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re-evaluation of tortella - Missouri Botanical Garden

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in the lamina, the var. fragilifolia is usually broken at<br />

the leaf tip.<br />

Dixon (1924) was also not su<strong>re</strong> <strong>of</strong> the<br />

distinction <strong>of</strong> Tortella fragilis from T. tortuosa. He<br />

would have liked to have been definite but, "I have,<br />

however, <strong>re</strong>ceived specimens from two or th<strong>re</strong>e<br />

localities in Labrador showing a distinct approach to [T.<br />

tortuosa]; the leaves on some <strong>of</strong> the plants, and even the<br />

lower leaves on some stems in which the distal ones a<strong>re</strong><br />

quite typical, being strongly contorted when dry, crisped<br />

and undulate at the margin, and when moist slightly<br />

flexuose and undulate; hardly, indeed, to be <strong>re</strong>cognised<br />

from the var. fragilifolia...except by the leaves<br />

somewhat firmer, the very shining nerve and the<br />

p<strong>re</strong>sence (usually) <strong>of</strong> some rigid and less curved<br />

leaves." He speculated that T. fragilis would be found to<br />

be close to T. tortuosa "by intermediate forms."<br />

Polunin (1947), when <strong>re</strong>porting Tortella<br />

fragilis for the Canadian Eastern Arctic, quoted the<br />

same lines from Dixon (1924), adding "This puzzling<br />

situation that Mr. Dixon has so ably elucidated is to be<br />

found also in the specimens from Baffin Island"<br />

although some <strong>of</strong> these specimens, upon examination,<br />

may turn out to be T. tortuosa var. arctica.<br />

Tortella tortuosa var. fragilifolia in North<br />

America appears to be a montane and bo<strong>re</strong>al taxon. The<br />

var. fragilifolia in North America is appa<strong>re</strong>ntly not in<br />

montane elevations much south <strong>of</strong> the latitude <strong>of</strong> the<br />

Canadian border with the conterminous United States.<br />

TORTELLA NITIDA IN NORTH AMERICA<br />

Sources <strong>of</strong> North American <strong>re</strong>ports <strong>of</strong> Tortella nitida<br />

perhaps began with Renauld and Cardot (1889: 99) (as<br />

Trichostomum nitidum) "United States, without locality,<br />

collected by James, communicated by Mr. Besche<strong>re</strong>lle."<br />

Barnes (1897: 275–276) next <strong>re</strong>ported the<br />

species, quoting Renauld and Cardot and basing his<br />

description on Limpricht (1890, 1: 581) whe<strong>re</strong> the<br />

description <strong>of</strong> the proximal cells indicates their being<br />

"less sharply marked <strong>of</strong>f from chlorophyllose cells."<br />

Assuming James, <strong>of</strong> the Renauld and Cardot quote, to<br />

be Thomas Potts James (1803–1882), a <strong>re</strong>quest was<br />

made to the Farlow Herbarium for a specimen that<br />

might cor<strong>re</strong>spond, but without success. A James<br />

specimen at NY collected by him in Canada, July 1850,<br />

was Tortella tortuosa var. tortuosa.<br />

Next, the<strong>re</strong> is the exsiccat specimen in the<br />

Canadian Musci by John Macoun, number 605, labeled<br />

as Barbula nitida. In 1892, Macoun and N. C. Kindberg<br />

published the label data (number 196, p. 52). Kindberg<br />

(1897) affirmed its occur<strong>re</strong>nce ("Can.: Macoun.),<br />

p<strong>re</strong>sumably from Macoun's exsiccat number 605.<br />

Macoun 605 is labeled as Barbula nitidum [sic] (Lindb.)<br />

Jur., <strong>of</strong> the Canadian Musci series, "on wet earth at<br />

Hector, Rocky Mountains, B.C. [British Columbia,<br />

Canada], Aug. 14th, 1890" and is Tortella tortuosa var.<br />

fragilifolia (one specimen at FH, two others at NY). It is<br />

as tall as the typical variety gets, has a broad stem<br />

section but no central strand. Some apices a<strong>re</strong> deciduous<br />

and the<strong>re</strong> is conspicuous fragility <strong>of</strong> the lamina. The<br />

