orthotrichum casasianum, a new epiphytic moss from humid forests of northern spain

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Published by Maney Publishing (c) British Bryological Society Journal of Bryology (1999) 21: 47-53 Orthotrichum casasianum, a new epiphytic moss from humid forests of Northern Spain v. MAZIMPAKA\ F. LARA l , R. GARILLETI 2 , M. INFANTE 3 , AND P. HERAs 3 1 Universidad Autonoma de Madrid, 2Universidad Europea de Madrid, and 3 Museo de Ciencias Naturales de A!ava, Spain SUMMARY A new epiphytic moss, Orthotrichum casasianum F. Lara, Garilleti & Mazimpaka, is described, which is characterized by leaves slightly flexuose when dry, linear lanceolate or linear-oblong, with margins recurved and apices obtuse to rounded, frequently concave and mucronate; upper cells with low papillae or almost smooth; capsules emergent, uniformly brown, cylindric-urceolate and strongly ribbed when dry except at base, stomata hemicryptoporous; exostome of 16 teeth fused into 8 pairs, reticulate on the external side and almost smooth on the internal one; endostome of 16 linear, smooth and markedly incurved segments; calyptra naked. The new species is only known from two well preserved riparian woods in Northern Spain, beyond the Mediterranean area. The new moss is taxonomically close to the complex group of holarctic taxa related with O. scanicum Gronv., but is also similar to O. pal/ens Brid. Characters discriminating the new moss from these species are discussed. KEYWORDS: Orthotrichum, Orthotrichaceae, Bryopsida, taxonomy, Spain, Iberian Peninsula INTRODUCTION When revising Orthotrichum specimens from the herbar- ium of the Alava Museum of Natural Sciences (VIT), we found fruiting specimens of a small elegant moss that had peculiar leaf apices, and peristomial characters concordant with those of subgenus Pulchel/a (Schimp.) Vitt. A comparison with the closest European species (0. pal/ens, O. scanicum, and O. lewinskyae F. Lara, Garilleti & Mazimpaka) suggests· that these specimens share a unique combination of characters warranting recognition at the species level. We wish to name the new moss in honour of Creu Casas, as a tribute to her great contribution to Iberian bryology. Besides her extensive work, she generously helped and encouraged younger generations of Spanish bryologists. DESCRIPTION Orthotrichum casasianum F. Lara, Garilleti & Mazimpaka, sp. nov. (Figs 1-9, 11-12, 15-16) © British Bryological Society 1999 Planta fere 10 mm alta, filidiis siccis parum flexuosis, linear-Ianceolatis vellinear-oblongis, marginibus recurvatis apicibusque obtusis ad rotundata, concavis mucronatis- que, in sicco incurvatis. Cellulae superiores laeves vel humilissime papillosae, parietibus incrassatis. Autoica. Capsula emergens, in sicco cylindrica-urceolata et valde suIcata, basi ventricosa concoloraque, stomatis hemicryp- toporis. Exostoma 16 dentibus geminatis, in externa pagina reticulatis vix papillosisque, interne quasi laevibus. Endostoma 16 segmentis tenuibus, insigniter incurvatis, laevibus et transversis parietibus incrassatis. Calyptra nuda,laevis. Holotype: Spain, Alava, Luquiano, Bayas river, 590 m, UTM grid 30TWN1153, corticolous on Sambucus nigra L. P. Heras, 20 October 1985, VIT 1527/85. Plants olive green, 0.6-1.0 mm tall, in small irregular cushions or patches up to 20 mIll. Stems variously branched, slender, loosely foliose, sometimes with the branch apices curved downwards, rhizoids gathered ... in the lower third-half. Leaves erect and more or less flexuose when dry, the Received 20 April 1998. Revision accepted 20 July 1998

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Journal of Bryology (1999) 21 47-53

Orthotrichum casasianum a new epiphytic moss from humid forestsof Northern Spain

v MAZIMPAKA F LARAl R GARILLETI2 M INFANTE3 AND P HERAs3

1Universidad Autonoma de Madrid2Universidad Europea de Madrid and

3Museo de Ciencias Naturales de Aava Spain

SUMMARY

A new epiphytic moss Orthotrichum casasianum F Lara Garilleti amp Mazimpaka is described which ischaracterized by leaves slightly flexuose when dry linear lanceolate or linear-oblong with margins recurvedand apices obtuse to rounded frequently concave and mucronate upper cells with low papillae or almostsmooth capsules emergent uniformly brown cylindric-urceolate and strongly ribbed when dry except atbase stomata hemicryptoporous exostome of 16 teeth fused into 8 pairs reticulate on the external side andalmost smooth on the internal one endostome of 16 linear smooth and markedly incurved segmentscalyptra naked The new species is only known from two well preserved riparian woods in Northern Spainbeyond the Mediterranean area The new moss is taxonomically close to the complex group of holarctictaxa related with O scanicum Gronv but is also similar to O palens Brid Characters discriminating thenew moss from these species are discussed

KEYWORDS Orthotrichum Orthotrichaceae Bryopsida taxonomy Spain Iberian Peninsula

INTRODUCTION

When revising Orthotrichum specimens from the herbar-ium of the Alava Museum of Natural Sciences (VIT) wefound fruiting specimens of a small elegant moss that hadpeculiar leaf apices and peristomial characters concordantwith those of subgenus Pulchela (Schimp) Vitt Acomparison with the closest European species (0 palensO scanicum and O lewinskyae F Lara Garilleti ampMazimpaka) suggestsmiddot that these specimens share a uniquecombination of characters warranting recognition at thespecies level

We wish to name the new moss in honour of CreuCasas as a tribute to her great contribution to Iberianbryology Besides her extensive work she generouslyhelped and encouraged younger generations of Spanishbryologists

DESCRIPTION

Orthotrichum casasianum F Lara Garilleti amp Mazimpakasp nov (Figs 1-9 11-12 15-16)

copy British Bryological Society 1999

Planta fere 10 mm alta filidiis siccis parum flexuosislinear-Ianceolatis vellinear-oblongis marginibus recurvatisapicibusque obtusis ad rotundata concavis mucronatis-que in sicco incurvatis Cellulae superiores laeves velhumilissime papillosae parietibus incrassatis AutoicaCapsula emergens in sicco cylindrica-urceolata et valdesuIcata basi ventricosa concoloraque stomatis hemicryp-toporis Exostoma 16 dentibus geminatis in externapagina reticulatis vix papillosisque interne quasi laevibusEndostoma 16 segmentis tenuibus insigniter incurvatislaevibus et transversis parietibus incrassatis Calyptranudalaevis

