A New Species of () from Western North America and a Regional Key to the Taxa Author(s): Richard H. Zander Source: The Bryologist, Vol. 102, No. 1 (Spring, 1999), pp. 112-115 Published by: American Bryological and Lichenological Society Stable URL: http://www.jstor.org/stable/3244470 Accessed: 09-06-2015 12:52 UTC

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This content downloaded from 192.104.39.2 on Tue, 09 Jun 2015 12:52:27 UTC All use subject to JSTOR Terms and Conditions The Bryologist 102(1), pp. 112-115 Copyright ? 1999 by the American Bryological and Lichenological Society, Inc.

A New Species of Didymodon (Bryopsida) from Western North America and a Regional Key to the Taxa

RICHARD H. ZANDER Clinton Herbarium, Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, NY 14211, U.S.A.

Abstract. Didymodon norrisii Zand. is a new species of found in California and Oregon, U.S.A., belonging to Didymodon sect. Rufidulus (Chen) Zand. It differs most saliently from con- geners by the combination of the deep brick or rose red color in KOH, costa covered by quadrate epidermal cells nearly to the leaf insertion on both sides, and apparent asexual reproduction through fragile branch apices. The new species is included in a key to the North American (north of Mexico) taxa.

The genus Didymodon Hedw. (Zander 1993) has (BUF, holotype; uc, isotype). U.S.A. OREGON.Jackson Co., near Siskiyou Pass, state boundary, W. B. Schofield 23349, a reputation for difficult , and it is a plea- May 23, 1964 (uBC, paratype). sure to describe as new to science a very distinctive taxon from California and Oregon of the United growing in a tight turf, lower stems often States. The new species is easily identified by the denuded of leaves, red-brown above, tan below. combination of the following traits: leaf base re- Stems 1.0-1.5 cm long, branching often; rounded- flexed as a short collar, proximal lamina cells not pentagonal in transverse section, hyalodermis ab- or weakly differentiated, epidermal cells of costa sent, sclerodermis present, thick, diameter of cen- from to near the leaf insertion quadrate apex very tral cylinder cells 15-20 Cjm in diameter, central on both branch and often fall- sides, apices fragile strand present, strong; rhizoids few; axillary hairs as a means of asexual en, probably reproduction, three-celled, basal cell brown. Cauline leaves ap- and leaves a brick or rose red color in 2% evincing pressed when dry, spreading when moist; ovate to KOH solution. ovate-lanceolate, reflexed above very short-sheath- ing basal collar, adaxial surface broadly concave DIDYMODON NORRISIIZand. sp. nov. FIGS. 1-11 across leaf and sometimes narrowly channeled near apex, 1.2-1.5 mm long; base broadly ovate, sheath- Plantae in caespitem arctum crescentes; caules inferni dillute brunneoli saepe foliis carentes, superni rubro-brun- ing at insertion; margins recurved in lower 2/3 to ?, nei. Caules 1.0-1.5 cm longi, identidem ramosi, hyalod- minutely crenulate by projecting walls; apex acute ermide carentes, sclerodermide atque filo centrali crasso, to short-acuminate, often broken off; costa strong, valdo axillares cellula basali praediti; pili tricellulares, percurrent, weakly spurred, adaxial outgrowths brunnea praediti. Folia in statu sicco appressa, madido pa- adaxial cells from to near tentia, ovata vel ovato-lanceolata, supra collarem basalem none, quadrate apex very brevi-vaginantem reflexa, trans superficiem adaxialem late insertion, in 6-10 rows, abaxial cells quadrate in concava, 1.2-1.5 mm longa; basis late ovata, ad inserti- distal 4/5 of leaf, transverse section rounded-elliptic, onem vaginans; margines in partem 2/3-? inferiorem re- adaxial epidermis present, adaxial stereid band ab- curvae, per parietes cellulares projectos crenulatae; apex sent, guide cells 4 + 1-2, in 2 layers, hydroid acutus vel brevi-acuminatus, saepe abrumpens; costa val- da, percurrens, cellulis adaxialibus e apice usque ad in- strand present, abaxial stereid band present, small, sertionem quadratis in 6-10 seriebus adaxialiter dispositis substereid, rounded in sectional shape, abaxial epi- et cellulis abaxialibus in partem distalem 4/5folii quadratis dermis present; proximal cells not differentiated or et strato stereidarum unico praedita; cellulae proximales of 2-5 rows of slightly larger cells, quadrate to haud distinctae vel series 2-5 cellularum parum majorum formantes; cellulae mediales distales rotundato-quadratae short-rectangular, to 12 Cjm wide, 1(-1.5):1, walls vel hexagonae, 9-13 psmlatae, 1 : 1(-2), in paginis am- of proximal cells evenly thickened; distal medial babus libera tumescentes, saepe prope margines foliares cells rounded-quadrate to hexagonal, 9-13 jLm inferiores transverse lamina ca 15-20 jm cras- elongatae; wide, 1:1(-2), bulging on both free sides, often sa; papillae e 2-4 semiglobosae eminentiae per cellulam near lower leaf lam- compositae, quum absentes tum cellulae in aspectu late transversely elongate margins, mamillosae videntur; parietes cellulares aeque incrassati, ina ca 15-20 jLm thick, papillae of 2-4 hemispheric in paginis ambabus laminae tumescentes. Apices foliares salients per cell or essentially absent and cells ap- caulini foliosi fortasse ad asexualiter atque apices fragiles pearing broadly mamillose, walls evenly thickened, Steriles. Sporophyta visa. Plantae in reproducens. non on both sides of lamina. Asexual lamina KOH latericiae vel roseae reagentes. bulging reproduc- TYPE: U.S.A. CALIFORNIA.Humboldt Co., near Knee- tion of fragile leaf apices and foliose stem tips, pos- land, Sec. 22, R2E, T4N, D. H. Norris 7884, Oct. 8, 1967 sibly serving for asexual reproduction. Sexuality:

