Dissertation fat the” (”glreé of‘Fli D ALA/:7 ’ ,'.,.-: I 32'; A " “A W , H’M'u» ,,....,/;J-»}‘A ,1 ,,,..-—., 7’“ . 5 M. r 0. r' m: In, 'x by lg: n xv m» jfflr A, A REVISION 0? TH: SPECIES OF sAemA I r . _,(_ «1.;mmguuf ' JAIMMI 4n “n ’4‘» .""'.'3‘¢ " xyn‘ri'l -‘_‘, ”If . ,Hzmmwma 4,; (gym/v w my ’IW, Ff .15 «flint/r ‘Tr’W/‘l l' 2;, 3/“; K 1:3- I) “1.4!! ”3542...:r; .13?" W: r“- ”war." rpm :7" . r ‘ r-rw ('0’: n ,1. .- .335: {my v ...4. «VIA-111580: pr» «val—mm flAr‘v 1 Allan 4p . :, ”w:.x.}j!§ , «3- rm": 3 .4. .- tflm ,A or». «r A I ”I'll!‘ I flrr_. 4r 1-!”- r ;- A a. .1 <. ”a” 51:1”, 1‘) diff m 15,51.” ”5...» 4:1in 1:;037 ”of”; w x ”i ' £5.13": £1:an 4_..' 5:“ 3;; This is to certify that the thesis entitled A REVISION OF THE NORTH AMERICAN SPECIES OF SAGINA (CARYOPHYLLACEAE) presented by Garrett E. Crow has been accepted towards fulfillment of the requirements for ____Bh_rD4_degree in W /g{,{/W Major professor Da A1). 11 5, 1974 1 - ,l 0—7639 ABSTRACT A REVISION OF THE NORTH AMERICAN SPECIES OF SAGINA (CARYOPHYLLACEAE) By Garrett E. Crow The genus gaging consists of perennial and annual herbs indigenous to the cool temperate regions of the Northern Hemisphere. About 15 species are now recognized in the genus, l0 of which occur in North America, either as native or adventive taxa. Two lines of divergence are recognized at the sectional level. These are section §agina, with a center of diversity in Eurasia, and section Magima, with a center of diversity in eastern Asia. The latter section is newly described. Two new combinations at the subspecific level have also been made. A phenetic diagram has been constructed to provide a graphic representation of the interspecific relationships of the North American taxa. An index of divergence has also been prepared for the taxa concerned. Sagina nodosa is regarded as the most primitive species in section_§agina and §: maéima is considered primitive in section Mgfijma. Most of the species appear to be polythetic and have been rec- ognized on the basis of combinations of various characters. For the most part the species of §agina_are inbreeders. Sterility barriers appear to be weak in the genus, while actual occurrence of hybrids is Vi Garrett E° Crow .55 to no . . . \ largely reduced due to the inbreeding reproductive system and by §\1 different ecological preferences or flowering times of the taxa. An analysis of the seed morphology, utilizing the scanning electron microscope, has been helpful in assessing relationships within the genus. This study included European and east Asian taxa as well as North American material. Two basic types of seeds occur and are diagnostic for sectional subdivisions of the genus. The saginoid seed type, characteristic of section §agina_is obliquely triangular, possesses a dorsal groove and has its lateral surfaces drawn inward. The crassuloid seed type, characteristic of section Magima, is more nearly reniform or globose, lacks a dorsal groove and its lateral surfaces remain full and plump. The geographical distribution of §agina_species have been evaluated in the light of effect of Pleistocene glaciation. An attempt has been made to determine where the taxa may have survived the glacial advances. A REVISION OF THE NORTH AMERICAN SPECIES OF SAGINA (CARYOPHYLLACEAE) By u°> Garrett E° Crow A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology 1974 ACKNOWLEDGMENTS I wish to express my sincere appreciation to my major professor, Dr. John H. Beaman, for his guidance and encouragement during this study. Drs. A. T. Cross, J. Harman, S. N. Stephenson and w. Tai provided helpful comments and criticism concerning the manuscript. Special thanks is also expressed to Dr. John H. Thomas who introduced me to the taxonomic problems in Sagina and provided the Stanford University specimens for a preliminary investigation. He has been a source of encouragement to me throughout the study. Interaction with my fellow graduate students has also been greatly appreciated. During my field work several persons took time to guide me to collecting sites: Drs. E. K. Longpre and H. Barclay in Colorado, Dr. J. H. Thomas in California and Dr. E. G. Voss in Ontario. Mr. Robert L. Watson accompanied me in the field. Special thanks is extended to Dr. H. Z. Snyder, who provided travel support for field work through the Snyder Foundation. Mr. William MacAfee, of the Electron Microscope Laboratory, Michigan State University, took the SEM photographs. Dr. J. K. Morton graciously provided me with his unpublished chromosome counts of North American material of Sagina. I also wish to thank the curators of the herbaria from which specimens have been used for this study. These herbaria are listed at the beginning of the taxonomic treatment. A special word of appreciation goes to my wife, Charlyn, whose love, patience, and encouragement has been an important factor in the completion of this dissertation. ii TABLE OF CONTENTS LIST OF TABLES ....... . .................. LIST OF FIGURES ......... INTRODUCTION ..... HISTORICAL ACCOUNT ........ CHROMOSOME NUMBERS ...... EVOLUTION ........ Divergence: section Sagina Divergence: section Maxima PHYTOGEOGRAPHY . SEED MORPHOLOGY . . . . ..... TAXONOMIC CONCEPTS ..... CONSTANCY OF CHARACTERS FLORAL MORPHOLOGY . ..... Inflorescence . Flower . . ...... 000000 0 O O 0 O 0 000000 O O u 0 O 0 0 O 0 0 O O 0 O O O O 0 0 0000000000 OOOOOOOOOO 0000000 0000000000 0000000000 0000000000 0000000000 0000000000 00000000 0000000000 000000 0000000000 00000000 U 0 O O U 0 O 0 0 0 u o o o o o o o o .- Caitx Jy. u o o o o o o ooooooooooooooooooooo Corolla . . . . ...... Androecium .......... Gynoekcj um u o 0 o o o a o o 000000 L) 0 0000000000 0000000000 0 O O 00000 0 0 o o 0 0 F ' ruit . . ......................... 0 Seed . . . . . . . ...... POLLINATION . . ..... O 0 U 0 0000000000 O O O 0 O O U 0 DISPERSAL . . . . . . . . . . . . . . . . . . . HYBRIDIZATION IN SAGINA . . . . . 0 0 o o O 0 15 22 24 34 55 56 58 58 58 63 63 64 64 69 59 72 74 75 SPECIMENS EXAMINED . . . .............. 0000000 TAXONOMIC TREATMENT ....................... Key to North American Species and Infraspecific Taxa of Sagina ...................... l. Sagina nodosa . . . . . . . . . . . . . . . . . .... la. Sagina nodosa var. nodosa . ............ 1b. Sagina nodosa var. pubescens ......... 2. Sagina saginoides . . ................. 3. Sagina procumbens ................... 4. Sagina subulata .......... . . ........ 5. Sagina nivalis .......... . . . ........ 6. Sagina caespitosa . . . . . .............. 7. Sagina decumbens . . .................. 7a. Sagina decumbens subsp. decumbens ......... 7b. Sagina decumbens subsp. ocCidéntalis 8. Sagina apetala ..................... 9. Sagina maxima ..................... 9a. Sagina maxima subsp. maxima ............ 9b. Sagina maxima subsp. crassrcaulis ......... TO. Sagina japonica . . . . . . . . . . . . . . . . . . . . LITERATURE CITED . ....... . . ........ iv 81 86 86 98 102 124 141 148 155 164 164 178 187 200 200 205 217 222 TABLE LIST OF TABLES Chromosome Numbers for Taxa of Sagina Occurring in North America ...................... Characters and Character States in Sagina ........ Tabulation of Character States for the North American Taxa of Sagina ...................... Computed Differences Between North American Taxa of Sa ina ,,,,,,,,,,,,,,,,,,,,,,,,,, Comparison of Primitive and Advanced Character States in Sagina ........................ Index of Divergence for North American Taxa of Sagina. Tabulation of Character States .............. Comparative Features of Sagina Sections Sagina and Max-Ema o o o o o o o o o o u o o o o o o u o o 9 0 0 0 “— Page 10 16 17 84 ..__-_ .._.. FIGURE 1. 10. 11. 12. 13. LIST OF FIGURES Phenetic diagram for taxa of Sagina occurring in North America ..... " ................. Index of divergence for North American taxa of Sa ina °°°°°°°°°°°°°°°°°°°°°°°°°° Presumed relationships of taxa of Sagina occurring in North America ..................... SEM photographs of Sa ina seeds, saginoid type. (a) S. saginoides, b S. prOCUmbens, (c) S. subulata, (d) S. decumbéns subsp.—occiHEntalis ........... SEM photographs of Sagina seeds, saginoid type. (a-d) S. decumbens subsp° decumbens ........... SEM photographs of Sa ina seeds, saginoid type. (a-c) S. apetala, (d S. maritima . . . . . . . . . . . . SEM photographs of Sagina seeds, saginoid type. (a) S. glabra, (b) S. caespitosa, (c-d) S. nivalis . . . SEM photographs of Sagina seeds, intermediate between saginoid type and crassuloid type. (a~b) S. nodosa, (cad) S. abyssinica . . . . . . . ............ SEM photographs of Sa ina seeds, crassuloid type. (a) S. maxima subsp. maX1ma, (b) S. maxima subsp. crassicaulis, (C).§° paupuana, (d7 S. subuletorum SEM photographs of Sagina seeds, crassuloid type. (a-d) é.” jalgonica D 0 0 0 0 0 0 0 0 u 0 0 0 O O o 0 9 J 0 Photographs of flowers of Sagina nodosa ......... SEM photographs of Sagina nodosa pedicel showing glandular hairs ,,,,,,,,,,,,,,,,,,,, SEM photographs of Sagina pollen grains. (6-b) §3 nodosa, (c-d S. nivalis ............ vi Page 18 23 42 44 46 48 50 52 54 6D 62 66 - Afi-“.__, m-__. Page FIGURE 14. Papillate stigmas. Sagina maxima subsp. crassicaulis ....................... 68 15. Seed types. (a) Saginoid, Sagina saginoides, (b) Crassuloid, S. maxima subsp.crassicaulis ...... 71 16. Photographs of Sagina nodosa var. nodosa ......... 89 17. Geographic distribution of Sagina nodosa var. fl9g9§a_ .......................... 97 18. Geographic distribution of Sagina nodosa var. m ........................ 104 19. Photographs of Sagina sagjnoides ............. 108 20. Geographic distribution of Sagina saginoides in North America ...................... 123 21. Photographs of Sagina procumbens. (a) Flowers showing cupped sepalsand papillate stigmas, (b) Adventitious roots produced on procumbent stem of weedy plant in growth chamber ...................... 127 l 22. Photographs of Sagina procumbens. (a-b) Habit ...... 129 f i 23. Geographic distribution of Sagina procumbens in ' North America ...................... 143 24. Photographs of Sagina subulata .............. 146 25. Photographs of Sagina nivalis .............. 151 1 26. Geographic distribution of Sagina nivalis in ' North America ...................... 157 27. Photographs of Sagina caespitosa ............. 160 28. Geographical distribution of Sagina caespitosa in North America ...................... 163 29. Photographs of Sagina decumbens subsp. decumbens ..... 167 30. Geographical distribution of Sagina decumbens subsp. skgagflgyyi ........................ 176 181 3l. Photographs of Sagina decumbens subSp° occidentalis vii Page FIGURE 32. Geographical distribution of Sagina decumbens subsp. occidentalis ....................... 185 33. Photograph of holotype of Sagina occidentalis Nats. (= Sagina dchmbens subsp.'occidentalis) ......... 189 34. Photographs of Sagina apetala. (a) Living specimen, (b) Close-up showing glandular trichomes on pedicel and cilia of leaf base .................. 193 35. Photographs of Sagina apetala. (a-b) Habit ........ 195 36. Geographical distribution of Sagina apetala in North America ...................... 199 37. Photographs of Sagina maxima subsp. maxima ........ 204 38. Geographical distribution of Sagina maxima subsp. maxima_in North America ................. 