24<br />

crucial juxtacostal bistratose character <strong>of</strong> the leaf<br />

cross section is p<strong>re</strong>sent. The cells <strong>of</strong> the lamina a<strong>re</strong><br />

unusual for the species, on some leaves <strong>re</strong>gularly ca. 14<br />

µm, some even to 17 µm in diameter, with distal<br />

proximal cells thick-walled and porose. Some leaves on<br />

the same stem have adaxial quadrate cells covering the<br />

costa nearly to the apex, in others the adaxial surface <strong>of</strong><br />

the costa is exposed in a narrow groove from base to<br />

apex.<br />

Subsequent <strong>re</strong>ports <strong>of</strong> Tortella nitida include<br />

Haring's (1938) t<strong>re</strong>atment in Grout's moss flora <strong>of</strong> North<br />

America, Flower's t<strong>re</strong>atment for the state <strong>of</strong> Utah<br />

(1973), Ketchledge's publication on the moss flora <strong>of</strong><br />

New York State (1957), Stee<strong>re</strong> and Scotter's (1978)<br />

<strong>re</strong>port for the Nahanni P<strong>re</strong>serve, in the Mackenzie<br />

District <strong>of</strong> the Northwest Territories, and the <strong>re</strong>cent<br />

checklist <strong>of</strong> Canadian mosses (I<strong>re</strong>land et al. 1987).<br />

Haring (1938) cited Macoun's number 605 in<br />

the Canadian Musci as Tortella nitida; also a specimen<br />

from Montana in NY, and one by Homer House, a<br />

specimen from Harris Lake, New York. Haring and<br />

perhaps Grout, we<strong>re</strong> strongly influenced in their<br />

expectations by the Macoun specimen. This was at a<br />

time when it was not yet accepted that some <strong>of</strong> Macoun<br />

and Kindberg's determinations we<strong>re</strong> perhaps doubtful.<br />

Kindberg's determinations <strong>of</strong> Macoun's specimens<br />

"we<strong>re</strong> <strong>of</strong>ten hasty and consequently wrong and<br />

Macoun's own determinations we<strong>re</strong> <strong>of</strong>ten based on<br />

superficial study and thus a<strong>re</strong> open to doubt" (Sch<strong>of</strong>ield<br />

1965), and yet Besche<strong>re</strong>lle and Dixon also made similar<br />

independent determinations.<br />

Examination <strong>of</strong> the specimen <strong>of</strong> Tortella nitida<br />

that Haring cited for Lake Harris (New York, Essex Co.,<br />

Newcomb, surface <strong>of</strong> boulder on sho<strong>re</strong> (perichaetiate:<br />

without sporophytes), July 10, 1927, H.D.House (NYS),<br />

shows this specimen also to be Tortella tortuosa var.<br />

fragilifolia. The Harris Lake specimen was fragile in<br />

some stems, but on the whole seems mo<strong>re</strong> typical <strong>of</strong> T.<br />

tortuosa than most examples <strong>of</strong> the variety fragilifolia<br />

(it had no stem central strand). The leaves on some<br />

stems, however, we<strong>re</strong> mo<strong>re</strong> broadly lanceolate with<br />

larger distal laminal cells (to 12 µm). Some <strong>of</strong> the leaf<br />

cross sections showed some bistratose a<strong>re</strong>as near the<br />

costa in the distal median portion <strong>of</strong> the leaf.<br />

The Harris Lake specimen showed no<br />

annotation by Haring or Grout. The<strong>re</strong> was a query on<br />

the label, and the note, "Doubtful !E.H.K" [Edwin H.<br />

Ketchledge] written on the packet. Ketchledge's<br />

subsequent publication on the moss flora <strong>of</strong> New York<br />

State (1957) listed Tortella nitida for the New York<br />

flora, but in the <strong>re</strong>vised edition (Ketchledge 1980), the<strong>re</strong><br />

was no citation <strong>of</strong> that species.<br />

Another specimen <strong>of</strong> the var. fragilifolia, from<br />

New York on limestone pavement in Jefferson Co. in<br />

association with Tortella rigens, is cited below in the<br />

t<strong>re</strong>atment <strong>of</strong> that species.<br />

Haring's third <strong>re</strong>p<strong>re</strong>sentative specimen was<br />

from Montana at NY. A specimen from Montana<br />

collected by R. S. Williams, 329 at NY was determined<br />

he<strong>re</strong> to be Tortella alpicola (see discussion <strong>of</strong> this

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