lanceolate, in upper part slit in two small branches, lower half more or less perforate, densely covered with fine papillae, apices rarely with rounded papillae, the outer lowest plates nearly smooth or with few fine papillae, trabeculae broad throughout, distant, strongly protruding, prostome more or less developed. Spores 11-16 ��m, finely granulose. Diagnostic characters. ��� Gametophyte. Leaves becoming longer towards the apex of the stem (Fig. 3B); costa at insertion and in leaf base strikingly thin and small compared to its stout appearance in the laminal part (Fig. 3F, G); hydroids or substereids present in upper part of costa. Sporophyte. Capsule with well-developed apophysis; peristome teeth with trabeculae broad, distant, protruding. Distribution, habitat and ecology. ��� Grimmia consobrina is essentially a south-temperate species known from southern Africa, Australia, New Zealand, and southern South America, but extending northwards along the Andes into Central America and the mountains of Mexico and California. In South Africa and Lesotho (Fig. 1 B) this species is common in the mountains of the Western Cape, where it occurs predominantly on nutrient-poor quartzitic sandstones in fynbos at altitudes from 450-2,000 m. It is especially common in the Cedarberg-Koue Bokkeveld-Groot Winterhoek area, extending eastward through the Cape Fold Mountains to the Swartberg, where it becomes more rare. Also known from two disjunct localities in the Drakensberg of Lesotho and adjacent South Africa, where it occurs on basalt rocks in alpine grasslands. Notes. ��� A total of 64 specimens were examined, of which 16 had sporophytes, two of which had capsules in a suitable state. The nature of the costa in G. consobrina, which is strikingly thin at the insertion compared to its prominence in the laminal part, is unique among Grimmia species. The auriculate leaf base is best seen in well-developed leaves that are carefully removed, one by one, as the alar cells are very fragile. The more or less rectangular cell rows at the margins of the leaf base appear as a hyaline zone that narrows towards the broadest part of the leaf. Plants with leaves that are predominantly muticous have been named G. imberbis M��ll. Hal. (a synonym of G. consobrina, see Maier, 2010). In the study area, G. consobrina has frequently been misidentified as G. trichophylla, a species that is absent from South Africa and Lesotho. Selected specimens examined. ��� South AfriCA . Prov. Western Cape: Cederberg, Zuurvlakte, c. 1150 m, 32��36���32���S 19��12���12���E���, 27.II.2000, Hedderson 13088 (BOL); Citrusdal Region, Cederberg State Forest Welbedacht Kloof, 900- 1300 m, 32��24���30���S 19��10���24���E, 17.II.2001, Hedderson 13678 (BOL); ibid. loc., E side of Langberg and slopes of Shadow Peak, 1550-1870 m, 32��23���20���S 19��10���25���E, 17.II.2001, Hedderson 13726 (BOL); Koue Bokkeveld Mountains, Twee Riviere, Suikerbossie Farm, c. 900 m, 32��40���30���S 19��16���02���E, 7.IV.2001, Hedderson 13754 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 205-207, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. (1999). A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Ann. Missouri Bot. Gard. 86: 118 - 191.","Maier, E. (2010). The genus Grimmia Hedw. (Musci, Grimmiaceae) - A morphological-anatomical study. Boissiera 63."]}

11. Grimmia pygmaea M��ll. Hal. , Syn. Musc. Frond. 1: 787. 1849 (Fig. 13). ��� Grimmia trichophylla var. australis Hampe in Syn. Musc. Frond. 1: 788. 1849. [nom. inval.]. Lectotypus (designated by Mu��oz & Pando, 2000: 67): AuStrAliA : ���Gr. trichoph.-australis Hamp. N. Holl.���, s.d., Anon. s.n. (H-SOL!). 5 Grimmia drakensbergensis Sim in Trans. Roy. Soc. South Africa 15: 209. 1926. Typus: South AfriCA . Prov. Natal: Top of Giants Castle. 