Holotype Spain Alava Luquiano Bayas river 590 mUTM grid 30TWN1153 corticolous on Sambucus nigra LP Heras 20 October 1985 VIT 152785

Plants olive green 06-10 mm tall in small irregularcushions or patches up to 20 mIll Stems variouslybranched slender loosely foliose sometimes with thebranch apices curved downwards rhizoids gathered inthe lower third-half

Leaves erect and more or less flexuose when dry the

Received 20 April 1998 Revision accepted 20 July 1998

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48 V MAZIMPAKA ET AL

Figures 1-7 Orthotrichum casasianum F Lara Garilleti amp Mazimpaka 1 Leaf apices 2 Areolation of basal (left) and upper leaf cells (right)3 Cross section of leaf 4 Leaf form 5 Habit of plant 6 Calyptra 7 Moist capsule (All from type)

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ORTHOTRCHUM CASASANUN SP NOV FROM SPAIN 49

Figures 8-9 Orthotrichum casasianum F Lara Garilleti amp Mazimpaka 8 SEM micrographs of leaf apices 9 Upper capsule portion andperistome (All from type)

upper ones imbricate and erect patent to erecto-patentwhen moist In shape linear-Ianceolate or linear-oblongwith wide ovate base and linear or lingulate laminakeeled and concave (15-)20-27(-28) x 04-06 mmwith the margins finely recurved from base to 23 or 45of leaf length Leaf apices variable in general obtuseand with a small 1(-2)-celled mucro plane or slightlycarinate with the costa ending near the apex in someleaves more or less concave keeled and with the costapercurrent in others often incurved when dry Upper andexternal perichaetial leaves more or less imbricate notflexuose 23-26 x 05-07 mm linear or lingulate withwide bases with the margins recurved in the upper 23apices rounded or obtuse concave and generally cucul-late without mucro but with the costa percurrent andsometimes with a channelled apiculus having inflexedmargins Internal perichaetial leaves smaller more or lessoblong obtuse and with margins plane Apical leaf cellsirregularly rounded or oblong (8-)11-13(-15) x (9-)12-14(-16) flm variously oriented slightly thick-walledsmooth or with 1-2 low papillae Median and upper leafcells rounded or shortly ovate (9-)12-15 9 x 15(-17) flm

regularly disposed and more thick-walled with 1-3(-4)small blunt but generally visible papillae on both leaffaces Basal cells 25-35(-50) x 8-14 flm rectangularhyaline with thin and straight walls becoming quadrateto 13 flm wide towards margin

Autoicous with sessile perigonial gemmules onbranches that form apical perichaetia and additionallateral branches that are exclusively male Seta short04~06 mm with the lower half surrounded by anochrea Capsule emergent 175-200 mm long uniformly

dark brown with stomata hemicryptoporous in theupper half cylindric-urceolate when dry homogeneouslyconstricted (sometimes the mouth less so) by 8 thin andprominent ribs that do not reach the capsule basewhich is ventricose when moist capsules long oblong toellipsoid Neck concolorous with the theca in somecases sharply narrowed to the seta in others graduallyso Exothecial bands 4-6(-7) cell wide formed bymoderately differentiated and little thick-walled cellsextending from the mouth to 34 or 45 of capsulelength Peristome double Basal remains of prostomeoccasionally present Exostome of 8 pairs of teeth 200-225 flm long light or dark brown sometimes splittingirregularly in old capsules Ornamentation of the exter-nal side (OPL) apparently homogeneous and denselypapillose in the lower half consisting of a wall reticu-lum that forms scattered low papillae in the upper onecomposed of blunt low and dense (though becomingsparser towards the apex) papillae anticlinal wallremains not incrassate On the internal side (PPL) thelower half smooth or nearly so at most with light longi-tudinal striae while the upper half is sparsely and irre-gularly ornamented with striae and papillae intermixedin variable proportions incrassate remains of anticlinallongitudinal and transverse walls present at the toothapex Endostome of 16 segments hyaline thin andstrongly incurved shorter than teeth smooth on bothsides or with thin longitudinal striae on the PPL on theinternal side (IPL) remains of transverse walls incrassateand frequently brown-orange tinged Main segments180-200 flm long internally uniseriate or irregularlybiseriate always with a wide and asymetrically biseriate

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Figure 10 Known distribution of Orthotrichum casasianum in Spain

base Intermediate segments smaller entirely uniseriatemore fragile than the main ones and frequently missingin old capsules Basal membrane smooth continuousand narrow 2(-3) cells high and shortly protrudingbeyond the capsule mouth Spores (12-)14-20 pmdensely papillose Brood bodies not seen Calyptraconic-oblong brown darker on the apex 11-13 mark-edly plicate naked and smooth Vaginula naked or withshort smooth and brown hairs

SPECIMENS EXAMINED (PARA TYPI)

SPAIN Alava Luquiano riberas del Bayas 590 m30TWN1153 corticicola en Sambucus nigra L B Albertoset al 25-1-1997 (VIT BCB NY MA and authors herb)Ibidem sobre rama de Corylus avellana L eiusdem 25-1-1997 (authors herb) Ibidem sobre Sambucus nigra L BAlbertos amp R Garilleti 12-IV-1997 (VIT) Ibidem sobreAlnus glutinosa Gaertn P Heras amp M Infante VIII-1997(authors herb) Alava Sendadiano rio Bayas 540 m30TWN0848 tronco de Fraxinus excelsior L B Albertoset a 27-1-1997 (VIT)

ECOLOGY AND DISTRIBUTION

Orthotrichum casasianum has been found in only twolocalities from Alava (North Spain) on the banks of

Bayas river It is an epiphyte growing on Alnus glutinosaGaertn Sambucus nigra L Fraxinus excelsior L andCorylus avellana L in well preserved humid riparianwoods (Alnus and Fraxinus groves) subject to a eurosiber-ian climate The range of the new moss is possibly largerthan is currently known but confirmation of this dependsupon investigations of similar environments in the Cantab-rian Cordillera and Pyrenees

