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Pseudosymblepharis and Oxystegus Species (Pottiaceae) New to Northwestern North America, and Refutation of Molecular Chionoloma

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Pseudosymblepharis and Oxystegus Species (Pottiaceae) New to Northwestern North America, and Refutation of Molecular Chionoloma This is a distributable version of the paper with the same pagination. The original published paper is available on request from the authors or from The Bryologist journal. Pseudosymblepharis and Oxystegus species (Pottiaceae) new to northwestern North America, and refutation of molecular Chionoloma Richard H. Zander1 and Patricia M. Eckel Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, MO 63110 U.S.A. ABSTRACT. Three species of Pottiaceae (Bryophyta) of limited distribution worldwide are reported for hyperoceanic northwestern North America. The Asian Pseudosymblepharis angustata is a genus and species new to North America from Alaska. Oxystegus daldinianus, previously known from Europe and the Southern Appalachians in the U.S.A., is reported as new to Canada from British Columbia. A new species is described in Oxystegus from British Columbia. These genera continue to be recognized here instead of the broad molecular systematics concept of Chionoloma in part because of lack of resolution of molecular cladograms reported by a recent study on molecular races in the Pottiaceae. KEYWORDS. Alaska, British Columbia, dissilient genera, phylogenetics, macroevolutionary systematics, molecular races. The classification of Pottiaceae subfamily Chiono- reason to abandon them except that they contradict lomoideae R.H.Zander and Trichostomoideae strict phylogenetic monophyly, a stricture that is (Bruch & Schimp.) Limpr. has recently been much arbitrary and enforced simply because use of shared inturmoil(e.g.,Alonsoetal.2018a,b;Kockinger¨ et traits alone in cladistics eliminates evaluation of al. 2010; Werner et al. 2005). In particular, Sollman serial evolution, i.e., taxon to taxon. Primary is the (2000, 2005) reduced 53 species in Chionoloma fact that cladistics is actually hierarchical cluster Dixon,Oxystegus(Limpr.) Hilp.,Pseudosymblepha- analysis using trait changes in a non-ultrametric ris Broth.,TrichostomumHedw., and other genera tree. Sets of traits of two or more species are split to Pseudosymblepharis bombayensis (Mull.¨ Hal.) into pairs and these into pairs again, and so on. This P.Sollman. This in our opinion was a massive loss is not a model of evolution, which should try to of important evolutionary distinctions. The prob- explain at least species to species descent with lem was corrected to a considerable extent by modification. Molecular systematics has additional Alonso et al. (2018a), who re-established several problems (Zander 2013: 58), mainly a lack of distinctive species, but recognized most of these in resolution of evolutionary differences, summarized Chionoloma. in the Discussion. New discoveries are here reported Alonso et al. (2016) merged the genera Oxy- following classical generic concepts. stegus and Pseudosymblepharis with Chionoloma based on putative phylogenetic monophyly in an MATERIALS AND METHODS analysis of nuclear ITS and plastid markers atpB- Herbarium specimens of Tortella (Mull.¨ Hal.) rbcL, trnG and trnL-F molecular sequences. They Limpr. (Pottiaceae) associated with the studies of also evaluated traits used by Zander (1993a) as the second author (Eckel 2007, 2010) that were on generic distinctions that matched classical classifi- loan from UBC were restudied in light of the recent cations, but found them less than convincing. We discovery of Asian species of Pottiaceae (Zander have re-examined these distinctions and find no 2017a, 2019a) in the North American hyperoceanic 1 Corresponding author’s e-mail: northwest. Reference slides from previous studies of [email protected] Tortella (Eckel 1990, 1998, 2007a, 2010; Hedenas¨ & DOI: 10.1639/0007-2745-122.4.568 Eckel 2011) and related genera (Eckel 2000a,b, The Bryologist 122(4), pp. 568–577 Published online: October 21 2019 Copyright Ó2019 by The American Bryological and Lichenological Society, Inc. Zander & Eckel: Pseudosymblepharis and Oxystegus 569 2007b; Zander 1993b, 2017b) plus extensive material Etymology. The epithet is a Latinization of in MO were used to re-evaluate species distinctions. Greek maragna, whip, and phyllon, leaf. Among this material were misidentified specimens Discussion. Oxystegus maragniphyllus is closely of Oxystegus and Pseudosymblepharis. related to O. tenuirostris (Hook. & Taylor) A.J.E.Sm. by the notched and fragile leaf limb, and the TAXONOMY broadened leaf base, and stem with central strand. It differs from O. tenuirostris by its stiff, very narrow Oxystegus maragniphyllus R.H.Zander & Eckel, sp. leaf limb, the base commonly fully sheathing the nov. Fig. 1 stem (best seen in stripped leaves with overlapping Caulis funiculo centrali misero praeditus. Folia basal margins), and very narrow leaf apex usually fragilia, limbo inciso, saepe abrupto, basi dilatata with a strongly excurrent costa of 15–30 smooth sed humeris distinctis carentia. Costa ut mucro rhomboidal cells in the very long-linear perichaetial cylindrica, acuta, basi 70–110 lm lata, filo leaves, and 7–15 