- Email: [email protected]
A survey of the Campylopus species from the Azores
145
Cryptogamie, Bryol., 1999, 20 (2): 145-152
A survey of the Campylopus Jan-Peter Botanisches
Institut
species from the Azores FRAHM
der Universitat, Meckenheimer [email protected]
(Received
7 September
1998, accepted
Allee
170, 53115 Bonn,
27 January
Germany;
1999)
Abstract-Nine species of Campylopus are reported from the Azores. A key is given for the species as well as comments on the taxonomy, nomenclature, the worldwide ranges and the distribution on the Azores. 0 ADAC / Elsevier, Paris
R&urn6 - Neuf especes de Campylopus sont signal&es aux Acores. Une cl6 de determination des espece est prCsent6e et leur taxonomie, nomenclature et distribution mondiale et locale sont prtcisees. 0 ADAC I Elsevier, Paris
INTRODUCTION The checklist of the bryophytes of the Macaronesian Islands by Eggers (1982) lists 11 speciesof Campylopus for the Azores. The determination of these speciesis not easy becauseCampylopus is regarded asa difficult genusand the speciesfrom the Azores are composed of very different phytogeographical elements (e.g. Caribbean, African, European, Subantarctic). Therefore descriptionsand keys facilitating the identification are only found in many various and scattered publications. Furthermore the checklist by Eggers dates back 16 years and since that time several taxonomic changeshave been made. For that reasonand following an idea of Rent! Schumacker, a short survey of the Campylopus flora of the Azores is presentedhere.
TAXONOMIC
AND NOMENCLATURAL
REMARKS
Eggers (1982) listed the following 11 speciesof Campylopus for the Azores: C. ampliretis (C. Mull.) Par., C. atroluteus (C. Mull.) Par., C. atrovirens De Not., C. brevipilus B., S. & G., C. carreiroanus Card., C.fragiEis (Brid.) B., S. & G., C. marginatulus Geh., C. paradoxus Wils. in Hardy, C. pilifer Brid., C. pyriformis (Schultz) Brid. and C. setaceus Card. Campylopus ampliretis was describedby C. Mtiller from South Africa from a collection madeby Rehmannat Montagu Pass.Under this nameit is treated in the South African mossflora (Magill, 1981). This speciesproved, however, to be synonymouswith C. flaccidus Ren. & Card. (Frahm, 1984) which is a widespreadtropical African species
146
J.P. Frahm
(for details see Frahm, 1985a). Following Sim (1926) and the authors of the Index Muscorum, C. ampliretis is identical to C. lepidophyllus (C. Mull.) Jaeg. The type of C. lepidophyllus is, however, identical to C. introjkxus (Hedw.) Brid. (Frahm, 1975) and thus it is a synonym of the latter. The recordsof Cumpy~lopus atroluteus from Madeira and the Azores are a highly confusing matter. Eggers (1982) lists C. bartrumiuceus (C. Mull.) Par., C. calvatus (Mitt. in Dix.) Luis. and C. dixonii Luis. as synonyms. In 1927, Luisier describeda new species named C. dixonii from Madeira and compared it with Dicrunum atroluteum and Dicranum bartramiaceum from South Africa. Theriot (1939) synonymized C. dixonii with C. bartrumiaceus. However, as pointed out by Frahm (1976), Sim (1926) had already synonymized C. bartramiaceus with C. atroluteus. This is the reason that C. dixonii was named as C. atroluteus by Eggers (1982). Campylopus calvatus was describedby Dixon. Dixon had received a specimenfrom Luisier collected on Madeira and statesits identity with a specimenin the herbarium of Mitten, collected on Tristan da Cunha, which Mitten had provisionally namedC. calvatus. Accordingly, Dixon validated this specieswith two syntypes, one from Tristan da Cunha and the other from Madeira. Frahm (1981) was able to show that C. calvatus was, however, identical to C. dixonii, also described from Madeira, which was named C. atroluteus, since Theriot had stated its identity with C’. bartramiaceus from South Africa, which was again a synonym of C. utroluteus following Sim (1926). For his treatment of the African species of Campylopus, the author (Frahm, 1985a) studied the types of C. atroluteus and C. bartramiaceus. This study revealed that the type of C. atroluteus had nothing in common with the material of C. calvatus/dtkonii/bartramiaceus/atroluteus from Madeira, but showed cucullate leaf apices, was related to C. bicolor (C. Mull.) Wils. in Hook. from Australia and New Zealand and thus placed as a subspeciesof the latter. Cumpylopus burtrumiaceus proved not to be identical with C. atroluteus but a separatespecies,but also not identical with the specimensfrom Madeira. This Madeira material could be identified as C. eximius Reichdt., a subantarctic speciesknown under various synonyms (C. alvareziunus Card., C. jluvoviridis Dus., C. falci$olius Mitt., C. rullieri Card.) from Australia, New Zealand, Ascension, St. Paul, Gough Islands and Patagonia. Later on, C. incrussatusC. Mull. was found as an older name (Frahm, 1987). Although described from Madeira, Sjogren (1978) reported this speciesfrom Sao Jorge. Campylupus atrovirens was only recorded from Terceira by Da Cunha & de Barros (1942). Collaborative evidence could not be located. Since this specieshasnever be found by any other bryologist in the Azores, this record seemsdoubtful. Therefore this speciesis omitted in the key. The records of Campylopus brevipilus are mainly basedon C. brevipilus var. marginatulus (C. marginatulus Geh.), which proved to be a synonym of Ditrichum punctulutum (for details seenote under C. marginatulus). Campylopuscarreiroanus was describedby Cardot as endemicin the Azores. It proved to be synonymous with C. shawii Wils. from Britain (Frahm, 1985b). Cumpyi’opusmurginatulus was describedby Geheeb in 1910. Kindberg (1888) had publishedthis speciesalready earlier asnomennudum C. madeirensisGeh. in Kindb. Theriot (1939) placed it as a variety to C. brevipilus. According to Seppelt in Frahm (1985a) this taxon representsDitrichum punctulatum Mitt., a circum-subantarctic species. Eggers(1982) supposesthat all records of C. brevipilus from the Azores might belong to C. marginutulus and thus to Ditrichum punctulatum, which is, however, not the case. Cumpylopusparadoxus Wils. in Hardy was introduced by Corley (1976) for C.~exuosus (Hedw.) Brid., because C. jlexuosus was not typified (as many species described by Hedwig). A lectotypification of C. frexuosus (Frahm & Geissler, 1985) allowed this name to be usedagain.