207 39. Geographical distribution of Sagina maxima subsp. crassicaulis ....................... 214 40. Photograph of holotype of Sagina crassicaulis Mats. (= Sagina_maxima subsp. crassicaulis)— .......... 216 41. Photographs of Sagina japonica .............. 220 viii INTRODUCTION Sagina (Pearlwort), a genus of the Charyophyllaceae, consists of about l5 species indigenous to the cool temperature regions of the Northern Hemisphere. The genus is well defined, although it is occasionally confused with superficially similar taxa of Spergularia and Arenaria. Confusion with taxa of Colgbanthus, the genus most closely related tongging (Pax and Hoffmann, 1934), seldom occurs because Colobanthus is a circumaustral genus. There has been, however, confusion within the genus regarding delineation of taxa. Wright (l935, p. l) commented l'I find among my i friends many who are unwilling to give a definite opinion on Saginas, regarding them difficult to determine. I think such Opinion arises from inadequate realisation of the extreme variability of these plants." The extreme variability within the genus has generated nomenclatural recognition of numerous variants, especially in Eurasia, the primary 1 center of diversity for the genus. Many of these taxa were based on characters which are variable. Previous work of a revisionary nature in Sagina_is limited. Those works include only William“s (1918) revision of the British species of Sggjna and a treatment of the species of Saging occurring in Japan by Mizushima (1960). My study was undertaken with the intention of assessing the variability within the genus and attempting to clarify interspecific and intraspecific relationships. Nearly 6000 herbarium specimens, Eurasian and east Asian as well as North American, were examined. Most of the North American taxa have also been studied in the field and grown in a growth chamber. An analysis of the seed morphology, utilizing the scanning electron microscope, has been helpful in assessing relationships within the genus. The revision should first of all contribute to a better understanding of the North American species. Secondarily, it may contribute to the knowledge of the entire genus and to a resolution of problems in the classification of other genera of the subfamily Alsinoideae. HISTORICAL ACCOUNT Botanical history of Sggjgg begins in the 17th Century with the finding of S. gggggg by John Goodyer on August 12, 1626, ”on the boggy ground below the red Well of Wellingborough in Northamptonshire" in Great Britain, recorded in Johnston's Herball in 1633 under the name "Saxifraga palustris alsine folia” (Williams, 1918; Druce, 1932). In 1719 Dillenius included the Pearlworts under the generic name Alsinella in his Catalogus Plantus, a pre-Linnean name perpetuated by Hill (1756) in The British Herbal and by Greene (1891) in Flora Franciscana. The generic name Sagina first appears in print in Linnaeus' Systema Naturae in 1735. His Genera Plantarum (1737) indicates it was based on S. procumbens. In_Species Plantarum (Linnaeus, 1753) the genus included two tetramerous members of the tribe Alsineae, Sagina‘procumbens and Moenchia erecta, and a third species, Bartonia virginica, now recognized as belonging to the Gentianaceae. Presl (1826), in his Flora Sicula, was the first to include any of the exstipulate Spergulas in the genus Sggigg. Dumortier (1827), on the other hand, recognized the non-stipulate Spergulas of Linnaeus as a distinct genus, Phglgg. Reichenbach (1827) likewise regarded this group as a separate genus, giving it the name Spergella. Fenzl's (1833) redefinition of Sggigg_retained only S. procumbens of Linnaeus' genus and incorporated Reichenbach”s Spergella. Koch (1837), in the first edition of Synopsis Florae Germanicae e§_Helveticae retained Reichenbach's Spergella in the genus Spergula as sect. Spergella. In the second edition (Koch, 1843) he transferred sect. Spergella to Sggjgg. thus erecting the subdivisions in Sggiflg_as sect. Saginella, including the 4-merous taxa, and sect. Spergella, comprising the 5-merous taxa. Although this subdivision is sometimes followed in European floristic works, it neither adequately reflects natural relationships, nor provides a practical basis for classification. Not only are there both 4- and 5-merous plants within taxa, but individual plants may have both types of flowers. Moss (1920), in this treatment of Saging_for the Cambridge British Flora regarded the species as being too closely allied to be meaningfully subdivided into distinct groups higher than series. Thus, he treated the British Saginas in four series: Nodosae, Subulatae, Procumbentes, and Apetalae. With the exception of ser. Apetalae his groupings are quite natural. However, they deal only with the British species. Mizushima (1960) likewise concluded that generic subdivision does not merit rank above series level and attempted to apply Moss' series to the east Asian Saginas, but found it necessary to redefine these series. Unfortunately he based these divisions on 4-merous taxa versus 5-merous taxa. CHROMOSOME NUMBERS Very little information is presently available on chromosome numbers in Sagjng. It appears from the published chromosome numbers that two basic chromosome numbers occur in the genus, xf=6 and xf=ll (Darlington and Wylie, 1956). In section ygximg_all the taxa are based on xf=ll. Both diploid and tetraploid p0pulations are known in Sagina maxima subsp. maxima (2gg=22, 44). A tetraploid count has been reported for S. japoinca (23==44) and a hexaploid count (2g =66) has been reported for S. mgxyma subsp. crassicaulis. § Section Sagina contains taxa based on xf=6 and xf=ll. The _._.A . taxa based on xf=6 are annuals and consist of Sagina_apetala, whose published count is diploid (2§;=l2) and S. decumbens, with an unpub— lished hexaploid count (2flf=36). The remaining taxa in section Sgging_ I are perennials whose chromosome numbers are based on 5 =11. Sggiflg_ i saginoides, S. procumbens and S. subulata are reported as being diploids (Zn.=22). The polyploids reported include S. nodosa (2nf=44, 56) and S. nivalis and S. caespitosa (both 2g =88). Table 1 summarizes the chromosome numbers known for the taxa of Sagina occurring in North America. A large portion of the published counts are based on European material. Dr. J. K. Morton has graciously provided his unpublished counts, all based on his collections of North American material. _______J TABLE 1. CHROMOSOME NUMBERS FOR TAXA 0F SAGINA OCCURRING IN NORTH AMERICA Reference Taxon 2g_Number S. nodosa 56 56 56 44 56 S. saginoides 22 22 22 22 22 S. procumbens 22 22 22 22 22 22 22 S. subulata 22 22 22 18 S. nivalis 84 88 c.88 S. caespitosa 88 88 100 (higher than) S. decumbens 36 suBsp. decumbens S. apetala 12 12 S. maxima 42 or 44 suEsp. maxima 44 22 S. maxima 66 su5$p. crassicaulis S. Japonica 42 or 44 Blackburn, in Tischler (1938) Blackburn and Morton (1957) Gadella and Kliphuis (1968) L6ve and L6ve (1956) Morton (unpublished) Blackburn, in Wright (1938) Blackburn and Morton (1957) Lave and L6ve (l956) Morton (unpublished) Packer (1968) Blackburn, in Tischler (1938) Blackburn and Morton (1957) Calder and Taylor (1968) Gadella and Kliphuis (1966, 197l) Morton (unpublished) Rohweder (1937, 1939) Wulff and Lindschau, in Tischler (1938) Blackburn and Morton (1957) Findlay and McNeill (1973) L6ve and Lfive (1956) Rohweder (l937, 1939) Blackburn and Morton (1957) L6ve and L6ve (1948, 1956) Morton (unpublished) Knaben (1950) L6ve and L6ve (1956) L6ve and Lave (1944) Morton (unpublished) Blackburn and Morton (l957) Diers (1961) Blackburn, in Wright (1940) Calder and Taylor (1968) Taylor (1967) Calder and Taylor (1968) Blackburn, in Wright (1940) EVOLUTION Two centers of diversity are present in Sggjflg. A primary center is located in Eurasia (sect. Sggjng) and a secondary center occurs in eastern Asia (sect. Mgéimg). In order to provide an assessment of interspecific relationships a phenetic diagram was constructed. The method is one described by Wiffin and Bierner (1972) which allows the use of characters without designating which is the primitive or derived state. It is, however, necessary to designate one taxon as the most primitive. The procedure consists of the following: 1. A listing of characters used and their two states (Table 2). 2. Tabulation of the character states for each taxon (Table 3). 3. Designation of one taxon as the most primitive. 4. Tabulation of the computed differences between taxa (Table 4). 5. Construction of a phenetic diagram. The diagram produced is constructed by listing the taxa in order of absolute difference, least to greatest, from the designated most primitive taxon, Sagina nodosa. Taxa are then joined to the develOping tree in order of difference and connected to the pathway of the taxon already in the tree which is least different. 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An intermediacy in seed type is further suggestive of the primitiveness of §Efliflé_2292§22 The frequency of seeds appearing saginoid, with a distinct dorsal groove, nearly equals that of the crassuloid seed type. In terms of vegetative reproduction, however,_§. Egggsg possesses the most advanced mechanism of the genus. By the formation l of axillary bulbils which are readily detached in late autumn (Wright, l935) the species has developed a disseminule which can be dispersed by the wind. The leaves of the bulbils have a xerophytic anatomical structure (Wright, l935), thus enhancing the chances of establishment for the propagules. On Baffin Island, N. N. T., the species is reported to reproduce exclusively by bulbils (Polunin, l959). §agina saginoides, §, subulata and §, procumbens form a species complex which remains little divergent from the ancestral stock. The three species of this complex appear to be capable of interbreeding but largely maintain their integrity through inbreeding, ecological preference, and geographical affinities. §ggina saginoides is the least divergent of these three species. §agina_subulata appears to be slightly more advanced. The latter's possession of subulate leaves and a tendency toward a filiform growth habit are correlated with its tolerance of dryish sandy, gravelly or rocky habitats. 