The new moss occurs in epiphytic communities domi-nated by different Orthotrichum species or pleurocarpousmosses either on tree bases subject to periodical inunda-tions or on shrub trunks or branches On tree bases itgrows together with O rivulare Turn O sprucei MontHypnum cupressiforme Hedw and Leskea polycarpa Hedwon strata not subject to inundation it lives among otherswith O pulchellum Brunt O consimile Mitt O affineBrid Hypnum cupressijorme Cryphaea heteromalla(Hedw) Mohr Metzgeria jurcata (L) Dum M temperataKuwak Frullania dilatata (L) Dum and Radula compla-nata (L) Dum This list of cohabitants emphazises theecological and chorological interest of these woods whichbesides giving shelter to the new taxon have been reportedto enclose other taxa rare in the Iberian Peninsula (Garilletiet al 1998Heras amp Infante 1998)

DISCUSSION

The new taxon is closely related to Orthotrichum scani-cum O holmenii Lewinsky- Haapasaari and O lew ins-kyae a complex of hoi arctic taxa belonging to subgPulchella (Schimp) Vitt section Pulchella Schimp Themost important characters that support this relationshipand adscription are (1) the capsule position (emergent)and its cylindric form with ventricose base (2) theendostome structure and ornamentation consisting of16 filiform segments ornamented by thin longitudinallines on the PPL (3) the connecting membrane protrud-ing beyond the capsule mouth despite its narrowness(4) the marked trend of teeth to splitting (5) the occa-sional presence of prostome remains (6) the slenderhabit with somewhat flexuose leaves and (7) therecurved leaf margins O casasianum is re~dily sepa-rated from these taxa by gametophytic and peristomialcharacters the obtuse or rounded not dentate andfrequently cucullate leaf apices coupled to the hyalineendostome without papillae are the characters affordingits quick recognition

However the new species could be easily confusedwith O pallens a variable taxon adscribed to sectionDiaphana Vent (Lewinsky 1993) Both species share anumber of characters commonly used in identificationkeys emergent and strongly ribbed capsules hemicryp-toporous stomata exostome of 8 pairs of apparentlypapillose teeth conic plicate and naked calyptrae Inaddition variability of O pallens involves a number ofcharacters that may be similar to those of O casasia-

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ORTHOTRICHUM CASASIANUM SP NOV FROM SPAIN 51

Figures 11-14 Comparison of Orthotrichum casasianum F Lara Garilleti amp Mazimpaka and O pallens Brid peristomes (OPL) 11 Tooth ofO casasianum 12 Detail of the median zone of tooth of O casasianum 13 Tooth of O pallens 14 Detail of the inferior half of O pallenstooth [11 and 12 from type 13 and 14 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

num For example O pallens is a taxon typicallycladautoicous although goniautoicous populations havealso been described (Nyholm 1956) endostomesegments may appear in number of 8 8 + 8 or 16 andare smooth or scarcely papillose leaf apices vary fromacute to obtuse apiculate or not and sometimessubmucronate (Bridel 1827) usually leaf cells havehigh and sometimes branched papillae but in somepopulations papillae are low and scarcely visible Thevariability of O pallens is so high that it is sometimesdifficult to separate it from close species like O pumilumSm (Lara Garilleti amp Mazimpaka 1994) moreoverseveral taxa described as independent species or vari-eties of O pallens have been reduced to synonyms ofthe latter (Vitt 1973 Lewinsky 1977 1993) based onthe fact that their differential characters were comprisedin the variability range of this species

O casasianum differs from O pallens by its slenderloosely foliose habit with narrow leaves whose apicesthough variable within the same individual are charac-teristically obtuse and concave often incurvate or cucul-late when dry and with a 1(-2)-celled hyaline apiculusor the costa shortly percurrent Also the capsule form isdifferent although cylindric the capsule is notconstricted at base but is instead ventricose and

narrows more or less sharply in the neck in additionthe mature capsule and the seta are uniformly darkbrown while in O palens the capsule base and the setaare pale brown (a consistent feature on which is basedthe name of the taxon) so that they contrast with thecolour of the theca The most important character ishowever the different ornamentation of the external sideof the exostome teeth (Figs 11-14) in O casasianumalthough the aspect on optical microscope is papillose itis in fact formed by a network of high walls on whichsparse papillae are irregularly disposed in O pallens bycontrast the dense papillosity observed on opticalmicroscope is real and the the basal pattern is markedlydifferent from that of the tooth apex where papilloselines are usually abundant Other peristomial differencesare the occasional presence of prostome in O casasia-num the minor papillosi ty of the exostome PPL and themajor bending of the endostome segments Table Isummarizes the differences between the species

ACKNOWLEDGEMENTS

Thanks are due to the staff of S especially to Lars Hede-nas whose loan of Orthotrichum pallens material made

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52 V MAZIMPAKA ET AL

Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

--------------~~-----__--- ------------------~--------- --- - --------~--_------~---- - -- --~~~

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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48 V MAZIMPAKA ET AL

Figures 1-7 Orthotrichum casasianum F Lara Garilleti amp Mazimpaka 1 Leaf apices 2 Areolation of basal (left) and upper leaf cells (right)3 Cross section of leaf 4 Leaf form 5 Habit of plant 6 Calyptra 7 Moist capsule (All from type)

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ORTHOTRCHUM CASASANUN SP NOV FROM SPAIN 49

Figures 8-9 Orthotrichum casasianum F Lara Garilleti amp Mazimpaka 8 SEM micrographs of leaf apices 9 Upper capsule portion andperistome (All from type)

upper ones imbricate and erect patent to erecto-patentwhen moist In shape linear-Ianceolate or linear-oblongwith wide ovate base and linear or lingulate laminakeeled and concave (15-)20-27(-28) x 04-06 mmwith the margins finely recurved from base to 23 or 45of leaf length Leaf apices variable in general obtuseand with a small 1(-2)-celled mucro plane or slightlycarinate with the costa ending near the apex in someleaves more or less concave keeled and with the costapercurrent in others often incurved when dry Upper andexternal perichaetial leaves more or less imbricate notflexuose 23-26 x 05-07 mm linear or lingulate withwide bases with the margins recurved in the upper 23apices rounded or obtuse concave and generally cucul-late without mucro but with the costa percurrent andsometimes with a channelled apiculus having inflexedmargins Internal perichaetial leaves smaller more or lessoblong obtuse and with margins plane Apical leaf cellsirregularly rounded or oblong (8-)11-13(-15) x (9-)12-14(-16) flm variously oriented slightly thick-walledsmooth or with 1-2 low papillae Median and upper leafcells rounded or shortly ovate (9-)12-15 9 x 15(-17) flm

regularly disposed and more thick-walled with 1-3(-4)small blunt but generally visible papillae on both leaffaces Basal cells 25-35(-50) x 8-14 flm rectangularhyaline with thin and straight walls becoming quadrateto 13 flm wide towards margin