A survey of the Campylopus species from the Azores
147
Campylopus pilifer Brid. hasproved to be the correct namefor C. polytrichoides De Not. All records of C. introfexus (Hedw.) Brid. from the Azores belong to this species. Campylopus pyriformis (Schultz) Brid. was originally describedas Campylopus azoricus Mitt. from the Azores. Thtriot placed it as a variety to C. jlexuosus. Corley (1976) stated its identity with C. pyriformis and transferred the variety to C. pyr$ormis. This variety differs by taller plants and seemsto be a modification of wetter habitats. It has also been found in Europe. Campylopus setaceus Card. was described as endemic from the Azores but proved to be synonymous with C. cygneus (Hedw.) Brid. from the Caribbean (Frahm, 198%).
KEY TO THE SPECIES OF CAMPYLOPUS FROM THE AZORES Subsequentrevisions after the publication of the checklist of bryophytes of the Macaronesian Islands by Eggers (1982) resulted in a reduction from 11 to 9 species,of which 5 proved to be synonymous with other speciesand received other names. I
1” 2
2” 3
3”
4 4” 5 5” 6 6* 7 7* 8
Basal laminal cells incrassate,chlorophyllose ..... .............. . . ............ . .......... . .. 2 Basal laminal cells thin walled, hyaline ................................................................ 4 Leaves ending in a hyaline hairpoint. Plants blackish, robust. Basal laminal cells short rectangular or subquadrate,pitted. Costa relatively narrow (l/3 of leaf width), with ventral and dorsal stereids . .... ... ... ... ... ... ... ... ... ... ... .. .... ... ... ... ... ... ... . C. flaccidus Leaves without hyaline hairpoint ........................................................................... 3 Leaves short, 3-7 mm long. Costa in transversesection with small hyalocysts which are as small or even smaller than the median deuter cells. Basal laminal cells rectangular, rarely pitted. Upper laminal cells rectangular, smooth. Plants dark green, often with flagelliform brood branches ... . . .............. .............. C. ,jexuosus Leaves long, 6-l 1 mm, ending in a long fine almost entire subula. Costa in transverse section with large ventral hyalocysts, occupying half of the leaf width. Basal laminal cells short rectangular to almost subquadrate................................ 4 Upper laminal cells quadrate.Alar cells protruding into the costa. Plants up to 7 cm high, glossy ............................................................................................ C. cygneus Upper laminal cells oval, sometimesporose.Alar cells not protruding into the costa. Plants yellowish green, very large, up to IO cm high, Dicranum-like . C. shawii Leaves ending in a hyaline hairpoint .................................................................... 6 Leaf tips concolorous ............................................................................................. 8 Upper laminal cells oval to elongate oval ....................................._...................... 7 Upper laminal cells more or lessquadrate. Upper part of the dorsal sideof the costa smooth, in transverse section without lamellae . .. .... ... .. ... .... ... ... .. .... C. incrassatus Upper part of the dorsal side of the costa in transverse section with lamellae 34 cells high. Upper laminal cells oval .. ... .... ... ... .. .... ... ... .. .... ... .. .... ... ... ... C. pilifer Upper part of the dorsal side of the costa smooth. Costa in transversesection with dorsal and ventral stereids .. ... ... .. .... ... ... .. .... ... ... ... ... ... ... .... .. ... ... ... ... ... .. C. brevipilus Leaves longly subulate,chanelled.Upper laminal cells short rectangular, Transverse section of costa with small ventral hyalocysts and dorsal groups of stereids.
J.P. Frahm
148
8*
Vegetative propagation rare on the Azores, by deciduous stem tips C. pyriformis Leaves shorter, pointed. Upper laminal cells quadrate. Transverse section of costa with large ventral hyalocysts and dorsal groups of stereids. Vegetative propagation by deciduous brood leaves produced on comal tufts ... .... ... .. ... .... ... .. ... ... C. fragilis
DISTRIBUTION
IN THE AZORES AND WORLDWIDE
Campylopus brevipilus B., S. 8s G. As indicated above, most recordsof this speciesprobably concern C. brevipilus var. marginatulus, which is a synonym of Ditrichum punctulatum. However, Frahm (1985a) lists one specimenkept in the Botanical Museum Stockholm (S) and collected from the Azores collected by Persson in 1937 on Terceira. Another specimen was collected by G. Schwab in Caldeiras, Sao Miguel, in 1981 at 200 m altitude. Additonal unrevised literature records are from Faial and Flores.
