Research Article

Algae 2015, 30(4): 247-263 http://dx.doi.org/10.4490/algae.2015.30.4.247 Open Access

Marine macroalgae of the : I. Bangiales

Sandra C. Lindstrom1,*, Mandy R. Lindeberg2 and Daniel A. Guthrie3 1Department of Botany and Beaty Biodiversity Museum, #3529-6270 University Blvd., University of British Columbia, Vancouver, BC V6T 1Z4, Canada 2Auke Bay Laboratories, Fisheries Science Center, 17109 Pt. Lena Loop Road, Juneau, AK 99801, USA 3W. M. Keck Science Department, Claremont McKenna, Scripps and Pitzer Colleges, 925 N. Mills Avenue, Claremont, CA 91711, USA

We sequenced the rbcL gene in more than 100 collections of foliose Bangiales made in the Aleutian Islands and west- ern Alaska Peninsula during the past 25 years. This work allows us to recognize four previously undescribed species, two in the genus Boreophyllum and two in Pyropia. Boreophyllum aleuticum appears to be endemic to the Aleutian Islands, whereas B. ambiguum is known to occur from the Yakutat area to the tip of the Alaska Peninsula. The two previously undescribed species of Pyropia are more broadly distributed. Pyropia taeniata, which was previously identified under the name Py. pseudolinearis, occurs from northern Southeast Alaska through the Aleutian Islands. Pyropia unabbottiae, which is sister to Py. abbottiae, occurs from southern Vancouver Island to . Collections throughout the Aleu- tian Islands allow us to document the distribution of another dozen species of foliose Bangiales in this region, including Boreophyllum aestivale, Fuscifolium tasa, Pyropia fallax, Py. fucicola, Py. gardneri, Py. kurogii, Py. nereocystis, Py. pseudol- anceolata, Py. torta, Wildemania amplissima, W. norrisii, and W. variegata. We were unable to confirm the occurrence of the following species previously recorded from the Aleutian Islands: Porphya ochotensis, Pyropia abbottiae, Py. perforata, Py. pseudolinearis, P. purpurea, P. umbilicalis, Py. yezoensis and Wildemania schizophylla. At least two undescribed fila- mentous Bangiales also occur in the Aleutian Islands.

Key Words: Aleutian Islands; Bangiales; Boreophyllum; Porphyra; Pyropia; rbcL

INTRODUCTION

The Aleutian Islands, an archipelago of more than 300 situated on continental crust, whereas those west of Sa- islands, stretch more than 1,600 km from the western tip malga Pass are distinctly oceanic. The islands are grouped of the Alaska Peninsula at 163° W latitude () in five major clusters, which are separated by increasingly to beyond the International Dateline at 172° E longitude wider and deeper passages from east to west (Stabeno et (Attu Island) and span over 425 km of latitude (between al. 2005). Ocean mixing and transport along the archipel- 51 and 55° N). This remote archipelago is uniquely situ- ago are driven by upwelling, tidal action through passes, ated between the American and Asian continents and be- and three major currents: the Alaska Coastal Current and tween two major bodies of water, the North Pacific Ocean the Alaska Stream flow westward on the Pacific side, and and the (Fig. 1). The easternmost islands are the Aleutian North Slope Current flows eastward on the

This is an Open Access article distributed under the Received October 15, 2015, Accepted December 5, 2015 terms of the Creative Commons Attribution Non-Com- Corresponding Author mercial License (http://creativecommons.org/licenses/by-nc/3.0/) which * permits unrestricted non-commercial use, distribution, and reproduction E-mail: [email protected] in any medium, provided the original work is properly cited. Tel: +1-250-655-1534, Fax: +1-604-822-6089

Copyright © 2015 The Korean Society of Phycology 247 http://e-algae.kr pISSN: 1226-2617 eISSN: 2093-0860 Algae 2015, 30(4): 247-263

Fig. 1. Map of the Aleutian Islands showing islands where Bangiales were collected in this study.

Bering side (Ladd et al. 2005). Sea surface temperatures gae thrive in the Aleutian littoral zone, including members range from 5 to 10°C in summer and drop to near 0 in of the Bangiales (Lebednik and Palmisano 1977). In fact, winter (Rodionov et al. 2005). the wide spray zone of the upper intertidal is ideal habi- The shores of the Aleutian archipelago offer some of tat for many species of Bangiales, which favor cool waters the most exposed habitats of any rocky coastline on earth with frequent disturbances that not only provide cleared and experience a cool, wet and windy marine climate. substrate for colonization but also challenge the tenac- Extreme winter storms are fueled by the notorious low- ity of invertebrate herbivores. Species of Bangiales often pressure area known as the Aleutian Low, and moisture- form a distinct band visible from offshore and can be an laden fog often blankets the coast in summer (Rodionov abiding feature of the Aleutian littoral zone. Although et al. 2005). Coastal areas are generally characterized by species of Bangiales are commonly found throughout high cliffs with rock ramps (≥100 m wide) or steep bed- the Aleutian Islands, their identity and distribution have rock, boulder, and cobble beaches. Substrates range from been poorly documented because of the lack of collection basaltic lava flows to tuffs, breccia, conglomerates, and due to remoteness and the challenging environment of sandstone (Gard 1977). Oceanic ground swells constantly the archipelago and the application of appropriate meth- carve shorelines, forming wave-cut channels, round- ods of determination. Wynne (1972) published the only ing boulders and cobble, eroding sea cliffs, and creating study that focused on foliose Bangiales in this region, but extensive spray zones. In addition to wind, weather and his efforts were limited to Island. Conway et al. waves, the islands are also home to 37 extant volcanoes, (1975) included a number of records of Bangiales from and eruptions impacting coastal habitats are common the Aleutian Islands in their work. Both of these studies, (Jewett and Drew 2014). The archipelago’s vast and com- however, were done at a time when morphology was still plex physical setting creates highly variable and isolating the main method for seaweed identification. Today, DNA conditions between island groups and even neighboring sequencing is essential for identification of species such islands. Endemism can be expected along this extensive as foliose Bangiales, which have few morphological char- but narrow chain of islands (Kawai et al. 2008, Jewett and acters to distinguish them (Sutherland et al. 2011). Lind- Clark 2011). strom (2008) and Lindstrom et al. (2015) included records Despite this forbidding environment, marine macroal- of foliose Bangiales from the Aleutian Islands, but that

http://dx.doi.org/10.4490/algae.2015.30.4.247 248 Lindstrom et al. Aleutian Islands Bangiales

region was not the focus of their studies. new to science, and we describe these below. We also de- We have now sequenced the rbcL gene of over 100 col- scribe a third Alaskan species of Boreophyllum; although lections of foliose Bangiales made in the Aleutian Islands it is thus far unknown from the Aleutians, it occurs nearby and the western Alaska Peninsula during the past 25 years, at the western end of the Alaska Peninsula. and we have examined several hundred additional speci- mens of Bangiales collected during this period. Below we Boreophyllum aleuticum S. C. Lindstrom & M. R. provide records of these species in the region, including Lindeberg sp. nov. (Fig. 2A) descriptions of previously unnamed or incorrectly iden- tified species. This baseline information will aid future This species was identified as Unknown #4 in Lind- studies in biodiversity and phytogeography, especially strom (2008). as this region faces climate-related changes (e.g., ocean Description. Thalli to 1 m in length and width although warming, acidification, and changing weather patterns). usually much smaller, broadly ovate, obovate, or nearly orbiculate, often lacerated or even lobed, with margin very slightly but deeply ruffled. Base cuneate to umbili- MATERIALS AND METHODS cate. Monostromatic, with a single chloroplast per cell. Dark red, brown to nearly black, oily-looking when alive; Specimens were collected in the Aleutian Islands dusky purple when dried. Adhering well to paper. Vegeta- in 1990, 1992, 1993, 1999, and 2000 by D. A. Guthrie, in tive thalli 50-85 µm thick. Male and female reproductive 2004, 2005, and 2008 by S.C. Lindstrom, and in 2006 and cells on separate plants. Spermatangia in irregular mar- 2007 by M. R. Lindeberg. Some of these were included ginal sections of thalli, mostly distal, occurring accord- in earlier publications (Lindstrom 2008, Lindstrom et al. ing to the Hus formula 2-4/a, 2-4/b, 8-16/c (Hus 1902); 2015); the rest are listed in Appendix 1. Specimens were spermatangial thalli 75-100 µm thick. Zygotosporangia in pressed fresh on herbarium paper with a small portion nearly inconspicuous reddish marginal sections, occur- of the 2004-2008 specimens desiccated in silica gel for ring according to the Hus formula 1-2/a, 1-2/b, 2-4/c; zy- later extraction of DNA. These herbarium vouchers are gotosporangial thalli ~75 µm thick. deposited in the UBC herbarium. DNA extraction, ampli- Holotype. UBC A90644, Haycock Rock, Is- fication and sequencing followed Lindstrom (2008) with land, low intertidal, Jun 30, 2007, Leg. M. R. Lindeberg the replacement of the forward primer F57 with KitoF1 ALEUT07_119. GenBank KT936157. The holotype con- (5′-TGTCTCAATCCGTAGAATCA-3′) and the use of Bangi- sists of two thalli, one spermatangial, the other zygoto- ales-specific primers for material not desiccated in silica sporangial. gel or otherwise difficult to amplify (Table 1). Isotype. ALEUT07_118 in UBC (A90643) consists of a single thallus, which is spermatangial. Etymology. Thus far, this species has been collected RESULTS only in the Aleutian Islands, hence its name. Habitat. On cobble or larger rock in the mid to low in- We identified 17 species of Bangiales in our Aleutian tertidal zone. collections based on DNA sequencing (Table 2). Twelve Phenology. Specimens have been collected only in of these have been described previously. Two Bangia-like May, June, July, and August (the only months when Aleu- species await descriptions of the genera in which they oc- tian sites were visited). Spermantangia were observed in cur before the species can be described (see Sutherland early June collections as well as later but zygotosporangia et al. 2011). Three foliose species from the Aleutians are not until late June.