Autoicous with sessile perigonial gemmules onbranches that form apical perichaetia and additionallateral branches that are exclusively male Seta short04~06 mm with the lower half surrounded by anochrea Capsule emergent 175-200 mm long uniformly

dark brown with stomata hemicryptoporous in theupper half cylindric-urceolate when dry homogeneouslyconstricted (sometimes the mouth less so) by 8 thin andprominent ribs that do not reach the capsule basewhich is ventricose when moist capsules long oblong toellipsoid Neck concolorous with the theca in somecases sharply narrowed to the seta in others graduallyso Exothecial bands 4-6(-7) cell wide formed bymoderately differentiated and little thick-walled cellsextending from the mouth to 34 or 45 of capsulelength Peristome double Basal remains of prostomeoccasionally present Exostome of 8 pairs of teeth 200-225 flm long light or dark brown sometimes splittingirregularly in old capsules Ornamentation of the exter-nal side (OPL) apparently homogeneous and denselypapillose in the lower half consisting of a wall reticu-lum that forms scattered low papillae in the upper onecomposed of blunt low and dense (though becomingsparser towards the apex) papillae anticlinal wallremains not incrassate On the internal side (PPL) thelower half smooth or nearly so at most with light longi-tudinal striae while the upper half is sparsely and irre-gularly ornamented with striae and papillae intermixedin variable proportions incrassate remains of anticlinallongitudinal and transverse walls present at the toothapex Endostome of 16 segments hyaline thin andstrongly incurved shorter than teeth smooth on bothsides or with thin longitudinal striae on the PPL on theinternal side (IPL) remains of transverse walls incrassateand frequently brown-orange tinged Main segments180-200 flm long internally uniseriate or irregularlybiseriate always with a wide and asymetrically biseriate

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Figure 10 Known distribution of Orthotrichum casasianum in Spain

base Intermediate segments smaller entirely uniseriatemore fragile than the main ones and frequently missingin old capsules Basal membrane smooth continuousand narrow 2(-3) cells high and shortly protrudingbeyond the capsule mouth Spores (12-)14-20 pmdensely papillose Brood bodies not seen Calyptraconic-oblong brown darker on the apex 11-13 mark-edly plicate naked and smooth Vaginula naked or withshort smooth and brown hairs

SPECIMENS EXAMINED (PARA TYPI)

SPAIN Alava Luquiano riberas del Bayas 590 m30TWN1153 corticicola en Sambucus nigra L B Albertoset al 25-1-1997 (VIT BCB NY MA and authors herb)Ibidem sobre rama de Corylus avellana L eiusdem 25-1-1997 (authors herb) Ibidem sobre Sambucus nigra L BAlbertos amp R Garilleti 12-IV-1997 (VIT) Ibidem sobreAlnus glutinosa Gaertn P Heras amp M Infante VIII-1997(authors herb) Alava Sendadiano rio Bayas 540 m30TWN0848 tronco de Fraxinus excelsior L B Albertoset a 27-1-1997 (VIT)

ECOLOGY AND DISTRIBUTION

Orthotrichum casasianum has been found in only twolocalities from Alava (North Spain) on the banks of

Bayas river It is an epiphyte growing on Alnus glutinosaGaertn Sambucus nigra L Fraxinus excelsior L andCorylus avellana L in well preserved humid riparianwoods (Alnus and Fraxinus groves) subject to a eurosiber-ian climate The range of the new moss is possibly largerthan is currently known but confirmation of this dependsupon investigations of similar environments in the Cantab-rian Cordillera and Pyrenees

The new moss occurs in epiphytic communities domi-nated by different Orthotrichum species or pleurocarpousmosses either on tree bases subject to periodical inunda-tions or on shrub trunks or branches On tree bases itgrows together with O rivulare Turn O sprucei MontHypnum cupressiforme Hedw and Leskea polycarpa Hedwon strata not subject to inundation it lives among otherswith O pulchellum Brunt O consimile Mitt O affineBrid Hypnum cupressijorme Cryphaea heteromalla(Hedw) Mohr Metzgeria jurcata (L) Dum M temperataKuwak Frullania dilatata (L) Dum and Radula compla-nata (L) Dum This list of cohabitants emphazises theecological and chorological interest of these woods whichbesides giving shelter to the new taxon have been reportedto enclose other taxa rare in the Iberian Peninsula (Garilletiet al 1998Heras amp Infante 1998)

DISCUSSION

The new taxon is closely related to Orthotrichum scani-cum O holmenii Lewinsky- Haapasaari and O lew ins-kyae a complex of hoi arctic taxa belonging to subgPulchella (Schimp) Vitt section Pulchella Schimp Themost important characters that support this relationshipand adscription are (1) the capsule position (emergent)and its cylindric form with ventricose base (2) theendostome structure and ornamentation consisting of16 filiform segments ornamented by thin longitudinallines on the PPL (3) the connecting membrane protrud-ing beyond the capsule mouth despite its narrowness(4) the marked trend of teeth to splitting (5) the occa-sional presence of prostome remains (6) the slenderhabit with somewhat flexuose leaves and (7) therecurved leaf margins O casasianum is re~dily sepa-rated from these taxa by gametophytic and peristomialcharacters the obtuse or rounded not dentate andfrequently cucullate leaf apices coupled to the hyalineendostome without papillae are the characters affordingits quick recognition

However the new species could be easily confusedwith O pallens a variable taxon adscribed to sectionDiaphana Vent (Lewinsky 1993) Both species share anumber of characters commonly used in identificationkeys emergent and strongly ribbed capsules hemicryp-toporous stomata exostome of 8 pairs of apparentlypapillose teeth conic plicate and naked calyptrae Inaddition variability of O pallens involves a number ofcharacters that may be similar to those of O casasia-