2440 m, 1912, Sim 9962 [leg. Symons s.n.] (holo-: PRE), synonymized by Maier (2010: 300). Gametophyte. Monoicous. Female: innermost perichaetial leaf up to 2 mm long, sheathing, costa stout, percurrent, hair-point scarcely denticulate; male: perigonia in leaf axils or on short branches originating there, several on a stem, usually far from the perichaetium, innermost perigonial leaf 0.8 mm long, hyaline except the apical part, costa percurrent. Growth form: cushions adhering to the substrate by rhizoids or desintegrating, occasionally young shoots originating from the stem base, apices of the leaflets appressed to stem, plants erect, stems from 15 mm up to 70 mm high, branched by innovations, leaves arranged in tiers, stem with well-developed central strand. Leaves 1.5-2.5 mm long, densely set, in dry state appressed to stem, slightly turned, scarcely moving when moistened, erecto-patent when wet, from elongate-ovate leaf base lanceolate, tapering to acute apex, hair-point weakly denticulate; leaf form in situ, at base concave, lower laminal part keeled, upper part narrowly so, margins weakly recurved on both sides from above leaf base to upper part of lamina, apical part of lamina plane; basal cells elongate-rectangular, walls thickened, weakly to strongly nodulose, variable from leaf to leaf on the same plant, near margin two or three rows of shorter cells with smooth walls appearing hyaline and differentiated from the elongated nodulose basal cells, in transitional part lamina cells narrow, elongate-rectangular with mostly strongly sinuose walls becoming shorter towards the margin, in apical part cells isodiametric or short-rectangular, walls thickened, more or less sinuose; leaf base, seen in transverse section, unistratose, lamina mostly unistratose throughout, in rare cases with bistratose patches, margins unistratose in leaf base and lower half of lamina, in upper part of lamina one to three marginal rows bi- or even tristratose. Costa, seen on dorsal side, of nearly uniform width, slightly thinner in leaf base, percurrent, costa, seen in transverse section, dorsally rounded, from above insertion to below apical part the costa being prominent, in laminal part exterior walls of dorsal cells markedly thickened, on ventral side in basal part widely channelled, in laminal part channelled, at insertion and at leaf base with 4 guide cells, in laminal part 2 guide cells, in mid-leaf small and elliptic in shape, mostly obliquely arranged to leaf axis, in leaf base and laminal part a centrally arranged group of hydroids or only one big star-shaped hydroid, vanishing in apical part, the dorsal costal cells with a small rounded lumen, transformed to substereids or stereids in the lower part of the leaf, in the prominent part of the costa a series of stereids arranged around the hydroids. Sporophyte. Seta to 2 mm long, arcuate or curved, vaginula 0.8 mm long, cylindrical. Capsule exserted, horizontal or slightly pendulose, obloid, ribbed, exothecial cells elongated, rectangular, penta- and hexagonal, walls thin, stomata numerous on short neck, annulus of 3 to 4 rows of cells detaching in fragments, cells at orifice of capsule with smooth, or rarely slightly crenulate outer walls. Calyptra mitrate, covering the upper third of capsule. Operculum conical with short beak, at rim some rows of cells with rounded lumina, in conical part rectangular or of irregular shape. Peristome teeth erect when dry, lanceolate, divided in the upper half into two divisions, the lower dorsal plates smooth, the subsequent plates with fine papillae, upper dorsal and ventral sides densely covered with rough papillae, trabeculae small to broad throughout, distant. Spores 11-15 ��m, nearly smooth. Diagnostic characters. ��� Gametophyte. Cells in leaf base elongate-rectangular with nodulose walls, lamina cells in transitional part elongate-rectangular with sinuose walls. Costa rounded except from above insertion up to below apical part where the costa is prominent and the exterior walls of dorsal cells are markedly thickened. In the same part, the guide cells are small and elliptic, mostly arranged obliquely to the leaf axis, whilst in the leaf base and laminal part a centrally arranged group of hydroids or one star-shaped hydroid can be observed, vanishing in the apical part. Distribution, habitat and ecology. ��� Grimmia pygmaea is a south-temperate species, known from New Zealand, Australia, Patagonia, and the Kerguelen Islands. In South Africa and Lesotho (Fig. 2 E) G. pygmaea is the most common species of Grimmia at high altitudes (>2,500 m) in the Drakensberg of Lesotho and adjacent South Africa, where it grows on basalt. Within the study area the vast majority of known populations are from this region. However, it is also known from a smattering of localities in the southern part of the Cape Fold Mountains, and from two intermediate stations along the Great Escarpment. In the western part of its range it mostly occurs on quartzitic sandstone at high altitude, but occasionally descends to near sea level (e.g. on the Cape Peninsula). Notes. ��� A total of 79 