20 §agina_procumbens expresses a greater amount of divergence than §f saginoides and §: subulata, especially in the context of reproductive biology. The trend toward obligate inbreeding is evidenced by reduction of the androecium to one whorl, reduction of the petals, and a tendency for flowers to remain closed even on fair days. Wright (l935) observed that nearly all flowers of the species produced during autumn were cleistogamous. The species is weedy and has become dis— persed widely in cool temperate regions of both hemispheres. §agina_nivalis and §, caespitosa comprise a complex of the high Arctic region for which the highest chromosome counts in the genus are 1 reported (see Table 1). Both taxa are densely caespitose. Inbreeding is well developed, although §, caespitosa, with petals slightly longer than the sepals, has a greater potential for outcrossing. Vegetative reproduction, however, plays an extremely important role in the sur— vival of this high latitude complex. John Schindler (personal commu- nication, l970) notes that in the region around the Naval Arctic Research Laboratory at Point Barrow, Alaska, §f hiyalj§_has been seen to set seed once in a ten-year period. Indeed, I found it difficult to obtain well-formed, mature seeds of either taxon from herbarium material for scanning electron microscope studies. §agiga nivalis appears to have greatest affinity with §f saginoides. Where disjunct populations of §f nivalis bring the otherwise allopatric species together in alpine situations in the Cordillera of Alberta, there appears to be some intergradation and species occur which are difficult to assign to either taxon. Zl §ggina_decumbens, with subspp. decumbens and occidentalis, and §, apetala form a complex of annuals which appear to be highly divergent from the ancestral stock. Divergence in this group appears to be related to adaptation to stress of moisture deficiency and is expressed by the presence of subulate leaves and a loose, slender, much branched habit. In each taxon the habitat is one subject to a relatively brief period of moisture availability in early spring. Moisture deficiencies of late spring require rapid completion of the life cycle. Plants of all three taxa grown in a growth chamber exhib— ited the capability of producing flowers only two weeks after seeds were sown. In portions of the geographical range where climate is milder during the winter months these taxa successfully prolong their life histories by functioning as winter annuals, whereby seeds germinate under moist conditions of fall and may pass the winter in a weakly developed rosette state. §agina apetala scores highest in the divergence index. In addition to having a loose, slender habit and possession of subulate leaves, this taxon shows the greatest amount of floral reduction and more nearly approaches obligate inbreeding and cleistogamy than any other species of §agjn§, Native to Eurasia, §, apetala is weedy and has become introduced as widely as Japan (Mizushima, l960), Chile, Argentina and North America. 22 Divergence: section Maxima Sagina maxima subsp. crassicaulis, a perennial of northern Pacific coastal bluffs and gravelly—sandy beaches, appears to be the least divergent member of section Maxima. Although the taxon can be successful as an inbreeder, I have observed that the flowers generally remain open, even under dull, cloudy days, thus encouraging outcrossing. l §agjna_maxjma_subsp. maxima_is primarily an east Asiatic taxon. Like its American counterpart, its flowers generally encourage outcross— ing, but also successfully self-pollinate. Divergence, however, has proceeded a step further in floral biology. Cleistogamous flowers may be produced in autumn and sometimes in winter in portions of the range where the climate is milder (Mizushima, l960). There is also a tendency in this taxon toward an annual life history. Divergence in section Maxima parallels that in section §agina and is more fully realized in §3 japonica. This inland species has developed an annual life history and a loose, slender, much branched habit, as has occurred in §. decumbens and §3 apetala, but the primitive linear, somewhat succulent leaf form has been retained. Floral struc- ture is §. japonica expresses a trend toward obligate inbreeding. The petals are reduced, the androecium is generally reduced to one whorl, and cleistogamous, as well as chasmogamous flowers occur. This species has greatest affinities with §f maxima subsp. maxima. I Figure 3 summarizes the presumed relationships of the taxa of Sagina occurring in North America. 23 .8205 filoz E mica—:80 a mo axB 05 3220528. 8539—; .m 3%: 339% .m 3.15:- 3on o u x 33.33 5255 ammw mwgwmmfiu .mmm manage an :35 . NN Va l EN I]! flags Goo . S n x 533 .8 v3.33 noon 1. an“ I H . x 8% Bum “5832. 108: g .w 25: Sun 28285. 8mm I 12:: 3 13.32. «Ends .93 um I aa 259...: 5” vegan—mono angina «IE 9:38 31.23 9.333% aauwugu hug—vacuum 3515-5 .m Hana—“unus— novflofiwmm .w 9335 .m .3 I an 301.333 103 £30352. H0312. flown: In: 1:53 «359% .w. 333535 vuun «mono: .Im. «cognac-u 38333 .w “:3: .m (55:3 <_m< zmwkm225; no 558 PHYTOGEOGRAPHY The modern distribution patterns of boreal and cool temperate species in North America reflect the effect of Pleistocene glaciation. The postglacial distributions of Sagina must, therefore, be evaluated in this light and an attempt made to determine where the taxa may have survived the glacial advances. Circumpolar Sagina nivalis is widespread in the North American Arctic and appears to have survived the Pleistocene in the Beringian refugium. A large portion of Alaska, including the Bering Straight region and the North Slope remained ice-free during glacial advances (Péwé et_al,, l965; Heusser, l965) and the geographical distributions of numerous arctic species suggest a "Beringia radiant” pattern (Hultén, 1937). . Although portions of the high Arctic Archipelago do not show signs of glaciation (Flint, l957; Savile, l96l), this region probably did not serve as a refugium. One would not expect a region like the northwest Elizabeth Islands, with a depauperate flora and stunted plant growth, to function as a survivium (Savile, l96l). Savile is of the opinion that the flora of this region is one recently derived. The distribution of Sagifla_nivalis shows a noticeable gap in the region of the northwest Queen Elizabeth Islands, occurring on Prince Patrick Island and Axel Heiberg Island, but not between. The gap 24 25 apparently does not reflect lack of collecting, for Savile (l96l), who has done much collecting in this region, notes that the distributions of a number of widespread arctic species exhibit this pattern. Rare disjunct populations of Saging_nivalis occur southward in the alpine habitat along the Cordilleran system to Alberta. It is possible that these populations are relictual. Another possible explanation is that these populations are a result of long distance dispersal of the diaspores which have niches in newly opened habitats in the glaciated alpine of the Cordilleran range. §§£fl33_caespitosa, a rare species, exhibits an amphi-Atlantic distribution, occurring as a coastal plant in the eastern Arctic, western and southern Greenland, and as a montane plant on Iceland and in Scandinavia. The existence of coastal mountain refugia, as described by Dahl (l946) provides the most plausible explanation of survival. Dahl notes that nunataks occurring in western Greenland support a relatively rich flora. The populations occurring in the eastern Arctic of North America have distinct affinity with populations of western Greenland, pubescence of pedicel being present in both, while plants in populations of southern Greenland, Iceland, Jan Mayan and Scandinavia are completely glabrous. The Circumpolar distribution of Sagina saginoides correlates almost entirely with montane regions of the Northern Hemisphere. Hultén (l958) notes that the Pleistocene fragmented many Circumpolar distributions. This seems to be the case here. In North America this species survived Pleistocene glaciation in the southern portion of the Cordillera. Although mountain glaciation 26 occurred during the Pleistocene, Weber (l965) points out that large areas in Colorado were free of ice. In the Sierra Nevada glaciers descended to l300—2200 meters elevation but numerous refugia existed throughout the range (Wahrhaftig and Birman, 1965). Recession of alpine glaciers in the Olympic Mountains occurred during centinental expansion of the Cordilleran ice sheet, thus refugia occurred nearby at the time the Puget Sound lobe reached its maximum (Crandell, l965). Mount Albert, on the Gaspé Peninsula of Quebec, is a well known locale whose serpentine summit supports an alpine flora which contains a curious Cordilleran element including S, saginoides. Fernald (l925) considered these disjuncts to be relictual, persisting through the glacial period on this unusual nunatak, but his theory has lost much credibility. Marie-Victorin (T938) does not consider this serpentine habitat as a nunatak, but rather regards it simply as a place where arctic plants can survive while others cannot. Dr. J. K. Morton (per- sonal'communication) has been able to show that a number of Fernald's relictual endemics are not good taxa in their own right, but merely environmentally induced growth forms of other taxa, thus further dis- crediting Fernald's nunatak theory. The presence of S3 saginoides at one locality on the eastern shore of Hudson Bay, one locality near Shefferville, Quebec, on the Labrador-Quebec Peninsula, and on the Gaspé Peninsula seems better explained as the result of long distance dispersal. The diaspores of the Saginas clearly have properties con- ducive to long distance dispersal by wind (Ldve, D., l963; Van der Pijl, I969). These isolated populations illustrate well "a case-in-point” in 27 support of ”Baker's Law“ regarding long distance dispersal which basically states that for self-compatible taxa a single propagule is sufficient to start a new colony and that establishment is much more likely than is the chance of establishment for self—incompatible taxa (Baker, 1955, 1967). Sagina maxima subsp. crassicaulis, a strictly coastal taxon, was largely unaffected in the lower and major portion of its range. To the north, however, its range was abruptly truncated by the Cordilleran ice sheet. Hultén (1937) regards the taxon as one of his ”Western America Coast Radiants.” Migration northward along the coast was likely rapid during the post-Pleistocene as coastal winds may have facilitated rapid dispersal for species with light disseminules (Calder and Savile, 1960; Savile, 1961). Such dispersal would have been enhanced during periods when sea level was lower (Heusser, 1960). Heusser (1960) notes that some disapores are quite capable of traveling long distances over water, citing as evidence the composition of the flora of Middleton Island, Alaska. Sagjga_mnggg_subsp. crassicaulis (as S. crassicaulis) is recorded in this flora (Thomas, 1957). The range of S, maxima subsp. crassicaulis reaches northward and westward to Attu Island in the Aleutians. With the retreat of the ice along the Alaskan coast, the Asiatic S. maxima subsp. maxima was likewise able to extend its range along the newly opened North American Pacific coast. Intergradation between the two taxa is in evidence on the Queen Charlotte Islands and Vancouver Island. 28 While the region south of the glacial boundary provided the major source of plants for the revegetation of western North America, nunataks along the coast of British Columbia and Alaska served as refugia during the Pleistocene. One of these was the Queen Charlotte Islands, an island complex which is noted to have both floristic endemics (Calder and Taylor, 1968) and faunal distinctions reflecting separatiOn from mainland relatives (Heusser, 1960). It is entirely possible that Sggjfla_mgximg_subsp. ngima or perhaps even both subsp. maxima and subsp. crassicaulis could have existed as refugees on this nunatak. Hultén (1937) considered S: ngigg_subsp. ngima (as S. litoralis Hult.) as a Beringia radiant. Locating a survivium for Sagina_nggg§a_presents a problem, as it occurs entirely within the glacial boundaries of eastern North America. A number of northern species display this distribution pattern, perhaps reflecting a periglacial element (Crow, 1969). Some southward migration of northern species was made possible in part by slightly cooler climatic conditions and in part by lack of competition on the newly exposed coastal terraces and alluvial deposits during withdrawal of the sea (Braun, 1947). Sagiga_flgg9§§_would be well adapted to the periglacial situation, for it is a successful pioneer plant on rocky shores and gravelly beaches. Its capacity for vegetative reproduction through the production of numerous bulbils in the leaf axils increases its effectiveness for rapid migration. Inability to compete well with later successional vegetation might explain its absence from unglaciated regions south of the glacial boundary. 