Table 1. Bangiales-specificrbc L primers used in this study Name Direction Sequence Tm (˚C) MW No. of bases F256 Forward 5′-GCAAAAGCRTAYMGWGTWGATCC-3′ 55.1 7,069.2 23 R256 Reverse 5′-GGATCWACWCKRTAYGCTTTTGC-3′ 55.1 7,017.6 23 R492 Reverse 5′-GGTCTGCCRAAYTTRTCCATRCG-3′ 57.0 7,007.1 23 F973 Forward 5′-GGTGTTGACCATATTCAYGCAGG-3′ 57.2 7,087.1 23 R973 Reverse 5′-CCTGCRTGAATATGGTCAAACACC-3′ 57.2 7,000.6 23 R1432 Reverse 5′-GGAGTCTCAACGAARTCAGC-3′ 54.4 6,143.0 20

249 http://e-algae.kr Algae 2015, 30(4): 247-263 f - - - - - g g g g g g g g b c, g b, g Attu f ------g g g Alaid ------g g g g g g g g Shemya ------g Buldir ------g e, g Kiska ------g g g Hawadax ------c g g a a a a, g Amchitka ------g Ogliuga Ogliuga f ------g Kanaga Kanaga ------c g g g g g g g g b, g Adak f ------Atka ------e e, g Carlisle ------d Chuginadak . ------e Kagamil Kagamil W. variegata W. ------e Umnak - - - e e e e e g g b e, g e, g e, g e, g b, e b, e, f e, b, Unalaska ------e e e e e e e, f e, g e, g Akutan ------e e e e e g g g e, g Akun in 1977. f f ------g g g g g g Avatanak Avatanak - - - - ( V. S. C. Foslie (S. C. De Toni De S. C. Lind (V. Krish (V. (C. L. Ander (G. M. Smith & (G. M. Smith S. C. Lindstrom (Yendo) S. C. (Yendo) (V. Krishnamur (V. (S. C. Lindstrom) (S. C. Lindstrom) 2015). (S. C. Lindstrom & (S. C. Lindstrom (V. Krishnamurthy) (V. Verified Aleutian Islands distributions of species from this and previous studies (locations listed from east to west) to east from listed studies (locations this and previous Aleutian Islands distributions of species from Verified 3’ sp. 3 3’ sp. 2’ sp. 1 2’ sp. Lindstrom & Fredericq) S. C. & Fredericq) Lindstrom Lindstrom & M. R. Lindeberg Lindstrom Lindstrom S. C. Lindstrom K. M. Cole) thy) S. C. Lindstrom S. C. Lindstrom Hollenberg) S. C. Lindstrom son) S. C. Lindstrom Krishnamurthy) S. C. Lindstrom S. C. Lindstrom strom namurthy) S. C. Lindstrom Bangia ‘Bangia ‘ aestivale Boreophyllum aleuticum Boreophyllum tasa Fuscifolium Pyropia fallax Pyropia fucicola Pyropia gardneri Pyropia kurogii Pyropia nereocystis Pyropia pseudolanceolata Pyropia taeniata Pyropia torta Pyropia unabbottiae amplissima Wildemania norrisii Wildemania variegata Wildemania Conway et al. (1975). et al. Conway UBC A70807, A70808, Leg. & Det. K. A. Miller include some UBC A70807, A70808, Leg. 4200, Jul 10, 1987, may This study. This Lindstrom (2008). Lindstrom Wynne (1972). Wynne UBC specimens annotated by M. W. Hawkes Hawkes W. M. by UBC specimens annotated Lindstrom et al. ( et al. Lindstrom Table 2. 2. Table of islands. location 1 for See Fig. a b c d e f g http://dx.doi.org/10.4490/algae.2015.30.4.247 250 Lindstrom et al. Aleutian Islands Bangiales

A B

C D

Fig. 2. (A) Holotype of Boreophyllum aleuticum, UBC A90644, Haycock Rock, Kiska Island, low intertidal, Jun 30, 2007, Leg. M. R. Lindeberg ALEUT07_119. (B) Holotype of Boreophyllum ambiguum, UBC A90188, head of Northeast Harbor, , Alaska, lower mid intertidal, on large boulder, Aug 31, 2012, Leg. S. C. Lindstrom 15128. (C) Holotype of Pyropia taeniata, UBC A90672, Sunshine Cove, Lynn Canal, Southeast Alaska, low intertidal bedrock, Apr 19, 2011, Leg. S. C. Lindstrom 14611. (D) Holotype of Pyropia unabbottiae, UBC A90681, Akun Bay, Akun Island, low intertidal boulder, Jun 13, 2008, Leg. S. C. Lindstrom 13780.