specimens were seen for this study and 38 of these had sporophytes, of which 3 were in a suitable state for examination, 9 were immature and 27 were decomposed. This species has been misunderstood, and largely neglected. Specimens from the study area were almost invariably identified as G. pulvinata. Whilst the two species are superficially similar, the paracostal cell differences (elongate-rectangular and nodulose in G. pygmaea versus shorter and smooth-walled in G. pulvinata) are diagnostic. Furthermore, G. pulvinata is predominantly found at lower elevations (not known from> c. 1,900 m), whilst most collections of G. pygmaea are from higher altitudes, although in the Western Cape it can occur at much lower elevations. The costal anatomy will also readily distinguish G. pygmaea from both G. orbicularis (also superficially similar, but ecologically very different) and G. pulvinata. Selected specimens examined. ��� leSotho : Mokhoapong Pass, along Mountain Road, 147 km E of Maseru, 2710 m, 30.XI.1977, Magill 4204 (BOL); Moseru District. Thaba-Putsoa (High Pass) on the road to Semonkong. 3090 m, 29��44���S 27��57���E, 20.IV.1994, Duckett & Matcham 1246a (herb. Matcham). South AfriCA . Prov. Natal: Drakensberg area, Umgatsheni valley between Sani Pass and Vergelegen, c. 2130 m, VII.1983, Esterhuysen 35934 (BOL); E. Cape, Lady Gray Distr., Witteberg, Joubert���s Pass, 2440 m, 18.I.1979, Hilliard & Burtt 12221 (BM)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 223-225, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. & F. Pando (2000). A world synopsis of the genus Grimmia (Musci, Grimmiaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 83.","Maier, E. (2010). The genus Grimmia Hedw. (Musci, Grimmiaceae) - A morphological-anatomical study. Boissiera 63."]}

7. Grimmia longirostris Hook. , Musc. Exot. 1: 62. 1818 (Fig. 9). Lectotypus (designated by Deguchi, 1984: 21): eCuAdor : sine loc., s.d., Humboldt 76 [H 2669] (BM [BM000671051]!; isolecto-: BM [BM000670951]!). Gametophyte. Mostly monoicous. Female: innermost perichaetial leaf up to 4 mm long, sheathing up to 2/3 of leaf length, leaf base elongated, in broadest part suddenly narrowed to subulate apical part, at margin, from insertion up to broadest part of leaf, a broad band of hyaline cells present, vanishing gradually in upper leaf, costa excurrent in denticulate hair-point; male: perigonia not seen. Growth form: in dense cushions, adherent to substrate with rhizoids, from which young shoots with scale-like leaflets originate, apices patent, muticous, acute, plants erect, branched, radiculose at base, stems up to 30 mm high, central strand well-developed. Lower leaves scale-like, upper stem leaves up to 3 mm long, loosely disposed on stem, apices may be contorted, imbricate when dry, bending backwards when moistened, erecto-patent when wet, from ovate base gradually narrowed to lanceolate lamina, tapering to acute apex, hair-point short, faintly to strongly denticulate; leaf form in situ, at base concave, one side plicate, in laminal part keeled, narrowly so below apex, margin on larger side from insertion up to mid-leaf recurved or revolute, other side mostly plane; basal paracostal cells elongate-rectangular, walls nodulose, thickened, transverse walls thin, often oblique, cells near margin quadrate to rectangular, walls smooth, transverse walls thickened, in some specimens hyaline up to broadest part of leaf, in transitional part paracostal cells short-rectangular, walls thickened, nodulose or smooth, upper lamina cells rounded-quadrate, walls thickened, faintly sinuose; seen in transverse section, basal cells unistratose, in transitional and lower laminal part uni- to bistratose, in upper part bistratose, at margin several rows of cells bistratose except at leaf base. Costa, seen on dorsal side, of nearly uniform breadth, slightly smaller at insertion, obscure in keeled part, percurrent, seen in transverse section, costa on dorsal side at insertion and leaf base widely rounded, in lower laminal part unevenly rounded, somewhat angulate, and slightly contracted at origin of lamina, becoming indistinct in upper laminal and keeled part, on ventral side at insertion and leaf base widely channelled, in laminal part channelled, at insertion and leaf base 6 guide cells, rarely only 4, in upper part 5 or 4, in indistinct part 2 guide cells, from insertion up to indistinct part a central band of stereids or substereids, occasionally