29 Sagina_decumbens subsp. occidentalis, a plant of lower elevations of the coastal mountains of the Northwest and of the Great Valley of California, occurs almost entirely within non—glaciated territory and reaches its northern limits at the southern border of British Columbia. The coastal mountains were free from ice, with the exception of a few high peaks in the northern portion which experienced cirque glaciers (Crandell, 1965; Wahrhaftig and Birman, 1965). In the Great Valley, deposition of large amounts of alluvial soil took place which was derived from erosional activity in surrounding mountain systems (Wahrhaftig and Birman, 1965). This disturbance probably provided numerous habitats for this taxon. Hultén (1937) considered the taxon as a "Western America Coast Radiant." However, specimens from his area identified as Sagina occidentalis (S, decumbens subsp. occidentalis) were misdeterminations of plants belonging to the Sagina maxima complex. The present distribution of Sagina decumbens subsp. decumbens does not readily reflect events of the Pleistocene as its distribution in eastern United States, primarily the Coastal Plain and Piedmont, is entirely within unglaciated territory. Introduced taxa The presence of Sggina procumbens in North America as a native of the flora is a matter open to speculation. In the Northeast the plant occurs frequently on coastal rocks and sands and sea cliffs, along stream banks, stream beds and rocks in streams and in springy places. Occurrence in such habitats might lead one to include it as 30 a native species. However, it appears to be equally frequent along roadsides, disturbed ground and around gardens, lawns and dwellings, and in cracks between bricks of sidewalks and pavement. Torrey and Gray (1838) questioned the status of the species as one native to North America, but subsequently Gray (1895-97), and later Fernald (1950), regarded the species as native. Hultén (1958) includes the species among his "amphi—Atlantic plants," but, in a questioning tone, he notes that the species is anthropochorous to a large extent and attributes a great portion of its Eurasian range and possibly its presence in North America to this type of dispersal. It is noteworthy that in North America the species is most widely estab- lished in areas of early settlement, New England and the Maritime Provinces, and is especially prevalent in Massachusetts. Man is certainly responsible for its occurrence in the Southern Hemisphere. Hooker (1847) observed Sagina procumbens to be abundant near the sea in the Falkland Islands and considered it most certainly native. However, he also noted that it was indistinguishable from European material. Is it any wonder? Port Louis had been settled in Berkeley Sound by Bougainville in 1763 (Godley, 1965), and ships from Europe regularly visited the island during the 79 years prior to Hooker"s visit to this locale. He searched for it carefully in Fuegia but did not encounter the plant. He also indicated he knew no other locality for the plant in the subantarctic region. Subsequent to Hooker's explorations in Fuegia, an Anglican mission and settlement of Ushuaia was established along the Beagle 31 Channel on Isla Grande, Tierra del Fuego. On an expedition to this region in the austral spring of 1971 I found S, procumbens to be very abundant in such disturbed sites as roadsides and grassy meadows of logged and burned sites in the Nothofagus pumilio forest region, but it also appeared very natural growing in a gravelly stream bed on the east side of Lago Fagnano. I must add that this latter site is along a gravel highway (the southern extent of the Pan American Highway). In contrast, the plant was not found on the uninhabited eastern tip of the island, Peninsula Mitre, nor on uninhabited Isla de los Estados, even though "weedy" species native to Fuegia were present. Sagina procumbens has become frequent throughout the Sub- Antarctic where man has been active. Although the species was not encountered by Hooker on either the Campbell Islands or Kuerguelen Islands, R. C. Harris (personal communication) reports that he found the plant to be frequent at both localities and that the growth was so lush in places on Kuerguelen that, on occasion, reindeer (also intro— duced) feed on it. Man's activities are most certainly responsible for the occurrence of the species in central and western North America. By the Gray, 1838) S. procumbens had appeared in the iron mining regions of the south shore of Lake Superior, presumably introduced from eastern North America. Introduction into the Pacific Northwest probably took place in the latter part of the 18005. The earliest collections I am aware of include one specimen from Oregon, collected by Elihu Hall in 1871 and 32 one specimen from Vancouver Island, collected by John Macoun, May 9, 1875. As of 1894 the species had apparently not reached San Francisco, 9f_th§_San_Francisco Bay. 1 am of the opinion that Sagiga procumbens became introduced into northeastern North America shortly after settlement and soon became naturalized. Introduction into the Northwest most likely came by way of ships sailing around Cape Horn and could have originated from plants from eastern North America or Europe. The early collections in the Northwest are associated with coastal civilization while more recent collections indicate the plant is becoming naturalized in more remote areas of this region. Sggjna_apetala is an alien which probably appeared in California during the rapid influx of civilization in the mid-18005. The earliest recorded specimen I have seen is that of Congdon, collected in April, 1883, in Mariposa Co., California, where the species was probably a well established weed by that time. California collections prior to 1900 include: Mariposa Co., Tuolumne Co., Tehama Co., Plumas Co., and San Joaquin Co. As early as 1892 the species had reached southern Jackson Co., Oregon. Greene (1891) described the plant as a new species (Alsinella ciliata) in his flgra_Franciscana. Gray (1895—97) indicated the presence of the species in the Middle Atlantic States region, especially near the coast. The specimens he cited, however, belong to S. decumbens subsp. decumbens. I have seen but three herbarium specimens from eastern North America referable to S, apetala, one from Maryland, one from Illinois and one from Louisiana. j.“ 33 Sagina_ngd9§a_var. pubescens appears to have been introduced from Europe prior to the mid—1800s. The earliest collections known were made by J. Blake at Cape Elizabeth, Maine in 1857, and collections prior to 1900 included localities from Massachusetts to Nova Scotia. Several introductions may have occurred but the taxon appears to be especially well established along the coast in Lincoln Co., Maine and in the region of Digby, Nova Scotia. The taxon is not weedy, and Seymour (1969) notes in Ihg_flgra_9f New England that the plant is "uncommon." The normal range of Sagina_ggxima_subsp. ngj£@_extends into North America by way of the Aleutian Islands down along the Pacific coast. This chiefly northeastern Asia taxon, however, also occurs in eastern North America. Here it is of incidental introduction, occurring sporadically and does not appear aggressive or spreading. Known localities include Toronto, Montreal and Quebec, Canada and Amherst, Massachusetts, where plants were found growing in damp places around buildings and along footpaths. The eastern Asian Sagina japonica has appeared at only three localities in western North America, all seaports. As early as 1889 Macoun collected the plant at Nanimo, Vancouver Island. Suksdorf made collections of the species at Portland, Oregon in 1899 and 1900. In 1939 the plant was found growing along a railway bed at Prince Ruppert, British Columbia. A single collection is known from eastern North America. This specimen was found as a weed in a botanical garden in Ottawa, Ontario. SEED MORPHOLOGY Although the seeds of Sagjn3_are minute, (0.25—)0.3-05 (~O.6)nm long, several manuals make use of seed characters in their attempt to distinguish between the various species of a particular geographical region. Fernald's (1950) key uses the ridged markings on the seeds of S. decumbens and the black color and pebbled surface feature of S, gggg§af Hitchcock gt a1. (1964) refers to seed size and luster in distinguishing S3 crassicaulis (S. maxima_subsp. crassicaulis) from S. procumbens. Komarov (1936) employs the somewhat papillose surface feature of S, litoralis to separate it from S. maxima (the two are conspecific). Ohwi (1965) finds the tuberculate seed surface of S, japonica the only reliable character to distinguish it from the smooth seeded S. maxima. It was such use of the seeds as ”key characters" which led me to an investigation of the seed morphology. The study was an attempt to describe the amount of variation within taxa of the genus and to assess the value of seed morphology in determining interrelationships. European and east Asian taxa, as well as North American material, were included in the study.l Seeds were examined with a Bausch and Lomb dissecting microscope at a magnification of 30X and representative samples were further examined with a Zeiss research microscope using epi-illumination at 34 35 20X to 80X and with a scanning electron microscope at magnification of 200x, lOOOX and 2000X. Seeds for the scanning electron microscope study were obtained from herbarium specimens on deposit at the following herbaria: CAN, CAS, COLO, DAO, F, GH, MICH, M0, MSC, NY, UC, US. All specimens from which seeds were obtained have been so annotated. The seeds were placed on specimen stubs with double stick cellophane tape. Specimens were coated with 50 angstroms of carbon followed with 100 angstroms of gold- palladium while spinning at a 45° angle to the vapor source in a Ladd vacuum evaporator. The specimens were observed and photographed with an AMR Model 900 High Resolution Scanning Electron Microscope at Michigan State University. On a morphological basis, two basic seed types can be recognized within Sagina, the saginoid form (as in S, saginoides) and the crassuloid form (as in S, ngjflg_subsp. crassicaulis). The saginoid seed type is obliquely triangular. The ventrally located hilum, often associated with a distinct notch, is subterminal in position. An invagination occurs along the dorsal surface forming a dorsal groove. The curved embryo lies along this angled dorsal sur- face. The lateral surfaces are typically flat or drawn inward. The seeds range in size from 0.3 mm to 0.5 mm long, remaining rather con- stant within taxa. Taxa having the saginoid seed form include: S, saginoides, S, procumbens, S, subulata, S, decumbens subspp. decumbens and occidentalis, S, apetala, S, maritima, S, glabra, S. nivalis and S, caespitosa (Figures 4-7). 