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FIGURES 1-11. Didymodon norrisii. - 1. Habit. - 2. Habit with fallen stem tips. - 3-7. Five leaves. - 8. Leaf apex. - 9. Leaf base. - 10-11. Transverse sections at midleaf. - Scales: a = 1 mm; b = 0.5 mm; c = 50 pm. sterile. Sporophytes unknown. KOH laminal color leaf cells, but differ in the sharply broadened leaf reaction deep brick or rose red. base, sinuose costa distally, rather flat distal laminal Habitat.--field, dry wash, dry sunny rocks, boul- cell surfaces, and distinct multiplex papillae. der, 700-1,500 m. Didymodon norrisii apparently belongs with Di- The new species is named for Daniel H. Norris, dymodon sect. Rufidulus (Chen) Zand., and is the of Hansville, Washington, and formerly of Hum- fourth member of this section for the New World. boldt State University, Arcata, California, an inde- The section, according to Chen (1941), is distin- fatigable bryophyte collector and fine systematist guished among other characters by the narrow cos- with considerable experience in the . ta, which ends before the apex, and distal laminal Species in the genus Bellibarbula Chen are sim- cells that. are mamillose-papillose. In the new spe- ilar in the red color in KOH and quadrate proximal cies, however, the costa is rather thick and is usu-