Campylopus cygneus (Hedw.) Brid. A Caribbeanspeciesoccurring on Cuba, Jamaica,Haiti, Hispaniola, Puerto Rico and the LesserAntilles (Frahm 1991). On the Azores found on seven of nine islands (Faial, Flores, Pica, Sao Jorge, Sao Miguel, Santa Maria, Terceira).
Campylopus flaccidus Ren. & Card A specieswidespread in tropical Africa (Frahm 1985a). From the Azores it was-according to Eggers(1982)-only known from Sao Miguel but later collected on Pica (comm. Schumacker).
Campylopus flexuosus Hedw. This speciesoccurs in western Europe, in North America in a single locality in the Appalachian Mts. and a few localities in British Columbia and is additionally widespreadin tropical montane regions in Central and South America and Africa. It is widespreadon the Azores (Faial, Flores, Pica, Sao Jorge, Sao Miguel, Santa Maria and Terceira).
Campylopus fragilis (Brid.) B., S. & G. It has a range comparableto C. flexuosus in western Europe, in North America at a single locality in Arkansas and somelocalities in British Columbia and the tropical mountainsof Central and South America and Africa. In the Azores it is quite widespread and was so far found on Faial, Flores, Pica, Sao Jorge, Sao Miguel and Terceira.
A survey of the CumpyEopus species from the Azores
149
Campylopus incrassatusC. Miill A speciesranging from New Zealand, Australia, South Africa to Patagoniaand many subantarctic andAtlantic islands(Ascension, Madeira, Azores). In the Azores it was only known from Sao Jorge but was recently collected on Pica and Terceira (comm. Schumacker). Campylopus pilifer Brid. Like C. flrxuosus and C. fragilis widespreadin the tropical mountains of Africa and America and also in SW Europe and SE North America, and like the preceding speciesalso widespread on the Azores (Faial, Flores, Graciosa, Pica, Sao Jorge, Sao Miguel, Santa Maria, Terceira). Campylopus pyriformis (Schuttz.) Brid. Distributed through the subantarctic to subtropical regions of the southern hemisphere and found also in Central Africa, western Europe and recently also in SE North America. On the Azores it was known from Flores, SaoMiguel and Terceira. An additional unpublishedrecord wasmadeby G. Schwab on Faial (SchwabSN 270, 17.6.81, hb. Frahm, BONN). The speciesis found in two varieties: var. pyriformis consists of small plants up to I cm high and grows preferably on rotten wood (of Culcita, Erica, Juniperus), var. azoricus (Mitt.) Corley consistsof large plants severalcm high, which are always sterile. Such plants are regarded as modifications from wet habitats. Campylopus shawii Wils. A specieshighly disjunct between the oceanic parts of Britain, the Azores and the Caribbean (Jamaica, Cuba, Dominican Republic), from where it was known as C. brittonae Williams and C. under-woo&i Williams (Frahm, 1985b). This kind of disjunction is hard to explain. It could be a relatively young introduction by man or long distance dispersalacrossthe Atlantic Ocean. The presencein Britain might also indicate a relict from the Tertiary, which has survived glaciation periods under a mild oceanic climate. This hypothesis is supportedby the presenceof C. setifolius Wils. in Britain, a very closely related species occurring on wet rocks instead of swamps and being differentiated by smaller ventral hyalocysts in transversesection of the costa and a serrate leaf apex. If C. set@diusis in fact endemic to Britain, it may have been derived from C. shawii by occupying another ecological niche. In the Azores it is known from Faial, Sao Miguel and Terceira. The Campylopus speciesof the Azores can be grouped into the following phytogeographical elements: Caribbean: cygneus African ampliretis European: brevipilus Subantarctic: incrassatus Tropical montane: pilifel; jexuosus, fragilis Disjunct Caribbean-Oceanic Europe: shawii Disjunct Subantarctic-W Europe: pyriformis
150
J.P. Frahm
Most ot‘ the speciesof Campylopus recorded from the Azores (five out of nine) are known from the Caribbean (C. cygneus, shawii, fragilis, pilifel; jlexuosus) and therefore the Caribbean Islands seemto be the main source for the colonization of these young volcanic islands. Subantarctic elements account for two species (incrassatus, pyriformis), African and European (C. jlaccidus and C. brevipilus) account for two more species. The distribution on the different islandsof the Azores (basedon Eggers (1982), completedby Sjogren (1990, 1993) and additional unpublishedrecordsby G. Schwab and R. Schumacker) is shown in the table.
Corvo
Faial
Graciosa
Pica
Sao Jorge
X
X
?
Sao Miguel
brevipilus
X
Campylopus
cygneus
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
,jlacciduJ
Campylupus
,jexuosw
Cumpylopus,frugilis
X
Campylupus
incrawurus
Campylopus
pilifer
X
Campylopu.~
pyr(formis
X
Campylopus
shawii
6
X X
X
6
5
X
X X
X
X
2
X
X
8
X X
X
5
Terceira
X
X
6
Santa Maria
X
Campylopus
Camp~~lopus
?
