KEGLO .\),o BAY
D. MUNRO, SP. NOV
32. STACHYS TENUIFOLIA WILLD
Stachys tenuifolia Willd., Sp. PI. 3: 100, 1800 (Pennsylvania), Muhlenberg (Holotype: B, mi-crofiche!, photographs in DAO!, and LA in UC!).
Stachys glabra Riddell, Suppl. Cat. Ohio PI. 16: 580, 1836. Ohio, low woods, Worthington, August, Riddell (Holotype: US!, photograph in DAO!).
Stachys palustris L. var. glabra (Riddell) Gray, Man.
Bot. Northern U.S., ed. 2, 317, 1856.
Stachys aspera Michx. var. glabra Gray, Synopt. FI.
N. Am. 2: 387, 1878.
Stachys cincinnatensis Kuntze, Rev. Gen. PI. 2: 531, 1891. Ohio, near Cincinnati, Kuntze (Holotype:
NY).
This wide ranging species occurs from New Jersey and Pennsylvania in the northeast to South Carolina in the southeast, to Texas in the south-west and to Kansas, Iowa and Wisconsin in the west and northwest. It is concentrated along the
Mississippi River drainage basin from Iowa to In-diana, Pennsylvania, West Virginia, south to Louisiana and Alabama. It is most common in moist locations, especially along the margins of woodlands.
2n = 34. INDIANA. Adams Co., D.B.M. (D.B.M. &
G.A.M. 2180); Huntington Co., D.B.M. (D.B.M. & G.A.M.
2183).—VIRGINIA. Greensville Co., D.B.M. (D.B.M. &
G.A.M. 2418). — LOUIS1ANA. West Baton Rouge Co., D.B.M. (D.B.M. & G.A.M. 2430). — KANSAS. Franklin Co., D.B.M. (D.B.M. & G.A.M. 2472); Miami Co., D.B.M.
(D.B.M. & G.A.M. 2473). — MISSOURI. Newton Co., D.B.M. (D.B.M. & G.A.M. 2474). — KENTUCKY. Trigg Co., D.B.M. (D.B.M. & G.A.M. 2477). — WEST VIR-GINIA. Marion Co., D.B.M. (D.B.M. & G.A.M. 2485).
2n = 68. WEST VIRGINIA. Meade Co., D.B.M.
(D. B. M. & G.A.M. 2480); Cabell Co., D.B.M. (D.B.M. &
G.A.M. 2482).
Lewis et al. (1962) reported n = 16 for plants of this species from Texas. We believe that this count should be n = 17. Gill (1980, 1981) pub-lished the number n = 32 for S. tenuifolia plants from one location in Québec and two in Ontario.
However, S. tenuifolia does not occur in Canada and the key in Gill (1980) clearly shows that his plants were S. hispida not S. tenuifolia. In addition, the number undoubtedly should be n = 34, not n = 32. We conclude that S. tenuifolia contains diploid and tetraploid populations with the base number x = 17. Although tetraploid plants of S. tenuifolia grown in the greenhouse in Ottawa tended to be more robust than diploid plants, we could not find any characters that consistently en-abled us to distinguish plants at the two chromo-some levels. The somatic chromochromo-somes of S. tenuifolia are almost equal in length.
Conclusions
Thirty-two species of Stachys occur in North America north of Mexico, 25 of them are native to this area and seven are introduced from Europe.
The very widespread native S. pilosa (2n = 68) has often been confused with S. palustris (2n = 102), an introduced plant with a more restricted distribution. In addition, to having a different chro-mosome number, being quite different morpho-logically and occupying different habitats (Mulligan et al., 1983), we have shown here that S. palustris has a seed testa topography quite distinct not only from S. pilosa but also from other Stachys native to our study area.
Our 25 native Stachys form two distinct groups of species. One group contains species with the basic chromosome number x = 17 or derivatives of it, and the other contains species with a wide range of other numbers and a unique pollen apo-colpia morphology.