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ORTHOTRICHUM CASASIANUM SP NOV FROM SPAIN 51

Figures 11-14 Comparison of Orthotrichum casasianum F Lara Garilleti amp Mazimpaka and O pallens Brid peristomes (OPL) 11 Tooth ofO casasianum 12 Detail of the median zone of tooth of O casasianum 13 Tooth of O pallens 14 Detail of the inferior half of O pallenstooth [11 and 12 from type 13 and 14 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

num For example O pallens is a taxon typicallycladautoicous although goniautoicous populations havealso been described (Nyholm 1956) endostomesegments may appear in number of 8 8 + 8 or 16 andare smooth or scarcely papillose leaf apices vary fromacute to obtuse apiculate or not and sometimessubmucronate (Bridel 1827) usually leaf cells havehigh and sometimes branched papillae but in somepopulations papillae are low and scarcely visible Thevariability of O pallens is so high that it is sometimesdifficult to separate it from close species like O pumilumSm (Lara Garilleti amp Mazimpaka 1994) moreoverseveral taxa described as independent species or vari-eties of O pallens have been reduced to synonyms ofthe latter (Vitt 1973 Lewinsky 1977 1993) based onthe fact that their differential characters were comprisedin the variability range of this species

O casasianum differs from O pallens by its slenderloosely foliose habit with narrow leaves whose apicesthough variable within the same individual are charac-teristically obtuse and concave often incurvate or cucul-late when dry and with a 1(-2)-celled hyaline apiculusor the costa shortly percurrent Also the capsule form isdifferent although cylindric the capsule is notconstricted at base but is instead ventricose and

narrows more or less sharply in the neck in additionthe mature capsule and the seta are uniformly darkbrown while in O palens the capsule base and the setaare pale brown (a consistent feature on which is basedthe name of the taxon) so that they contrast with thecolour of the theca The most important character ishowever the different ornamentation of the external sideof the exostome teeth (Figs 11-14) in O casasianumalthough the aspect on optical microscope is papillose itis in fact formed by a network of high walls on whichsparse papillae are irregularly disposed in O pallens bycontrast the dense papillosity observed on opticalmicroscope is real and the the basal pattern is markedlydifferent from that of the tooth apex where papilloselines are usually abundant Other peristomial differencesare the occasional presence of prostome in O casasia-num the minor papillosi ty of the exostome PPL and themajor bending of the endostome segments Table Isummarizes the differences between the species

ACKNOWLEDGEMENTS

Thanks are due to the staff of S especially to Lars Hede-nas whose loan of Orthotrichum pallens material made

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Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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ORTHOTRCHUM CASASANUN SP NOV FROM SPAIN 49

Figures 8-9 Orthotrichum casasianum F Lara Garilleti amp Mazimpaka 8 SEM micrographs of leaf apices 9 Upper capsule portion andperistome (All from type)

upper ones imbricate and erect patent to erecto-patentwhen moist In shape linear-Ianceolate or linear-oblongwith wide ovate base and linear or lingulate laminakeeled and concave (15-)20-27(-28) x 04-06 mmwith the margins finely recurved from base to 23 or 45of leaf length Leaf apices variable in general obtuseand with a small 1(-2)-celled mucro plane or slightlycarinate with the costa ending near the apex in someleaves more or less concave keeled and with the costapercurrent in others often incurved when dry Upper andexternal perichaetial leaves more or less imbricate notflexuose 23-26 x 05-07 mm linear or lingulate withwide bases with the margins recurved in the upper 23apices rounded or obtuse concave and generally cucul-late without mucro but with the costa percurrent andsometimes with a channelled apiculus having inflexedmargins Internal perichaetial leaves smaller more or lessoblong obtuse and with margins plane Apical leaf cellsirregularly rounded or oblong (8-)11-13(-15) x (9-)12-14(-16) flm variously oriented slightly thick-walledsmooth or with 1-2 low papillae Median and upper leafcells rounded or shortly ovate (9-)12-15 9 x 15(-17) flm

regularly disposed and more thick-walled with 1-3(-4)small blunt but generally visible papillae on both leaffaces Basal cells 25-35(-50) x 8-14 flm rectangularhyaline with thin and straight walls becoming quadrateto 13 flm wide towards margin

Autoicous with sessile perigonial gemmules onbranches that form apical perichaetia and additionallateral branches that are exclusively male Seta short04~06 mm with the lower half surrounded by anochrea Capsule emergent 175-200 mm long uniformly

dark brown with stomata hemicryptoporous in theupper half cylindric-urceolate when dry homogeneouslyconstricted (sometimes the mouth less so) by 8 thin andprominent ribs that do not reach the capsule basewhich is ventricose when moist capsules long oblong toellipsoid Neck concolorous with the theca in somecases sharply narrowed to the seta in others graduallyso Exothecial bands 4-6(-7) cell wide formed bymoderately differentiated and little thick-walled cellsextending from the mouth to 34 or 45 of capsulelength Peristome double Basal remains of prostomeoccasionally present Exostome of 8 pairs of teeth 200-225 flm long light or dark brown sometimes splittingirregularly in old capsules Ornamentation of the exter-nal side (OPL) apparently homogeneous and denselypapillose in the lower half consisting of a wall reticu-lum that forms scattered low papillae in the upper onecomposed of blunt low and dense (though becomingsparser towards the apex) papillae anticlinal wallremains not incrassate On the internal side (PPL) thelower half smooth or nearly so at most with light longi-tudinal striae while the upper half is sparsely and irre-gularly ornamented with striae and papillae intermixedin variable proportions incrassate remains of anticlinallongitudinal and transverse walls present at the toothapex Endostome of 16 segments hyaline thin andstrongly incurved shorter than teeth smooth on bothsides or with thin longitudinal striae on the PPL on theinternal side (IPL) remains of transverse walls incrassateand frequently brown-orange tinged Main segments180-200 flm long internally uniseriate or irregularlybiseriate always with a wide and asymetrically biseriate

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Figure 10 Known distribution of Orthotrichum casasianum in Spain

base Intermediate segments smaller entirely uniseriatemore fragile than the main ones and frequently missingin old capsules Basal membrane smooth continuousand narrow 2(-3) cells high and shortly protrudingbeyond the capsule mouth Spores (12-)14-20 pmdensely papillose Brood bodies not seen Calyptraconic-oblong brown darker on the apex 11-13 mark-edly plicate naked and smooth Vaginula naked or withshort smooth and brown hairs