Flares
X
3
R
Eight out of nine speciesare found on Sao Miguel and on Terceira. Faial, Corvo and Pica have six, Flores and Sao Jorgehave five species,Santa Maria three and Graciosa only two. It is conspicuous that the specieson the islands with few speciesnumbers belong to widespread species. Rare species are found on islands with high species numbers. The species with the widest ranges are most widespread on the Azores. Campylopus pilijk was collected on all islands, on which Campylopus species occur, C.$exuosus and C. fragilis on eight out of nine. The Caribbean speciesare in part widespread(C. cygneus: eight of nine islands), in part rare (C. shawii: three out of nine islands).The rarest speciesare C. JIaccidus (two islands),followed by C. incrassatus and C. brevipilus (three islands) and C. pyrtformis (five islands). However, the floristic exploration might still be insufftcient, as expressedby the fact that Eggers (1982) listed no speciesof Campylopus from the island of Corvo, but a few years later Sjijgren (1993) listed six species. For the Cape Verde Islands, a distinct correlation was stated between the elevation and the speciesnumber of bryophytes (Frahm et al., 1996). In the case of the Azores, this trend is not very distinct (Fig. 1) with regard to the genus Campylopus. In contrast, the correlation between the area of the islands and species numbers of Campylopus (Fig. 2) is more pronounced, with the exception of the island of Corvo. Acknowledgements. I would like to thank R. Schumacker for suggestions and additions to the manuscript. E. Sjiigren provided valuable comments and literature.
A survey of the
151
species from the Azores
Campylopus
r-- ~ ; :-- -~~
elevation
-.*.-
species
(m) number
,
2500
- 7 2000
7 6 \
i
‘,S,
r
. 5
1500
1 4 1 3
1000
2 500 1 0
Fig. 1. Correlation their elevation.
between the species numbers of
Campylopus
on the islands of the Azores and
Cumpylopus
on the islands of the Azores and
600t 500 1~ 400 i j
Fig. 2. Correlation their
size.
between the species numbers of
J.P. Frahm
152
REFERENCES CORLEY
M.F.V., 1976 - The taxonomy of Campylopus pyriformis (Schultz) Brid. and related species. Journal of Bryology 9: 193-212. DA CUNHA A.G., DE BARROS G., 1942 - Algunas especies de musgos da Terceira nova para OS Acores ou para a ilha. Boletim da Sociedade Portuguesa de Ciencias Naturais 13: 156-157. EGGERS J., 1982 - Artenliste der Moose Makaronesiens. Cryptogamie, Bryologie-Lichhologie 3 (4):
FRAHM
283-335.
J.-P., 1975 -
FRAHM FRAHM
Taxonomische Notizen zur Gattung Campylopus. Revue bryologique et 41 (3): 321-332. J.-P, 1976 - Taxonomische Notizen zur Gattung Campylopus II. Revue bryologique et Zichenologique 42 (1): 603-616. J.-P, 1981 - Taxonomische Notizen zur Gattung Campylopus VIII. Herzogia 5: 517-529. J.-P., 1984 - Campylopus Brid. 1. Subg. Thysanomitrion (Schwaegr.) Kindb. emend. J.-P. Frahm. Nova Hedwigia 39: 585-621. J.-P., 1985a - Afrikanische Campylopus-Arten. Bryophytorum Bibliotheca 31: I-216. J.-P, 1985b - The world-range of Campylopus shawii Wils. Journal of Bryology 13:
FRAHM
J.-P, 1985~ -
lichenologique
FRAHM FRAHM FRAHM
359-367.
Taxonomische
Notizen zur Gattung Campylopus
XIV. Nova
Hedwigia
41:
273-277.
FRAHM
J.-P., 1987 -
A revised list of the Campylopus species of the world. Bryologische I-117. J.-P, 1991 - Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica Monograph 54, 238 pp., New York. J.-P & GEISSLER P., 1985 - Lectotypification von Dicranum flexuosum Hedw. (Campylopus jlexuosus [Hedw.] Brid.) Cryptogamie Bryologie-Lichenologie. 6 (3): Beitrage
FRAHM FRAHM
7:
287-290.
FRAHM
J.-P., LINDLAR A., SOLLMAN P., FISCHER E., 1996 - Bryophytes from the Cape Verde Islands. Tropical Bryology 12: 123-154. KINDBERG N., 1888 - Enumeratio bryinearum exoticarum quam alphabetice disposuit N. Conr. Kindberg. Linkoping. MAGILL R.E., 1981 - Flora of South Africa, Bryophyta, Part 1 Mosses, Fast. 1 Sphagnaceae Grimmiaceae. Pretoria. SIM T.R., 1926 -- The Bryophyta of South Africa Transactions of the Royal Society of South Africa
SJGGREN
I 5.
E., 1978 Broteriana
26:
Bryophyte
vegetation in the Azores Islands. Memorias
de Sociedade
l-273.
SJGGREN E., 1990 - Bryophyte flora and vegetation on the island of Graciosa (Azores), with remarks on floristic diversity of the Azorean Islands. Arquipelago, Life and Earth Sciences
SJGGREN
8: 63-96.
E., 1993 - Bryophyte flora and vegetation on the island of Corvo (Azores). Arquipclago, L$e and Marine Sciences 1IA: 17-48. THERIOT I., 1939 - Campylopodioidtes des Iles Acores recolt&es par P. Allorge and H. Persson. Revue brvologique et lichenologique I 1: IO@-109.
