Bryological Monograph An annotated checklist of the mosses of ...

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MOSSES OF EUROPE AND MACARONESIA 237usual non-fertile plants would be impossible to namewith certainty. However, European bryologists havelong recognized A. concinnatum without greatdifficulty from non-fertile material, so this form issurely best treated at species rank as advocated byLoeske.266. Sérgio, Cros & Brugués (1996) re-examined Anomobryumlusitanicum in detail, retaining it in the genusAnomobryum.267. Brachymenium commutatum is known in Europe onlyfrom the Sierra Nevada. It was first reported byThériot (1932) as B. commutatum var. hispanicumThér. and rediscovered in 1997 (Brugués et al., 2003).European plants are sterile and less robust than var.attenuatum Thér. & Trab., which was described fromthe Hoggar mountains, Algeria. Small forms are alsofound in the Hoggar (Thériot, 1932). Ochi (1972,1973) did not examine any of the type material of B.commutatum during his review of African Bryoideae.However, a specimen assigned to B. commutatum var.attenuatum from Sudan and material from Eritreawere reidentified by him as the widespread B. exileDozy & Molk. It appears very likely that B.commutatum should be placed as a synonym of B.exile.268. Brachymenium notarisii was placed in the genusHaplodontium for many years, within which it is thetype of subgenus Ateleobryum (Mitt.) Broth. (Wijk,Margadant & Florschütz, 1962). It was transferredfrom Haplodontium to Brachymenium by Shaw (1987),who commented that ‘it certainly does not belong inthe Mielichhoferioideae’.269. Podpěra (1954) identified Mielichhoferia paradoxawith ‘Bryum splachnoides C. M.’, i.e. B. cellulare.However, Shaw (1987) reidentified it as a distinctspecies known only from Macedonia that he placed inBrachymenium, an arrangement that was followed byCorley et al. (1991). The recent checklist of mosses ofthe former Yugoslavia (Pavletic, Martincic & Düll,1999) continues to treat Brachymenium paradoxumerroneously as Bryum cellulare.270. Brachymenium philonotula is known in our area onlyfrom Madeira (Rocha da Pena, 1928, leg. C. Barreto,S). According to Ochi (1972) it is perhaps conspecificwith B. exile Dozy & Molk.271. Bryum pamirense H.Philib. ex Broth. was reportednew for Europe from Komi Republic in northernRussia (Zheleznova & Shubina, 1998). However,Ignatov & Afonina (1992) marked it with ?, to denotea ‘particularly poorly known and putatively ill-definedspecies’. Bryum pamirense is apparently close to B.savicziae and B. uliginosum (Savicz-Ljubitzkaja &Smirnova, 1970), but differs from both of these inbeing synoicous rather than autoicous. Like B.savicziae, it has the cilia of the endostome welldeveloped and appendiculate, whereas B. uliginosumhas short cilia. It might be a form of B. uliginosum orpossibly a hybrid.272. Bryum savicziae Schljakov was described from theKola peninsula in Russia in 1951 but overlooked byIndex Muscorum (Wijk, Margadant & Florschütz,1959, 1969). It is known only from the type-locality. Itresembles B. uliginosum but has the cilia of theendostome well developed and appendiculate. Sporesizes of 19–31 mm given by Savicz-Ljubitzkaja &Smirnova (1970) seem rather variable and perhapsthis suggests a hybrid is involved. Ignatov & Afonina(1992) also reported B. ekstamii L.I.Savicz, B.labradorense H.Philib., B. umbratum I.Hagen and B.zemliae Arnell & Jaderh. from the north of EuropeanRussia. These little-known species require furtherstudy.273. Bryum vermigerum Arnell & C.E.O.Jensen wasdiscovered at Hardanger, Norway in 1915 and1916; it is otherwise known only by a recent reportfrom Iceland. Bryum vermigerum was treated as avalid species by Nyholm (1993) and its combinationof characters is undoubtedly different from that inany other European Bryum species. Nevertheless,Nyholm reported an extraordinarily wide range ofspore sizes (10–20 mm) and, although one capsuleexamined by Holyoak (at S) contained spores ofless variable size (10–14 mm), these included aproportion of small shrunken and presumablyabortive spores. The rarity of B. vermigerum, itspeculiar characters and the variable and sometimesabortive spores may suggest that an interspecifichybrid is involved.274. Bryum colombii Meyl. and B. mesodon J.J.Amann arelisted in the current Swiss list http://www.bryolich.ch/,but their identity is uncertain. Bryum murmanicumBroth. is on the Red List for the Murmansk Oblast ofRussia as Data Deficient; its identity is also uncertain.Bryum geheebii Müll.Hal. was treated as probably asynonym of B. alpinum by Corley et al. (1981), as aspecies by Düll (1985), but very doubtful by Düll(1992); its identity is uncertain. The identity of B. teresLindb. is also uncertain; it was treated by Düll (1985)as a synonym of B. nitidulum (here treated as asynonym of B. intermedium), but this synonymy wasquestioned by Nyholm (1993).275. Bryum gerwigii (Müll.Hal.) Limpr. is known from itstype-locality on limestone rocks above the Rheinfallesnear Schaffhausen in Switzerland and from a fewother reports from central Europe. As with othersubmerged Bryum, it is apparently so greatly modifiedby growth in water (and perhaps by low light levels)that it is difficult to judge which species was itsimmediate progenitor. This uncertainty providesinsufficient reason for treating it as a valid species.Bryum gemmiparum seems closest in view of the leafshape, occurrence of bulbils and the habitat.
