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Corresponding author: Juan Larraín ( musgoschiloe@gmail.com ) Academic editor: Matt von Konrat
© 2016 Juan Larraín.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Larraín J (2016) The mosses (Bryophyta) of Capitán Prat Province, Aisén Region, southern Chile. PhytoKeys 68: 91-116. https://doi.org/10.3897/phytokeys.68.9181
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The bryophytes of Capitán Prat province have remained one of the least explored in Chile. The eventual construction of several dams on the rivers Baker and Pascua required prospection of all groups of organisms including bryophytes, work that was facilitated by the recent construction of vehicular roads that now offer easy access to previously almost unaccessible locations. The results of intense bryophyte collecting during the austral summer of 2007 are here presented. A total of 260 moss taxa are reported for the province, corresponding to 256 species and four infraspecific taxa, of which 211 are new records for the province, 54 are new for Aisén Region, and two are new records for continental Chile (Pohlia longicollis (Hedw.) Lindb. and Rigodium toxarion var. robustum (Broth.) Zomlefer). Twelve species extend their known distribution ranges to the north, whereas 49 extend them to the south.
Biodiversity, checklist, range extensions, new records, southern South America
The Aisén Region is divided into four administrative provinces, including Capitán Prat as the southernmost province of the Region, bordering by the south with Última Esperanza Province in the Magallanes Region. It is of particular botanical interest due to: (i) the diversity of climates, ranging from the per-humid temperate rainforests along the coast line and the western archipelago, to the extremely dry steppe in the eastern border with Argentina, and (ii) it is the southernmost ice-free area of continental Chile before the beginning of the massive Southern Patagonian Ice Field that partially isolates it from the neighboring Magallanes Region.
Although the Aisén Region has received considerable exploration by bryologists, and there is even a moss flora already published (
Map of Aisén Region showing the four provinces (AI = Aisén; CO = Coihaique; GC = General Carrera; CP = Capitán Prat) and the itinerary of bryological expeditions. A Expeditions that collected mosses previous to this study: 1. Charles Darwin, 1834; 2. Per Dusén, 1896–97; 3. Carl Skottsberg, 1908–09; 4. Federico Reichert and Cristóbal Hicken, 1921; 5. Heikii Roivainen, 1929; 6. Arturo Donat, 1933; 7. Gerhard Schwabe, 1939–41; 8. Rolf Santesson, 1940–41; 9. Tarow Seki, 1967; 10. Hironori Deguchi, 1981 B Explored area during the 2007 expedition where collections reported here were made. Inset: map of Chile showing in red the location of Aisén Region. (dotted line = regional border; continuous line = provincial limit).
Unfortunately, in his checklist of Chilean mosses,
Capitan Prat Province lies in the southern portion of Aisén Region (XI) in southern Chile, spanning between lat. 46°57'-49°09'S and long. 71°51'-75°36'W, with a total area of 37200 km². The province is crossed by three major river basins from north to south (the rivers Baker, Bravo and Pascua), with Baker river the largest river in Chile in terms of the volume of water. The province shows a remarkable heterogeneity of landscapes and climates (Fig.
In terms of the vegetation that dominates the landscape, there is a clearly marked gradient from east to west, mostly determined by the extreme variation in rainfall as explained above. In the easternmost parts of the province the vegetation fits into
Descending in altitude, and towards the west, the vegetation is dominated by deciduous forests and shrublands, inserted in what
Moving further towards the west, the vegetation changes into what
Two field trips were made during the Austral summer of 2007, making up a total of 13 full days of collecting. During these expeditions a total of 1283 bryophyte collections were made throughout the area between the localities of Puerto Bertrand and Villa O’Higgins, most of them adjacent to the main vehicular roads (Route 7, the road to Tortel, and the road to Villa O’Higgins) spanning the whole basin of Baker River and some sites along the Bravo and Pascua River basins (Fig.
The collecting localities were arbitrarily chosen, attempting to include the largest number of different floristic associations and landscapes, both in the driest and the wettest areas of the province and throughout all the gradients in between, from the coast line to about 700 m a.s.l.