Cryptogamie, Bryol., 1999, 20 (2): 145-152
A survey of the Campylopus Jan-Peter Botanisches
Institut
species from the Azores FRAHM
der Universitat, Meckenheimer [email protected]
(Received
7 September
1998, accepted
Allee
170, 53115 Bonn,
27 January
Germany;
1999)
Abstract-Nine species of Campylopus are reported from the Azores. A key is given for the species as well as comments on the taxonomy, nomenclature, the worldwide ranges and the distribution on the Azores. 0 ADAC / Elsevier, Paris
R&urn6 - Neuf especes de Campylopus sont signal&es aux Acores. Une cl6 de determination des espece est prCsent6e et leur taxonomie, nomenclature et distribution mondiale et locale sont prtcisees. 0 ADAC I Elsevier, Paris
INTRODUCTION The checklist of the bryophytes of the Macaronesian Islands by Eggers (1982) lists 11 speciesof Campylopus for the Azores. The determination of these speciesis not easy becauseCampylopus is regarded asa difficult genusand the speciesfrom the Azores are composed of very different phytogeographical elements (e.g. Caribbean, African, European, Subantarctic). Therefore descriptionsand keys facilitating the identification are only found in many various and scattered publications. Furthermore the checklist by Eggers dates back 16 years and since that time several taxonomic changeshave been made. For that reasonand following an idea of Rent! Schumacker, a short survey of the Campylopus flora of the Azores is presentedhere.
TAXONOMIC
AND NOMENCLATURAL
REMARKS
Eggers (1982) listed the following 11 speciesof Campylopus for the Azores: C. ampliretis (C. Mull.) Par., C. atroluteus (C. Mull.) Par., C. atrovirens De Not., C. brevipilus B., S. & G., C. carreiroanus Card., C.fragiEis (Brid.) B., S. & G., C. marginatulus Geh., C. paradoxus Wils. in Hardy, C. pilifer Brid., C. pyriformis (Schultz) Brid. and C. setaceus Card. Campylopus ampliretis was describedby C. Mtiller from South Africa from a collection madeby Rehmannat Montagu Pass.Under this nameit is treated in the South African mossflora (Magill, 1981). This speciesproved, however, to be synonymouswith C. flaccidus Ren. & Card. (Frahm, 1984) which is a widespreadtropical African species
146
J.P. Frahm
(for details see Frahm, 1985a). Following Sim (1926) and the authors of the Index Muscorum, C. ampliretis is identical to C. lepidophyllus (C. Mull.) Jaeg. The type of C. lepidophyllus is, however, identical to C. introjkxus (Hedw.) Brid. (Frahm, 1975) and thus it is a synonym of the latter. The recordsof Cumpy~lopus atroluteus from Madeira and the Azores are a highly confusing matter. Eggers (1982) lists C. bartrumiuceus (C. Mull.) Par., C. calvatus (Mitt. in Dix.) Luis. and C. dixonii Luis. as synonyms. In 1927, Luisier describeda new species named C. dixonii from Madeira and compared it with Dicrunum atroluteum and Dicranum bartramiaceum from South Africa. Theriot (1939) synonymized C. dixonii with C. bartrumiaceus. However, as pointed out by Frahm (1976), Sim (1926) had already synonymized C. bartramiaceus with C. atroluteus. This is the reason that C. dixonii was named as C. atroluteus by Eggers (1982). Campylopus calvatus was describedby Dixon. Dixon had received a specimenfrom Luisier collected on Madeira and statesits identity with a specimenin the herbarium of Mitten, collected on Tristan da Cunha, which Mitten had provisionally namedC. calvatus. Accordingly, Dixon validated this specieswith two syntypes, one from Tristan da Cunha and the other from Madeira. Frahm (1981) was able to show that C. calvatus was, however, identical to C. dixonii, also described from Madeira, which was named C. atroluteus, since Theriot had stated its identity with C’. bartramiaceus from South Africa, which was again a synonym of C. utroluteus following Sim (1926). For his treatment of the African species of Campylopus, the author (Frahm, 1985a) studied the types of C. atroluteus and C. bartramiaceus. This study revealed that the type of C. atroluteus had nothing in common with the material of C. calvatus/dtkonii/bartramiaceus/atroluteus from Madeira, but showed cucullate leaf apices, was related to C. bicolor (C. Mull.) Wils. in Hook. from Australia and New Zealand and thus placed as a subspeciesof the latter. Cumpylopus burtrumiaceus proved not to be identical with C. atroluteus but a separatespecies,but also not identical with the specimensfrom Madeira. This Madeira material could be identified as C. eximius Reichdt., a subantarctic speciesknown under various synonyms (C. alvareziunus Card., C. jluvoviridis Dus., C. falci$olius Mitt., C. rullieri Card.) from Australia, New Zealand, Ascension, St. Paul, Gough Islands and Patagonia. Later on, C. incrussatusC. Mull. was found as an older name (Frahm, 1987). Although described from Madeira, Sjogren (1978) reported this speciesfrom Sao Jorge. Campylupus atrovirens was only recorded from Terceira by Da Cunha & de Barros (1942). Collaborative evidence could not be located. Since this specieshasnever be found by any other bryologist in the Azores, this record seemsdoubtful. Therefore this speciesis omitted in the key. The records of Campylopus brevipilus are mainly basedon C. brevipilus var. marginatulus (C. marginatulus Geh.), which proved to be a synonym of Ditrichum punctulutum (for details seenote under C. marginatulus). Campylopuscarreiroanus was describedby Cardot as endemicin the Azores. It proved to be synonymous with C. shawii Wils. from Britain (Frahm, 1985b). Cumpyi’opusmurginatulus was describedby Geheeb in 1910. Kindberg (1888) had publishedthis speciesalready earlier asnomennudum C. madeirensisGeh. in Kindb. Theriot (1939) placed it as a variety to C. brevipilus. According to Seppelt in Frahm (1985a) this taxon representsDitrichum punctulatum Mitt., a circum-subantarctic species. Eggers(1982) supposesthat all records of C. brevipilus from the Azores might belong to C. marginutulus and thus to Ditrichum punctulatum, which is, however, not the case. Cumpylopusparadoxus Wils. in Hardy was introduced by Corley (1976) for C.~exuosus (Hedw.) Brid., because C. jlexuosus was not typified (as many species described by Hedwig). A lectotypification of C. frexuosus (Frahm & Geissler, 1985) allowed this name to be usedagain.