in cauline leaves. It is known from adaxiali stereidarum e cellulis paucioribus quam only one specimen, but may turn up in other abaxiali constante. Cellulae laminales distalliter hyperoceanic areas, particularly in Asia. Oxystegus 10–13 lm latae, secus margines foliares indiscrete maragniphyllus is similar to O. hibernicus (Mitt.) ascendentes. Hilp. but the former differs in the narrower leaf limb, longer and broader leaf base, and very fragile TYPE: CANADA. BRITISH COLUMBIA: West Vancouver leaves. Crum & Anderson (1958) discuss a very Island, Ocluelet, limestone outcrop in forest, narrow-leaved form of O. tenuirostris (as Trichosto- May 7, 1969, W. B. Schofield 89947 (UBC: mum cylindricum Hedw.) in the Southern Appala- holotype). chians, but their text and illustration concerns a Description. Plants in a dense turf, yellow- population with non-fragile leaf limbs and narrow green, 2–2.5 cm tall. Stem with weak hyalodermis, leaf bases, not broadly ovate and sheathing as is the weakly thickened sclerodermis, central cells all of case with O. maragniphyllus. Although this species is similar size, and central strand weak or rarely recognized from only one specimen, some traits are strong in same plant, this often red just below qualitative, not quantitative; the range of variation archegonia. Axillary hairs of ca. 8–10 cells, the in all related species is such that there is no overlap distal cells hyaline and basal 4–5 thick-walled or of traits; and distinctiveness is greater between the brownish. Leaves linear-lanceolate from an ovate or new species and related species than between each of elliptical base, fragile and often broken off at mid- those related species. limb, when dry inwardly hooked, incurved-twisted Pseudosymblepharis new to North America and tubulose, when wet spreading at ca. 458 from from Alaska (Fig. 2). The discovery of the Asian sheathing base, apex narrow, gradually merging Pseudosymblepharis angustata (Mitt.) Hilp. in Alaska into a long-pointed, sharp mucro, mid-limb often is not unexpected given that Alaska harbors isolated notched or cracked at margins; costa with adaxial populations of the Asian Pottiaceae species of stereid band smaller than the abaxial; leaf base similar morphology: Anoectangium sikkimense Aziz much broadened, sheathing the stem, shoulders & Vohra (Zander 2017a) and Molendoa hornschuchi- weakorabsent,notdecurrentatinsertion.Leaf ana (Hook.) Lindb. ex Limpr. (Zander 1978, 2007), cells at mid-leaf quadrate to short-rectangular, 10– both differing in being pleurocarpous. Pseudosym- 13 lmwide,papillaebifid,solid,crowded;basal blepharis angustata is described and illustrated by cells not sharply distinguished from the distal but Saito (1975) and by Li et al. (2001), and an grading through thick-walled, smooth rectangular illustration of the Alaskan plant is provided here. cells, hyaline and thin-walled, ca. 13 lmwide,3– It has leaves that are less strongly squarrose or 5:1, rising up the margins in a narrow line. shouldered than the Latin American P. schimperiana Perichaetial leaves gradually longer and narrower (Paris) H.A.Crum. The European Oxystegus hiber- than the cauline, with longer cusp and more nicus is similar to P. angustata in the large, very elongated sheathing base. Sporophytes and peri- long-lanceolate narrowly acuminate leaves with goniate plants not seen. widened leaf base and plane margins, distal marginal 570 The Bryologist 122(4): 2019 Figure 1. Oxystegus maragniphyllus. A. Habit, moist. B. Habit, dry. C–E. Leaves. F. Stem section. G. Leaf section at mid-leaf. H. Section at leaf base. I. Distal lamina with crack. J. Leaf base. K. Mucro of cauline leaf. L. Mucro of perichaetial leaf. Scale bars: a ¼ 2 mm for A–B, 60 lm for F; b ¼ 60 lm for G–L; c ¼ 50 lm for C–D. Zander & Eckel: Pseudosymblepharis and Oxystegus 571 Figure 2. Pseudosymblepharis angustata. A. Habit, moist. B. Habit, dry. C–E. Leaves. F. Section of stem. G. Leaf section near apex. H. Section at mid-leaf. I. Section at leaf base. J. Basal laminal cells. K. Cells at leaf shoulder. L. Leaf mucro. Scale bars: a ¼ 1 mm for C–E; b ¼ 2 mm for A–B, 90 lm for F; c ¼ 50 lm for G–L. 572 The Bryologist 122(4): 2019 cells with oval lumens with walls weakly projecting identified by M. Alonso, M. J. Cano and J. A. as a minute crenulation, and distal marginal cells Jimenez´ (unpubl.). entire (not even distantly denticulate), but O. CANADA. BRITISH COLUMBIA. Vancouver Island, hibernicus differs in that the costal adaxial stereid Indian Arm, Wigwam Creek, cliff crevices, 15 May band is smaller than the abaxial, the basal cells do 1963, W.B. Schofield 20536 (UBC); mountain just S of not rise into the limb, the basal cells are restricted to Harrison Lake, Skeena River, wet cliff, 13 Jun. 1963, below the shoulders of the sheathing base, and there W.B. Schofield 21266 (UBC); Haida Gwaii, NW is a region of thick-walled rectangular cells between Moresby Island, Paul Inlet Rd., ca. 5 km E of Inlet, the thin-walled extreme basal cells and the distal dry cliff, 22 Aug. 1961, W.B. Schofield 15218 (UBC). laminal cells. Distal laminal cells of O. hibernicus are KEY TO OXYSTEGUS AND PSEUDOSYMBLEPHARIS IN NORTH commonly 10–13 lm wide and short-rectangular, AMERICA NORTH OF MEXICO but those of P.
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