a central group of hydroids, vanishing in upper part of leaf. Sporophyte. Seta straight, 1-5 mm long, vaginula 0.9- 1.2 mm long, cylindrical, with ochrea. Capsule exserted, emergent or immersed, erect, cylindrical, oblong-ovoid, with short neck, narrowed at orifice, smooth, in immature state shrivelled, exothecial cells multiform, elongated, walls curvilinear, stomata at transition above neck and at base of capsule, annulus of three or four rows of cells, detaching in a spiral. Calyptra mitrate, mostly with a single slit, covering upper third of capsule. Operculum rostellate, beak straight or slightly oblique, margin crenulate, one or two rows of rounded marginal cells, in conical part rhombic, rounded-rectangular, thick-walled. Peristome erect when dry, teeth lanceolate, entire or split into two or three branches, on dorsal side two or three plates smooth, the consequent dorsal part covered with fine papillae, the upper third of dorsal side and ventral side densely covered with conspicuous papillae, trabeculae small in lower part, thin in upper part. Spores 10-12 ��m, smooth. Diagnostic characters. ��� Gametophyte. Leaf apically keeled, margin recurved on broader side. Costa, in lower part of lamina, unevenly rounded, somewhat angulate. Distribution, habitat and ecology. ��� Grimmia longirostris is a very widely distributed species that is often associated with montane or sub-montane habitats in temperate and arctic regions. This species is found on all continents except Antarctica. It also occurs widely throughout the tropics in higher mountain systems. In South Africa and Lesotho (Fig. 2 A) this species is restricted to the high Drakensberg (1,860-3,370 m) and associated ranges, where it occurs on basalt in high-altitude grasslands. Notes. ��� A total of 56 specimens were seen, 16 with sporophytes but only 4 with capsules in a suitable state for examination, 3 were immature and 9 were decomposed. Misidentifications of this species appear largely to be responsible D for the persistent records of G. ovalis (Hedw.) Lindb. from South Africa, (see under Excluded species for South Africa and Lesotho below). A syntype of Grimmia longirostris is extant at G [G00281576]). Selected specimens examined. ��� South AfriCA . Prov. Natal: Bergville Dist., Mnweni area, Drakensberg, 2740 m, VII.1953, Esterhuysen 21637 (BOL); Bethlehem Division, Drakensberg, Mont-aux-Sources, Crows��� Nest, VII.1947, Schelpe 2116 (BOL [3 specimens]); Drakensberg: Umgatsheni valley S aspect bank, c. 2130 m, s.d., Esterhuysen 35931 (BOL)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 215-216, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Deguchi, H. (1984). Studies on some Patagonian species of Grimmiaceae (Musci, Bryophyta). In: Inoue, H. (ed), Studies on Cryptogams in Southern Chile: 17 - 72. Tokyo."]}

Key to Grimmia species from South Africa and Lesotho As plants are often collected without sporophytes, and when present the sporophytes are often in poor condition, we have utilised only gametophytic characters. (Plants should be examined in the wet state). 1. Cross section of costa at insertion with 7 or more guide cells (see Fig. 8 G, H), in lamina with 2 guide cells sunken into narrow channel, their adaxial walls strongly thickened, at insertion with a small median band of substereids, interrupted by 3 groups or 1 large central group of hydroids that vanish in the apical part.............................. 6. G. laevigata 1a. Costa cross section at insertion with fewer than 7 guide cells............................................................................... 2 2. Costa cross section at insertion with 6 guide cells (see Fig. 9 F); in lower lamina costa unevenly rounded and somewhat angulate as seen in surface view, indistinct in apical part of leaf; basal cells with transverse walls markedly thinner than longitudinal...... 7. G. longirostris 2a. Costa at insertion with 4 guide cells or with 4 guide cells of which the outer two are contiguous with the basal cells.............................................................................. 3 3. Costa at insertion with 4 guide cells, outer guide cells non-contiguous with the laminal cells......................... 4 3a. Costa at insertion with 4 guide cells, the two outer ones contiguous with the laminal cells (see Fig. 3F, G; 6G, H)....................................................................... 