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Known distribution. Thus far known only from the Boreophyllum aestivale and B. ambiguum differ by 7 bp Aleutian Islands: Akutan, Adak. Ogliuga, Amchitka, and (0.5%) fixed differences. These differences are evident Kiska (Table 2, Appendix 1) although probably occurring even between specimens that co-occur on the same rock. on additional islands in the chain. Specimens of B. ambiguum are often quite large and Comments. This species clearly belongs in the genus nearly orbiculate in contrast to the usual broadly lanceo- Boreophyllum based on rbcL and 18SrRNA gene sequenc- late form of B. aestivale. This difference, however, may es (Lindstrom 2008, Sutherland et al. 2011). It is sister to merely reflect that the former has usually been found in the clade containing the other species of Boreophyllum fairly protected habitats, whereas the latter is often seen that have been described (B. aestivale, B. birdiae, and B. on exposed shores. pseudocrassa). Pyropia taeniata S. C. Lindstrom sp. nov. (Fig. 2C) Boreophyllum ambiguum S. C. Lindstrom sp. nov. (Fig. 2B) This species was identified as Unknown #2 in Lind- strom (2008). Description. Thalli to at least 50 cm in length and width Description. Thalli lanceolate to linear, usually taper- although usually much smaller, broadly ovate, obovate, ing to a point when young, with finely ruffled margins or nearly orbiculate, often lacerated or even lobed, with (male thalli occasionally planar), reaching at least 44 cm margin moderately ruffled, particularly proximally. Base in length and 14 cm in width although usually shorter (20- umbilicate. Monostromatic, with a single chloroplast per 40 cm) and much narrower (1.2-3.0 cm); often distinctly cell. Dusky purplish rose when dried. Mostly adhering stipitate. Monostromatic, with a single chloroplast per well to paper. Male and female reproductive cells on the cell. Adhering well to paper. Color usually reddish pink, same thalli but on separate “halves,” separated by a very reddish brown, or reddish purple. Male and female repro- irregular, often inconspicuous vertical line. Spermatangia ductive cells usually on separate plants; when on the same in more or less solid smears across the distal end of thal- thallus, sectored by a vertical line (UBC A90660) or even lus, occurring according to the Hus formula 2-4/a, 2-4/b, monoecious, with marginal spermatangial streaks adja- 8/c; spermatangial thalli 52-65 µm thick. Zygotosporan- cent to zygotosporangial patches (UBC A90661, A90669, gia in mottled, very slightly rosaceous smears, occurring A91706, A91709, A91711, A91742, A91743, A91744, and according to the Hus formula 1-2/a, 2/b, 4/c, with clearly A91752). Spermatangia in a narrow band along the mar- visible cell walls separating packets; zygotosporangial gin, occurring according to the Hus formula 2 /a, 2-4/b, thalli 55-85 µm thick. 8/c; spermatangial thalli 45-70 µm thick. Zygotosporan- Holotype. UBC A90188, head of Northeast Harbor, gia in reddish distal ends and along distal margins, be- Sanak Island, Alaska, lower mid intertidal, on large boul- coming hieroglyph-like or mottled as spores are released, der, Aug 31, 2012, Leg. S. C. Lindstrom 15128. GenBank arranged according to the Hus formula 2-4/a, 2-4/b, 4/c; KT936164. zygotosporangial thalli 45-50 µm thick. Isotypes. SCL 15130 in UBC (A90189). GenBank KT93 Holotype. UBC A90672, Sunshine Cove, Lynn Canal, 6165. Southeast Alaska, low intertidal bedrock, Apr 19, 2011, Etymology. The species epithet derives from the diffi- Leg. S. C. Lindstrom 14611. GenBank KT936189. The ho- culty in differentiating this species from the other species lotype consists of two thalli, one spermatangial, the other of Boreophyllum in the Alaska flora. zygotosporangial. Habitat. Mid intertidal boulder, cobble, and pebble. Isotypes. There are no isotypes, but there is abundant Phenology. Specimens have been collected in July and topotype material in UBC and ALAJ, the herbarium of the August; so, like its sister taxon B. aestivale, this appears to University of Alaska Southeast, Juneau (see Comments be a summer species, likely extending into autumn. below). Known distribution. Confirmed to occur from the Etymology. The specific epithet refers to the fine, rib- eastern (La Chausee Spit) to the tip of the bon-like nature of the thallus. Alaska Peninsula (Cold Bay, Sanak Island) (Appendix 1). Habitat. This species occurs on low intertidal rock, Comments. An earlier record of this species from usually bedrock or boulders, except in the western Aleu- Prince William Sound was published as B. aestivale (S. tian Islands, where it occurs in the mid to high intertidal C. Lindstrom & Fredericq) S. C. Lindstrom (UBC A84436, and may also occur epiphytically. GenBank AF452425) (Lindstrom and Fredericq 2003). Phenology. This is a spring species, extending into

http://dx.doi.org/10.4490/algae.2015.30.4.247 252 Lindstrom et al. Aleutian Islands Bangiales

summer in colder regions. Most specimens were collect- matangia occurring according to the Hus formula 2-4/a, ed in April, May, and June, but July and August collections 4/b ,8/c, zygotosporangia, 2-4/a, 2-4/a, 4/c. were also made in the Aleutian and . Holotype. UBC A90681, Akun Bay, Akun Island, low in- Known distribution. Northern Southeast Alaska (near tertidal boulder, Jun 13, 2008, Leg. S. C. Lindstrom 13780. Juneau), Gulf of Alaska west through the Aleutian Islands GenBank KT936200. The holotype consists of two thalli, to Kamchatka (Lindstrom 2008). both of which are fertile. Comments. This Alaskan species has been identified Isotypes. SCL 13781 in UBC (A90677). GenBank KT93 as Porphra pseudolinearis Ueda since the work of Wyn- 6203. ne (1972). Indeed, Py. pseudolinearis (Ueda) N. Kikuchi, Etymology. The name is a combination of Unalaska, M. Miyata, M. S. Hwang & H. G. Choi and Py. taeniata where many specimens were first collected and iden- are related although they are not sister taxa according to tified, andabbottiae , which is the species to which it is Sutherland et al. (2011) despite their morphological simi- most closely related (they differ by 8 out of ~1,400 bp in larity. Subsequent studies in Alaska of P. pseudolinearis the rbcL gene, or 0.6%). have rather been of Py. taeniata (Stekoll et al. 1999, Lin Habitat. This species usually occurs on lower mid in- and Stekoll 2007, Lin et al. 2008, Lindstrom et al. 2008). tertidal bedrock or boulders usually in areas subjected to These studies were carried out primarily on material from some oceanic swell. It occasionally can be found higher Sunshine Cove, hence the decision to select a specimen or lower on the shore. It can also occur epiphytically on from that location rather than the Aleutian Islands as the Fucus and possibly other mid intertidal algal hosts. type of the species. Lindstrom (2008) called this species Phenology. Like its sister species, Py. abbottiae, this Unknown #2, and Lin and Stekoll (2011), Porphyra sp. species appears to be a spring-summer species. Speci- This species is very closely related to the North Atlan- mens have been collected in May, June, and July. tic Py. thulae (Munda & P. M. Pedersen) Neefus, differing Known distribution. Vancouver Island (near Victoria from it by only 6 bp fixed differences (0.4%). That said, and Malcolm Island), Kenai Peninsula (Chugach Bay), within the northeast Pacific region, this is the most ge- and the Aleutian Islands (Akun Island to Attu Island) netically variable species of foliose Bangiales, with varia- (Lindstrom 2008) (Table 2, Appendix 1). tion in at least 22 positions, all of which are third codons. Comments. When co-occurring with Py. abbottiae, this species feels “softer.” Also, in contrast to Py. abbot- Pyropia unabbottiae S. C. Lindstrom sp. nov. (Fig. tiae, this species can occasionally be epiphytic; Py. abbot- 2D) tiae has never been reported to be epiphytic. Kucera and Saunders (2012) also reported this species from the Vic- This species was identified as Unknown #1 in Lind- toria area of southern Vancouver Island (JN029000 and strom (2008). JN029001, both as Pyropia sp. 1POR). Description. Thallus broadly lanceolate to nearly or- biculate, to at least 30 cm tall and 20 cm wide but usually smaller. Margin very strongly ruffled, especially near the DISCUSSION base (the distal end is often quite flat), the ruffles extend- ing nearly to the center of the thallus, especially mid thal- Recent studies of the Bangiales on a global scale lus. Center of proximal part also planar, often greenish, (Sutherland et al. 2011) as well as more localized stud- whereas rest of blade is dusky pink or purplish. Older thal- ies in the northeast Pacific (Lindstrom 2008, Kucera and li often extremely perforated (presumably from littorine Saunders 2012, Lindstrom et al. 2015) allow us to update grazing and subsequent growth of the blade). Monostro- the distributions of the following species of Bangiales matic, with one chloroplast per cell. Adhering well to pa- confirmed herein to occur in the Aleutian Islands: per. Vegetative and reproductive thalli c. 50-60 µm thick. 1) Filamentous Bangiales. Two species of filamentous Male and female structures intermixed on same thallus, Bangiales have been confirmed from the Aleutian Islands. occurring along a broad marginal area in the distal region One, in ‘Bangia 2’ (sensu Sutherland et al. 2011), has been of the thallus. Spermatangia in streaks or distinctly pat- confirmed for the eastern Aleutian Islands. This species terned patches amid solid red to pale zygotosporangial (‘Bangia 2’ sp. 1) is widely distributed in the northeast areas, the solid red areas reminiscent of Py. abbottiae ( V. Pacific from southern British Columbia to the Eastern Krishnamurthy) S. C. Lindstrom, the pale areas of Py. fal- Aleutian Islands (Lindstrom in preparation). The other lax (S. C. Lindstrom & K. M. Cole) S. C. Lindstrom. Sper- species, ‘Bangia 3’ sp. 3, has been confirmed from Attu I.