The specimens were identified with the monographs or taxonomic revisions currently available for each group, and comparing with reference material kept at
The species list is given below in alphabetical order, indicating the new records for Chile (***), for Aisén Region (**) and for Capitán Prat Province (*). A systematic arrangement of the taxa, including habitat information, altitude where taxa were found, frequency in the studied area, global and Chilean distribution, and a list of studied specimens, is presented in the Suppl. material
For the analysis of the distribution patterns, the many different patterns observed were reduced into five major categories for simplification: (1) Wide distribution (WD), meaning cosmopolitan or subcosmopolitan species present in both hemispheres and in both tropical and temperate areas of the planet; (2) Bipolar, meaning species distributed in the temperate areas of both hemispheres with eventual isolated populations at high altitudes in the tropics; (3) Austral, meaning species with various distribution patterns along the Southern Hemisphere - this is the most heterogeneous group because it includes both species with very wide circumsubantarctic distributions to narrow “sub-endemics” maybe recorded from a few populations in southern South America and South Africa, for example; (4) Andean, meaning species mostly distributed throughout the Andes but incidentally extending into Antarctica, Africa or some subantarctic Islands; and (5) Endemic, meaning species only known from southern South America and eventually extending into Juan Fernández or the Falkland and South Georgia Islands.
A total of 260 moss taxa belonging to 256 species and 4 infraspecific taxa, in 42 botanical families are reported for Capitán Prat Province (Suppl. material
*Achrophyllum anomalum var. anomalum
Achrophyllum anomalum var. pallidum (Cardot & Broth.) S.He
*Achrophyllum haesselianum (Matteri) Matteri
*Achrophyllum magellanicum (Besch.) Matteri (Figure
Acrocladium auriculatum (Mont.) Mitt. (Figure
**Acroschisma wilsonii (Hook.f.) A.Jaeger
*Amphidium tortuosum (Hornsch.) Cufod.
*Ancistrodes genuflexa (Müll.Hal.) Crosby
Andreaea alpina Hedw.
Andreaea fuegiana (Cardot) S.W.Greene
Andreaea regularis Müll.Hal.
Andreaea vaginalis Herzog
Arbusculohypopterygium arbuscula (Brid.) M.Stech, T.Pfeiff. & W.Frey
*Bartramia ithyphylloides Schimp. ex Müll.Hal.
Bartramia mossmaniana Müll.Hal.
Bartramia patens Brid.
*Bartramia robusta Hook.f. & Wilson
*Bartramia stricta Brid.
*Blindia contecta (Hook.f. & Wilson) Müll.Hal.
Blindia magellanica Schimp.
Brachytheciastrum paradoxum (Hook.f. & Wilson) Ignatov & Huttunen
*Brachythecium albicans (Hedw.) Schimp.
*Brachythecium austroglareosum (Müll.Hal.) Paris
*Brachythecium austrosalebrosum (Müll.Hal.) Paris
Brachythecium subpilosum (Hook.f. & Wilson) A.Jaeger
*Brachythecium subplicatum (Hampe) A.Jaeger
*Breutelia angustiretis E.B.Bartram
*Breutelia aureola (Besch. ex Müll.Hal.) Besch.
*Breutelia dumosa Mitt.
Breutelia integrifolia (Taylor) A.Jaeger
*Breutelia plicata Mitt.
*Breutelia subplicata Broth.
**Bryum algovicum Sendtn. ex Müll.Hal.
**Bryum archangelicum Bruch & Schimp.
*Bryum australe Hampe
**Bryum billarderi Schwägr. (Figure
*Bryum caespiticium Hedw.
**Bryum canariense Brid.
*Bryum capillare Hedw.
Bryum clavatum (Schimp.) Müll.Hal.
*Bryum coronatum Schwägr.
Bryum crassinervium Lorentz
**Bryum dichotomum Hedw.
Bryum donatii Thér.
Bryum gemmatum Müll.Hal.
*Bryum laevigatum Hook.f. & Wilson
*Bryum macrophyllum Cardot & Broth.
*Bryum perlimbatum Cardot
**Bryum platyphyllum (Schwägr.) Müll.Hal.
Bryum pseudotriquetrum (Hedw.) Schwägr.