36 The crassuloid form, on the other hand, is more nearly reniform. The hilum is subterminal, although the notch is less distinct. The embryo lies along the convex, dorsal surface. No dorsal groove is present. The lateral surfaces appear full and plump in contrast to the drawn lateral surfaces of most of the taxa with the saginoid form. Seed size is quite consistently 0.5 mm long with the exception of that of Sagina_japonica, which is 0.4-0.5 mm long. Taxa of the crassuloid seed type include: S, ngima_subsp. maxima and crassicaulis,_S. paupuana, S, subuletorum and S, japonica (Figures 9 and 10). Two taxa have seeds which appear intermediate between the two types. These are S, n9g9§a_and S, abyssinica (Figure 8). The seeds of both the saginoid type and the crassuloid type have a crassuloid appearance during the developmental stages. With capsular dehiscence, the mature seeds become dissociated with the placenta and upon air drying an invagination occurs along the dorsal surface of the seeds of the saginoid type, while the dorsal surface of the more rigid seed coat of the craSSuloid seed type remains without a groove. However, air drying of the crassuloid type seed prior to seed coat maturation will cause dorsal invagination. At higher magnifications with the scanning electron microscope the testa in both morphological forms appears to be constructed of interdigitating cog-like cells. This pattern is apparent throughout all the species, although quite obscure in S, decumbens subsp. decumbens (Figure 5). At ZOOOX magnification the pattern is revealed in this taxon also, although the pattern may remain obscure even at this magnification (Figure 5d). 37 The cog—like cells vary in shape from orbicular to elongate. The elongate cells occur most abundantly on the lateral surfaces of the seed, are oriented toward the hilum, and tend to become more elongate nearer the hilum. The interdigitating cog teeth range from long and acute to short and blunt, the feature being inconsistent within taxa. Topography of the seed surface takes on an array of appearances associated with the amount of uplifting of the central portion of the epidermal cells. The surface of the cog—like cells may be very flat, as seen on the lateral surfaces of some seeds of S, maxflna subsp. crassicaulis (Figure 9b). Seeds of S, maxima subsp. nggj£§_with a pebbled appearance show an increased uplifting of the central portion of the cell (Figure 9a). The most pronounced protu— berances are the knobbed tuberculae of S, decumbens subsp. decumbens (Figure 5) and S, japonica (Figure 10). There is some variation in surface features within certain taxa. In S, apetala the seed surface texture ranges from pebbled to densely papillose. The pebbled surface of the seeds of S. flgdgga ranges from weak to strong. In S, procumbens the pebbled surfaces of the seed can i be so slight as to appear “smooth.“ Sagina_saginoides can be smooth 1 or pebbled. Sagjflg_decumbens subsp. decumbens seeds possess a tuber- culate feature of well developed knobbed protuberances. Occasionally the tuberculae are less well developed and the surface appears papillose. In about 40% of the specimens examined, however, the tuberculae were 3 absent. Both tuberculate and non-tuberculate seeds are consistent in the possession of a delicate reticulate ridge pattern. The western 38 equivalent, S, decumbens subsp. occidentalis, is characteristically smooth seeded, but the surface sometimes appears slightly pebbled. Seeds of the S, maxima_complex are smooth to pebbled. Sggjna_japonica is characteristically densely tuberculate, but less well developed tuberculae give a papillose and, infrequently, a pebbled appearance approaching the seeds of the closely related S, maxima subsp. maxima. In spite of this variation the surface features have proved to be very helpful in determining relationships between some taxa. At first glance the tuberculate feature seen in S, decumbens subsp. decumbens (Figure 4), a Coastal Plain and Piedmont species of the eastern U.S. and that of S, japonica (Figure 10), a plant of equiv- alent habitat in eastern Asia, may suggest a close relati0nship. The scanning electron microscope, however, reveals that the tuberculae are of distinctly different origin. The tuberculae of S, japonica occur as knobbed or rounded protuberances of the center of individual cells. These tuberculae are as many as the number of cells, thus the distri— bution appears even and dense. On the other hand, the tuberculae of S, decumbens occur associated with a ridge system reticulate in pattern. The tuberculae are not always evenly distributed and there is variability in density (Figure 5a and 5c). The fact that the seed of S, japonica is crassu- loid and that of S. decumbens is saginoid further discourages one from placing the two taxa close together. The papillose seed form of S, apetala (Figure 6a) superficially approaches the tuberculate feature. However, the SEM micrographs reveal 39 the projections to be distinctly nipple-like (Figure 6b). This mammillate condition is not uniform over the surface of a single seed, much less within the species. In contrast to the situation where relationship is negated, the elongate ridges present in Sagina_nivalis and S, caespitosa are identical. The ridges of the lateral surfaces seen under the dis— 5 secting microscope are clearly seen in SEM photographs as elongate cells, with the central portion raised. This feature, as well as other characters held in common, suggest that the two should be regarded as very closely related taxa. The following key to the North American taxa, using the morphological features of seeds, summarizes differences in the seed characteristics. a. Seed crassuloid (dorsal groove lacking; reniform to nearly globose) . . . b b. Seed distinctly reniform; surface smooth to pebbled; reddish-brown ......... S, maxima subsp. maxima subsp. craSSicaulis b. Seed nearly globose; surface distinctly pebbled, papillose or tuberculate; dark brown . . . c c. Seed densely tuberculate or papillose . . . . S, japonica c. Seed merely pebbled .............. S, ngggga a. Seed saginoid (dorsal groove present: obliquely triangular; lateral ; surfaces drawn in) . . . d d. Seed tan or light brown; surface smooth or tuberculate . . . e 40 Seeds tan; delicate reticulate ridge pattern present; frequently tuberculate (60%) . S, decumbens subsp. decumbens Seeds light brown; never tuberculate, reticulate ridge pattern lacking ..... _S. decumbens subsp. occidentalis d. Seed brown; surface smooth to pebbled (sometimes papillose in S, apetala) . . . f. . f. f. Seeds with elongate ridge pattern (non—reticulate) . . . S. nivalis s. S g s. :. apetala . sa inoides . procumbens subulata LN Seeds not ridged .............. Figure 4. 41 SEM photographs of Sagina seeds, saginoid type. (a) S. saginoides, oblique dorsal view, showing dorsal groove and smooth seed surface; Sierra Nevada Range, California (Crow 1162, MSC). (b) S. procumbens, oblique lateral View, shBWThE‘dErsal groove and EEEBTTTEEEH surface; Washington (Crow 1106, MSC). (c) S. subulata, lateral view, show1ng dorsal groove, elongate—epidermal cells of lateral surface, smooth seed surface; Sweden (Weimarck s.n., 17 June 1943, DAO)- (d) §, ggggmbgfl§_subsp. occ13entaTis, oblique lateral view, showing blunt interdigitating—EEEth_of cog—like epi- dermal cells and pebbled seed surface; California (Howell 32536, CA5). A11 X 200. 42 Figure 4 Figure 5. 43 SEM photographs of Sagina seeds, saginoid type. a) S. decumbens subsp. decumbens, dorsal view, showing dorsal groove and densely tuberculate seed surface, X 200; North Carolina (Ahles 40187, GH). (b) S. decumbens subsp decumbens, detail of knobbed tuberculae located on ridges, X 1000; Florida (i 1ggins 19395, 05). (c) S. decumbens subsp. decumbens, ventral view, showing less dense distribution of tuberculae borne on reticulate ridges, X 200; North Carolina (Biltmore Herbarium No. 1300, F). (d) S. decumbens subsp. decumbens, oblique lateral view, showing dorsal groove, non- -tubercu1ate surface with reticulate ridge pattern ' present, cog-like epidermal cells obscure, X 200; Missour1 (Steyermark 8482, M0). 44 Figure 5 45 Figure 6. SEM photographs of Sagina seeds, saginoid type. (a) S. apetala, oblique lateral view, showing dorsal groove and mam1 late papillae, X 200; California'(Tragy‘12200, DAO). Eb) S, a etala, showing detail of mamillate papillae, X 1000 same as 5a). (c) S, apetala, oblique dorsal view, showing dorsal groove and pebbled surface, X 200; California (Howell 29079, CAS). (d) S. maritima, oblique lateral view, showing orsa groove, blunt interdigitating teeth, and smooth seed surface, X 200; Sweden (Asplund 757, NY). 6 4 a“ 1—9 Figure 6 Figure 7. 47 SEM photographs of Sagina seeds, saginoid type. a) S, glabra, oblique lateral view, showing dorsal groove and slightly pebbled seed surface, X 200; France uu50cc 0.20u __. .I..v\d I I‘ II ‘p. u . I c .n I. .. . u41Un . V. l n i .e \ s _ R a _ P L} _ ./. .1 ./ _ CEHS 104 leaf margins. In more pubescent plants the trichomes are more frequent along the veins on the abaxial surface. The leaves may be glabrous. Variety pubescens appears to be introduced in North America and is most firmly established along the Atlantic Coast at Digby, Nova Scotia and Franklin Co., Maine. There is little overlap with var. ggggsa, which is rare south of Quebec Province. However, intergrades between the two taxa are apparent where p0pulations of both taxa occur together. 2. Sagina saginoides (L.) Karst. Sagina saginoides (L.) Karst., Deutsch. Fl. p. 539. 1880—1883. Spergula saginoides L., Sp. Pl. 1: 441. 1753. Alsine saginoides (L.) Crantz, Inst. 2: 408. 1766. Phaloe saginoides (L.) Dumort., Fl. Belgica, p. 110. 1827. Sagina Linnaei Presl, Rel. Haenk 2: 14. 1831. Nom. illeg. Spergella saginoides (L.) Reichb., F1. Germ. Excurs., p. 794. 1832. Sagina spergella (L.) Fenzl, Ver. Verbr. Alsin., tab. ad. p. 18. 1833. Nom. illeg. Sagina saginoides (L.) Dalla Torre, Anteit. Beob. Alpenfl. p. 75 in Hartinger, Atlas der Alpenfl. 1882. Alsinella saginoides (L.) Greene, F1. Fran. p. 125. 1891. Sagina saginoides (L.) Britton, Mem. Torrey Bot. Club 5: 151. 1894. Alsine Linnaei (Presl) Krause, in Sturms, F1. Deutschl. 2 ed., 5: 35. 1901. Nom. illeg. Type: not seen. "Habitat in Gallia.” Sagina saginoides var. hesperia Fern., Rhodora 27: 131. 1925. Type: Crandall g2) 9500 ft., Chambers Lake, Colorado. (GH, h010tYPe!) 105 Spergula saxatilis Wimm., Fl. Schles. p. 193. 1832. Sagina saxatilis (Wimm.) Wimm. Fl. Schles. p. 75. 1841. Spergella saxatilis (Wimm.) Schur, Enum. P1. Transs. p. 109. 1866. Type: not seen. Original material: grass-covered rocky sites in mountains, near Waldenburg and Einsiedel, Silesia (Germany). Spergella macrocarpa_Reichb., Ic. Fl. Germ. 5: 26. 1841-1844. Sagina macrocarpa(Reichb.) Maly, Enum. P1. Phan. p. 292. 1848. Sagina Linnaei var. macrocarpa (Reichb.) Beck, F1. Nied.