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This species is widely distributed in northern waters, hav- and P. umbilicalis are North Atlantic species; the former ing been recorded for the Sitka area in Southeast Alaska has been recorded in the northeast Pacific, where it oc- and from eastern Canada, Greenland, and the northeast curs in harbors (Stiller and Waaland 1996 and Lindstrom US (Müller et al. 1998). 2008, as P. rediviva J. W. Stiller & J. R. Waaland; we have 2) Foliose Bangiales. Boreophyllum. We recognize two also collected it in Homer harbor: UBC A88285, GenBank species of this genus in the Aleutian Islands, B. aesti- KT936235, Homer, Alaska, S. C. Lindstrom 13403). Wynne’s vale, which is confirmed only from the eastern Aleutian record of P. purpurea from Amchitka Island was likely a Is.: Akun I., Akutan I. and Unalaska I., and B. aleuticum, species of Boreophyllum, which can be morphologically which is widely distributed along the chain (Table 2). indistinguishable from P. purpurea. We have verified the 3) Fuscifolium. A single species, F. tasa (Yendo) S. C. occurrence of Py. abbottiae as far west as Sanak Island, Lindstrom, has been recorded across the Aleutian Islands. just east of the Aleutian Islands (SCL 15206, GenBank This species was originally reported by Wynne (1972) as KT936246, Eagle Rock, Northeast Harbor, Sanak I., Sep Porphyra tasa (Yendo) Ueda, which has a type locality of 2, 2012). All specimens previously identified asPorphyra Onetokan Island, Kurile Islands, Russia (Yendo 1920). The or Wildemania schizophylla from Alaska are currently in- identity of the Aleutian species with Kurile Islands mate- cluded in W. norrisii (Sutherland et al. 2011; this species rial has not been confirmed. was identified as Unknown #5 in Lindstrom 2008). 4) Pyropia. Nine species have been confirmed to occur There is a possibility that some of the Aleutian Ban- in the Aleutian Islands: Py. fallax, Py. fucicola, Py. gard- giales are the same as some of the poorly known spe- neri, Py. kurogii, Py. nereocystis, Py. pseudolanceolata, Py. cies from the northwestern Pacific Ocean. The northern taeniata, Py. torta, and Py. unabbottiae (Table 2, Appendix Japanese species of Porphyra bulbopes (Yendo) Ueda, P. 1). Most of these are distributed throughout the islands of ochotensis and P. punctata Yamada & Mikami have yet the chain. However, P. kurogii has been reported only as to be studied in a molecular context (Kurogi 1972), as is far west as Adak I., and P. nereocystis only to Umnak I., the also the case with the Russian P. inaequicrassa Perestenko same as its host, Nereocystis luetkeana. (1980). Klochkova et al. (2009) provide descriptions and 5) Wildemania. Three species of Wildemania are con- illustrations of a dozen foliose Bangiales from the Kam- firmed to occur in the Aleutian Islands:W. amplissima chatka region of Russia but without reference to type ma- [also sometimes identified asW. cuneiformis (Setchell & terial or molecular sequencing. Hus) S. C. Lindstrom], W. norrisii, and W. variegata. These Although the Aleutian Islands offer a near-ideal habi- too have been found throughout the chain. tat for species of Bangiales (cold, high disturbance, lim- We were unable to confirm the occurrence of the fol- ited herbivory), the numbers of species we were able to lowing species in the Aleutian Islands: Porphya ochotensis confirm is surprisingly low (2 filamentous and 15 foliose Nagai, Pyropia perforata (J. Agardh) S. C. Lindstrom, Py. species). We suspect that a number of species have yet to pseudolinearis, P. purpurea (Roth) C. Agardh, P. umbilicalis be uncovered in the flora. Collecting opportunities are Kützing, Py. yezoensis (Ueda) M. S. Hwang & H. G. Choi (all largely limited to summer months (May to August), and recorded by Wynne 1972 at Amchitka I. as Porphyra spp.) this may in part be responsible for these low numbers. and Py. abbottiae and Wildemania schizophylla (Hollen- However, the possibility remains that these numbers re- berg) S. C. Lindstrom (recorded by Conway et al. 1975 at flect the true diversity of Bangiales in the Aleutian Islands. various islands as P. abbottiae V. Krishnamurthy and P. Only further intense collecting will determine which is schizophylla Hollenberg, respectively). We attempted to the case. amplify DNA from several of Wynne’s original collections, but only Lebednik AM343 in MICH (970724), originally identified by Wynne asPorphyra yezoensis Ueda, provid- ACKNOWLEDGEMENTS ed an unambiguous sequence, which indicated it to be Boreophyllum aleuticum. The identity of P. ochotensis has Collection of the specimens recorded herein would not yet to be determined; its description suggests it may be a have been possible without the help or support of the fol- species of Boreophyllum. Pyropia perforata has been con- lowing people: John Cubit, Jochen Halfor, Jon Houghton, firmed to occur as far west as Kodiak I., Alaska (Lindstrom Paul Tate, and David Ward. Michael Wynne kindly sup- 2008). Pyropia pseudolinearis is restricted to the north- plied fragments of some of his collections in MICH, for west Pacific, where at least two species are identified -un which we are most grateful. Expenses associated with der this name (Sutherland et al. 2011). Porphyra purpurea DNA sequencing were defrayed by the Natural Sciences

http://dx.doi.org/10.4490/algae.2015.30.4.247 254 Lindstrom et al. Aleutian Islands Bangiales

and Engineering Research Council of Canada, the Census and respiration of the conchocelis stage of Alaskan Por- of Marine Life, and Emilie D. Lindstrom. phyra (Bangiales, Rhodophyta) species in response to environmental variables. J. Phycol. 44:573-583. Lin, R. & Stekoll, M. S. 2007. Effects of plant growth sub- REFERENCES stances on the conchocelis phase of Alaskan Porphyra (Bangiales, Rhodophyta) species in conjunction with Conway, E., Mumford, T. F. Jr. & Scagel, R. F. 1975. The genus environmental variables. J. Phycol. 43:1094-1103. Porphyra in British Columbia and Washington. Syesis Lin, R. & Stekoll, M. S. 2011. Phycobilin content of the con- 8:185-244. chocelis phase of Alaskan Porphyra (Bangiales, Rho- Gard, L. M. Jr. 1977. Geologic history. In Merritt, M. L. & dophyta) species: responses to environmental variables. Fuller, R. G. (Eds.) The Environment of Amchitka Island, J. Phycol. 47:208-214. Alaska. TID-26712. Technical Information Center, U.S. Lindstrom, S. C. 2008. Cryptic diversity, biogeography and Energy Research and Development Administration, genetic variation in Northeast Pacific species ofPorphy - Springfield, VA, pp. 13-34. ra sensu lato (Bangiales, Rhodophyta). J. Appl. Phycol. Hus, H. T. A. 1902. An account of the species of Porphyra 20:951-962. found on the Pacific coast of North America. Proc. Calif. Lindstrom, S. C., Conitz, J. M., Hall, S. & Stekoll, M. S. 2008. Acad. Sci. 2:173-241. Induction of conchospore release: ecotypic variation in Jewett, S. C. & Clark, R. N. 2011. Discoveries of new marine northeast Pacific species ofPorphyra . J. Appl. Phycol. species of the Aleutian Islands. In Pollock, N. W. (Ed.) 20:331-340. Diving for Science 2011. Proc. 30th Am. Acad. Underwa- Lindstrom, S. C. & Fredericq, S. 2003. rbcL gene sequences ter Sci. Symp., Dauphin Island, AL, pp. 91-109. reveal relationships among north-east Pacific species of Jewett, S. C. & Drew, G. S. 2014. Recolonization of the in- Porphyra (Bangiales, Rhodophyta) and a new species, P. tertidal and shallow subtidal community following the aestivalis. Phycol. Res. 51:211-224. 2008 eruption of Alaska’s Kasatochi Volcano. Biogeosci. Lindstrom, S. C., Hughey, J. R. & Aguilar Rosas, L. E. 2015. Discuss. 11:3799-3836. Four new species of Pyropia (Bangiales, Rhodophyta) Kawai, H., Hanyuda, T., Lindeberg, M. & Lindstrom, S. C. from the west coast of North America: the Pyropia lan- 2008. Morphology and molecular phylogeny of Aureo- ceolata species complex updated. PhytoKeys 52:1-22. phycus aleuticus gen. et sp. nov. (Laminariales, Phaeo- Müller, K. M., Sheath, R. G., Vis, M. L., Crease, T. J. & Cole, K. phyceae) from the Aleutian Islands. J. Phycol. 44:1013- M. 1998. Biogeography and systematics of Bangia (Ban- 1021. giales, Rhodophyta) based on the Rubisco spacer, rbcL Klochkova, N. G., Korolyova, T. N. & Kusidi, A. E. 2009. Atlas of gene and 18S rRNA gene sequences and morphometric marine algae of Kamchatka and surrounding areas. Vol. analyses. I. North America. Phycologia 37:195-207. 2. Red seaweeds. KamchatNIRO Press, Petropavlovsk- Perestenko, L. P. 1980. Vodorosli Zaliva Petra Velikogo [The Kamchatsky, 301 pp. seaweeds of Peter the Great Bay]. Akademia NAUKA Kucera, H. & Saunders, G. W. 2012. A survey of Bangiales SSSR, Leningrad, 232 pp. (Rhodophyta) based on multiple molecular markers re- Rodionov, S. N., Overland, J. E. & Bond, N. A. 2005. Spatial veals cryptic diversity. J. Phycol. 48:869-882. and temporal variability of the Aleutian climate. Fish. Kurogi, M. 1972. Systematics of Porphyra in Japan. In Abbott, Oceanogr. 14(Suppl. 1):3-21. I. A. & Kurogi, M. (Eds). Contributions to the Systematics Stabeno, P. J., Kachel, D. G., Kachel, N. B. & Sullivan, M. E. of the Benthic Marine Algae of the North Pacific. Japa- 2005. Observations from moorings in the Aleutian Pass- nese Society of Phycology, Kobe, pp. 167-192. es: temperature, salinity and transport. Fish. Oceanogr. Ladd, C., Hunt, G. L. Jr., Mordy, C. W., Salo, S. A. & Stabeno, P. 14(Suppl. 1):39-54. J. 2005. Marine environment of the eastern and central Stekoll, M. S., Lin, R. & Lindstrom, S. C. 1999. Porphyra culti- Aleutian Islands. Fish. Oceanogr. 14(Suppl. 1):22-38. vation in Alaska: conchocelis growth of three indigenous Lebednik, P. A. & Palmisano, J. F. 1977. Ecology of marine al- species. Hydrobiologia 398/399:291-297. gae. In Merritt, M. L. & Fuller, R. G. (Eds.) The Environ- Stiller, J. W. & Waaland, J. R. 1996. Porphyra rediviva sp. nov. ment of Amchitka Island, Alaska. TID-26712. Technical (Rhodophyta): a new species from northeast Pacific salt Information Center, U.S. Energy Research and Develop- marshes. J. Phycol. 32:323-332. ment Administration, Springfield, VA, pp. 353-393. Sutherland, J. E., Lindstrom, S. C., Nelson, W. A., Brodie, J., Lin, R., Lindstrom, S. C. & Stekoll, M. S. 2008. Photosynthesis Lynch, M. D. J., Hwang, M. S., Choi, H. -G., Miyata, M.,