SPECIMENS EXAMINED (PARA TYPI)

SPAIN Alava Luquiano riberas del Bayas 590 m30TWN1153 corticicola en Sambucus nigra L B Albertoset al 25-1-1997 (VIT BCB NY MA and authors herb)Ibidem sobre rama de Corylus avellana L eiusdem 25-1-1997 (authors herb) Ibidem sobre Sambucus nigra L BAlbertos amp R Garilleti 12-IV-1997 (VIT) Ibidem sobreAlnus glutinosa Gaertn P Heras amp M Infante VIII-1997(authors herb) Alava Sendadiano rio Bayas 540 m30TWN0848 tronco de Fraxinus excelsior L B Albertoset a 27-1-1997 (VIT)

ECOLOGY AND DISTRIBUTION

Orthotrichum casasianum has been found in only twolocalities from Alava (North Spain) on the banks of

Bayas river It is an epiphyte growing on Alnus glutinosaGaertn Sambucus nigra L Fraxinus excelsior L andCorylus avellana L in well preserved humid riparianwoods (Alnus and Fraxinus groves) subject to a eurosiber-ian climate The range of the new moss is possibly largerthan is currently known but confirmation of this dependsupon investigations of similar environments in the Cantab-rian Cordillera and Pyrenees

The new moss occurs in epiphytic communities domi-nated by different Orthotrichum species or pleurocarpousmosses either on tree bases subject to periodical inunda-tions or on shrub trunks or branches On tree bases itgrows together with O rivulare Turn O sprucei MontHypnum cupressiforme Hedw and Leskea polycarpa Hedwon strata not subject to inundation it lives among otherswith O pulchellum Brunt O consimile Mitt O affineBrid Hypnum cupressijorme Cryphaea heteromalla(Hedw) Mohr Metzgeria jurcata (L) Dum M temperataKuwak Frullania dilatata (L) Dum and Radula compla-nata (L) Dum This list of cohabitants emphazises theecological and chorological interest of these woods whichbesides giving shelter to the new taxon have been reportedto enclose other taxa rare in the Iberian Peninsula (Garilletiet al 1998Heras amp Infante 1998)

DISCUSSION

The new taxon is closely related to Orthotrichum scani-cum O holmenii Lewinsky- Haapasaari and O lew ins-kyae a complex of hoi arctic taxa belonging to subgPulchella (Schimp) Vitt section Pulchella Schimp Themost important characters that support this relationshipand adscription are (1) the capsule position (emergent)and its cylindric form with ventricose base (2) theendostome structure and ornamentation consisting of16 filiform segments ornamented by thin longitudinallines on the PPL (3) the connecting membrane protrud-ing beyond the capsule mouth despite its narrowness(4) the marked trend of teeth to splitting (5) the occa-sional presence of prostome remains (6) the slenderhabit with somewhat flexuose leaves and (7) therecurved leaf margins O casasianum is re~dily sepa-rated from these taxa by gametophytic and peristomialcharacters the obtuse or rounded not dentate andfrequently cucullate leaf apices coupled to the hyalineendostome without papillae are the characters affordingits quick recognition

However the new species could be easily confusedwith O pallens a variable taxon adscribed to sectionDiaphana Vent (Lewinsky 1993) Both species share anumber of characters commonly used in identificationkeys emergent and strongly ribbed capsules hemicryp-toporous stomata exostome of 8 pairs of apparentlypapillose teeth conic plicate and naked calyptrae Inaddition variability of O pallens involves a number ofcharacters that may be similar to those of O casasia-

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ORTHOTRICHUM CASASIANUM SP NOV FROM SPAIN 51

Figures 11-14 Comparison of Orthotrichum casasianum F Lara Garilleti amp Mazimpaka and O pallens Brid peristomes (OPL) 11 Tooth ofO casasianum 12 Detail of the median zone of tooth of O casasianum 13 Tooth of O pallens 14 Detail of the inferior half of O pallenstooth [11 and 12 from type 13 and 14 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

num For example O pallens is a taxon typicallycladautoicous although goniautoicous populations havealso been described (Nyholm 1956) endostomesegments may appear in number of 8 8 + 8 or 16 andare smooth or scarcely papillose leaf apices vary fromacute to obtuse apiculate or not and sometimessubmucronate (Bridel 1827) usually leaf cells havehigh and sometimes branched papillae but in somepopulations papillae are low and scarcely visible Thevariability of O pallens is so high that it is sometimesdifficult to separate it from close species like O pumilumSm (Lara Garilleti amp Mazimpaka 1994) moreoverseveral taxa described as independent species or vari-eties of O pallens have been reduced to synonyms ofthe latter (Vitt 1973 Lewinsky 1977 1993) based onthe fact that their differential characters were comprisedin the variability range of this species

O casasianum differs from O pallens by its slenderloosely foliose habit with narrow leaves whose apicesthough variable within the same individual are charac-teristically obtuse and concave often incurvate or cucul-late when dry and with a 1(-2)-celled hyaline apiculusor the costa shortly percurrent Also the capsule form isdifferent although cylindric the capsule is notconstricted at base but is instead ventricose and

narrows more or less sharply in the neck in additionthe mature capsule and the seta are uniformly darkbrown while in O palens the capsule base and the setaare pale brown (a consistent feature on which is basedthe name of the taxon) so that they contrast with thecolour of the theca The most important character ishowever the different ornamentation of the external sideof the exostome teeth (Figs 11-14) in O casasianumalthough the aspect on optical microscope is papillose itis in fact formed by a network of high walls on whichsparse papillae are irregularly disposed in O pallens bycontrast the dense papillosity observed on opticalmicroscope is real and the the basal pattern is markedlydifferent from that of the tooth apex where papilloselines are usually abundant Other peristomial differencesare the occasional presence of prostome in O casasia-num the minor papillosi ty of the exostome PPL and themajor bending of the endostome segments Table Isummarizes the differences between the species

ACKNOWLEDGEMENTS

Thanks are due to the staff of S especially to Lars Hede-nas whose loan of Orthotrichum pallens material made

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52 V MAZIMPAKA ET AL

Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

--------------~~-----__--- ------------------~--------- --- - --------~--_------~---- - -- --~~~