A survey of the Campylopus species from the Azores
147
Campylopus pilifer Brid. hasproved to be the correct namefor C. polytrichoides De Not. All records of C. introfexus (Hedw.) Brid. from the Azores belong to this species. Campylopus pyriformis (Schultz) Brid. was originally describedas Campylopus azoricus Mitt. from the Azores. Thtriot placed it as a variety to C. jlexuosus. Corley (1976) stated its identity with C. pyriformis and transferred the variety to C. pyr$ormis. This variety differs by taller plants and seemsto be a modification of wetter habitats. It has also been found in Europe. Campylopus setaceus Card. was described as endemic from the Azores but proved to be synonymous with C. cygneus (Hedw.) Brid. from the Caribbean (Frahm, 198%).
KEY TO THE SPECIES OF CAMPYLOPUS FROM THE AZORES Subsequentrevisions after the publication of the checklist of bryophytes of the Macaronesian Islands by Eggers (1982) resulted in a reduction from 11 to 9 species,of which 5 proved to be synonymous with other speciesand received other names. I
1” 2
2” 3
3”
4 4” 5 5” 6 6* 7 7* 8
Basal laminal cells incrassate,chlorophyllose ..... .............. . . ............ . .......... . .. 2 Basal laminal cells thin walled, hyaline ................................................................ 4 Leaves ending in a hyaline hairpoint. Plants blackish, robust. Basal laminal cells short rectangular or subquadrate,pitted. Costa relatively narrow (l/3 of leaf width), with ventral and dorsal stereids . .... ... ... ... ... ... ... ... ... ... ... .. .... ... ... ... ... ... ... . C. flaccidus Leaves without hyaline hairpoint ........................................................................... 3 Leaves short, 3-7 mm long. Costa in transversesection with small hyalocysts which are as small or even smaller than the median deuter cells. Basal laminal cells rectangular, rarely pitted. Upper laminal cells rectangular, smooth. Plants dark green, often with flagelliform brood branches ... . . .............. .............. C. ,jexuosus Leaves long, 6-l 1 mm, ending in a long fine almost entire subula. Costa in transverse section with large ventral hyalocysts, occupying half of the leaf width. Basal laminal cells short rectangular to almost subquadrate................................ 4 Upper laminal cells quadrate.Alar cells protruding into the costa. Plants up to 7 cm high, glossy ............................................................................................ C. cygneus Upper laminal cells oval, sometimesporose.Alar cells not protruding into the costa. Plants yellowish green, very large, up to IO cm high, Dicranum-like . C. shawii Leaves ending in a hyaline hairpoint .................................................................... 6 Leaf tips concolorous ............................................................................................. 8 Upper laminal cells oval to elongate oval ....................................._...................... 7 Upper laminal cells more or lessquadrate. Upper part of the dorsal sideof the costa smooth, in transverse section without lamellae . .. .... ... .. ... .... ... ... .. .... C. incrassatus Upper part of the dorsal side of the costa in transverse section with lamellae 34 cells high. Upper laminal cells oval .. ... .... ... ... .. .... ... ... .. .... ... .. .... ... ... ... C. pilifer Upper part of the dorsal side of the costa smooth. Costa in transversesection with dorsal and ventral stereids .. ... ... .. .... ... ... .. .... ... ... ... ... ... ... .... .. ... ... ... ... ... .. C. brevipilus Leaves longly subulate,chanelled.Upper laminal cells short rectangular, Transverse section of costa with small ventral hyalocysts and dorsal groups of stereids.
J.P. Frahm
148
8*
Vegetative propagation rare on the Azores, by deciduous stem tips C. pyriformis Leaves shorter, pointed. Upper laminal cells quadrate. Transverse section of costa with large ventral hyalocysts and dorsal groups of stereids. Vegetative propagation by deciduous brood leaves produced on comal tufts ... .... ... .. ... .... ... .. ... ... C. fragilis
DISTRIBUTION
IN THE AZORES AND WORLDWIDE
Campylopus brevipilus B., S. 8s G. As indicated above, most recordsof this speciesprobably concern C. brevipilus var. marginatulus, which is a synonym of Ditrichum punctulatum. However, Frahm (1985a) lists one specimenkept in the Botanical Museum Stockholm (S) and collected from the Azores collected by Persson in 1937 on Terceira. Another specimen was collected by G. Schwab in Caldeiras, Sao Miguel, in 1981 at 200 m altitude. Additonal unrevised literature records are from Faial and Flores.