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Kikuchi, N., Oliveira, M. C., Farr, T., Neefus, C., Mols- Aleutians. In Kazutosi (Ed.) Proc. 7th Int. Seaweed Mortensen, A., Milstein, D. & Müller, K. M. 2011. A new Symp., University of Tokyo Press, Tokyo, pp. 100-104. look at an ancient order: generic revision of the Bangia- Yendo, K. 1920. Novae algae Japonicae, I-III. Bot. Mag. 34:1- les (Rhodophyta). J. Phycol. 47:1131-1151. 12. Wynne, M. J. 1972. The genus Porphyra at Amchitka Island,

http://dx.doi.org/10.4490/algae.2015.30.4.247 256 Lindstrom et al. Aleutian Islands Bangiales - S. C. Lindstrom 12141 S. C. Lindstrom s/n S. C. Lindstrom 12247 S .C. Lindstrom 13744 S. C. Lindstrom 13436 S. C. Lindstrom 15129 S. C. Lindstrom ALEUT07_118 [121] M. R. Lindeberg ALEUT07_119 [121] M. R. Lindeberg 13695 S. C. Lindstrom A. Lebednik AM343 P. s/n, 1104 Guthrie D. 1137, 1142 Guthrie D. 1331 Guthrie D. H-31, 32 Guthrie D. A Guthrie D. C. Lees s/n D. G. J. Mueller 14175 S. C. Lindstrom 14176 S. C. Lindstrom 14177 S. C. Lindstrom 15128 S. C. Lindstrom 15130 S. C. Lindstrom 13450 S. C. Lindstrom ALEUT07_115 [124] M. R. Lindeberg M. R. Lindeberg 13631 S. C. Lindstrom 13639 S. C. Lindstrom ALEUT07_211 [214] M. R. Lindeberg Collector and No. Collector Jun 3, 2005 Jun 5, 2005 Jun 7, 2005 Jun 12, 2008 Jun 4, 2008 Jun 31, 2012 Aug 30, 2007 Jun 30, 2007 Jun 9, 2008 Jun 3, 1969 Jun 19, 1993 May 20, 1993 May 22, 1993 May 25, 1993 May 14, 1999 Aug 8, 2000 Aug 6, 1971 Aug 9, 2009 Aug 9, 2009 Aug 9, 2009 Aug 31, 2012 Aug 31, 2012 Aug 4, 2008 Jun 30, 2007 Jun 29, 2007 Jun 7, 2008 Jun 8, 2008 Jun 3, 2007 Jul Date collected Date a Sedanka Pt, Unalaska I. Unalaska Pt, Sedanka I. Unalaska Cannery Bay, I. Unalaska Bay, Skan I. Avatanak I. Attu Chichagof Harbor, I. Sanak Harbor, Northeast Kiska I. Rock, Haycock Kiska I. Rock, Haycock I. Ogliuga Amchitka I. I. Adak Bay, Kuluk I. of Clam Lagoon, Adak Mouth I. Adak Bay, Shagak I. Adak Bay, Kuluk Akutan I. La Chausee Spit Inside Bay Cold Bay Cold Bay Cold I. Sanak Harbor, Northeast I. Sanak Harbor, Northeast I. Attu Chichagof Harbor, Kiska I. Rock, Haycock Kiska I. Pt, Harpoon Alaid Island Island Rat Amchitka I. Pt, Midden Collection location Collection - KT936247 - KT936248 KT936244 KT936245 KT936157 KT936157 KT936158 KT936159 KU248444 KT936160 - KU248441, KU248443 KU248442 - KT936163 KT936161 - KT936162 KT936164 KT936165 KT936207 KT936204 KT936206 KT936213 KT936206 KT936205 GenBank No. GenBank - S. C. Lindstrom sp. nov. sp. S. C. Lindstrom S. C. Lindstrom & M. R. S. C. Lindstrom (S. C. Lindstrom & Fred (S. C. Lindstrom (Yendo) S. C. Lindstrom (Yendo) Newly reported specimens of Bangiales from the Aleutian Islands plus records of the newly described species from outside the Aleutian Islands region beyond those already pub those already beyond outside the Aleutian Islands region of the newly described species from the Aleutian Islands plus records Newly reported from specimens of Bangiales 2’ sensu Sutherland et al. (2011) sp. 1 et al. (2011) sp. 2’ sensu Sutherland 3’ sensu Sutherland et al. (2011) sp. 3 et al. (2011) sp. 3’ sensu Sutherland UBC A87130 - UBC A87145 UBC A87381 SCL 13436 UBC A90190 UBC A90643 UBC A90644 UBC A90642 MICH 970724 UBC A91899, A91784 UBC A91788, A91789 UBC A91789 UBC A91896, A91695 UBC A91890, A91891 UBC A84389 ALA 1608 UBC A89054 UBC A89053 UBC A89052 UBC A90188 UBC A90189 UBC A87883 UBC A90022 ALEUT07_135 [099] UBC A87881 UBC A87340 UBC A89553 ericq) S. C. Lindstrom (this species was identified as nov. sp. Lindeberg 2008) #4 in Lindstrom Unknown Bangia Bangia ‘ ‘ aestivale Boreophyllum aleuticum Boreophyllum ambiguum Boreophyllum tasa Fuscifolium Accession / Collection No. / Collection Accession Appendix 1. Appendix (2008) (2003) and Lindstrom and Fredericq lished in Lindstrom