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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Figure 10 Known distribution of Orthotrichum casasianum in Spain

base Intermediate segments smaller entirely uniseriatemore fragile than the main ones and frequently missingin old capsules Basal membrane smooth continuousand narrow 2(-3) cells high and shortly protrudingbeyond the capsule mouth Spores (12-)14-20 pmdensely papillose Brood bodies not seen Calyptraconic-oblong brown darker on the apex 11-13 mark-edly plicate naked and smooth Vaginula naked or withshort smooth and brown hairs

SPECIMENS EXAMINED (PARA TYPI)

SPAIN Alava Luquiano riberas del Bayas 590 m30TWN1153 corticicola en Sambucus nigra L B Albertoset al 25-1-1997 (VIT BCB NY MA and authors herb)Ibidem sobre rama de Corylus avellana L eiusdem 25-1-1997 (authors herb) Ibidem sobre Sambucus nigra L BAlbertos amp R Garilleti 12-IV-1997 (VIT) Ibidem sobreAlnus glutinosa Gaertn P Heras amp M Infante VIII-1997(authors herb) Alava Sendadiano rio Bayas 540 m30TWN0848 tronco de Fraxinus excelsior L B Albertoset a 27-1-1997 (VIT)

ECOLOGY AND DISTRIBUTION

Orthotrichum casasianum has been found in only twolocalities from Alava (North Spain) on the banks of

Bayas river It is an epiphyte growing on Alnus glutinosaGaertn Sambucus nigra L Fraxinus excelsior L andCorylus avellana L in well preserved humid riparianwoods (Alnus and Fraxinus groves) subject to a eurosiber-ian climate The range of the new moss is possibly largerthan is currently known but confirmation of this dependsupon investigations of similar environments in the Cantab-rian Cordillera and Pyrenees

The new moss occurs in epiphytic communities domi-nated by different Orthotrichum species or pleurocarpousmosses either on tree bases subject to periodical inunda-tions or on shrub trunks or branches On tree bases itgrows together with O rivulare Turn O sprucei MontHypnum cupressiforme Hedw and Leskea polycarpa Hedwon strata not subject to inundation it lives among otherswith O pulchellum Brunt O consimile Mitt O affineBrid Hypnum cupressijorme Cryphaea heteromalla(Hedw) Mohr Metzgeria jurcata (L) Dum M temperataKuwak Frullania dilatata (L) Dum and Radula compla-nata (L) Dum This list of cohabitants emphazises theecological and chorological interest of these woods whichbesides giving shelter to the new taxon have been reportedto enclose other taxa rare in the Iberian Peninsula (Garilletiet al 1998Heras amp Infante 1998)

DISCUSSION

The new taxon is closely related to Orthotrichum scani-cum O holmenii Lewinsky- Haapasaari and O lew ins-kyae a complex of hoi arctic taxa belonging to subgPulchella (Schimp) Vitt section Pulchella Schimp Themost important characters that support this relationshipand adscription are (1) the capsule position (emergent)and its cylindric form with ventricose base (2) theendostome structure and ornamentation consisting of16 filiform segments ornamented by thin longitudinallines on the PPL (3) the connecting membrane protrud-ing beyond the capsule mouth despite its narrowness(4) the marked trend of teeth to splitting (5) the occa-sional presence of prostome remains (6) the slenderhabit with somewhat flexuose leaves and (7) therecurved leaf margins O casasianum is re~dily sepa-rated from these taxa by gametophytic and peristomialcharacters the obtuse or rounded not dentate andfrequently cucullate leaf apices coupled to the hyalineendostome without papillae are the characters affordingits quick recognition

However the new species could be easily confusedwith O pallens a variable taxon adscribed to sectionDiaphana Vent (Lewinsky 1993) Both species share anumber of characters commonly used in identificationkeys emergent and strongly ribbed capsules hemicryp-toporous stomata exostome of 8 pairs of apparentlypapillose teeth conic plicate and naked calyptrae Inaddition variability of O pallens involves a number ofcharacters that may be similar to those of O casasia-

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ORTHOTRICHUM CASASIANUM SP NOV FROM SPAIN 51

Figures 11-14 Comparison of Orthotrichum casasianum F Lara Garilleti amp Mazimpaka and O pallens Brid peristomes (OPL) 11 Tooth ofO casasianum 12 Detail of the median zone of tooth of O casasianum 13 Tooth of O pallens 14 Detail of the inferior half of O pallenstooth [11 and 12 from type 13 and 14 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

num For example O pallens is a taxon typicallycladautoicous although goniautoicous populations havealso been described (Nyholm 1956) endostomesegments may appear in number of 8 8 + 8 or 16 andare smooth or scarcely papillose leaf apices vary fromacute to obtuse apiculate or not and sometimessubmucronate (Bridel 1827) usually leaf cells havehigh and sometimes branched papillae but in somepopulations papillae are low and scarcely visible Thevariability of O pallens is so high that it is sometimesdifficult to separate it from close species like O pumilumSm (Lara Garilleti amp Mazimpaka 1994) moreoverseveral taxa described as independent species or vari-eties of O pallens have been reduced to synonyms ofthe latter (Vitt 1973 Lewinsky 1977 1993) based onthe fact that their differential characters were comprisedin the variability range of this species

O casasianum differs from O pallens by its slenderloosely foliose habit with narrow leaves whose apicesthough variable within the same individual are charac-teristically obtuse and concave often incurvate or cucul-late when dry and with a 1(-2)-celled hyaline apiculusor the costa shortly percurrent Also the capsule form isdifferent although cylindric the capsule is notconstricted at base but is instead ventricose and

narrows more or less sharply in the neck in additionthe mature capsule and the seta are uniformly darkbrown while in O palens the capsule base and the setaare pale brown (a consistent feature on which is basedthe name of the taxon) so that they contrast with thecolour of the theca The most important character ishowever the different ornamentation of the external sideof the exostome teeth (Figs 11-14) in O casasianumalthough the aspect on optical microscope is papillose itis in fact formed by a network of high walls on whichsparse papillae are irregularly disposed in O pallens bycontrast the dense papillosity observed on opticalmicroscope is real and the the basal pattern is markedlydifferent from that of the tooth apex where papilloselines are usually abundant Other peristomial differencesare the occasional presence of prostome in O casasia-num the minor papillosi ty of the exostome PPL and themajor bending of the endostome segments Table Isummarizes the differences between the species