12 4. Basal paracostal cells (excluding marginal cells) nodulose....................................................................... 5 4a. Basal paracostal cells (including marginal cells) smooth..................................................................................... 8 5. Transverse section of leaf base not concave, leaf base with a marginal border (see Fig. 5 F) of narrowly elongaterectangular, hyaline, thin walled cells in 3-4 rows, gradually transitioning to short-rectangular to quadrate cells but with outermost row ascending to above broadest part of leaf; margin recurved on one side from insertion to mid-leaf (see Fig. 5 C, D); lamina unistratose (see Fig. 5 H), rarely with bistratose patches..... 3. G. elongata 5a. Transverse section of leaf base concave, plants otherwise... 6 6. Transverse section of leaf above broadest part broadly keeled (see Fig. 7 I), unistratose at insertion and leaf base, partially bistratose in laminal part, bistratose in apex; margin with several rows of bistratose cells; costa in transverse section (see Fig. 7 I), with or without hydroids, cells in transitional part of leaf with walls smooth or sinuose (see Fig. 7 H).................................. 5. G. kidderi 6a. Transverse section of leaf above broadest part not broadly keeled; plants lacking the above combination of characters............................................................................... 7 7. Transverse section of costa at insertion with dorsal cell walls slightly bulging (see Fig. 13 J, K), below mid-leaf with 2 narrowly elliptical median guide cells arranged obliquely to leaf axis; marginal cells partly bi-tristratose in 1-3 rows above the leaf base; laminal cells in transitional zone elongate-rectangular (see Fig. 13 G, I), walls sinuose................................................... 11. G. pygmaea 7a. Transverse section of costa at insertion with dorsal cell walls bulging (see Fig. 11 H), the guide cells rounded, arranged horizontally; margin unistratose or at most bistratose in 1 cell row on one side in apex; lamina cells in transitional zone short-rectangular or isodiametric with walls smooth or slightly sinuose (see Fig. 11 G)............................................................................ 9. G. orbicularis 8. Cells in leaf base elongate-rectangular......................... 9 8a. Cells in leaf base short-rectangular............................ 11 9. Basal cells of upper stem leaves elongate-rectangular (see Fig. 14 D), with thickened transverse walls throughout; lower stem leaves with inner elongate rectangular and evenly thickened walls (see Fig. 14 E), but with 2-3 marginal rows short-rectangular to quadrate cells and with thickened transverse walls; margin (transverse section) (see Fig. 14 C) plane or occasionally recurved on one side from insertion to transitional zone (see Fig. 14 F)................................................ 12. G. sessitana 9a. Cells of leaf base elongate-rectangular, hyaline with cell walls evenly thin, margins plane................................. 10 10. Costa in transverse section markedly stout in the lamina (see Fig. 15 H), indistinct at apex, without hydroids but with abundant stereids; leaf forming a v-shape in crosssection in upper leaf................................ 13. G. tortuosa 10a. Costa in transverse section not markedly stout in laminal part, distinct at apex; transverse section with hydroids, stereids absent but substereids usually present; leaf section otherwise in upper leaf............... 2. G. donniana 11. Leaves abruptly lanceolate from ovate base (see Fig. 10 B), thus with distinct shoulders, keeled (transverse section) from mid-leaf to apex (see Fig. 10 H, I); in transverse section margin gradually incurved from base to apex (see Fig. 10 D, I), lamina bistratose (see Fig. 10 H, I), costa prominent above leaf base....................... 8. G. montana 11a. Leaves broad-lanceolate or lanceolate from a short, ovate leaf base, keeled from leaf base to apex (transverse section) (see Fig. 12 H, I), thus lacking distinct shoulders, in transverse section margin recurved on one side to above midleaf (see Fig. 12 C), rarely on both sides and then with one side recurved only at middle of leaf; lamina unistratose from insertion to apex (see Fig. 12 H), usually with marginal 1-2 cell rows that are bi- tristratose, occasionally only on one side and rarely unistratose (Fig. 12 I), costa prominent throughout..................................... 