50 LE NATURALISTE CANADIEN, VOL 6, 1989
There are eleven native species with the base number of x = 17: 6 are diploid (S. bergii, S. eplingii, S. floridana, S. hyssopifolia, S. nuttallii and S. rothrockii), 2 species have both diploid and tetraploid plants (S. latidens and S. tenuifolia), and 3 species are only tetraploid (S. aspera, S. hispida and S. pilosa). Ail of these, except S. bergii, S. pilosa and S. rothrockii, are restricted to the eastern half of our study area and ail of them have 34 and/or 68 somatic chromosomes that are about equal in length. Stachys bergii is a local endemic on serpentine deposits in Del Norte County, Ca-lifornia, whereas S. pilosa, a very widespread species, is absent from most of California and the western parts of Oregon, Washington and British Columbia. Stachys rothrockii is localized in Ari-zona, New Mexico and Utah.
There are eight native Stachys with 66 somatic chromosomes: Stachys ajugoides, S. albens, S. bullata, S. mexicana, S. pycnantha, S. rigida, S. stebbinsii and S. stricta. Sixty-four of their chromosomes are about the same size as in spe-cies with 34 and/or 68 somatic chromosomes, and the remaining two are about twice this size. In ad-dition, there is one species, S. chamissonis, that has 64 somatic chromosomes, 60 of them about the same size as those found in species with the base number of x = 17 and four chromosomes approximately twice this size. Ail the species with 66 chromosomes are native to California with some species occurring northward, in extreme western North America, to Annette Island in Alaska. Sta-chys chamissonis is a coastal species that is native from California to southwestern British Columbia.
We believe that native species of Stachys with the base number of x = 17 are closely related, with tetraploids derived from diploids. In generai, te-traploids, with this base number, have a much wider range than diploids. It seems likely that Sta-chys species with the chromosome numbers 2n 66 and 2n = 64 are derived from tetraploid ancestors with the base number of x = 17, and that the reduction of somatic chromosome num-bers to 66 and 64 are the resuit of the fusion of material from pairs of somatic chromosomes. It is interesting that S. chamissonis and its variety cooleyae, both with the reduced chromosome number of 2n = 64, are the only taxa in this group of species that are commonly pollinated by hum-mingbirds (Grant & Grant, 1968) and that have a large number of punctae per lumen on the me-socolpia of their pollen (Bassett & Munro, 1986).
The other Stachys examined cytologically have quite different chromosome numbers: S. agraria, 2n = 32; S. coccinea, 2n = 84; S. drummondii, 2n = ca 80-82. These three species, along with S. bigelovii, have the northern part of their distri-bution in the south-central United States. Bassett
& Munro (1986) reported that these species, along with six others native from Mexico southward into South America, have pollen with tectate-perforate apocolpia. These six species are; S. boraginoides Cham. & Schl., S. costaricensis Briq., S. eriantha Benth., S. grahami Benth., S. petiolosa Briq. and S. tenerrima Epl. The pollen that they examined in material of 38 other Stachys received from North American, European and Japanese sources ail had microreticulate apocolpia. Bassett & Munro (1986) suggested that the ten species of southern Stachys with unique tectate-perforate apocolpia may be part of a large group of species that is quite distinct and should be recognized in some formai taxonomic manner. Further studies on the chro-mosome numbers, pollen morphology and other characteristics of species native to south-central United States southward into Mexico, Central America and South America are needed to further clarify this situation.
Acknowledgments
We would like to thank the many persons who sup-plied us with living material, herbarium specimens, pho-tographs and gave us other assistance. We especially want to acknowledge the valuable advice and other contributions of Dr J. B. Nelson. We also thank Walter Wojtaà for the preparation of many slides for chromo-somal studies, Dr Jacques Cayouette for assisting us with the Latin description for S. stebbinsii, and Dr Guy Bail-largeon for the French résumé.