**Bryum puconense Herzog & Thér.
*Bryum subapiculatum Hampe
*Calyptopogon mnioides (Schwägr.) Broth.
*Camptodontium cryptodon (Mont.) Reimers
**Campylium stellatum (Hedw.) C.E.O.Jensen
*Campylopodium euchlorum (Mont.) Matteri
*Campylopodium medium (Duby) Giese & J.-P.Frahm
*Campylopus acuminatus Mitt.
*Campylopus clavatus (R.Br.) Wilson
*Campylopus incrassatus Müll.Hal.
*Campylopus introflexus (Hedw.) Brid.
Campylopus purpureocaulis Dusén
*Campylopus pyriformis (Schultz) Brid.
*Campylopus vesticaulis Mitt.
*Catagonium nitens (Brid.) Cardot
*Ceratodon purpureus subsp. purpureus
*Ceratodon purpureus subsp. convolutus (Reichardt) Burley
Chorisodontium aciphyllum (Hook.f. & Wilson) Broth.
*Chorisodontium dicranellatum (Dusén) Roiv.
*Chorisodontium spegazzinii (Müll.Hal.) Roiv.
**Chrysoblastella chilensis (Mont.) Reimers
*Conostomum pentastichum (Brid.) Lindb.
Cratoneuropsis relaxa subsp. minor (Hook.f. & Wilson) Ochyra
*Cryphaea consimilis Mont.
*Cryphaeophilum molle (Dusén) M.Fleisch.
*Daltonia gracilis Mitt.
*Daltonia trachyodonta Mitt.
*Dendroligotrichum dendroides (Hedw.) Brid.
Dendroligotrichum squamosum (Hook.f. & Wilson) Broth. ex Cardot
*Dicranella campylophylla (Taylor) A.Jaeger
*Dicranella hookeri (Müll.Hal.) Cardot
**Dicranella pseudorufescens Cardot & Broth.
Dicranoloma billardieri (Brid.) Paris
*Dicranoloma chilense (De Not.) Ochyra & Matteri
**Dicranoloma imponens (Mont.) Renauld
*Dicranoloma menziesii (Hook.f. & Wilson) Paris
*Dicranoloma perremotifolium (Dusén) Broth.
Dicranoloma robustum (Hook.f. & Wilson) Paris
*Didymodon andreaeoides Cardot & Broth.
*Didymodon australasiae (Hook. & Grev.) R.H.Zander
*Didymodon fuscus (Müll.Hal.) J.A.Jiménez & M.J.Cano
Distichium capillaceum (Hedw.) Bruch & Schimp.
*Distichophyllum dicksonii (Hook. & Grev.) Mitt.
*Ditrichum cylindricarpum (Müll.Hal.) F.Muell.
*Ditrichum difficile (Duby) M.Fleisch.
*Ditrichum ditrichoideum (Cardot) Ochyra
*Ditrichum heteromallum (Hedw.) E.Britton
**Drepanocladus aduncus (Hedw.) Warnst.
*Drepanocladus longifolius (Mitt.) Broth. ex Paris
*Drepanocladus polygamus (Schimp.) Hedenäs
*Dryptodon austrofunalis (Müll.Hal.) Ochyra & Żarnowiec
*Dryptodon humilis (Mitt.) Ochyra & Żarnowiec
*Dryptodon navicularis (Herzog) Ochyra & Żarnowiec
Dryptodon trichophyllus (Grev.) Brid.
*Encalypta ciliata Hedw.
**Encalypta rhaptocarpa Schwägr.
*Eriodon conostomus Mont.
*Eucamptodon perichaetialis (Mont.) Mont.
*Eurhynchiella acanthophylla (Mont.) M.Fleisch.
*Eustichia longirostris (Brid.) Brid
**Fabronia jamesonii Taylor
*Fissidens curvatus Hornsch.
*Fissidens rigidulus Hook.f. & Wilson
*Funaria hygrometrica Hedw.
*Glyphothecium sciuroides (Hook.) Hampe
*Gymnostomum calcareum Nees & Hornsch.