257 http://e-algae.kr Algae 2015, 30(4): 247-263 2116, 2124, 2125, 2126, 2128 S. C. Lindstrom 13670 S. C. Lindstrom 13733 S. C. Lindstrom 13734 S. C. Lindstrom 13735 S. C. Lindstrom 13747 S. C. Lindstrom 13770 S. C. Lindstrom 1247, 1248 Guthrie D. 1263, 1264, 1267 Guthrie D. 1368 Guthrie D. 1430 Guthrie D. 1432, 1433 Guthrie D. 1441 Guthrie D. 1737 Guthrie D. 2111, 2112, 2114, 2115, Guthrie D. s/n Guthrie D. 15 Guthrie D. 11503 S. C. Lindstrom 11568 S. C. Lindstrom 11615 S. C. Lindstrom 11617 S. C. Lindstrom 11618 S. C. Lindstrom 11619 S. C. Lindstrom 13508 S. C. Lindstrom 13706 S. C. Lindstrom 13707 S. C. Lindstrom 13708 S. C. Lindstrom 13710, 13711 S. C. Lindstrom 13738 S. C. Lindstrom 13741 S. C. Lindstrom ALEUT07_031 [005] M. R. Lindeberg 13462 S. C. Lindstrom 13484 S. C. Lindstrom 13606 S. C. Lindstrom 13742 S. C. Lindstrom Collector and No. Collector Jun 8, 2008 Jun 12, 2008 Jun 12, 2008 Jun 12, 2008 Jun 2008 Jun12, 13, 2008 Jun 21, 1993 May 21, 1993 May 23, 1993 May 24, 1993 May 24, 1993 May 24, 1993 May 1, 1993 Jun 29, 1993 Sep 9, 1993 Oct 3, 2000 Jun 14, 1972 Aug 30, 2004 Jul 30, 2004 Jul 3, 2004 Aug 3, 2004 Aug 3, 2004 Aug 3, 2004 Aug 4, 2008 Jun 11, 2008 Jun 11, 2008 Jun 11, 2008 Jun 11, 2008 Jun 11, 2008 Jun 12, 2008 Jun 25, 2007 Jun 4, 2008 Jun 4, 2008 Jun 6, 2008 Jun 12, 2008 Jun Date collected Date Amchitka Island Island Avatanak Island Avatanak Island Avatanak Island Avatanak Akun I. Akun Bay, I. Adak Sound, Sitkin I. Adak Beach, Station Loran I. Adak Bay, Kuluk I. Adak Bay, Kuluk I. Adak Beach, Station Loran I. Adak Bay, Andrew I. Shemya Shore, North I. Attu Antenna Beach, I. Attu Antenna Beach, I. Attu Beach, South Amchitka I. Bay, Makarius St. Akutan I. Akutan Harbor, Akutan I. Akutan Harbor, I. Unalaska Harbor, Dutch I. Unalaska Harbor, Dutch I. Unalaska Harbor, Dutch I. Unalaska Harbor, Dutch I. Attu Chichagof Harbor, I. Unalaska Bay, Surveyor I. Unalaska Bay, Surveyor I. Unalaska Bay, Surveyor I. Unalaska Bay, Surveyor I. Unalaska Bay, Surveyor Island Avatanak I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Point, Murder Island Avatanak Collection location Collection b b b KT936209 - - - KT936210, KT936211 KT936212 ------P742 - P385 - - - - KT936243 - - P545 - KT936242 KT936241 KT936221 - KT936222 KT936223, KT936224 KT936225 GenBank No. GenBank (Yendo) S. C. Lindstrom (Yendo) (V. Krishnamurthy) S. C. Lindstrom (V. (S. C. Lindstrom et K. M. Cole) S. C. et K. M. Cole) (S. C. Lindstrom Continued A91879, A91880, A91881, A91882 SCL 13670 UBC A87472 UBC A87473 SCL 13735 UBC A87384 UBC A87379 UBC A91798, A91799 UBC A91800, A91801, A91803 UBC A91809 UBC A91817 UBC A91820, A91821 UBC A91823 UBC A91849 UBC A91874, A91875, A91876, A91877, A91878, UBC A91901, A91902 UBC A91687 AM512 in MICH UBC A86964 UBC A85657 UBC A86669 UBC A86992 UBC A86669 UBC A86992 UBC A87469 UBC A87453 SCL 13707 SCL 13708 UBC A87451 SCL 13738 UBC A87388 UBC A89550 UBC A90693 UBC A87337 UBC A87321 UBC A90694 Lindstrom Fuscifolium tasa Fuscifolium Pyropia fallax Pyropia fucicola Accession / Collection No. / Collection Accession Appendix 1. Appendix

http://dx.doi.org/10.4490/algae.2015.30.4.247 258 Lindstrom et al. Aleutian Islands Bangiales D. Guthrie 489 Guthrie D. 1443, 1444, 1446 Guthrie D. 1473 Guthrie D. 1574 Guthrie D. 56 Guthrie D. 57 Guthrie D. A. Lebednik AM-394 P. 2087 Guthrie D. H-21, H-22 Guthrie D. 12084 S. C. Lindstrom 12240 S. C. Lindstrom 4018 K. A. Miller s/n Guthrie D. 329, 338 Guthrie D. 1244, 1245, 1246 Guthrie D. 1426, 1435 Guthrie D. 1900, 1903 Guthrie D. M. H. Dick 11510 S. C. Lindstrom 11614 S. C. Lindstrom 12279 S. C. Lindstrom M. R. Lindeberg 12278 S. C. Lindstrom ALEUT06_263 [264] M. R. Lindeberg ALEUT07_044 [011] M. R. Lindeberg ALEUT07_030 [003] M. R. Lindeberg ALEUT07_287 [309] M. R. Lindeberg 13449 S. C. Lindstrom 13451 S. C. Lindstrom 13562 S. C. Lindstrom 13580 S. C. Lindstrom Collector and No. Collector May 28, 1992 May 24, 1993 May 25, 1993 May 28, 1993 May 4, 2000 Jun 5, 2000 Jun 27, 1969 Jun 29, 1993 Sep 22, 1993 May 2, 2005 Jun 7, 2005 Jun 16, 1987 Jun 1990 May 22, 1992 May 21, 1993 May 24, 1993 May 5, 1993 Jun 27, 1974 May 30, 2004 Jul 3, 2004 Aug 9, 2005 Jun 8, 2006 Jul 9, 2005 Jun 15, 2006 Jul 26, 2007 Jun 25, 2007 Jun 8, 2007 Jul 4, 2008 Jun 4, 2008 Jun 6, 2008 Jun 6, 2008 Jun Date collected Date South Beach, Attu I. Attu Beach, South I. Adak Bay, Andrew I. Adak Bay, Kuluk I. Shemya Shore, South I. Attu Alexi Point, I. Attu Beach, South Amchitka I. Cape, Bird I. Attu Beach, Pratincole I. Adak Bay, Shagak I. Unalaska Alimuda Bay, I. Unalaska Bay, Skan I. Attu I. Attu I. Attu Beach, South I. Adak Sound, Sitkin I. Adak Bay, Andrew I. Shemya Shore, South I. Buldir Akutan I. Akutan Harbor, I. Unalaska Harbor, Dutch I. Unalaska Bay, Volcano I. Unalaska Bay, Driftwood I. Unalaska Bay, Volcano Island Carlisle I. Attu Bay, Temnac I., Savage I. Attu Chichagof Harbor, I. Kanaga Sound, Kanaga I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Point, Murder I. Attu Point, Murder Collection location Collection KT936226 - - - KU248440 KT936227 KT936249 - KT936250 ------KT936190 KU248445 - KT936166, KT936167 KT936168, KT936169 KT936172, KT936173 KT936174 KT936175 KT936176 KT936177 KT936179 KT936180 KT936181 KT936184 GenBank No. GenBank - (V. Krishnamurthy) S. C. (V. (G. M. Smith & Hollenberg) S. C. & Hollenberg) (G. M. Smith S. C. Lindstrom sp. nov. (this spe nov. sp. S. C. Lindstrom (V. Krishnamurthy) S. C. Lindstrom (V. (S. C. Lindstrom) S. C. Lindstrom (S. C. Lindstrom) Continued UBC A91765 UBC A91824, A91825, A91826 UBC A91828 UBC A91838 UBC A91714 UBC A91716 UBC A54202 UBC A91873 UBC A91894, A91895 UBC A85155 UBC A85159 UBC A70820 UBC A91904 UBC A91746, A91747 UBC A91795, A91796, A91797 UBC A91815, A91822 UBC A91867, A91869 ALA 1602 UBC A90658 UBC A90664 UBC A85162 ALEUT06_137 [138] UBC A85161 UBC A85402 UBC A90651 UBC A90650 UBC A90649 UBC A90659 UBC A90660 UBC A90669 UBC A90665 Lindstrom Lindstrom #2 in Lindstrom cies was identified as unknown 2008) Pyropia fucicola Pyropia gardneri Pyropia kurogii Pyropia pseudolanceolata Pyropia taeniata Accession / Collection No. / Collection Accession Appendix 1. Appendix