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ARCTIC ALGAL COMMUNITIES IN THE REGION OF THE NUVUK ISLANDS, NORTHEASTERN HUDSON BAY, CANADA'
D. W. KEATS2, J. M. GREEN and R. G. HOOPER Department of Biology and Ocean Sciences Centre
Memorial University of Newfoundland, St. John's (Newfoundland) Al B 3X9 Résumé
Les communautés d'algues marines benthiques de la région des îles Nuvuk, près de l'extrémité sud-ouest du détroit d'Hudson, ont été étudiées par plongée (SCUBA) en juillet 1982 et 1983, et en août 1984. Seize espèces de chlorophytes, 29 de phéophytes, et 29 de rho-dophytes ont été identifiées. La zone littorale supérieure et moyenne, soumise à une forte abrasion par les glaces, est occupée par une bande d'Utothrix flacca (Dillwyn) Thuret in Le Jol. et autres micro-algues. Une ceinture dense de Fucus evanescens Agardh se rencontre dans le littoral inférieur. Sur les substrats durs, l'infralittoral supérieur est soumis à une forte abrasion par les glaces et est généralement occupé par une communauté d'algues annuelles.
Aux étages inférieurs de l'infralittoral, on trouve des forêts de laminariales caractérisées par une richesse spécifique élevée, une forte productivité et une grande abondance d'invertébrés.
Abstract
Benthic marine algal communities were studied using SCUBA diving in the vicinity of the Nuvuk Islands near the southwestern entrance to Hudson Strait. Studies were conducted in July 1982 and 1983, and August 1984. Sixteen chlorophytes, 29 phaeophytes and 29 rho-dophytes were identified. The high to mid intertidal zone was heavily ice-scoured, and was covered by a green band of Ulothrix flacca (Dillwyn) Thuret in Le Jol. and other microalgae.
The low intertidal zone had a dense band of Fucus evanescens Agardh. On hard substrate, the upper subtidal zone was heavily ice-scoured, and was generally covered by a dense growth of ephemeral algae. Below this, kelp beds characterized by a high species richness, apparently high productivity, and an abundance of invertebrates, predominated.
Introduction
Floristics and community composition of benthic marine algae of the Canadian Arctic are very poorly known. Lee (1980) published a checklist sum-marizing previously published distribution records as well as his own collections. Wilce (1959) and Hooper & Whittick (1984) described the marine algal flora of the Labrador region, and Dunton (1984) analyzed the structure of western arctic kelp beds. These reports indicate that rich, high-bio-mass algal communities are common in the Arctic where substrate and oceanographic conditions are suitable. In northwestern Québec the only pub-lished report is of vine seaweed species collected from Ivugivik by Lee (1980). He did not describe seaweed communities.
' Contributions of the Ocean Sciences Centre no. 75.
2 Present address: Botany Department, University of the Western Cape, P. Bag X17, Bellville 7530, South Africa.
A series of field trips to the Nuvuk Island region of northeastern Hudson Bay in 1982, 1983, and 1984 provided the opportunity to examine marine algal communities there in some detail. The pur-pose of this paper is to describe the species com-position and dominant zonation patterns of marine algae in this region.
Materials and Methods STUDY AREA
The Nuvuk Islands are located in southwestern Hudson Strait (Fig. 1) and consist of a series of small islands interspersed with narrow, shallow marine channels approximately 100 m in width and less than 10 m in depth. The maximum tidal am-plitude is about 2 m. Strong semidiurnal tidal cur-rents sweep through the channels between islands. Substrates are silty with mid-channel re-gions of unsorted gravel, cobble and scattered small and large boulders. The sides of channels
54 LE NATURALISTE CANADIEN, VOL. 116, 1989
consist of exposed bedrock, gravel and, in some cases, soft mud. Gaston et al. (1984) described the general oceanographic features and naturel history of the region, and Gilkinson et al. (1986) provided a description of the shallow water marine benthic molluscs of the study area. While the sites described by Gilkinson et al. (1986) are the same as those described in this paper, the site numbers used in the present study have been arranged in order of decreasing exposure to wave action.
FIELD COLLECTIONS AND OBSERVATIONS Observations and collections of marine benthic algae were made during 40-50 person hours of SCUBA diving at sites in the region of the Nuvuk Islands in July 1982 and 1983. In 1984 more in-tensive collections and observations were made during ca 20 person hours of diving at six sites (Fig. 1). Observations and collections were made while systematically crossing the sites from the intertidal zone to the maximum depth, and these were supplemented by casuel observations and collections made by divers who were collecting fish. Species composition was determined
ac-65.