Campylopus cygneus (Hedw.) Brid. A Caribbeanspeciesoccurring on Cuba, Jamaica,Haiti, Hispaniola, Puerto Rico and the LesserAntilles (Frahm 1991). On the Azores found on seven of nine islands (Faial, Flores, Pica, Sao Jorge, Sao Miguel, Santa Maria, Terceira).
Campylopus flaccidus Ren. & Card A specieswidespread in tropical Africa (Frahm 1985a). From the Azores it was-according to Eggers(1982)-only known from Sao Miguel but later collected on Pica (comm. Schumacker).
Campylopus flexuosus Hedw. This speciesoccurs in western Europe, in North America in a single locality in the Appalachian Mts. and a few localities in British Columbia and is additionally widespreadin tropical montane regions in Central and South America and Africa. It is widespreadon the Azores (Faial, Flores, Pica, Sao Jorge, Sao Miguel, Santa Maria and Terceira).
Campylopus fragilis (Brid.) B., S. & G. It has a range comparableto C. flexuosus in western Europe, in North America at a single locality in Arkansas and somelocalities in British Columbia and the tropical mountainsof Central and South America and Africa. In the Azores it is quite widespread and was so far found on Faial, Flores, Pica, Sao Jorge, Sao Miguel and Terceira.
A survey of the CumpyEopus species from the Azores
149
Campylopus incrassatusC. Miill A speciesranging from New Zealand, Australia, South Africa to Patagoniaand many subantarctic andAtlantic islands(Ascension, Madeira, Azores). In the Azores it was only known from Sao Jorge but was recently collected on Pica and Terceira (comm. Schumacker). Campylopus pilifer Brid. Like C. flrxuosus and C. fragilis widespreadin the tropical mountains of Africa and America and also in SW Europe and SE North America, and like the preceding speciesalso widespread on the Azores (Faial, Flores, Graciosa, Pica, Sao Jorge, Sao Miguel, Santa Maria, Terceira). Campylopus pyriformis (Schuttz.) Brid. Distributed through the subantarctic to subtropical regions of the southern hemisphere and found also in Central Africa, western Europe and recently also in SE North America. On the Azores it was known from Flores, SaoMiguel and Terceira. An additional unpublishedrecord wasmadeby G. Schwab on Faial (SchwabSN 270, 17.6.81, hb. Frahm, BONN). The speciesis found in two varieties: var. pyriformis consists of small plants up to I cm high and grows preferably on rotten wood (of Culcita, Erica, Juniperus), var. azoricus (Mitt.) Corley consistsof large plants severalcm high, which are always sterile. Such plants are regarded as modifications from wet habitats. Campylopus shawii Wils. A specieshighly disjunct between the oceanic parts of Britain, the Azores and the Caribbean (Jamaica, Cuba, Dominican Republic), from where it was known as C. brittonae Williams and C. under-woo&i Williams (Frahm, 1985b). This kind of disjunction is hard to explain. It could be a relatively young introduction by man or long distance dispersalacrossthe Atlantic Ocean. The presencein Britain might also indicate a relict from the Tertiary, which has survived glaciation periods under a mild oceanic climate. This hypothesis is supportedby the presenceof C. setifolius Wils. in Britain, a very closely related species occurring on wet rocks instead of swamps and being differentiated by smaller ventral hyalocysts in transversesection of the costa and a serrate leaf apex. If C. set@diusis in fact endemic to Britain, it may have been derived from C. shawii by occupying another ecological niche. In the Azores it is known from Faial, Sao Miguel and Terceira. The Campylopus speciesof the Azores can be grouped into the following phytogeographical elements: Caribbean: cygneus African ampliretis European: brevipilus Subantarctic: incrassatus Tropical montane: pilifel; jexuosus, fragilis Disjunct Caribbean-Oceanic Europe: shawii Disjunct Subantarctic-W Europe: pyriformis
150
J.P. Frahm
Most ot‘ the speciesof Campylopus recorded from the Azores (five out of nine) are known from the Caribbean (C. cygneus, shawii, fragilis, pilifel; jlexuosus) and therefore the Caribbean Islands seemto be the main source for the colonization of these young volcanic islands. Subantarctic elements account for two species (incrassatus, pyriformis), African and European (C. jlaccidus and C. brevipilus) account for two more species. The distribution on the different islandsof the Azores (basedon Eggers (1982), completedby Sjogren (1990, 1993) and additional unpublishedrecordsby G. Schwab and R. Schumacker) is shown in the table.
Corvo
Faial
Graciosa
Pica
Sao Jorge
X
X
?
Sao Miguel
brevipilus
X
Campylopus
cygneus
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
,jlacciduJ
Campylupus
,jexuosw
Cumpylopus,frugilis
X
Campylupus
incrawurus
Campylopus
pilifer
X
Campylopu.~
pyr(formis
X
Campylopus
shawii
6
X X
X
6
5
X
X X
X
X
2
X
X
8
X X
X
5
Terceira
X
X
6
Santa Maria
X
Campylopus
Camp~~lopus
?