259 http://e-algae.kr Algae 2015, 30(4): 247-263 S. C. Lindstrom 13581 S. C. Lindstrom 13582 S. C. Lindstrom 13629 S. C. Lindstrom 13763 S. C. Lindstrom 13783 S. C. Lindstrom 9089 S. C. Lindstrom S. C. Lindstrom 11797 S. C. Lindstrom 14001 S. C. Lindstrom 14611 S. C. Lindstrom s/n Guthrie D. 390 Guthrie D. 409 Guthrie D. 1238 Guthrie D. 1273 Guthrie D. 1414, 1415, 1417, 1418 Guthrie D. 1495 Guthrie D. 1519, 1520, 1576, 1579 Guthrie D. 1645, 1647 Guthrie D. 1747, 1780, 1792 Guthrie D. 1854, 1870 Guthrie D. 1265 Guthrie D. 52, 53, 54 Guthrie D. 222, 224, 238 Guthrie D. 277 Guthrie D. 387, 391, 392 Guthrie D. 400, 403, 411 Guthrie D. 1416 Guthrie D. 1855 Guthrie D. 1429, 1431 Guthrie D. 12085 S. C. Lindstrom 12086 S. C. Lindstrom 12138 S. C. Lindstrom 13448 S. C. Lindstrom 13464 S. C. Lindstrom Collector and No. Collector Jun 6, 2008 Jun 6, 2008 Jun 7, 2008 Jun 12, 2008 Jun 2008 Jun12, 13, 1995 Apr 1995 May1, 19, 2005 May 27, 2009 Apr 19, 2011 Apr 1990 May 25, 1992 May 26, 1992 May 21, 1993 May 21, 1993 May 24, 1993 May 27, 1993 May 28, 1993 May 31, 1993 May 2, 1993 Jun 4, 1993 Jun 19, 1993 May 4, 2000 Jun 18, 1992 May 21, 1992 May 25, 1992 May 26, 1992 May 24, 1993 May 4, 1993 Jun 24, 1993 May 2, 2005 Jun 2, 2005 Jun 3, 2005 Jun 4, 2008 Jun 4, 2008 Jun Date collected Date Murder Point, Attu I. Attu Point, Murder I. Attu Point, Murder Alaid Island Akun I. Akun Bay, Akun I. Akun Bay, Cove Sunshine Cove Sunshine I. Kodiak Bay, Pasagshak Cove Sunshine Cove Sunshine I. Attu I. Attu Ridge, Gilbert I. Attu Ridge, Gilbert I. Adak Sound, Sitkin I. Adak Bay, Andrew I. Adak Sound, Sitkin Island Shemya End, West I. Shemya Shore, South I. Shemya Beach, South I. Shemya Shore, South I. Shemya Beach, South I. Adak Beach, Station Loran I. Attu Alexi Point, I. Attu Beach, Pratincole I. Attu Beach, South I. Attu Ridge, Gilbert I. Attu Ridge, Gilbert I. Adak Sound, Sitkin I. Shemya Beach, South I. Adak Beach, Station Loran I. Unalaska Alimuda Bay, I. Unalaska Alimuda Bay, I. Unalaska Point, Sedanka I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, Collection location Collection KT936183 KT936182 KT936185 KT936186 KT936188 - KT936170, KT936171 EU223179 KT936186 KT936189 ------KT936191 KT936192 KT936193, KT936196 KT936194 KT936195 - - - KT936197, KU248439 - - - KT936236 KT936237 GenBank No. GenBank - S. C. Lindstrom sp. nov. (this spe nov. sp. S. C. Lindstrom (V. Krishnamurthy) S. C. Lindstrom (V. Continued UBC A90661 UBC A90668 UBC A90670 UBC A90656 UBC A90654 UBC A90652 s/n S. C. Lindstrom UBC A90667 UBC A90653 UBC A90672 UBC A91905 UBC A91749 UBC A91755 UBC A91794 UBC A91804 UBC A91810, A91811, A91813, A91814 UBC A91830 UBC A91832, A91833, A91839, A91840 UBC A91842, A91844 UBC A91850, A91854, A91856 UBC A91861, A91865 UBC A91802 UBC A91706, A91709, A91711 UBC A91742, A91743, A91744 277 Guthrie D. UBC A91748, A91750, A91751 UBC A91752, A91754, A91756 UBC A91812 UBC A91862 UBC A91816, A91819 UBC A85151 UBC A85180 UBC A85181 SCL 13448 SCL 13464 cies was identified as Unknown #2 in Lindstrom #2 in Lindstrom Unknown cies was identified as 2008) Pyropia taeniata Pyropia torta Accession / Collection No. / Collection Accession Appendix 1. Appendix

http://dx.doi.org/10.4490/algae.2015.30.4.247 260 Lindstrom et al. Aleutian Islands Bangiales S. C. Lindstrom 13628 S. C. Lindstrom 13649 S. C. Lindstrom 13731 S. C. Lindstrom 13784 S. C. Lindstrom 1567, 1568 Guthrie D. 13455 S. C. Lindstrom 13463 S. C. Lindstrom 13482 S. C. Lindstrom 13704 S. C. Lindstrom 13705 S. C. Lindstrom 13728 S. C. Lindstrom 13780 S. C. Lindstrom 13781 S. C. Lindstrom 14136 S. C. Lindstrom 14139 S. C. Lindstrom 15250 S. C. Lindstrom s/n Guthrie D. 190 Guthrie D. 402 Guthrie D. 486, 487 Guthrie D. 52 Guthrie D. 532 Guthrie D. 68, 69, 657 Guthrie D. 723 Guthrie D. 1430 Guthrie D. 25, 1496 Guthrie D. 1706, 1707 Guthrie D. 1770 Guthrie D. 1827, 1843 Guthrie D. S60 Guthrie D. 28, 37 Guthrie D. 53 Guthrie D. Collector and No. Collector Jun 7, 2008 Jun 8, 2008 Jun 12, 2008 Jun 13, 2008 Jun 28, 1993 May 4, 2008 Jun 4, 2008 Jun 4, 2008 Jun 11, 2008 Jun 11, 2008 Jun 12, 2008 Jun 13, 2008 Jun 13, 2008 Jun 28, 2009 May 28, 2009 May 27, 2013 Apr 1990 May 16, 1992 May 26, 1992 May 28, 1992 May 28, 1992 May 29, 1992 May 2, 1992 Jun 4, 1992 Jun 24, 1993 May 27, 1993 May 31, 1993 May 2, 1993 Jun 3, 1993 Jun 4, 1993 Jun 1, 2000 Jun 5, 2000 Jun Date collected Date a Alaid Island Island Rat Island Avatanak Akun I. Akun Bay, I. Shemya Shore, South I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Unalaska Chernofski Hbr, I. Unalaska Chernofski Hbr, Island Avatanak Akun I. Akun Bay, Akun I. Akun Bay, I., BC Malcolm I., BC Malcolm Lagoon, BC Witty’s Outside Island Attu I. Attu Beach, South I. Attu Ridge, Gilbert I. Attu Beach, South I. Attu Beach, Pratincole I. Attu Beach, South I. Attu Debris Beach, I. Attu Beach, South I. Adak Bay, Andrew I. Shemya End, West I. Shemya Beach, South I. Shemya Shore, South I. Shemya Shore, South I. Shemya Shore, South I. Attu Beach, South I. Attu Beach, South Collection location Collection KT936238 KT936240 KT936239 - - KT936198 KT936201 - KT936202 - KT936199 KT936200 KT936203 KU248446 KU248447 KU248448 ------GenBank No. GenBank - S. C. Lindstrom sp. nov. (this nov. sp. S. C. Lindstrom (V. Krishnamurthy) S. C. Lindstrom (V. Continued UBC A87361 UBC A87884 UBC A87389 UBC A87378 UBC A91836, A91837 UBC A90675 UBC A90674 SCL 13482 UBC A90676 SCL 13705 UBC A90673 UBC A90681 UBC A90677 UBC A90680 UBC A87894 SCL 15250 UBC A91907 UBC A91738 UBC A91753 UBC A91762, A91763 UBC A91707 UBC A91766 UBC A91720, A91722, A91781 UBC A91782 UBC A91817 UBC A91889, A91831 UBC A91845, A91846 UBC A91852 UBC A91858, A91860 UBC A91917 UBC A91691, A91699 UBC A91710 species was identified as Unknown #1 in Lind Unknown species was identified as 2008) strom Pyropia torta Pyropia unabbottiae Accession / Collection No. / Collection Accession Appendix 1. Appendix