**Hebantia rigida (Lorentz) G.L.Merr.
Hedwigia ciliata var. nivalis Hampe (Figure
**Hennediella antarctica (Ångström) Ochyra & Matteri
*Hennediella arenae (Besch.) R.H.Zander
**Hennediella heimii (Hedw.) R.H.Zander
*Hymenodontopsis mnioides (Hook.) N.E.Bell, A.E.Newton & D.Quandt (Figure
Hymenoloma crispulum (Hedw.) Ochyra
Hymenoloma turpe (Cardot) Cardot & Broth.
*Hymenostylium recurvirostrum (Hedw.) Dixon
*Hypnodendron microstictum Mitt. ex A.Jaeger & Sauerb.
*Hypnum chrysogaster Müll.Hal.
*Hypnum cupressiforme Hedw. var. cupressiforme
**Hypnum cupressiforme var. filiforme Brid.
*Hypnum cupressiforme var. mossmanianum (Müll.Hal.) Ando
*Hypnum skottsbergii Ando
*Hypopterygium didictyon Müll.Hal.
*Kiaeria pumila (Mitt.) Ochyra
*Leptobryum pyriforme (Hedw.) Wilson
**Leptodontium longicaule var. microruncinatum (Dusén) R.H.Zander
**Leptostomum menziesii R.Br.
Leptotheca gaudichaudii Schwäger.
*Lepyrodon hexastichus (Mont.) Wijk & Margad.
Lepyrodon lagurus (Hook.) Mitt.
Lepyrodon patagonicus (Cardot & Broth.) B.H.Allen
*Lepyrodon tomentosus (Hook.) Mitt.
**Looseria orbiculata (Thér.) D.Quandt, Huttunen, Tangney & M.Stech
*Lopidium concinnum (Hook.) Wilson
**Macromitrium pertriste Müll.Hal.
**Mahua enervis W.R.Buck
*Matteria gracillima (Besch.) Goffinet
*Matteria papillosula (Thér.) Goffinet
*Neckera scabridens Müll.Hal.
*Notoligotrichum minimum (Cardot) G.L.Sm. (Figure
*Notoligotrichum trichodon (Hook.f. & Wilson) G.L.Sm.
*Oligotrichum austroaligerum G.L.Sm.
Ombronesus stuvensis N.E.Bell, N.Pedersen & A.E.Newton
**Orthodontium lineare Schwägr.
*Orthotrichum assimile Müll.Hal.
*Orthotrichum brotheri Dusén ex Lewinsky
*Orthotrichum elegantulum Schimp. ex Mitt.
**Orthotrichum hortense Bosw.
*Orthotrichum incanum Müll.Hal.
*Orthotrichum ludificans Lewinsky
**Orthotrichum pariatum Mitt.
*Orthotrichum rupestre Schleich. ex Schwägr.
*Pararhacocarpus patagonicus (Broth.) J.-P.Frahm
Philonotis brevifolia Herzog
Philonotis esquelensis Matteri
*Philonotis krausei (Müll.Hal.) Broth.
**Philonotis polymorpha (Müll.Hal.) Kindb.
**Philonotis scabrifolia (Hook.f. & Wilson) Braithw.
Philonotis vagans (Hook.f. & Wilson) Mitt.
**Pilopogon schilleri Herzog & Thér.
**Plagiothecium lucidum (Hook.f. & Wilson) Paris
*Platyneuron praealtum (Mitt.) Ochyra & Bednarek-Ochyra (Figure
Pohlia cruda (Hedw.) Lindb.
***Pohlia longicollis (Hedw.) Lindb.
*Pohlia nutans (Hedw.) Lindb. (Figure
*Pohlia wahlenbergii (F.Weber & D.Mohr) A.L.Andrews (Figure
*Polytrichadelphus magellanicus (Hedw.) Mitt. (Figure
**Polytrichastrum alpinum (Hedw.) G.L.Sm.
*Polytrichastrum longisetum (Sw. ex Brid.) G.L.Sm.
*Polytrichum juniperinum Hedw.