Flares
X
3
R
Eight out of nine speciesare found on Sao Miguel and on Terceira. Faial, Corvo and Pica have six, Flores and Sao Jorgehave five species,Santa Maria three and Graciosa only two. It is conspicuous that the specieson the islands with few speciesnumbers belong to widespread species. Rare species are found on islands with high species numbers. The species with the widest ranges are most widespread on the Azores. Campylopus pilijk was collected on all islands, on which Campylopus species occur, C.$exuosus and C. fragilis on eight out of nine. The Caribbean speciesare in part widespread(C. cygneus: eight of nine islands), in part rare (C. shawii: three out of nine islands).The rarest speciesare C. JIaccidus (two islands),followed by C. incrassatus and C. brevipilus (three islands) and C. pyrtformis (five islands). However, the floristic exploration might still be insufftcient, as expressedby the fact that Eggers (1982) listed no speciesof Campylopus from the island of Corvo, but a few years later Sjijgren (1993) listed six species. For the Cape Verde Islands, a distinct correlation was stated between the elevation and the speciesnumber of bryophytes (Frahm et al., 1996). In the case of the Azores, this trend is not very distinct (Fig. 1) with regard to the genus Campylopus. In contrast, the correlation between the area of the islands and species numbers of Campylopus (Fig. 2) is more pronounced, with the exception of the island of Corvo. Acknowledgements. I would like to thank R. Schumacker for suggestions and additions to the manuscript. E. Sjiigren provided valuable comments and literature.
A survey of the
151
species from the Azores
Campylopus
r-- ~ ; :-- -~~
elevation
-.*.-
species
(m) number
,
2500
- 7 2000
7 6 \
i
‘,S,
r
. 5
1500
1 4 1 3
1000
2 500 1 0
Fig. 1. Correlation their elevation.
between the species numbers of
Campylopus
on the islands of the Azores and
Cumpylopus
on the islands of the Azores and
600t 500 1~ 400 i j
Fig. 2. Correlation their
size.
between the species numbers of
J.P. Frahm
152
REFERENCES CORLEY
M.F.V., 1976 - The taxonomy of Campylopus pyriformis (Schultz) Brid. and related species. Journal of Bryology 9: 193-212. DA CUNHA A.G., DE BARROS G., 1942 - Algunas especies de musgos da Terceira nova para OS Acores ou para a ilha. Boletim da Sociedade Portuguesa de Ciencias Naturais 13: 156-157. EGGERS J., 1982 - Artenliste der Moose Makaronesiens. Cryptogamie, Bryologie-Lichhologie 3 (4):
FRAHM
283-335.
J.-P., 1975 -
FRAHM FRAHM
Taxonomische Notizen zur Gattung Campylopus. Revue bryologique et 41 (3): 321-332. J.-P, 1976 - Taxonomische Notizen zur Gattung Campylopus II. Revue bryologique et Zichenologique 42 (1): 603-616. J.-P, 1981 - Taxonomische Notizen zur Gattung Campylopus VIII. Herzogia 5: 517-529. J.-P., 1984 - Campylopus Brid. 1. Subg. Thysanomitrion (Schwaegr.) Kindb. emend. J.-P. Frahm. Nova Hedwigia 39: 585-621. J.-P., 1985a - Afrikanische Campylopus-Arten. Bryophytorum Bibliotheca 31: I-216. J.-P, 1985b - The world-range of Campylopus shawii Wils. Journal of Bryology 13:
FRAHM
J.-P, 1985~ -
lichenologique
FRAHM FRAHM FRAHM
359-367.
Taxonomische
Notizen zur Gattung Campylopus
XIV. Nova
Hedwigia
41:
273-277.
FRAHM
J.-P., 1987 -
A revised list of the Campylopus species of the world. Bryologische I-117. J.-P, 1991 - Dicranaceae: Campylopodioideae, Paraleucobryoideae. Flora Neotropica Monograph 54, 238 pp., New York. J.-P & GEISSLER P., 1985 - Lectotypification von Dicranum flexuosum Hedw. (Campylopus jlexuosus [Hedw.] Brid.) Cryptogamie Bryologie-Lichenologie. 6 (3): Beitrage
FRAHM FRAHM
7:
287-290.
FRAHM
J.-P., LINDLAR A., SOLLMAN P., FISCHER E., 1996 - Bryophytes from the Cape Verde Islands. Tropical Bryology 12: 123-154. KINDBERG N., 1888 - Enumeratio bryinearum exoticarum quam alphabetice disposuit N. Conr. Kindberg. Linkoping. MAGILL R.E., 1981 - Flora of South Africa, Bryophyta, Part 1 Mosses, Fast. 1 Sphagnaceae Grimmiaceae. Pretoria. SIM T.R., 1926 -- The Bryophyta of South Africa Transactions of the Royal Society of South Africa
SJGGREN
I 5.
E., 1978 Broteriana
26:
Bryophyte
vegetation in the Azores Islands. Memorias
de Sociedade
l-273.
SJGGREN E., 1990 - Bryophyte flora and vegetation on the island of Graciosa (Azores), with remarks on floristic diversity of the Azorean Islands. Arquipelago, Life and Earth Sciences
SJGGREN
8: 63-96.
E., 1993 - Bryophyte flora and vegetation on the island of Corvo (Azores). Arquipclago, L$e and Marine Sciences 1IA: 17-48. THERIOT I., 1939 - Campylopodioidtes des Iles Acores recolt&es par P. Allorge and H. Persson. Revue brvologique et lichenologique I 1: IO@-109.