10. G. pulvinata 12. Costa in dorsal view strikingly small and thin from insertion to broadest part of leaf (Fig. 3B), becoming stout and prominent towards the apex; transverse sections of costa in the upper stout portion with a group of hydroids (see Fig. 3F, G); leaf bases auriculate, decurrent (see Fig. 3D)................................................ 1. G. consobrina 12a. Costa in dorsal view weak at insertion and leaf base but enlarged throughout laminal part (Fig. 6 B); above the broadest part of leaf the 2 median guide cells become narrowly elliptical and obliquely oriented to leaf axis (see Fig. 6 G, H); hydroids sometimes present in leaf base (see Fig. 6 H); leaf bases neither auriculate nor decurrent........................................................ 4. G. fuscolutea, Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 203-204, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344

2. Grimmia donniana Sm. in Engl. Bot. 18: 1259. 1804 (Fig. 4). Lectotypus (designated by Mu��oz, 1998: 378): united Kingdom [WAleS]: Pales at Beddgelart [Beddgelert], VII.1802, Turner s.n. (BM [BM000918087!]). Gametophyte. Monoicous. Female: innermost perichaetial leaf to 2.5 mm long, sheathing to mid-leaf, not very different from stem leaves in shape and cell pattern, lower part of leaf hyaline, costa percurrent, hair-point long, denticulate; male: perigonia on short subperichaetial branches as buds on short stalks in leaf axils, innermost perigonial leaf 0.8 mm long, sheathing to broadest part, ovate, obtuse or mucronate, muticous or with acute hyaline end cell, in lower part or from base to below leaf tip hyaline, costa weak, percurrent, antheridia with few paraphyses. Growth form: dense, compact cushions, adherent to substrate by rhizoids, young shoots originating from rhizoids or older stem parts, leaflets muticous, apex acute, stems to 20 mm long, plants erect, branched, radiculose at base, central strand developed. Leaves in lower part of stem small, muticous, in upper part to 2.4 mm long, loosely arranged on stem, apices slightly bent towards stem when dry, longer leaves bending backwards, shorter leaves moving slightly when moistened, erecto-patent when wet, from narrow base elongate-lanceolate, tapering to rounded or obtuse apex, hair-point elongate, smoothly denticulate; leaf form in situ, at insertion and in leaf base concave, lower laminal part keeled, upper part narrowly so, margins plane throughout; basal paracostal cells elongate-rectangular, walls smooth, towards margin some rows of elongate-rectangular, hyaline cells present, longitudinal and transverse walls evenly thin, smooth, the rows gradually vanishing, outermost row reaching to above broadest part of leaf, thus forming a delimitation between hyaline and thicker walled chlorophyllose cells running obliquely from costa to margin, cells in transitional part near costa rectangular, walls sinuose, towards margin some hyaline cells with finely sinuose walls, lower lamina cells short-rectangular, walls more or less sinuose, thickened, in upper laminal part cells mostly isodiametric, walls thickened, smooth or sinuose; seen in transverse section, lamina unistratose or bistratose in places, occasionally totally bistratose, margin unistratose in leaf base, in laminal part several rows bi- or tristratose. Costa, seen on dorsal side, small in leaf base, becoming gradually broader in laminal part, indistinct at apex, percurrent or excurrent, seen in transverse section, on dorsal side at insertion and in leaf base angulate or rounded, in laminal part rounded or occasionally prominent, on ventral side at insertion nearly plane, in leaf base widely channelled, in lower part of lamina channelled, in upper part narrowly so, at insertion and in leaf base 4 guide cells, from transitional part to apical part 2 guide cells, from insertion to upper laminal part a median group of hydroids, at insertion and in leaf base costal cells substereid. Sporophyte. Seta of different lengths, 0.8-3.0 mm long, straight, occasionally curved, vaginula 0.8 mm long, cylindrical, ochrea broad. Capsule erect, horizontal at curved seta, ovoid, smooth, bright-coloured when mature, with age turning brownish, exothecial cells irregular, quadrate, hexagonal, elongate, thin-walled, seen in transverse section of capsule, exterior walls smooth, stomata large above capsule base, neck lacking, annulus of 2 rows of cells, if detaching then singly, seen in