261 http://e-algae.kr Algae 2015, 30(4): 247-263 D. Guthrie 55 Guthrie D. 123 Guthrie D. s/n Guthrie D. H98? Guthrie D. ALEUT07_037 [004] M. R. Lindeberg ALEUT07_430 [403] M. R. Lindeberg 13465 S. C. Lindstrom 13585 S. C. Lindstrom 13701 S. C. Lindstrom 13730 S. C. Lindstrom 13743 S. C. Lindstrom 15201 S. C. Lindstrom 15197 S. C. Lindstrom 1860, 1861 Guthrie D. (SCL 12120) Fukuyama A. F. ALEUT07_318 [301] M. R. Lindeberg 13472 S. C. Lindstrom 13505 S. C. Lindstrom 13509 S. C. Lindstrom 13514 S. C. Lindstrom 13579 S. C. Lindstrom 13774 S. C. Lindstrom s/n Guthrie D. 52 Guthrie D. 2, 117, 119, 122 Guthrie D. 140 Guthrie D. 146, 152, 159 Guthrie D. 189 Guthrie D. 34, 35, 318 Guthrie D. 424 Guthrie D. 48, 49 Guthrie D. 488 Guthrie D. 56, 58, 565, 573, 574, 578, 581 Guthrie D. 582 Guthrie D. Collector and No. Collector May 30, 1992 May 24, 1993 Sep 12, 1999 Aug 21, 1993 May 25, 2007 Jun 18, 2007 Jul 4, 2008 Jun 6, 2008 Jun 11, 2008 Jun 12, 2008 Jun 12, 2008 Jun 2, 2012 Sep 2, 2012 Sep 4, 1993 Jun 2, 2005 Jun 10, 2007 Jul 4, 2008 Jun 4, 2008 Jun 4, 2008 Jun 4, 2008 Jun 6, 2008 Jun 13, 2008 Jun 1990 May 27, 1991 May 29, 1991 May 30, 1991 May 14, 1992 May 16, 1992 May 22, 1992 May 26, 1992 May 27, 1992 May 28, 1992 May 30, 1992 May 1, 1992 Jun Date collected Date Casco Cove, Attu I. Attu Cove, Casco I. Attu Antenna Beach, I. Unalaska Bay, Summer I. Adak Bay, Andrew I. Attu Chichagof Harbor, I. Idalug, Cape I. Attu Chichagof Harbor, I. Attu Point, Murder I. Unalaska Chernofski Hbr, Island Avatanak Island Avatanak I. Sanak Eagle Rock, I. Sanak Eagle Rock, I. Shemya Beach, South I. Unalaska Alimuda Bay, I. Adak Strait, Kagalaska I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Chichagof Harbor, I. Attu Point, Murder Akun I. Akun Bay, I. Attu I. Attu Beach, town Navy I. Attu Alexi Point, I. Attu Beach, Pratincole I. Attu Beach, Navy I. Attu Beach, South I. Attu Beach, South I. Attu Ridge, Gilbert I. Attu Beach, Pratincole I. Attu Beach, South I. Attu Cove, Casco I. Attu Beach, town Navy Collection location Collection b - - - - KT936228 KT936229 KT936231 - KT936230 KT936232 - KT936233 KT936234 P946 - KT936214 KT936220 KT936215 KT936219 KT936217 KT936216 KT936218 ------GenBank No. GenBank - Foslie (Kjellman) Toni De (V. Krishnamurthy) S. C. Lind (V. Continued A91776, A91777 UBC A91713 UBC A91731 UBC A91908, A91909 UBC A91898 UBC A89552 UBC A89551 UBC A87882 SCL 13585 UBC A87697 UBC A89940 SCL 13743 UBC A90099 UBC A90208 UBC A91863, A91864 UBC A85184 UBC A90049 UBC A87407 UBC A87367 UBC A87347 UBC A87328 UBC A87319 UBC A87373 UBC A91913, A91914 UBC A91708 UBC A91683, A91728, A91729, A91730 UBC A91732 UBC A91733, A91734, A91735 UBC A91737 UBC A91696, A91697, A91745 UBC A91759 UBC A91704, A91705 UBC A91764 UBC A91715, A91718, A91769, A91774, A91775, UBC A91778 strom (this species was identified as Unknown #5 Unknown (this species was identified as strom 2008) in Lindstrom Wildemania amplissima Wildemania norrisii Wildemania variegata Wildemania Accession / Collection No. / Collection Accession Appendix 1. Appendix

http://dx.doi.org/10.4490/algae.2015.30.4.247 262 Lindstrom et al. Aleutian Islands Bangiales - 1901, 1941 D. Guthrie 607 Guthrie D. 67, 653 Guthrie D. 73, 76 Guthrie D. 749 Guthrie D. H-6, 1121, 1134 Guthrie D. 1175, 1202, 1203, 1213 Guthrie D. 1358, 1360 Guthrie D. 35, 1450, 1475 Guthrie D. 54, 1534, 1535 Guthrie D. 5-13, 1611 Guthrie D. 5-16 Guthrie D. 5-19 Guthrie D. 1646 Guthrie D. 1717, 1718 Guthrie D. 5-38, 1765, 1775, 1788, 1793 Guthrie D. 548, 549, 1829 Guthrie D. 565, 566, 569, 570, 1897, Guthrie D. s/n Guthrie D. A Guthrie D. 2147 Guthrie D. 4 Guthrie D. 20, 22, 28 Guthrie D. 29 Guthrie D. 42 Guthrie D. 45, 46 Guthrie D. 57 Guthrie D. 68 Guthrie D. 84 Guthrie D. Collector and No. Collector Jun 1, 1992 Jun 2, 1992 Jun 3, 1992 Jun 4, 1992 Jun 19, 1993 May 20, 1993 May 23, 1993 May 25, 1993 May 28, 1993 May 29, 1993 May 29, 1993 May 30, 1993 May 31, 1993 May 1, 1993 Jun 2, 1993 Jun 3, 1993 Jun 5, 1993 Jun 12, 1999 Aug 14, 1999 Aug 1, 1993 Oct 28, 2000 May 30, 2000 May 1, 2000 Jun 3, 2000 Jun 4, 2000 Jun 5, 2000 Jun 6, 2000 Jun 7, 2000 Jun Date collected Date Debris Beach, Attu I. Attu Debris Beach, I. Attu Debris Beach, I. Attu Alexi Point, I. Attu Beach, South I. Adak Bay, Kuluk I. Adak Bay, Kuluk I. Adak Bay, Kuluk I. Adak Bay, Kuluk I. Shemya Shore, South I. Shemya Shore, South I. Shemya shore, North I. Shemya shore, North I. Shemya Shore, South I. Shemya shore, South I. Shemya Shore, South I. Shemya Shore, South I. Shemya Shore, South I. Unalaska Bay, Summer Akutan I. I. Attu Beach, South I. Attu Beach, South I. Attu Point, Pratincole-Murder I. Attu Beach, South I. Attu Beach, South I. Attu Alexi Point, I. Attu Beach, South I. Attu Beach, Tower Loran I. Attu Beach, South Collection location Collection were published in Lindstrom et al. (2015). et al. published in Lindstrom were ------GenBank No. GenBank P. nereocystis P. , P. pseudolanceolata P. , Pyropia fallax (Kjellman) Toni De Continued A91868, A91870 UBC A91779 UBC A91719, A91780 UBC A91723, A91724 UBC A91783 UBC A91897, A91786, A91787 UBC A91790, A91791, A91792, A91793 UBC A91807, A91808 UBC A91698, A91827, A91829 UBC A91712, A91834, A91835 UBC A91885, A91841 UBC A91886 UBC A91887 UBC A91843 UBC A91847, A91848 UBC A91884, A91851, A91853, A91855, A91857 UBC A91767, A91768, A91859 UBC A91770, A91771, A91772, A91773, A91866, UBC A91915 UBC A91892 UBC A91883 UBC A91684 UBC A91689, A91690, A91691 UBC A91693 UBC A91700 UBC A91701, A91702 UBC A91717 UBC A91721 UBC A91725 Wildemania variegata variegata Wildemania Accession / Collection No. / Collection Accession Extract number. Sequence not accessioned in GenBank. not accessioned Sequence Extract number. In 2012, the name of Rat Island was changed to its native name, , after rats were eradicated from the island in late 2008. the island in late from eradicated were after rats Island, Hawadax name, In its native 2012, the name of Rat changed to Island was numbers in brackets refer to silica gel material, which was numbered separately from the herbarium vouchers. Additional collecting details can be found for each specimen in the UBC algal for collecting details can be found Additional the herbarium vouchers. from separately numbered which was silica gel material, to refer ALEUT numbers in brackets her UBC the into accessioned be to yet collectors´ have numbers the by only identified Specimens herbarium(http://bridge.botany.ubc.ca/herbarium/search.php?db=ubcalgae.fmp12). database of records barium. Additional a b Appendix 1. Appendix

263 http://e-algae.kr