This content downloaded from 192.104.39.2 on Tue, 09 Jun 2015 12:52:27 UTC All use subject to JSTOR Terms and Conditions 114 THE BRYOLOGIST [VOL. 102 ally percurrent, and the distal leaf cells are multi- leaves present on fragile branchlets or portions of some stems----Didymodon subandreaeoides papillose over a strongly bulging free laminal sur- 7. Leaves monomorphic------8 face. It further differs from the three Asian species 8. Plants often fruiting, leaf apices acute, pro- of the section by the little-differentiated proximal pagula absent- ---Didymodon nigrescens cells. Didymodon nigrescens (Mitt.) Saito, found in 8. Plants sterile, leaf apices obtuse, clusters of coastal regions of northwestern North America unicellular propagula in the leaf axils (Zander 1978) as well as central and eastern Asia, Didymodon perobtusus 9. Costa with elongate superficial ventral cells 10 is also red in but differs from the new deep KOH, 9. Costa with quadrate or occasionally short-rectan- species in having thick-walled stem central cylinder gular superficial ventral cells, or, if elongate, then cells, crowded and lanceolate leaves, leaf bases not distal laminal cells bistratose------16 strongly reflexed, rectangular basal cells, a thin cos- 10. Leaf base auriculate or weakly winged at in- sertion, often ta only two cells in width distally, and the hydroid apex whip-like, very long-acu- minate Didymodon leskeoides strand absent from the costa. Given the and bulging 10. Leaf base------...... gradually or quickly narrowed to massively unipapillose cells of D. nigrescens and the insertion, not flaring, apex obtuse to acu- its two close relatives D. subandreaeoides (Kindb.) minate------11 Zand. and D. perobtusus Broth., the crenulations on 11. Leaves ovate to long-elliptic, apex often obtuse, costa often before the distal leaf margin, and the unusually thin costae ending apex of the these three should foregoing species, species 11. Leaves short-to long-lanceolate, apex always be certainly be recognized along with D. norrisii in acute, costa subpercurrent to short-excurrent 12 sect. Rufidulus. 12. Plants with axillary gemmae, leaves mostly The absence of a costal adaxial stereid band, and 0.9-1.1 mm long, catenulate when dry the guide cells occurring in two layers are further -Didymodon michiganensis 12. Plants in leaf axils, leaves distinctive characters that to relate the new lacking gemmae appear usually 1.2-5.0 mm long, appressed-incurved species to Didymodon vinealis (Brid.) Zand. and its to weakly spreading when dry----- 13 relatives, but those species lack the strongly mam- 13. Leaves when moist spreading to weakly recurved, illose cells. The most similar of the D. vinealis- usually lying flat, costa usually distinctly widened at base------14 related species, D. nevadensis Zand. (Zander et al. 13. Leaves when moist recurved and keeled, differs in the revolute and strongly 1995) strongly margin lying on their sides, costa little widened at base reproduction by axillary gemmae, although leaf 15 shapes and costal sections are alike. The following 14. Leaves 0.8-1.7(-2.5) mm, acuminate, distal key is modified from that of Zander (1998). cell walls little thickened or irregularly thick- ened and lumens angular, trigones absent or weakly developed ---- Didymodon fallax KEY TO DIDYMODONIN NORTH AMERICAINCLUDING 14. Leaves usually 2.0(-3.5) mm, distal cell walls MEXICO irregularly thickened and trigonous, trigones as large as the lumens or nearly so 1. Leaf apices caducous or fragile---- 2 Didymodon giganteus 1. Leaf apices intact or only occasionally broken 5 15. Stems to 2.5 cm, leaves usually 0.8-2.0 mm long .. 2. Leaf apices not swollen, usually evenly nar- Didymodon ferrugineus rowing------3 15. Stems usually more than 3 cm, leaves mostly 2.0- 2. Leaf swollen a 4 apices apically as propagulum ___ 2.5 mm long ------Didymodon maximus 3. Cells of leaf smooth apex Didymodon rigidulus 16. Leaves ventrally with a narrow medial chan- (var. icmadophilus and var. subulatus) nel about the width of costa at least at leaf 3. Cells of leaf apex weakly conic-mamillose apex, apex mostly apiculate by one or more Didymodon sinuosus conic cells, costa 4. Distal laminal cells 13-15 jim wide usually percurrent, margins usually recurved, often to near apex, laminal Didymodon johansenii color reaction to KOH oc- 4. Distal laminal cells 8-10 jm wide usually brick-red, casionally orange------17 ------Didymodon anserinocapitatus 16. Leaves channeled me- 5. Leaves not keeled, not highly recurved, margins ventrally very widely finely crenulate by bulging cell walls--- 6 dially or merely slightly concave across leaf, 5. Leaves sometimes keeled or highly recurved, mar- apex seldom apiculate by a conic cell, costa gins usually entire or dentate but not minutely percurrent or excurrent as a multicellular, crenulate------9 stout mucro, margins plane to recurved below 6. Plants in nature brick or rose red, costa thick, midleaf, laminal color reaction to KOH usu- 6-10 cells wide above midleaf, laminal pa- ally negative, yellow or orange, seldom brick- pillae absent or hemispheric and several per red 21 lumen------Didymodon norrisii 17. Entire leaf------.. strongly both reflexed backwards and 6. Plants in nature red- to black-brown, costa strongly keeled when moist, papillae when present thin, 2-3 cells wide above midleaf, laminal simple, stem central strand usually absent papillae absent or low, massive and lens- Didymodon asperifolius shaped------7 17. Entire leaf spreading to weakly reflexed back- 7. Leaves dimorphic: cochleariform, epapillose wards and weakly keeled when moist, papillae