surface view cells round with small lumina. Calyptra mitrate, lobed, covering operculum. Operculum conical, mammillate, base smooth, cells rounded, thick-walled. Peristome teeth incurved or recurved when dry, broad at base, mostly entire at apex, separated down to insertion or at least to the lowermost plates, dorsal side in lower half covered with minute papillae, upper half of dorsal side and ventral side densely covered with fine papillae, trabeculae small, close together or distant, prominent in lower half, scarcely marked in upper part. Spores 8-9 ��m, smooth. Diagnostic characters. ��� Gametophyte. Leaf base with marginal rows of elongate-rectangular, hyaline cells, with longitudinal and transverse walls that are evenly thin, extending to broadest part of leaf (see Fig. 4 D); margin never recurved; calyptra mitrate. Distribution, habitat and ecology. ��� Grimmia donniana is disjunctively distributed in boreo-montane areas of the northern hemisphere, with extensions southwards in alpine areas. This species is also known in the southern hemisphere from New Zealand, southern Africa and Bolivia. In South Africa and Lesotho (Fig. 1 C) G. donniana is rare and known only from two high-altitude (> 3,000 m) sites in Lesotho, where it occurs on basalt in alpine grasslands. Specimens examined. ��� leSotho: Leribe, slopes and cliffs N of Bokong Headwaters, 3100 m, 29��5���S 28��21���E, 16.I.1995, Duckett, Hodgetts & Matcham 3274a (herb. Matcham); Kotisephola Pass, between Sani Top and Mokhotlong, 3240 m, II.1987, van Rooy 3407 (PRE)., Published as part of Eva Maier, Michelle J. Price & Terry A. Hedderson, 2017, A revision of Grimmia (Grimmiaceae) from South Africa and Lesotho, pp. 199-230 in Candollea 72 (1) on pages 207-208, DOI: 10.15553/c2017v721a12, http://zenodo.org/record/888344, {"references":["Munoz, J. (1998). A taxonomic revision of Grimmia subgenus Orthogrimmia (Musci, Grimmiaceae). Ann. Missouri Bot. Gard. 85: 367 - 403."]}

8. Grimmia montanaBruch & Schimp. in Bruch et al., Bryol. Eur. 3: 128. 1845 (Fig. 10). Lectotypus (designated by Cao & Vitt, 1986: 161): germAny : “ Felsen auf dem Donnersberg ”, IV.1843, Gümbel s.n. (BM [BM000670468]!). 5 Grimmia argyrotricha Müll. Hal. in Flora 73: 485. 1890. Neotypus (designated by Maier, 2010: 224): tAnzAniA : “Kilimandscharo, unter Schnee in Felsspalten nach W. vorspringendem Grat der Mawenzi- Spitze”, 4600 m, 31.X.1893, Volkens 1347 (H-BR!), synonymized by Maier (2010: 224). Gametophyte. Dioicous. Female: innermost perichaetial leaf 2.7-3.5 mm long, sheathing up to broadest part, at lower third of leaf cells transparent, some rows of hyaline marginal cells vanishing at broadest part, costa small at leaf base, stout in upper part, excurrent to long-excurrent and forming a denticulate hair-point; male plants in separate cushions, perigonia as multifoliose buds terminal or in leaf-axils, often several in one plant, innermost perigonial leaf 1 mm long, strongly sheathing, concave, apex acute or blunt, costa percurrent, paraphyses numerous. Growth form: cushion compact, interwoven with young shoots originating from rhizoids or older stem parts, leaflets appressed to stem, apices slightly patent, plants erect, radiculose at base, stems up to 20 mm high, central strand developed. Lower leaves small, muticous, suddenly longer, 1.8-2.5 mm long, lower part appressed to stem, apical part of longer leaves slightly twisted when dry, slowly bending backwards when moistened, erectopatent when wet, at transitional part from broad, ovate leaf base restricted to lanceolate laminal part, thus forming shoulder, tapering to acuminate apex, hairpoints of variable length, densely and bluntly denticulate; leaf form in situ, concave from leaf base up to upper part of leaf, upper third keeled, margins in laminal part from leaf base up to apical part gradually becomng plane or incurved; basal cells near costa rectangular, towards margin two or three rows of short-rectangular or quadrate cells, walls smooth, transverse walls markedly thicker than longitudinal walls, at margin some rows of hyaline cells, above shoulder at margin a longitudinal row of transversely oval cells, lamina cells small, more or less isodiametric, lumina rounded, lamina cells, seen in transverse section, short-rectangular, exterior cell walls smooth; leaf base unistratose, in transitional part lamina bistratose in places, in upper part bistratose, occasionally unistratose near costa, margin