-0$t. p. 358. 1890. Nom. illeg. Sagina saginoides var. macrocarpa (Reichb.) Moss, in Jour. Bot. 52: 60. 1914. Type: Icones 4963.b. (W. lectotype.) Sagina Baumgarteni Simonkai, Enum. F. Transs. p. 134. 1886. Type: not seen. Perennial. Plants tufted, branches ascending or sometimes procumbent, becoming caespitose in alpine habitats. Entire plant glabrous. Rosettes of linear leaves frequently present, 9-45 mm in diameter, or replaced by a tuft of ascending linear leaves. Cauline leaves linear, sometimes linear-subulate in caespitose plants. Connate leaf bases not conSpicuous, rarely appearing inflated and then so only in caespitose plants. Axillary fascicles of linear leaves frequently on procumbent stems. Flowers axillary or terminal. Pedicels generally long, filiform, mean length 14.5 mm, recurved during capsular develop— ment, becoming erect at maturity. Flowers 5-merous, very rarely 4- merous. Sepals elliptical, hyaline margins white, rarely purple in alpine specimens, 2.0-2.5 mm long. Stamens 10, or less frequently 5, filaments (1.05) 1.5 mm long, anthers 0.25 mm long. Capsules 1.5-2 times 106 the length of the sepals; capsule valves thin, 2.5-3.0 (-3.5) mm long, dehiscing to base. Sepals remaining appressed following capsule dehiscence. Seeds brown, obliquely triangular, with distinct dorsal groove, surface smooth to slightly pebbled, 0.3-0.4 mm long. Chromosome number: 2£f=22. Figure 19. Ecology and distribution: A montane species growing in the open or in light shade in wet places on lake margins, along streams and seep- ages in rock ledges and roadcuts, often in subalpine and alpine zones. From Alaska, south to Arizona and New Mexico. Its occurrence is rare in eastern North America. I have seen collections only from Richmond Gulf on Hudson Bay, the Shefferville area of the Labrador-Quebec Peninsula and on the Gaspé Peninsula, Quebec. Circumpolar. Flowering June to August. Figure 20. Representative specimens: CANADA: ALBERTA: Bertha Lake, Waterton Lakes National Park, elev. 6000 ft., Breitung 16239 (NY). Near Cameron Lake, Waterton Lakes National Park, elev. 5450 ft., Breitung l6178a (NY). Malique Lake, near headwaters of the Saskatchewan and Athabasca Rivers, Brown 1176 (GH, MO, NY). Little Beehive Mt., vicinity of Lake Louise, Hunnewell 3529 (MIN). About 5 mi, east north- east of Bow Peak, Banff National Park, Hitchcock §_Martin 7756 (COLO, DS, GH, MO, NY, OSC, RM, UC, WS, WTU). Crow's Nest Forest Reserve, Malte s.n., 10-24 August 1915 (CAN). Crow's Nest Pass, summit of Turtle Mt., Macoun 18290 (CAN). Mt. Edith Cavell, Jasper National Park, Scamman 3189 (GH). BRITISH COLUMBIA: Asulkan Valley, Glacier, Selkirk Mts., Brown 581 (GH, MO, US). About 8 mi. southeast of Barkerville, 107 Figure 19. Photographs of Sagina saginoides° (a) Habit. (b) Close-up of fruiting material. 108 p" u re) Figure 19 109 elev. 5500 ft., Calder, Savile §_Ferguson'14237 (DAO). North of Ft. St. James, Wolverine Lake, 55°41'N., 124°26'W., Calder, Savile g Ferguson 13645 (DAO).‘ Between Baldy Mt. and Dunn Peak, ca. 7 1/2 mi. east northeast of Littlefort, ca. 51°27'N., 120°03'W., elev. 7100 ft., Calder, Parmelee §_Taylor 19908 (DAO, UC). 8 mi. southeast of Nelson along road to COpper Mt., elev. 5400-5700 ft., Calder §_Savi1e 10993 (DAO). Mt. Apex, southwest of Penticton, elev. 6800 ft., Calder E Savile 11731 (DAO). Mt. Thornhill, near Terrace, elev. 3800 ft., Calder, Savile §_Ferguson 14871 (DAO). Mt. Sir Donald, Glacier, Fletcher §_(NY). Selkirk Range, Vicinity of Glacier, Hunnewell 4251 (MIN). Near Rogers Pass, Selkirk Mts., Heacock 4400 (GH, MIN, MO, NY, US). Victoria Lake, ca. 11 mi. west of Revelstoke, elev. 1785 ft., Hitchcock §_Martin 7592 (DS, WTU). Battle Mt., Wells Gray Park, 50°N., 120°W., Ahti §_Ahti 7003 (WTU). Chilliwack River, Macoun 34034 (CAN), Lake House, Skagit River, Macoun 79583 (CAN, NY). Upper Canyon Creek, Golden, Taylor 6727 (MICH). Sheep Mt., Elk River Valley, 34 mi. north of Natal, Weber 2347 (COLO, GH, UC, WS). VANCOUVER ISLAND: Below Mt. Burman, near Burman Lake, 49037'N., 125°44”W., elev. 5000 ft., Calder §_MacKay 32567 (DAO). Along Elk River, Strathcona Provincial Park, 49°46“N., 125°51’W., elev. 2800 ft., Calder §_MacKay 31643 (DAO). Moat Lake, Forbidden Plateau, ca. 49°41'N., 125°24’W., elev. 4100 ft., Calder §_MacKay 3228OA (DAO). Mt. Arrowsmith, Anderson §_F1etcher s.n., 7 August 1901 (DAO). Vicinity of Victoria, Macoun s.n., 23 May 1893 (MSC). DISTRICT OF MACKENZIE: Vicinity of Brintnell Lake, ca. 62°05'N., 127°359W., Raup §_Soper 9159 (GH). YUKON TERRITORY: Canol Rd., upper south fork of MacMillan River, opposite mile 280, Porsild §_Breitung 110 11300 (CAN). Canol Rd., Rose—Lapie River Pass, mile 98—99, elev. 4000 ft., Porsild g Breitung 11900 (GH, UC, US). Canol Rd., slopes of Mt. Sheldon, opposite mile 222, elev. 6000 ft., Porsild §_Breitung 11742 (CAN). Vicinity of Mackintosh, mile 1022 Alaska Highway, slopes of Mt. Decoeli, elev. 4000 ft., Schofield §_Crum 8049 (CAN). QUEBEC: Fishing Lake Creek, Richmond Gulf, east coast of Hudson Bay, ca. 56°N., 76°W., Abbe, Abbe §_Marr 4396 (CAN, DAO, MIN, US). Knob Lake, Scheffer- ville area, Quebec-Labrador Peninsula, ca. 54°45”N., 66°40'W., Hustich _&_Kallio 752 (CAN). GASPE CO.: Mt. Albert, elev. 950 m, Collins _§._ Fernald 14 (CAN, GH, MIN, MSC, NY, UC); 1 mi. above Marten River, River: Ste~Anne-des—Monts, Collins é Fernald s.n., 3~17 August 1905 (GH). UNITED STATES: ALASKA: Thompson Pass, Richardson Highway, Anderson 2777 (CAN). Unalaska, Anderson 4117 (GH). Falls Creek Mine, near Kenai Lake, Kenai Peninsula, 60°26'N., 149°17“W., Calder 6088 (CAS, DAO). Head of Resurrection Bay, Seward, Kenai Peninsula, 60°07‘N., 149”25°W., Calder 7018 (DAO). Isabel Pass, mile 199 Richardson Highway, 63¢32"N., 145°52”W., Cody §_Webster 5824 (DAO). Tangle Lakes area, mountain east of Landmark Gap, Alaska Range, Gjaerevoll 1293 (CAN). Smith”s Dry Lake, Attu Island, Aleutian Islands, Hardy 385 (GH). Juneau Ice Field, Heusser 212 (OSC). Mountains southeast of Texas Lake, 20 mi. northwest of Hyder, elev. 4600 ft., McCabe 8428 (UC). About 15 mi, due east of Berners Bay at Vaughan Lewis glacier, 25 mi. north of Juneau, Alaska, elev. 2800 ft., Miller 1745 (MSC). Peaceful Valley, Attu Island, Aleutian Islands, Soule 185 (WTU). Along Salmon River road 3 mi. north of Hyder, Whited 1208 1/2 (MO, WS). ARIZONA: COCONINO CO.: San Francisco Mts., elev. 11500 ft., Knowlton 134 (US); Kaibab Basin, elev. 8200 ft., Merkle 586 (CAS); Little Park, North Rim Grand Canyon, elev. 8800 Ft., Merkle 601 (CAS). CALIFORNIA: ALPINE CO.: Carson Pass, elev. 8200 ft., Jepson 8116 (UC); 2 mi. west of Ebbet's Pass, elev. 8500 ft., Tracy 16642.(UC). AMADOR CO.: Silver Lake, elev. 8000 ft., Newell s.n., 24 August 1929 (CAS). BUTTE CO.: along Butte Creek, vicinity of Jonesville, Copeland s.n., 18 July 1931 (MIN, RM); Butte Meadows, Heller 14688 (DS, MO, US). CALAVERAS CO.: Big Meadows, Stanislaus National Forest, Crow 1175 (MSC); Dorrington, Jepson 10201 (UC). DEL NORTE CO.: High Prairie Creek, Jepson 9346 (JEPS). FRESNO CO.: Pine Ridge, elev. 5300 ft., Hall §_Chandler 135 (MIN, NY, UC, US); Huntington Lake, elev. 6900 ft., Jepson 13057 (UC); Huckleberry Meadows, King's River region, elev. 6500 ft., Newlon 212 (UC). GLENN CO.: Sheetiron Mt., elev. 5950 ft., Bacigalupi 4677 (UC); 1/2 mi. on old road from Plaskett Meadows Ranger Station, Baker 10279 (UC). HUMBOLDT CO.: Trinity Summit, near Box Camp, elev. 5500 ft., Tracy 17918 (UC, US, WTU). INYO CO.: Lone Pine Creek Canyon, east Slope of Sierra Nevada, elev. 6950 ft., Alexander §_Kellogg 2907 (DS, UC, US); Sweetwater'Creek, Sweetwater Mts., elev. 8200 ft., Munz 21172 (UC); Pine Canyon, east of Mount Muir, vicinity of Mt. Whitney, elev. . 12200 ft., Sharsmith 3313 (UC). KERN CO.: about 1/2 mi. northeast of Evans Flat, Greenhorn Mts., elev. 5925~2950 ft., Smith 652 (UC); Bitter Creek, Twisselmann 12596 (GAS). LASSEN CO.: Martin Springs, Eagle Lake, Brown §_Wieslander 4§_(JEPS); Lassen Butte, Eastwood 1772 (CAS). MADERA CO.: Long Meadow, elev. 6800 ft., Hawkes 5204 (UC); near Garnet Lake, elev. 9700 Ft., Howell 16776 (CAS). MARIPOSA CO.: Summit of 112 Chowchilla Mt., Congdon s.n., 29 June 1885; Yosemite Valley, elev. 3900 ft., Hall 8879 (DS, US); near Wawona, elev. 4000 ft., Howell 431 (CAS); near Bridal Veil Creek, above.Brida1 Veil Falls, Yosemite Valley, elev. 4800 ft., Sharsmith 2175 (UC). MODOC CO.: East Creek, about 1 mi. east of Patterson Ranger Station, Warner Mts., elev. 7100 ft., Ferris g Lorraine 10630 (DS); 15 mi. southeast of Alturas, elev. 5500 ft., Hitchcock 6708 (NY, UC, WTU); mouth of Dry Creek at Parker Creek, Warner Mts., southeast of Alturas, Payne 791 (UC). MONO CO.: along California Rt. 120, 5 1/2 mi. West of Ranger Station, Inyo National Forest, elev. 8500 ft., Crow 1161 (MSC); Twin Lakes, west of Bridgeport, elev. 7400 ft., Ferris §_Lorraine 10983 (DS); Tioga Crest, east of Saddlebag Lake, Tioga Pass region, elev. 11000 ft., Mason 11470 (UC); north Slope of Excelsior Mt., near Summit Lake, elev. 10500 ft., Sharsmith 4145 (DS, GH, UC). NEVADA CO.: Lytton Creek, Norden, Jorgesen 501 (DS, WTU); Lake Van Norden, elev. 6900 ft., Raven 2436 (WS). PLACE CO.: Summit Valley, Howell 18582 (CAS). PLUMAS CO.: slopes across from the Devils Kitchen, Lassen Volcanic National Park, Gillett 84 (UC); Middle Fork Feather River, Rich Point, Jepson 10619a (UC). “- RIVERSIDE CO.: Deer Springs, San Jacinto Mts., elev. 9000 ft., Meyer 541 (UC); Strawberry Valley, San Jacinto Mts., elev. 5400 ft., Reed 2454 (UC). SAN BERNARDINO CO.: Big Bear Lake, San Bernardino Mts., Breitung 15504 (DAO); Dry Lake, San Bernardino Mts., Howell 23626 (CAS). SHASTA CO.: Brokeoff Meadows, Lassen Volcanic National Park, elev. 6300 ft., Gillett 112 (UC); Manzanita Lake, Lassen Volcanic National Park, elev. 5850 ft., Rose 45193 (CAS, GH). SISKIYOU CO.: Horse Camp Springs, Mt. Shasta, elev. 8250 ft., Cooke 11474 (UC); Wagon Camp, 113 Mt. Shasta, elev. 6000 ft., Crow.1206.(MSC); Compton”s Prairie, Mt. Eddy, elev. 3800 ft., Heller 13267.