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when present bifid to multiplex, stem central leaf cells quadrate strand present------18 Didymodon rigidulus (var. icmadophilus) 18. Leaves deltoid to short-lanceolate, to 1.5 or 27. Distal lamina entirely or mostly bistratose 28 rarely to 2.0 mm long, margins recurved or 27. Distal lamina bistratose along margins----29 revolute to near apex, propagula sometimes 28. Peristome teeth long-triangular; leaves long- present------19 triangular to ovate-lanceolate, distal cells het- 18. Leaves short- to long-lanceolate or long-tri- erogeneous in size angular, to 4.0 mm long, margins recurved Didymodon incrassatolimbatus near base or up to lower 2/3of leaf, propagula 28. Peristome teeth rudimentary, elliptic, leaves rare------20 long-lanceolate, distal cells homogeneous in 19. Costa often with an apical conic cell, costa grad- size Didymodon rigidulus (var. subulatus) ually narrowing distally, ventral surface nearly flat 29. Proximal laminal cells with firm, weakly to and not forming a wide pad of cells (but costa strongly thickened walls, differentiated usually occasionally thickened and bulging ventrally), only medially ------30 guide cells in 1 layer, leaf margins recurved to 29. Proximal laminal cells thin-walled and usually tightly revolute, gemmae often present in leaf ax- somewhat inflated, often bulging-rectangular, dif------ils------Didymodon brachyphyllus ferentiated across leaf base 31 19. Costa usually lacking an apical conic cell, costa 30. Leaves long-ligulate to lanceolate wider at midleaf than below, with bulging ventral D. rigidulus (var. rigidulus) 30. Leaves short-ovate surface forming a long-elliptic unistratose pad of D. revolutus cells, guide cells in 2(-3) layers, leaf margins 31. Leaves smooth or loosely revolute, tubers occasional on rhizoids - long-lanceolate, usually weakly Didymodon nevadensis papillose, marginal proximal cells narrowly rect- in 2-4 ventral cells of 20. Leaves long-ovate to broadly lanceolate, apex angular rows, superficial costa stem with blunt to broadly acute, distal marginal cells usually elongate, hyalodermis __ umbrosus bistratose marginally, throughout or or occa- -Didymodon 31. Leaves short-lanceolate, smooth to sionally in patches, laminal color reaction to strongly papil- lose, cells not or dif- KOH deep yellow or orange-brown to red- marginal proximal weakly ferentiated from the medial, ventral brown------Didymodon nicholsonii superficial cells of costa quadrate, stem lacking hyalodermis 20. Leaves triangular to narrowly lanceolate, or this weakly differentiated---D. australasiae apex narrowly acute, distal marginal cells uni- stratose or occasionally bistratose in patches, laminal color reaction to KOH deep red to The authorities for taxa in the key are given by Zander red-brown------Didymodon vinealis (1993). 21. Axillary gemmae present------22 21. Axillary gemmae absent------24 ACKNOWLEDGMENTS 22. Propagula all multicellular, leaf apex acute Didymodon rigidulus (var. rigidulus) The author appreciates the loan of specimens by the 22. Propagula mostly unicellular, leaf apex broad- curators at ALA,NY, and UBC.Patricia M. Eckel contributed ly obtuse------23 the Latin description and illustration. Lloyd Stark and an reviewer comments. 23. Costa narrow, usually 2 cells wide at midleaf, not anonymous provided helpful strongly spurred-----Didymodon perobtusus 23. Costa broad, 4-6 cells wide at midleaf and often LITERATURECITED strongly spurred by rows of lateral cells ending CHEN, 1941. Studien uiber die ostasiatischen Arten abruptly in the lamina- --Didymodon revolutus P.-c. der Pottiaceae, Hedwigia 80: 1-76; 141-322. 24. Distal lamina unistratose or bi- I-II. occasionally ZANDER,R. H. 1978. New combinations in Didymodon stratose in small ------25 patches (Musci) and a key to the taxa in North America north 24. Distal lamina bistratose totally or just along of Mexico. Phytologia 41: 11-32. margins------27 1993. Genera of the Pottiaceae: of Harsh 25. Plants flagellate, leaves strongly appressed when Environments. Bulletin of the Buffalo Society of Nat- dry, linear-lanceolate, costa long-excurrent ural Sciences 32: vi + 378 pp., 113 pls. .. Didymodon rigidulus (var. ditrichoides) S1998. A phylogrammatic evolutionary analysis 25. Plants not flagellate, leaves appressed-incurved to of the moss genus Didymodon in North America North weakly twisted and weakly spreading when dry, of Mexico. Bulletin of the Buffalo Society of Natural short- to long-lanceolate, costa short- to long-ex- Sciences 36: 81-115. ------current 26 -, L. R. STARK& G. MARRS-SMITH.1995. Didy- 26. Leaf base rectangular and gradually widened, modon nevadensis, a new species for North America, costa short-excurrent, unbroken, proximal with comments on phenology. THE BRYOLOGIST98: leaf cells short-rectangular 590-595. ----Didymodon rigidulus (var. gracilis) 26. Leaf base ovate and often abruptly widened, costa long-excurrent, often fragile, proximal ms. submitted June 24, 1998; accepted Nov. 5, 1998.

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