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Taxonomy Campylopus Introflexus and C. Pilifer

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Taxonomy Campylopus Introflexus and C. Pilifer Taxonomy and World Distribution of Campylopus introflexus and C. pilifer (= C. polytrichoides ): New a Synthesis1 2 2 S.R. GradsteinI AND H.J. M. SipmanI Abstract. introflexus Brid. lato Campylopus (Hedw.) sensu comprises two widespread, closely related species: tropical and warm-temperate C. pilifer Brid. (= C. polytrichoides DeNot.) and temperatesouthern hemispheric C. introflexus which introduced Main (Hedw.) Brid., was recently in Europe. in the the dorsal lamellae the in differences are height of of leaves, spore size and in seta length. In C. pilifer lamellae are more pronounced in tropical mountains than in lowland An extreme with lamellae to areas. form up seven cells high is C. pilifer var. lamellatus (Mont.) comb. nov. from Bolivia. The recent invasion of Campylopus introflexus (Hedw.) Brid. in Europe (Prahm, has renewed the of this 1972) interest in taxonomy species. Traditionally it was con- sidered an almost cosmopolitan species until Giacomini (1955) showed that two species are at hand: C. introflexus s.str., occurring in temperateregions of the southern of and that hemisphere, large parts America (at time!) one locality in Europe (Bre- tagne), and C. polytrichoides DeNot., occurring in southwestern Europe, Africa and southern Asia. The characteristics used to distinguish the two species, particularly those of the have been gametophyte, questioned by several authors, e.g. Richards Barkman and Mabelis and (1963), (1968) Jacques and Lambinon (1968). Therefore a renewed ofthe ofthe investigation taxonomy species was undertaken, simultaneously, Frahm and the authors. Frahm’s studies confined by J.-P, (Duisburg) present were to characteristics of the gametophyte, whereas ours also included characteristics of the sporophyte. As a result of his investigations, Frahm (1974, 1975) showed that: 1. The only reliable differences the between the the in gametophyte two species are found in —the of the dorsal the number nerve height lamellae, of stereids in each group and the width oftheir lumen. These characteristics are best studied in transverse sections of the of upper part the leaf, near the center of the nerve. Leaves should be taken from just below the comal heads of the stems, since comal leaves (“Hochblätter”) of С. resemble leaves of The distribution of the polytrichoides may C. introflexus. 2. two Giacomini also the species as given by is incorrect since С. polytrichoides occurs in ‘This paper is dedicated to the memory of the late Dr. P. A. Florschiitz (f 1976), whose guidance and expert knowledge of Campylopus has been ofgreat help. We thank the of the in text the loan of For curators herbaria cited the for specimens. valuable informationwe express our gratitude to Dr. J. J. Barkman (Wijster), Dr. G. C. S. Clarke (London), Dr. A. C. Crundwell (Glasgow), Dr. E. Hegewald (Dortmund) and especially to Dr. J. -P. Frahm (Duis- close with burg) in cooperation whom the work was carried out. 2 Institute for Systematic Botany, State University of Utrecht, Heidelberglaan 2, Utrecht, Netherlands. 0007-2745/78/114-121$1,05/0 1978] GRADSTEIN & SIPMAN: TAXONOMY OF CAMPYLOPUS 115 Europe (2.5-)4.1(-5.2) (15-)30(-45)(15-)30M5) (50-)60(-70) (3.0-)4.5(-6.6) (2-)3(-4) (3-)4(-5) (14-)17(-19) 19(5) variation. SW geographical America pilifer (30-)50(-65) (2-)3(-7) (3-)4(-5.5) (10-)13(-16) 38(11)38(11) C. (1.8-)3.3(-5.6) (10-)35(-50) (30-)50(-65) (2.0-)4.1(-6.8) (10-)13(-16) showing Tropical , pilifer Africa C. (l-)2(-4) (4-)5(-9) 16(10) and Tropical (2.4-)3.7(-5.2)(23-)35(-45) (35-)55(-68) (2.6-)4.0(-5.2) (11-)15(-17) introflexus 2) temperateHemisphere (1.8-)3.3(-4.8)(30-)35(-60) (40-)55(-75) (2.6-)3.5(-5.2)(2.6-)3.5(-5.2) K-2)1(— (5-)6-7(-9)(5_)6-7(-9) (10-)12(-15)(10-)12(-I5) 17(10) Campylopus S troflexus of introflexus in C. 4.8) 2) Europe K-2) (20-)35(-55) (40-)55(-75) 1(— (5-)7(-10) (10-)12(-14) 22(7) W (2.2-)3.3(-4.8) (2.4-)3.6(-5.4)(2.4-)3.6(-5.4) characteristics W (2.2—)3.3(— significant cells) of of with a group)group) per (number hairpoint)“hairpoint) length) (number Measurements leaf number height brackets) of width) ’ 61 1. (without (% in leaf Examined of (average Table Base Lamellae, Diameter Length (% Length-Length" sporophytes mm. )im. in in Leaf Hyaline Nerve Stereids Dorsal Seta Spore Specimens - b 116 THE BRYOLOGIST [Volume 81 U.S.A. and Central and South America, whereas C. introflexus is restricted to the of the southern and and Central The temperate zone hemisphere West Europe. two species overlap in distributionin Brazil, Uruguay, Argentina, South Africa and, since recent times, in western Europe. Our studies, based on examination of over 150 collections from all over the world, the distributionof confirm Frahm’s conclusions on the two taxa. We also recognize the two taxa as species, although we find fewer gametophytic differences. Measurements of in significant characteristics are given Table I. Some “classical” characteristics, e.g. reflexion ofhairpoints, leaf auricles, widthof stereid lumina and length of the convo- luted ofthe omittedbecause be less also part leaf, are they proved to significant, as was shown by Frahm (1974). Table 1 shows that material assigned to C. introflexus (Hedw.) Brid.