Polytrichum piliferum Hedw. (Figure
**Polytrichum strictum Menzies ex Brid.
*Porotrichum arbusculans (Müll.Hal.) Ochyra
*Pseudocrossidium crinitum (Schultz) R.H.Zander
Ptychomniella ptychocarpon (Schwägr.) W.R.Buck, C.J.Cox, A.J.Shaw & Goffinet
*Ptychomnion cygnisetum (Müll.Hal.) Kindb.
*Racomitrium didymum (Mont.) Lorentz (Figure
Racomitrium geronticum Müll.Hal. (Figure
*Racomitrium heterostichoides Cardot
Racomitrium laevigatum A.Jaeger
*Racomitrium lamprocarpum (Müll.Hal.) A.Jaeger (Figure
*Racomitrium orthotrichaceum (Müll.Hal.) Paris
*Racomitrium pachydictyon Cardot
*Racomitrium rupestre (Hook.f. & Wilson) Wilson & Hook.f.
*Racomitrium subcrispipilum (Müll.Hal.) A.Jaeger
*Rhacocarpus purpurascens (Brid.) Paris
**Rhaphidorrhynchium amoenum (Hedw.) M.Fleisch.
*Rhaphidorrhynchium callidum (Mont.) Broth. (Figure
*Rhaphidorrhynchium dendroligotrichum (Dusén) Broth.
*Rigodium adpressum Zomlefer
*Rigodium brachypodium (Müll.Hal.) Paris
*Rigodium pseudothuidium Dusén
*Rigodium tamarix Müll.Hal.
*Rigodium toxarion var. toxarion
***Rigodium toxarion var. robustum (Broth.) Zomlefer
*Sanionia uncinata (Hedw.) Loeske
*Sauloma tenella (Hook.f. & Wilson) Mitt.
Schistidium andinum (Mitt.) Herzog
**Sciurohypnum plumosum (Hedw.) Ignatov & Huttunen
**Scorpidium revolvens (Sw.) Rubers
Scouleria patagonica (Mitt.) A.Jaeger
*Sematophyllum scorpiurus (Mont.) Mitt.
*Sphagnum falcatulum Besch.
Sphagnum fimbriatum Wilson
Sphagnum magellanicum Brid.
Straminergon stramineum (Dicks. ex Brid.) Hedenäs
Syntrichia anderssonii (Ångström) R.H.Zander
**Syntrichia costesii (Thér.) R.H.Zander
**Syntrichia epilosa (Broth. ex Dusén) R.H.Zander
**Syntrichia fragilis (Taylor) Ochyra
**Syntrichia glacialis (Kunze ex Müll.Hal.) R.H.Zander
*Syntrichia lithophila (Dusén) Ochyra & R.H.Zander
*Syntrichia magellanica (Mont.) R.H.Zander
*Syntrichia princeps (De Not.) Mitt.
*Syntrichia pseudorobusta (Dusén) R.H.Zander
**Syntrichia ruralis (Hedw.) F.Weber & D.Mohr
**Syntrichia scabrella (Dusén) R.H.Zander
*Tayloria dubyi Broth.
**Tayloria magellanica (Brid.) Mitt.
**Tayloria stenophysata (Herzog) A.K.Kop.
*Tetraplodon fuegianus Besch. (Figure
**Tortella knightii (Mitt.) Broth.
**Tortula atrovirens (Sm.) Lindb.
Ulota billbuckii Garilleti, Mazimpaka & F.Lara
**Ulota carinata Mitt.
*Ulota fuegiana Mitt.
*Ulota germana (Mont.) Mitt.
Ulota larrainii Garilleti, Mazimpaka & F.Lara
*Ulota luteola (Hook.f. & Wilson) Wijk & Marg.
**Ulota macrocalycina Mitt.
*Ulota macrodontia Dixon & Malta
*Ulota magellanica (Mont.) A.Jaeger
**Ulota phyllantha Brid.
**Ulota pusilla Malta
**Ulota pycnophylla Dusén ex Malta
*Ulota pygmaeothecia (Müll.Hal.) Kindb.