ACKNOWLEDGEMENTS

Thanks are due to the staff of S especially to Lars Hede-nas whose loan of Orthotrichum pallens material made

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Man

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52 V MAZIMPAKA ET AL

Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

--------------~~-----__--- ------------------~--------- --- - --------~--_------~---- - -- --~~~

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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ORTHOTRICHUM CASASIANUM SP NOV FROM SPAIN 51

Figures 11-14 Comparison of Orthotrichum casasianum F Lara Garilleti amp Mazimpaka and O pallens Brid peristomes (OPL) 11 Tooth ofO casasianum 12 Detail of the median zone of tooth of O casasianum 13 Tooth of O pallens 14 Detail of the inferior half of O pallenstooth [11 and 12 from type 13 and 14 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

num For example O pallens is a taxon typicallycladautoicous although goniautoicous populations havealso been described (Nyholm 1956) endostomesegments may appear in number of 8 8 + 8 or 16 andare smooth or scarcely papillose leaf apices vary fromacute to obtuse apiculate or not and sometimessubmucronate (Bridel 1827) usually leaf cells havehigh and sometimes branched papillae but in somepopulations papillae are low and scarcely visible Thevariability of O pallens is so high that it is sometimesdifficult to separate it from close species like O pumilumSm (Lara Garilleti amp Mazimpaka 1994) moreoverseveral taxa described as independent species or vari-eties of O pallens have been reduced to synonyms ofthe latter (Vitt 1973 Lewinsky 1977 1993) based onthe fact that their differential characters were comprisedin the variability range of this species

O casasianum differs from O pallens by its slenderloosely foliose habit with narrow leaves whose apicesthough variable within the same individual are charac-teristically obtuse and concave often incurvate or cucul-late when dry and with a 1(-2)-celled hyaline apiculusor the costa shortly percurrent Also the capsule form isdifferent although cylindric the capsule is notconstricted at base but is instead ventricose and

narrows more or less sharply in the neck in additionthe mature capsule and the seta are uniformly darkbrown while in O palens the capsule base and the setaare pale brown (a consistent feature on which is basedthe name of the taxon) so that they contrast with thecolour of the theca The most important character ishowever the different ornamentation of the external sideof the exostome teeth (Figs 11-14) in O casasianumalthough the aspect on optical microscope is papillose itis in fact formed by a network of high walls on whichsparse papillae are irregularly disposed in O pallens bycontrast the dense papillosity observed on opticalmicroscope is real and the the basal pattern is markedlydifferent from that of the tooth apex where papilloselines are usually abundant Other peristomial differencesare the occasional presence of prostome in O casasia-num the minor papillosi ty of the exostome PPL and themajor bending of the endostome segments Table Isummarizes the differences between the species

ACKNOWLEDGEMENTS

Thanks are due to the staff of S especially to Lars Hede-nas whose loan of Orthotrichum pallens material made

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Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

--------------~~-----__--- ------------------~--------- --- - --------~--_------~---- - -- --~~~

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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52 V MAZIMPAKA ET AL

Figures 15-18 Comparison of Orthotrichum casasianum and O pallens peristomes 15 PPL view of O casasianum 16 Partial view of O casasianumendostome with the connecting membrane an intermediate segment (left) and a principal one (right) 17 PPL view of O pallens 18 Partial view ofO pallens endostome [15 and 16 from type 17 and 18 from Sweden Gotland Klinte sn Klinteberget G Een s n (S)]

Table 1 Summary of main differences between Orthotrichum casasianum and O pallens

Orthotrichum casasianum

Plants slender thinly foliose leaves somewhatflexuose

Leaf apices concave-carinate often cucullateincurved when dry

Capsules cylindric-urceolate with ventricose basemore or less sharply narrowed to the neck Neckand theca the same colour

OPL scarcely papillose reticulate

16 endostome segments linear strongly incurved

Prostome occasionally present

this study possible Financial support was partly providedby the Spanish National Institute of Meteorology and theDGES of the Spanish Ministry of Education and Culturegrant no PB96-1111-C02-0 1

TAXONOMIC ADDITIONS AND CHANGES Orthotrichumcasasianum F Lara Garilleti amp Mazimpaka sp nov

Orthotrichum pallens

Plants compact densely foliose leaves not flexuose

Leaf apices not concave erect when dry

Capsule cylindric less contracted at the mouthneck gradually narrowed and paler than the theca

OPL denseley papillose reticulum not visible

8 8 + 8 16 segments at least the intermediate erector slightly incurved

Prostome absent

REFERENCES

Bridel SE 1827 Bryologia universa vol 1(2) LeipzigGarilleti R Lara F Mazimpaka V Albertos B Heras P Infante M

1998 On the presence of Orthotrichum pulchellum Sm in SpainJournal of Bryology 20 246-249

Heras P Infante M 1998 Orthotrichum sprucei Mont and Orthotri-chum rivulare Turn in Northern Spain Cryptogamie BryologieLichenologie 19 77-81

--------------~~-----__--- ------------------~--------- --- - --------~--_------~---- - -- --~~~

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain

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ORTHOTRCHUM CASASANUM SP NOV FROM SPAIN 53

Lara F Garilleti R Mazimpaka V 1994 Clave y notas para laidentificaci6n de las especies epifitas del genera OrthotrichumHedw en la Pennsula Iberica Boletin de la Sodedad Espanolade Briologia 5 1-6

Lewinsky J 1977 The family Orthotrichaceae in Greenland Ataxonomic revision Lindbergia 4 57-103

Lewinsky J 1993 A synopsis of the genus Orthotrichum Hedw(Musci Orthotrichaceae) Bryobrothera 2 1-59

Nyholm E 1956 Illustrated moss flora of Fennoscandia Fascicle 4Lund Gleerup

Vitt DH 1973 A revision of the genus Orthotrichum in NorthAmerica North of Mexico Bryophytorum Bibliotheca 1 1-108

V MAZIMPAKAand F LARADepartamento de Biologia (Botanica) Facultad de Ciencias Universidad Aut6noma de Madrid E-28049 Madrid SpainR GARILLETIDepartamento de Medio Ambiente Facultad de Ciencias Universidad Europea de Madrid E-28670 Villaviciosa de Od6n Spain

M Infante and P Heras Museo de Ciencias Naturales de Alava Fundadora de las Siervas de Jesus 24 E-OlOOI Vitoria Spain