238 M. O. HILL ET AL.276. Crundwell (1970) recognized the widespread synoicousforms of Bryum algovicum as var. rutheanum andthe less common autoicous ones as var. algovicum (ofwhich he treated var. compactum as a synonym).Unpublished studies by Holyoak suggest that autoicousplants appear to occur mainly if not entirely onhigh ground in the Alps, whereas synoicous plantsoccur throughout the lowlands, from central Europenorth to Svalbard. The autoicous plants are oftensmall and densely tufted, they sometimes have narrowcapsules, and the endostome processes have narrowperforations. They might merit treatment as a separatespecies rather than merely as a variety.277. Bryum apiculatum has a Pantropical range extendinginto temperate areas, with a single confirmed record inour region, from Tenerife (Ochi, 1972). The synonymyof B. apiculatum follows Ochi (1994).278. Taxonomic treatment of several Bryum species followsHolyoak (2004).279. Bryum veronense was treated as a subspecies of B.argenteum by Podpěra (1954) and Urmi (1987), as ‘aweak species’ by Düll (1992) who noted that verificationis needed for most specimens, and as a distinctspecies allied to B. argenteum by Nyholm (1993).Some of the material referred to B. veronense is amodified form of B. argenteum, including plants witha proportion of achlorophyllose leaf tips thatundoubtedly intergrade with that species, and otherwisetypical B. veronense may have the papilloserhizoids of B. argenteum. However, other materialplaced as B. veronense in herbaria differs in variousrespects and may comprise similarly reduced, ratherfeatureless, modified forms of other Bryum species,e.g. diminutive B. pallens.280. Longton (1981) studied intraspecific variation inmorphology and physiology of Bryum argenteum ona global scale by means of cultures grown on agarunder standardized laboratory conditions. He establishedthat var. lanatum is of doubtful taxonomic valuebecause the excurrent nerve was not maintained incultivation by clones originating in Canada andHawaii. Ochi (1994) also noted ‘gradations with thetypical variety’. Nevertheless, lanatum was treated as adistinct species by Frahm (2002) and Spence &Ramsay (2002), on the basis of its longer acuminateleaf point, excurrent nerve and differing habitatpreferences, but without any mention of Longton’sdetailed study. Unless better evidence for the geneticdistinctness of lanatum can be found it should bedismissed as an inconstant form of B. argenteum.281. Bryum oblongum and B. blindii are treated as separatespecies based on the detailed study by Shaw (1981).282. Bryum bornholmense was redescribed by Crundwell &Whitehouse (2001).283. Corley et al. (1981) suggested that Bryum caespiticiumis probably an aggregate species, within which B.comense and B. kunzei may be good segregates. Thelatter is treated as a valid species here, but the case formaintaining B. comense appears weak. Although Düll(1985) listed B. comense as a species, he later (Düll,1992) expressed doubt. Nyholm (1993) treated it as asynonym of B. caespiticium, as we do here.284. Demaret & Wilczek (1982) examined the types of B.canariense and B. provinciale and concluded that theyrepresented different species, but Corley et al. (1991)did not follow them because ‘their work was based toorigidly on the study of types alone’. Studies byHolyoak support the conclusions of Ochi (1972),who noted that ‘The form called B. provinciale issynoicous, but there are also dioicous plants whichhave no other clear-cut differences from such synoicousones. These two forms should be better includedin a single taxon’.285. Bryum caucasicum is known in Europe only from theholotype from the Caucasus (S). It was transferredfrom Mielichhoferia to Bryum by Cox & Hedderson(2003).286. Bryum demaretianum was described from Belgium byArts (1992).287. Treatment of the Bryum bicolor complex followsHolyoak (2003), who described B. dyffrynense as anew species from Britain.288. Bryum kunzei, Pohlia acuminata and P. polymorphawere described by Hornschuch (1819), not by Hoppe& Hornschuch as commonly stated.289. Bryum miniatum is a predominantly North Americanspecies known in Europe only from the FaeroeIslands.290. Bryum minii was mainly overlooked until the paper bySérgio et al. (1999).291. Bryum muehlenbeckii is probably best regarded as aform of B. alpinum that grows where intermittentlysubmerged in cold water, since a substantial proportionof European specimens intergrade with thatspecies (Holyoak, unpublished). Some specimens havethe characteristic leaves of both B. muehlenbeckii andB. alpinum on different parts of the same stem,apparently as a result of growth under wet or dryconditions, respectively.292. Bryum neodamense is almost certainly a recurrentform of B. pseudotriquetrum, since intermediate plantsoccur intermixed with most populations of neodamense,often in large numbers (Holyoak, in preparation).Occurrence of the neodamense morphotypeappears to be closely associated with intermittentflooding by hard water, most often on lake shores, butalso in basic fens and dune-slacks. Detailed morphologicalanalyses and molecular studies have beencarried out (Holyoak & Hedenäs, in preparation).293. Bryum aeneum was later emended to B. oeneum; seeWijk et al. (1959).294. Zolotov (2000) treats the polyoicous (partly synoicous)B. lonchocaulon as a species distinct from theautoicous B. pallescens. However, he examined few
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