unistratose at leaf base, in laminal part bistratose, below apex occasionally tristratose. Costa, seen on dorsal side, small at leaf base, enlarged, prominent in laminal part, excurrent, costa, seen in transverse section, on dorsal side at leaf base rounded, in lower laminal part prominent, slightly angulate, in upper laminal part rounded, on ventral side at insertion widely channelled, in laminal part channelled or narrowly channelled, at insertion and leaf base 4 guide cells, occasionally enlarged to 5 cells, in laminal part reduced to 2, from insertion to mid-leaf with a central median group of hydroids, transformed to substereids in upper part, vanishing in apical part. Sporophyte. Seta up to 3.5 mm long, straight, vaginula 0.8 mm long, cylindrical. Capsule emergent, erect, obloid, occasionally slightly asymmetric, smooth, exothecial cells mostly elongated, rarely pentagonal, walls thin or thick, depending on orientation and focusing, curvilinear, stomata none, at orifice several rows of transversely rectangular cells, the two upper rows suggesting a persistent annulus. Calyptra cucullate, covering part of capsule. Operculum conical, rostrate or rostellate, beak straight or oblique, blunt, margin uneven, some rows of small, rounded marginal cells, in conical part larger oval or rounded cells. Peristome teeth erect when dry, lanceolate, irregularly perforated or slit to two unequal branches, lower dorsal part smooth, subsequent plates sparsely covered with fine papillae, upper dorsal and ventral sides densely covered with fine papillae, trabeculae thin, in lower part close together, in upper part distant, slightly protruding. Spores 10-12 µm, smooth. Diagnostic characters. – Gametophyte. Leaves with cells at basal margin short-rectangular to quadrate, with transverse walls strongly thickened, above shoulder margins with a row of transversely oval cells; in transverse section lamina cells shortrectangular, with exterior cell walls smooth; margins in laminal part plane or incurved. Distribution, habitat and ecology. – Grimmia montana is widely, but highly disjunctively distributed. In the northern hemisphere, it occurs in the warm and dry western interior of North America, extending south to Mexico and north to a few localities in the Yukon and Alaska, and with a few localities in eastern Canada and Greenland. It also occurs in Europe and adjacent North Africa, including Macaronesia, western and central Asia, and Hawaii. In the southern hemisphere it occurs only in Africa (Tanzania and southern Africa) and New Zealand. In South Africa and Lesotho (Fig. 2 B) most of the known populations of Grimmia montana are on seasonally wet quartzitic sandstone, often on slabs that receive water from melting snow, in the higher peaks (mostly above 1,500 m, but rarely down to 1,300 m) in the Cape Fold Mountains from the Hantamsberg and Northern Cederberg, south and eastwards to the Swartberg. Two disjunct populations occur on basalt at high altitude (>3,200 m) in the Drakensberg. This species is also likely to be found on the higher peaks of the Great Escarpment in the Eastern Cape; these areas remain bryologically poorly explored. Notes. – A total of 27 specimens were seen all of which were sterile. In female plants the basal cells may be elongated, and the loss of chlorophyll may give the leaf base a hyaline aspect. In the local specimens available for this study, no sporophytes were found in appropriate condition for the description of their characters. The name Grimmia argyrotricha Müll. Hal. has been used in various works on the bryophyte flora of Africa, e.g. O’Shea (2006: 92) and Van Rooy & Perold (2006: 15). The type specimen of G. argyrotricha, collected by Dr. Hans Meyer in the Kilimandscharo Massif in 1880 was destroyed at B and a neotype has been selected by Maier (2010: 232). Selected specimens examined. – South AfriCA . Prov. Western Cape: Citrusdal Area, Cederberg Apex Peak area, 1500 m. 32°35’34”S 19°12’27”E, 9.I.2001, Hedderson 13645 (BOL); Cederberg, Cederberg State Forest, E side of Langberg, 1550-1870 m 32°23’20”S 19°10’25”E, 17.II.2001, Hedderson 13706 (BOL); Cederberg, Sneeuberg, Ridge to NW of main peak, 1500-1700 m, 32°29’58”S 19°08’51”E, 7.IV.2002, Hedderson 14482 (BOL, G); ibid. loc., Hedderson 14491 (BOL); ibid. loc., Hedderson 14501 (BOL); Cederberg Wilderness Area, slopes on West side of Shadow Peak, 1850 m, 32°23’20”S 19°10’25”E, 22.II.2003, Hedderson 14990 (BOL).