60
cording to depth, substrate. and locality. Speci-mens were identified usina the classification and nomenclature of South & Tittley (1986). Voucher specimens of most species are deposited in the Phycological Herbarium of Memorial University of Newfoundland (NFLD).
PHYSICAL DESCRIPTION OF SITES
Site 1 is located outside of the main channels, and is fully exposed from the southwest through west to the north. Towards the shore there is an extensive shallow area (2 m) protected by an off-shore shoal. Beyond the shoal the slope is mod-erate to a depth of ca 10 m. At 10-15 m, the bedrock and boulder substrate gives way to hard packed sand.
Site 2 is moderately exposed to swells from the northeast. It has a near vertical bedrock slope to the maximum diving depth of 30 m. The bedrock is broken by crevices and ledges, and an occa-sional boulder.
Site 3 is partially exposed to swells from the northeast, but is still relatively sheltered. The slope
Figure 1. Map of the study area. Numbers 1-6 show collecting sites.
KEATS ET AL ARCTIC ALGAL COMMUNITIES 55 is very steep (>45°) consisting primarily of bedrock
down to about 8 m, and bedrock with very large boulders to the maximum depth of 22 m.
Site 4 is a moderately exposed cove on the western side of Nuvuk Island. It is exposed to swells from the northwest, but is relatively pro-tected from other directions. The head of the cove is a beach, made up of sand and Pleistocene shell deposits. The sides of the cove consist of steeply sloping bedrock down to about 4-5 m. The centre of the cove consists of large boulders mixed with sand, which give way to small rocks and sand to-ward the beach.
At site 5, which is fully sheltered, collections were made from the main island, across the channel to the smaller island to the south. On the north side of the channel, the substrate is mainly bedrock in the intertidal and upper subtidal zones, although there is a boulder beach near the col-lecting site. This slopes steeply to ca 3 m, where there are mainly boulders and large rocks. Below this (ca 4 m) there is a relatively bare, muddy, ice-scoured area, with boulders and large rocks along the mid channel side. The central channel area (maximum depth of 7 m) consists mainly of boul-ders, large rocks and gravel interspersed. The lower intertidal zone along the south side of the channel consists of a boulder barricade, above which there is mainly a mixture of gravel and mud.
Strong semidiurnal tidal currents sweep through the channel.
At site 6, which is also fully sheltered, the sub-strate is mainly muddy silt with scattered cobble and boulders. The slope is quite gentle, and a maximum depth of 7 m was reached. The upper intertidal zone is mainly relatively bare mud.
Semidiurnal tidal currents occur through this channel, but they are not as strong as those at site 5.
Results
We recorded 16 species of Chlorophyta, 29 Phaeophyta and 29 Rhodophyta (Table I). We also recorded one chrysophyte; other microalgae were not identified. One common terete dichotomously branched rhodophyte could not be positively identified because it was sterile, and is probably an undescribed species.
SPECIES COMPOSITION OF SITES Site 1
The upper to mid-intertidal zone had a green band, composed mainly of Ulothrix flacca, mixed with other filamentous microscopic green algae and diatoms. Below this was a dense zone of
Fu-cus evanescens. Behind the protection of the shoal, Fucus extended well into the subtidal region, where it was mixed with Alaria esculenta, Lami-naria digitata, L. longicruris, L. solidungula, Sac-corhiza dermatodea, and Agarum cribrosum.
There were also abundant understorey rhodo-phytes in the shallow area, especially Devaleraea ramentacea, Odonthalia dentata, and Dilsea integra.
Beyond the shoal, the above community gave way to a Laminaria longicruris bed. The L. longi-cruris was a narrow, short, hollow stiped form, which formed dense impenetrable thickets, sep-arated by patches of L. solidungula, A. cribrosum, and other algae. The latter kelps were also mixed with L. longicruris, and L. solidungula was common even in the dense thickets. The understorey rho-dophytes mentioned above were still prevalent.
On isolated patches of hard substrate within the sand-dominated bottom 10-15 m in depth, L. so-lidungula and A. cribrosum were prevalent, and understorey rhodophytes were abundant.
Sites 2 and 3
These sites showed the same intertidal
These sites showed the same intertidal