*Ulota rufula (Mitt.) A.Jaeger
Ulota streptodon Garilleti, Mazimpaka & F.Lara
*Vittia pachyloma (Mont.) Ochyra
*Warnstorfia exannulata (Schimp.) Loeske
**Warnstorfia fluitans (Hedw.) Loeske
*Warnstorfia fontinialopsis (Müll.Hal.) Ochyra
*Weymouthia cochlearifolia (Schwägr.) Dixon
*Weymouthia mollis (Hedw.) Broth.
**Zygodon hookeri var. hookeri
*Zygodon hookeri var. leptobolax (Müll.Hal.) Calabrese
*Zygodon magellanicus Dusén ex Malta
*Zygodon papillatus Mont.
*Zygodon pentastichus (Mont.) Müll.Hal.
*Zygodon pichinchensis (Taylor) Mitt.
Zygodon reinwardtii (Hornsch.) A.Braun
From the analysis of the five simplified distribution patterns, 38.08% (n=99) of the taxa are actually endemics of southern South America, followed by the “austral” taxa with 30.77% (n=80). The rest of the taxa were found to be either of “wide distribution” (13.85%, n=36), “bipolar” (9.62%, n=25), or “Andean” (7.69%, n=20).
The known diversity of Capitán Prat Province was incremented from 49 to 260 taxa, representing a 531% increment. This is remarkable and demonstrates the very little attention this area has historically received in terms of its bryophyte diversity. The current number of species reported for the province makes sense with the numbers known from the adjacent provinces of Aisén (311) and Última Esperanza (250). The number of moss taxa known from adjacent General Carrera province [27 in
Three taxa were described as new based on the collections gathered during this study in Capitán Prat (Ulota billbuckii, U. larraini and U. streptodon), and four others were found to be new records for Chile (Hedwigia ciliata var. nivalis, Philonotis esquelensis, Pohlia longicollis and Rigodium toxarion var. robustum), the latter previously known to be an endemic of the Juan Fernández Islands. The number of new species and new records for Chile might increment even more after study of the material that still remains unidentified.
The high level of endemic taxa in southern South America temperate rainforests has already been reported in the literature for neighboring areas (i.e.,
Several species were found farther south from their previously known distribution ranges. This is interesting as it would appear to suggest that many of these taxa are apparently not able to disperse south of the Southern Patagonia Ice Field. This ice field would act as a natural barrier to the dispersion of Valdivian rainforest endemics such as Ancistrodes genuflexa, Cryphaea consimilis, Cryphaeophilum molle, Eriodon conostomus, Eurhynchiella acanthophylla, Rhaphidorrhynchium dendroligotrichum, Rigodium tamarix, and Weymouthia cochlearifolia, among others, preventing their colonization of Magallanes Region. This might be due also to less collecting effort in the more wet areas of Magallanes, because of the logistical difficulties to access the most western islands, where rainfall is similar to what can be found in the wettest areas of Capitán Prat.
It is important to mention that not a single locality above 700 m was visited. Although most of the area above this altitude is covered in ice and snow all year round, it would be interesting to visit higher altitude spots and glaciers, since several of the species found by Donat around the glaciers of Lago O’Higgins were not found during this expedition. Covering these sites would maybe increase even more the number of moss taxa known from Capitán Prat Province.
This work was made as a consultancy for Hidroaysén hydroelectric project within the framework of the baseline study prior to the environmental-impact study as required by the Chilean authority. I am specially acknowledged to the staff at the botany department of Universidad de Concepción, and to the many colleagues that helped with the determination of difficult groups of mosses: María Jesús Cano, Mayte Gallego, Ricardo Garilleti, Paco Lara, Lars Hedenäs, John Spence, Barbara Murray, Guille Suárez, Sol Jimenez and Barbara Andreas. Reinaldo Vargas and Götz Palfner are acknowledged for company during the fieldwork. Thanks also to Alicia Marticorena, Alejandra Jiménez, Ernesto Teneb and Roberto Rodríguez for making their collections from the area available for my study. I acknowledge the staff at CONAF office in Cochrane for allowing me to collect inside Tamango National Reserve. Finally I would like to thank the staff at
Systematic list of taxa and additional information
Data type: specimens data