(CAS, DS, MO, NY, US, WIS); South Fork of Samon River, 9 1/2 mi. above Cecilville, elev. 3500 ft., Wiggins 13445 (CAS, DS, NY, RM, US). TEHAMA CO.: South Yollo Bolly, Jepson s.n., 24 July 1897 (JEPS); north slope of North Yollo Bolly Peak, Munz 16679 (CAS, DS, NY). TRINITY CO.:. Preacher Camp Grounds, 4 mi. west of Scott Ranch, Cantelow 1543 (CAS); South Fork Mt., Tracy 19055 (UC). TULARE CO.: headwaters of Freeman Creek, north slope of The Needles, Sequoia National Forest,.Bacigalupi,.Wiggins §_Ferris 2569 (DS, WTU); Mineral King, Howell 17121 (CAS); South Fork of Kern River, Bakeoven Meadows, elev. 8100 ft., Howell 26886 (CAS). TUOLUMNE CO.: Mather, Keck 5318 (CAS, DS, US); Smoky Jack Meadows, above Glen Aulin, elev. 8600 ft., Sharsmith 4080 (CAN, GH, NY, UC); Iceberg Meadow along Clark Fork of the Stanislaus River, Wiggins 20193 (DS); Niagra Creek Public Camp, Sonora Pass Road, elev. 6550 ft., Wiggins 8975 (CAS). VENTURA CO.: Seymour Creek, Pierson s.n., in 1922(?) (US). COLORADO: BOULDER CO.: Green Lakes Valley, north of Kiowa Peak, elev. 11500-12000 ft., Weber §_ Dahl 8589 (COLO). CLEAR CREEK CO.: Berthoud Pass, near Idaho Springs, Degener g Peiler 16768 (RM); Cold Spring, Yankee Creek, above Brookvale, Churchill s.n., 22 June 1918 (M0): Pendleton Mt., above Leavenworth Gulch, Silver Plume region, elev. 11000 ft., Ewan 14472 (CAS); mountains about the headwaters of Clear Creek, valley near Empire, Patterson 173 (COLO, F, MICH, MIN, MO, Msc, UC, US); Mt. Evans, slope above Summit Lake, elev. 12800~13400 ft., Porsild §_Weber.22869 (CAN); north slope Gray's Peak, elev. 12500 ft., Weber 5927 (COLO). CONEJOS CO.: west of 114 Platoro, near head.of Adams Fork,.elev. 11500 ft., Harrington 1719 (RM); just south of Platoro, elev. 9900 ft., Weber 7864 (CAS, COLO, DAO, MIN, RM, WS, WTU). COSTILLA CO.: slope of old Baldy Peak, 8 mi. from Fort Garland, elev. 9700 ft., Mattoon 179 (COLO); Buena Vista, Harper s.n., in 1886 (WIS). FREMONT CO.:.Lake.Creek, 3 mi. west of Hillside, Sangre de Cristo Mts., Erlanson 1416 (MICH)...GILPIN CO.: Eldora to Baltimore, elev. 8500-9500 ft., Tweedy 5535 (RM, NY). GRAND CO.: East St. Louis Creek, Fraser.Experimental Forest, southwest of Fraser, elev. 9500 ft., Weber 8616 (COLO,.DAO, WTU); East Ten Lakes Park, elev° 11800 ft., Willard 61228 (COLO). GUNNISON.CO.: along Spring Creek, 5 mi. north of Spring Creek campground, Gunnison National Forest, Crow 1146 (MSC); just west of Schofield Pass in Elko Park, 6 mi. northwest of Gothic, Crow 1147 (MSC); vicinity of Mt. Carbon, Kebbler Pass, Eggleston 5727 (US)° LAKE CO.: Twin Lakes, Clements 421 (NY); Leadville, Trelease s.n., 8 July 1886 (M0). LA PLATA CO.: near La Plata, elev. 9000 ft., Baker, Earle §_Tracy 675 (F, MIN, MO, NY, RM, US). La Plata Mts., elev. 11500 ft., Tweedy 430 (US). LARIMER CO.: Chamber's Lake, elev. 9500 ft., Crandall 29_(GH, NY);.Beaver Creek, 50 mi. west of Fort Collins, elev. 9500-10000 ft., Crandall s.n., 19'20 JUIY 1898 (RM); Cameron Pass, Osterhout 5024 (M0, MSC, RM); Loch Vale Trail, about 1/2 mi. north of The Loch, elev. 9900 ft., Willard 6027 (COLO); Trail Ridge, terrace south of Iceberg Lake, elev. 12000 ft., Willard 61131 (COLO). PARK CO.: vicinity of Georgia Pass, South Park, elev. 10500 ft., Walter 581 (COLO); trail from Platte Gulch to Wheeler.Lake, ca. 5 mi. northwest of Alma, elev. 11500 ft., Weber 8747 (COLO, DAO, MIN, 115 WTU); Duck Lake, Zobel s.n., 18 July 1934 (M0). ROUTT CO.: mountains east of Streamboat Springs, Osterhout 2685 (RM); trail from Columbine to summit of Hahn's Peak, elev. 8400-10800 ft., Weber 6913 (COLO). SAN JUAN CO.: 1/2 mi. south of South Mineral Campground, elev. 10200 ft., Douglass 54-427 (COLO); Needle Mt., Tenmile Basin, ca. 18 mi. southeast of Silverton, elev. 12800 ft., Michener 830 (COLO). SUMMIT CO.: near Breckenridge, elev. 9800 ft., Mackenzie 234 (MO, NY, RM, WIS); south side Monte Cristo Creek, just north of Hoosier Pass, elev. 11000 ft., Weber, Rollins §_Livingston 6495 (COLO). IDAHO: BLAINE CO.: 14 mi. north of Ketchum, Wood River, Christ 15818 (NY); Boulder Creek Canyon, Sawtooth Mts., elev. 8000 ft., Thompson 14142 1/2 (WTU). BOISE CO.: headwaters of S. Fork Payette River above Sacajawea Hot Springs, 3 mi. north of Elk Lake, Sawtooth Primitive Area, Hitchcock §_Muhlick 2861 (NY, WTU). BOUNDARY CO.: Continental Mt., Christ 1697 (NY); north slope of Mt. Rootnaah, Daubenmire 44379 (WS). CUSTER CO.: Stanley Basin, near Cape Horn Ranger Station, Christ 9683 (NY); on banks of Yankee Fork, near Custer, Christ 11345 (NY); North Fork Big LOSt River, 9 mi. west of Wild Horse Ranger Station, Christ 16003 (NY); Ryan Peak, Boulder Mts., elev. 9000 ft., Hitchcock §_Muh1ick 10638 (WTU). FRANKLIN CO.: Bear River Range, Franklin Basin, Maguire 21627 (NY). FREEMONT CO.: Red Rock Pass, Christ 5767 (NY). IDAHO CO.: Callender, west of orogrande, Christ 11623 (NY); west side of Seven Devils Divide, Seven Devils Mts., Christ 12493 (NY, US). LEMHI CO.: South Fork Camas Creek near Sleeping Deer Mt., 4 mi. northwest of Challis, Hitchcock g Muhlick 11345 (WTU). SHOSHONE CO.: near Sohons Pass, region of the Coeur 116 D‘Alene Mts., elev. 1500 m, Leiberg.1425 (GH, NY, US). TETON CO.: Game Creek, Victor, Christ 5145 (NY). Victor, Merrill §_Wi1cox 893 (GH, NY, RM, US). VALLEY CO.: valley of Monumental Creek,.near old town of Roosevelt, 21 mi. east of Stibnite, Christ §_Ward 1464 (NY). MONTANA: DEER LODGE CO.:. 4.mi. west of Storm Lake, Anaconda Mts., Hitchcock §_Muh1ick 14868 (NY, WIS, WS, WTU). FLATHEAD CO.: vicinity of Lake McDonald, 4 mi. north of Belton, Holzinger §l_(US); near White- fish divide on Yakinikak Creek, Mooar 10736 (MSC); vicinity of Lake McDermott, elev. 1450-1740 m, Glacier National Park, Standley 15640 (US). MISSOULA CO.: near Lagoon Lake, above Glacier Lake, Mission Mts. Primitive Area, elev. 6500 ft., Crow 918 (MSC); near headwaters of the South Fork of the Jooko River, 21 mi. northeast of Missoula, elev. 5700 ft., Thomas 11243 (DS). GALLATIN CO.: Spanish Basin, elev. 6500 ft., Rydberg §_Bessey 4034 (CAN, NY). GLACIER CO.: Many Glacier, Glacier National Park, Jones 5329 (GH); Hidden Lake Overlook, Logan Pass, Glacier National Park, Harvey 7024 (MONTU); vicinity of Going-to-the- Sun Chalets on St. Mary Lake, Glacier National Park, e1ev° 1350 m, Standley 17124 (US). LINCOLN CO.: Big Cherry Creek, east of Libby, Harvey 229§_(MONTU). PARK CO.: Big Muddy Creek, near Wilsall, Suksdorf .222_(WS); Cooke Guard Station, about 2 mi. east of Cooke City, elev. 8000 ft., EEEE_EZ§9_(CAS, COLO, DAO, MIN, NY, UC, WIS, WS, WTU). RAVALLI CO.: Watchtower Creek.Trai1, Bitterroot Mts., elev. 5600 ft., Lackschewitz é Pageraas 633 (MONTU). SWEET GRASS CO.: Sweet Grass Canyon, Crazy Mts., elev. 6000-7000 ft., Flodman 447 (MO, NY); along East Fork Boulder River, 3 mi. south of Rainbow Lakes, Hitchcock 16490 117 (WTU). Montana, county uncertain, Long.Ba1dy, Little Belt Mts., elev. 7000-8000 ft., Flodman 446 (NY, MO); Belt Mts., Williams 493 (MIN). NEVADA: DOUGLAS CO.: .2.mi. east of junction of Kingsbury & Clear Creek Grades, Train 3178 (UC). .ELKO CO.: Jarbidge River, 2 mi, south of Jarbidge,.Baker 8636A.(WTU). ESMERALD CO.: Chiatovitch Creek, Duran 2801 (UC). HUMBOLDT CO.:. valley of Lawance Creek, Santa Rosa Mts., elev. 6000 ft., Archer 246 (MICH). .NYE CO.: North Twin River, Toyabe Mts., Linsdale §_Linsdale 658.(CAS). ORMSBY.CO.: head of Fall Creek, elev. 2460 m, Baker 1332 (GH, MO, MSC, NY, UC, US). WASHOE CO.: Galena Creek, 8 mi. west of Reno Hot Springs, elev. 6100 ft., Archer 5469 (UC); Third Creek, near Mt. Rose, elev. 8500 ft., Howell 14055 (CAS, WTU). NEW MEXICO: RIO ARRIBA.CO.: vicinity of Brazos Canyon, Standley g Bollman 11043 (US). SAN MIGUEL CO.: Winsor's Ranch, Pecos River National Forest, elev. 8400 ft., Standley 4170.(GH, MO, NY, US). SANTA FE CO.: Santa Clara Canyon, Marcelline 1911 (F). OREGON: BAKER CO.: Hudson Creek, R44E, T68, sec. 9., Head 1594 (OSC). CROOK CO.: vicinity of Laidlaw, Whited 3216 1/2 (US). DESCHUTES CO.: Paulina Lake, Howell 7076 (CAS); Diller Glacier, Three sisters Mts. elev. 7500 ft., Van— Vechten 248 (WTU). GRANT CO.: Dixie Mt., Blue Mts., Henderson 5536 (CAS, GH); Strawberrert., Blue Mts., elev. 8000 ft., Maguire §_Holmgren 26843 (GH, NY). HARNEY CO.: White Horse Mts., Griffiths §_Morris 464 (US); Steens Mts., 12 1/2 mi. east & 9 1/4 mi. south of Frenchglen, Hansen 877 (OSC). HOOD RIVER CO.: White River, S. Mt. Hood; 10 mi. southeast of Mt. Hood, Lloyd s.n., July 1894 (NY). JACKSON CO.: northern slopes of Mt. Asland, Rossbach 599 (DS). JOSEPHINE CO.: 118 near Bolan Lake, Siskiyou.Mts., Hitchcock E Martin 5238 (DS, NY, UC, WS, WTU); 4 mi. southeast of Oregon Caves, Siskiyou Mts., Peck 8288 (GH); Grayback Area, Siskiyou Mts., Whittacker s.n., 24 July 1949 (WS). KLAMATH CO.: Kerr Notch,.Crater Lake National Park, Baker 7127 (WTU); Crater Lake, Hawkins s.n., 30 July 1917 (WIS); 15 mi. north of Fort Klamath, Peck 9367 (DS, GH, MO,.NY). LAKE CO.: Cogswell Creek, 8 mi. south of Lakeview, Peck 15564 (DS, WTU). LANE CO.: Scott Lake, McKenzie Pass, Jones 5769 (WTU); west of North Sister, elev. 6500 ft., VanVechten 221 (OSC). UMATILLA CO.: 2 mi. north of Tollgate, Blue Mts., elev. 5000 ft., Crow 1235 (MSC). UNION CO.: Indian Creek, Blue Mts., Darlington 146 (CAS). WALLOWA CO.: Lake Wallowa, Powder River Mts., Ferris §_Dulthie 1387 (DS); Mirror.Lake, Wallowa Mts., 7550 ft., Mason 1957 (OSC); Imnaha Canyon, 24 mi. above Imnaha, Peck 18379 (DS, NY, WTU). WASCO CO.: 15 Mile Meadow, Mt. Hood National Forest, elev. 4500 ft., Jones 4135 (GH, UC, WTU). UTAH:..BEAVER CO.: Big Flat Ranger Station, Tushar Mts., Eggleston 10461 (US); vicinity of Big John Flats, Beaver River headwaters, elev. 9000-10000 ft., Maguire 19834 (GH, WTU). BOX ELDER CO.: Dunn Canyon, Raft River Range, elev. 6500 ft., Maguire 3 Holmgren 22176 (GH, NY, UC, US). CACHE CO.: mountains near Logan, Shear s.n., 9 August 1895 (NY). DAGGETT CO.: Green Lakes, elev. 7500 ft., Hermann 4824 (MO)° DUCHESNE.CO.: Moon Lake, Ashley Forest, elev. 8100 ft., Harrison §_Larsen 7704 (M0); Krebs Basin, above Chain Lake, Mt. Emmons, elev. 11300 ft., Hermann 4938.(MO). ELKO CO.: Verdi Lake, Ruby Mts., elev. 10450 ft., Mills §_Beach 1575 (UC). GRAND CO.: north base of Haystack Mt., La Sal Mts., elev. 9300 ft., Maguire, Richards, 119 Maguire §_Hammond 17965 (CAN, WTU). JUAB.CO.: Granite Creek, Deep Creek Range, elev. 7000 ft.,.Maguire §_Holmgren 21865 (GH, NY, UC, US, WTU). PIUTE CO.: Tate Mine, near Maryville, Jones 5855 (MSC, NY, RM, UC). RICH CO.: Laketown, elev. 6300 ft., Harrison §_Larsen 7956 (M0)» SALT LAKE CO.: Silver Lake, Big Cottonwood Canyon, Clemens s.n., 30 September 1909 (RM, UC). SAN JUAN CO.: southeast part of La Sal Mts., elev. 10000 ft., Goldman gynitchcock 1473 (M0); Abajo Mts., elev. 3000—3300 m, Rydberg §_Garrett.9746 (NY, US). SANPETE CO.: vicinity of Clayton Peak, Wasatch Mts., elev. 9000 ft., Stokes s.n., 12—26 August 1903 (M0). SEVIER CO.: Fish Creek Canyon, Garrett 2596 (NY). SUMMIT CO.: Mill Creek, southwest base of Mt. Elizabeth, Uinta Mts., elev. 8500 ft., Hermann 5897 (M0, RM); Burntfork Creek, below Thompson Creek, elev. 7500 ft., Jensen s.n., 24 July 1942 (UC, WTU); South Blanchard Lake, Henry's Fork Basin, Kings Peaks-Gilbert Peak region, Uinta Mts., elev. 11100 ft., Maguire, Hobson §_Maguire 14616 (WTU). UINTAH CO.: Paradise Park, Uinta Basin, elev. 10000 ft., Graham 10056 (GH, MO); east slopes of Leidy Peak, Uinta Mts., elev. 10000 ft., Maguire 17669 (WTU). UTAH CO.: American Fork Canyon, elev. 8000 ft., Jones 1362 (CAS, F, MICH, MSC, NY, UC, US: WS, WTU). WAYNE CO.: Blind Lake, Aquarius Plateau, elev. 10000 ft., Dixon 758 (F); Fremont Reservoir, head of Fremont River, elev. 8500 ft., Dixon- 549 (F). WASHINGTON: ASOTIN CO.: Blue Mts., Jones 1876 (WS). CLALLAM CO.: east face of Obstruction Point, elev. 5600 ft., Meyer 1258 (M0). CHELAN CO.: northeast side of Snow Lake, Stuart Range, Wenatchee Mts., southwest of Leavenworth, elev. 5000 ft., Crow 1108 120 (MSC). COLUMBIA CO.: Indian Corral, Blue Mts., Carlington 146 (WS). JEFFERSON CO.: Mt. Olympus, eleV. 5000 ft., Flett 3043 (WTU); Skokomish River, Olympic Mts., Piper s.n., August 1895. KLICKITAT CO.: Bingen, Suksdorf s.n., 18 April 1895 (WS). OKANOGAN CO.: head of Middle Fork of Pasayten River, north of Harts Pass, Ownbey §_Meyer 2312 (DS, MO, NY, UC, WS, WTU); Salmon Creek, near Conconully, Thompson 6967 (MIN). PEND OREILLE CO.: near Gypsy Meadow, elev. 4800 ft., Layser 918 (WS). PIERCE CO.: Nisqually Bridge, Mt. Rainier, elev. 3900 ft., Jones 9917 (GH, NY); road