—from the temperate southern hemisphere and from western Europe—is morphologically rather uniform. The remaining material, assigned to C. pilifer Brid, (=C. polytrichoides DeNot.) is morphologically less uniform, but differs significantly from C. introflexus by the height of the dorsal lamellae, the length of the seta and the diameter of the summarized in the to Frahm spores. The differences are key given below. Contraiy (1974) we cannot accept characteristics derived from the stereids as diagnostic, since a considerable all is overlap among populations present. in section with of in Nerve transverse dorsal lamellae composed 2-4 cell rows (5-7 cell rows var. lamellatus comb, 3-5.5 in East Africa!); 12-19 (Mont.) nov.); seta mm (-9 mm tropical spores ¿rm and in diam. Tropical warm-temperate: North and South America, Europe, Africa, India C. pilifer Brid. transverse with of seta Nerve in. section dorsal lamellae composed l(-2) cell rows; 5-8(-10) mm 4 10-15 long (only mm in var. brachycarpa Giac.); spores ßm in diam. Temperate southern hemisphere, neophytic in West and Central Europe C. introflexus (Hedw.) Brid. Campylopus pilifer Brid. “In Insula Ischia ad 1806” “In Lectotype: Italy, rupes, Auguste (b). Syntype: France, Sylva prope Fontainebleau, Octobre 1807” (b). Synonyms (see Frahm, 1975, for complete listing): Dicranum capitiflorum P. Beauv., Prodr. 53. 1805. Type: “Bourbon” (Reunion), Bory St. Vincent (pc, in herb. L. C. Richard); nonCampylopus capitiflorus Mont., Syll. 42. 1856. Campylopus longipilus Brid., Bryol. Univ. 1: 447. 1826 nom. illeg. Campylopus polytrichoides DeNot., Syll. Muse. 222. 1838. cordobaensis Campylopus introflexus (Hedw.) Brid. var. Then, in Bauer, Musci Eur. Am. El exsicc. ser. 45, nr. 2204. 1934 syn. nov. Type: Argentina, Cordoba, Durazno, Hossaeus 149 (isotypes in BM, herb. Frahrn). Description and illustrations; See Grout (1937: 93-94), Giacomini (1955: 55, fig. 1-6), Gan- gulee (1971: 301-303, fig. 141), Frahrn (1974: fig. 6). Distribution (Fig. 2) and Ecology. —Campylopus« pilifer is widespread between 40°N and 35°S in North and South America, Africa, India and in Europe, where the has western-Mediterranean-Atlanticdistributionand reaches north species a to 57°N. Presumed occurrence in Indomalesia(cf. Gangulee, 1971, map LXXXVI) could not be confirmed. Records from the Netherlands proved to belong to C. introflexus (Hedw.) Brid. low eleva- (Sipman, 1977). In warm-temperate regions the species occurs at tions, whereas in tropical regions it is mainly found at higher altitudes, reaching to 3500 m in the mountains of Central Africa and the Andes of northern South America. in The species occurs regions with an, at least periodically, humid climate, on dry, It acid, poor mineral soil. often grows onthin soil covering granite or schistose rock, or in in bases oftrees rotten rock crevices, open woods, heath-islands, etc.; rarely on or GRADSTEIN & SIPMAN: TAXONOMY OF CAMPYLOPUS 117 117 Figure 1. Campylopus pilifer var. lamellatus (Mont.) comb. nov. Transverse section from of 200x the upper region leaf, ca. (from type collection). wood. Richards has According to (1963) C. pilifer a narrower ecological tolerance than C. introflexus in Britain and is more thermophytic. Nomenclature.—When reestablishing Campylopus polytrichoides as a species, Giacomini three older in its without at- (1955) placed species names synonymy paying tention to priority; Dicranum capitiflorum P.Beauv. 1805 from La Reunion and Brid. 1819 and C. Brid. 1826from have Campylopus pilifer longipilus Europe. We ex- amined type material of these binomials and found them to fit Giacomini’s concept of C. polytrichoides. material of Type Dicranum capitiflorum was found in the herbarium of L. C. Richard where (PC-Gen.), it was erroneously insertedunder Campylopus capitiflorus Mont, The plants are fairly robust, and fruiting stems have distinct comal heads with long, slightly decurved hairpoints, resembling those of C. introflexus. However, the cell 2-3 high lamellae, the 5-5.5 mm setae and, particularly, the spores (16-17 ¿im) indicate typical “C. polytrichoides.” Campylopus pilifer was described by Bridel (1819) on the basis oftwo collections, from one Italy and the other from France: “In insula Ischia 1806, et in saxosis sylvae Fontainebleau prope 1807 caespitibus densis pulchre variegatis crescentem sed sine theca inveni” (Mant. Muse., p. 72), He recognized a close relationship of his new from based species to Dicranum flexuosum var. piliferum Turn. 1804, Great Britain, from Wales and Ireland and on piliferous specimens (probably C.
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