to Sunrise glacier, Mt. Rainier, Crow, 1236 (MSC); trail above White River Campground, Inter Fork of White River, Mt. Rainier, elev. 4500 ft., Raven 8963 (CAS); "Ellerbach," Mt. Adams, Suksdorf 334 (WS). WHATCOM CO.: Winchester Mt., elev. 4500 ft., St. John 8964 (WS); Bagley Lake, near Mt. Baker Lodge, elev. 4300 ft., Thompson 5700 (DS, MO, WTU); below glacier on Heliotrop Ridge, Mt. Baker, elev. 7500 ft., Thompson 11253 (US, WTU). YAKIMA CO.: Wodan's Vale, Mt. Adams, elev. 2000 m, Suksdorf 6829 (COLO, DS, NY, UC, WS, WTU). WYOMING: ALBANY CO.: Medicine Bow, above Nash Forks, off route 130 near Centennial, Churchill s.n., 13 July 1958 (MSC); Centennial, Nelson 7728 (GH, MIN, MO, NY, RM, US); Trail Creek, near Sand Lake, Medicine Bow Range, elev. 9700 ft., Porter §_Porter 7927 (DS, UC. WTU)» CARBON CO.: South Spring Creek, Hayden Forest, Eggleston 11277 (US); Battle Lake Mt., Nelson 4241 (RM). FREMONT CO.: Brooks Lake, near Dubois, Churchill s.n., 18 July 1958 (MSC). LINCOLN CO.: near junction of Box Canyon Creek and Grey"s River, elev. 7800 ft., Goodman 5133 (RM). PARK CO.: Beartooth Pass, Beartooth Range, elev. 9200 ft., Porsild, Johnson 121 E Darling 22753 (CAN); Old Faithful, Hawkins 580d (US); Canyon Junction, Langdon 160 (OSC); Yellowstone Lake, Rydberg §_Bessey 4037 (NY). TETON CO.: Teton Creek, Targhee National Forest, 11 mi. east of Driggs, Anderson 415 (NY); Teton Pass, Merrill 340 (US); "Jackson Hole," vicinity of Jackson Lake, near Moran, elev. 7500-3500 ft., Yuncker 5290 (F, NY); vicinity of Holback Canyon, elev. 7500 ft., Williams §_Pierson 735 (CAS, GH, MO, NY, RM). Wyoming, county unknown, vicinity of Big Horn Mts., Williams s.n., July-August 1897 (RM). In reorganizing the Specimens of Sagina in the Gray Herbarium, Fernald (1925) noted that the typical phase of §, saginoides of the Arctic and Eurasia occurred locally throughout the North American range of the species, but that most of the material of western America was found having sepals and capsules of shorter length than the typical. This American extreme he described as var. hesperia. The Eurasian specimens exhibit considerable variability, the range for sepal and capsule length being continuous (sepals 2.0-3.0 mm long; capsules 2.5u5.0 mm long). Frequently plants of lower elevations tend to be more robust, producing slightly larger flowers, while plants of higher elevations are generally smaller. The more robust, larger flowered plants, with capsules up to 5.0 mm long were first recognized at the specific level as Spergella.macrocarpa Reichb., and later at the varietal level as Sagina saginoides var. macrocarpa (Reichb.) Moss. Moss (l9l4) admits that the discontinuity between the two varieties is trifling, but continues to recognize two varieties in §, saginoides Figure 20. 122 Geographic distribution of Sagina saginoides in North America. 123 ,__ -‘m- “\. ._,_—- __. . H_ __ _ - - —- — “L -:¥J -}O :0 '00 9C .0 Ill" ”‘4199. r -_ - 'H— —E__— LANCIQT S AZ'UU7“AL [QUAL'AQEA 9°C-[ 4' 3% u. .. -, ' .. g'.‘ .. ’-'vl - . . . . . - o . o . - a" I.‘- 0‘ ' ._ {-7 0.. -.‘Itl’l‘-\ Figure 20 124 (Moss, 1920). The treatment in Flora Europaea (Clapham and Jardin, in Tutin gt 21,, 1964) notes that the larger flowered plants cannot be clearly separated as a taxon distinct from typical §, saginoides. Discontinuity in this variable species appears to be lacking between those plants which were regarded as.var. macrocarpa_and those which were regarded as var. hesperia. I am therefore not recognizing infraspecific categories in the species. Earlier collections of Reichenbach, which were housed at the Zwinger Museum in Dresden, Germany, were destroyed by fire in May l849 (Stafleu, l967). However, the original plates of his Icgfl§§_ Florae Germanicae §t_Helveticae are preserved at Vienna (N). I therefore designate Icones 4963.b. as the lectotype of Spergella macrocarpa Reichb° 3. Sagina prpcumbens L. Sagina procumbens L., Sp. Pl. l: 128. l753. Alsine procumbens (L.) Crantz, Inst. 2: 404. l766. Sagina pentamera Rouy & Foucaud, Fl. France 3: 286. 1896. Nom. nud. Alsine procumbens (L.) Krause, in Sturms, Fl. Deutschl° 2 ed., 5: 36. l901. Type: not seen. “Habitat in Europae, pascuis Sterilibus uliginosis aridis.” Sagina procumbens var. compacta Lange, Meddel. Groenl. 3: 242. l887. Type: not seen. Original material: Igaliko, Greenland. Collected by Vahl? Sagina muscosa Jord., Pugill. Pl. Nov. p. 32. l852. Type: not seen. Original material: on mossy cliffs of woodlands and in 125 subalpine meadows of mountains, Gerbier de Jones and Mont Pila, near Lyon, France. Perennial. Plants totally glabrous° Stems ascending or, more frequently, procumbent. Rosettes of linear leaves frequent in younger plants, 9-35 (-55) mm in diameter. Procumbent stems with axillary fascicles giving rise to secondary tufts or, less frequently, secondary rosettes, rooting at the nodes. Cauline leaves linear, lower leaves 4~l5 mm long, becoming shorter toward the apex, upper leaves 2.5-6.0 mm long. Leaf margins entire, rarely with minute glandular cilia. Leaf tips apiculate to aristate. Connate leaf bases not conspicuous, never forming an inflated cup. Pedicels generally long, filiform, recurved during capsule development, becoming erect at maturity. Flowers 4- merous, occasionally 4- and 5-merous. Pedicels long, filiform. Sepals elliptical to orbicular, 1.5-2.0 (-2.5) mm long, hyaline margins white, never purple tinged. Petals 4 or fewer or sometimes absent, minute, 0.7541.0 («1.5) mm long, orbicular to elliptical, clawed. Stamens 4, occasionally 8, filaments l.0sl.5 mm long, anthers 0.25 mm long. Cap- sules slightly exceeding sepals. Capsule valves thin, (l.5-) 2.0-2.5 («3.0) mm long. Sepals appressed during capsular development, divergent following dehiscence. Seeds brown, obliquely triangular, with a distinct dorsal groove, (0.3-) 0.4 (—O.5) mm long. Chromosome number: Zflf=22. Figures 2] and 22. EOOZOQy and distribution: A weedy species, growing in wet or damp gravelly or sandy soils along roadsides, sidewalk cracks, and margins of paths or lawns. Also frequent along pond and lake margins, coastal rocks and sands and sea cliffs. The species is sometimes Figure 21. 126 Photographs of Sagina procumbens. (a) Flowers showing CUPPEd sepals and papillate stigmas (emasculated), Natcom Co., Washington (Crow 1238, MSC). (b) Adventitious roots produced on procumbent stem of weedy plant in grOWth chamber. FlgWe 22. Photographs of Sagina procumbens. (a) Habit, Bolton Pass, QUEbeC (Marie-Victorm, ROI land—Germaine, Raymond and MWM Habit, St. John's Avalon Peninsula. Newfoundland (Bassett sfl, DAO) o 129 name 1 2 3 4 5 nhn Figure 22 130 cultivated as a ground cover. In eastern North American from Newfoundland, west to New York and eastern Pennsylvania, and the southwest shore of Lake Superior. In western North America. from the San Francisco area north to Washington and infrequently northward to the Queen Charlotte Islands, and the Aleutian Islands. Single collections are known from Detroit, Michigan; Columbus, Ohio; Pulaski Co., Arkansas; Missouri (no specific locality cited); and Marysvale, Utah. Occasionally at high elevations in Mexico and Central America. Also introduced in eastern Asia and the Southern Hemisphere. Native to Eurasia. Flowering May to September. Figure 23. figgggsentative specimens: CANADA: BRITISH COLUMBIA: Salt- spring Island (Straight of Georgia), Ashlee §;§., in 1959 (DAO). Between Prince Rupert and Galloway Rapids, Calder, Savile 9 Ferguson 19999 (DAO). Near east end of Summit Lake on road from Nakusp to New Denver, Calder §_Savi1e 19919 (DAO, NY). Vancouver, Eastham 9999 (UC). Road to Silver King Mine, Nelson, Eastham 19999 (UC). New Westminster, 92951 9199 (RM). Langley Prairie, Ergh 993., 26 July 1939 (DAO). Hecate Island, MaCabe 1194 (UC). East shore of Kootnay Lake, 15 mi. south of Boswell, Senn_§_Frankton 5999 (DAO). Queen Charlotte Islands: Summit, between Gillatt Arm of Cumshewa Inlet and Peel Inlet, Moresby Island, Calder §_Taylor 95119_(DAO); approximately 3 mi. from east end of Skidegate Lake along Copper Creek, Moresby Island, Calder E Taylor 99921 (DAO). Vancouver Island: Vicinity of Victoria, Macoun E19,, 23 May 1893; Quarantine Lake, about 20 mi. west of Victoria, McCabe 5647 (UC); Nanimo, Malte s.n., 30 June 1914 (CAN); Duncans—Cowichan 131 Lake road, Rosendah11758 (GH, NY, UC, US). NEW BRUNSWICK: CHARLOTTE CO.: Deer Island, Chrysler 629§_(GH); Mill Cove, Campobello Island, Malte 971/29 (CAN, GH); Digidiquash, about 20 mi. northeast of St. Andrews, Malte 648/29 (CAN); west side of Whale Cove, Grand Manan Island, Weatherby §_Weatherby 5492 (CAN, MO, US). GLOUCHESTER CO.: Grande Anse, Blake 5533 (CAS, GH, NY, US, WTU); Caraquet, Dore, Senn §_Gorham 45.593A (DAO). KENT CO.: Bass River, Fowler s.n., August 1875 (WIS). KINGS CO.: Sussex, Svenson §_Fassett 2000 (GH). NORTH- UMBERLAND CO.: Little Branch, Miramichi, Fowler s.n., 24 August 1894 (F, US, WIS). RESTIGOUCHE CO.: Eel River, near Dalhousie, Malte 456 (CAN). SUNBURY CO.: Oromocho River at Fredericton Junction, Roberts §_Bateman 64-2069 (DAO). WESTMORLAND CO.: Shediac, Hubbard s.n., 7 July 1914 (GH); about 15 mi. northeast of Sackville, Scoggan 12243 (CAN). YORK CO.: Fredericton, Scoggan 11867 (CAN). LABRADOR: Perquet Island, 51°26'N., Allen §§_(GH: NY). Red ISlandSr near Turnavik, 53°48'N., 56°46'W., Bishop 285 (CAN, US). Forteau, on Strait of Belle Isle, Donly s.n., ca. 20 August 1964 (DAO). NEWFOUNDLAND: Gander, Bassett 483 (DAO, MO). St. John's, Avalon Peninsula, Bassett 541 (DAO). Ferryland, Brooks s.n., 20 July 1937 (GH, MO). Tompkins, Edgerton §_Pease s.n., 4 July 1939 (GH). Birchy Cove, Curling, Bay of Islands, Fernald §_Wiegand 3334 (CAN, GH, NY). Cow Head, north of St. Paul's Bay, Fernald 9 Wiegand 3335 (GH). Deer Pond Brook, highlands of St. John, northwestern Newfoundland, Fernald §_Long 28154 (GH). Dildo Run, southern shores of Notre Dame Bay, Fernald §_Wiegand 5378 (F, GH, MO) . Glenwood, valley of Gander River, Fernald 9 Wiegand 4376 (GH, US). 132 Grand Bruit, district of Burgeo and La Poile, Fernald, Long & Fogg 247 (GH). Grand Falls, Fernald 9 Wiegand 4375 (GH). Port Saunders, region of Ingorachoix Bay, Fernald.9_Wiegand.3336 (GH). Sacred Island, Straits of Belle Isle, Fernald E Long 28150 (GH). Ship Cove, Sacred Bay, Fernald, Wiegand §_Long 28152.(GH). Tilt Cove, northern shores of Notre Dame Bay, Fernald §_Wiegand 5379 (NY, RM, UC). Bell Island, near TOpsail, Conception Bay, Howe 9_Lang 1308 (NY). Channel, Howe 9_Lang 932 (NY). Frenchman's Cove, Bay of Islands, Mackenzie §_Griscom 10263 (GH, US). St. John's, Robinson §_Schrenk 218 (CAN, DAO, F, GH, MIN, MO, NY, US). Hughes Brook, Humber District, Rouleau 1662 (CAN, DAO, NY, US). Hughes Brook, Humber District, Rouleau 1662 (CAN, DAO, NY, US). East Port, Bonavista Bay, Smith, Smith §_Squires 303 (DAO). Fogo Island, Sornborger s.n., 7 August 1903 (CAN, GH, NY, US). Terra Nova National Park, Stirrett 1244 (DAO). NOVA SCOTIA: CAPE BRETON CO.: Sydney, Cape Breton Island, Barnhart 832 (NY); N. W. Cove. Sactari Island. Smith. Schofield, Sampson §_Bent 5351 (DAO). COLCHESTER CO.: Lynn, Dore 45.1095 (DAO); west of Folley Lake, Fassett 19040 (GH, WIS); London- derry Bridge, Great Village River, Smith, Curry §_C1attenburg 19184 (DAO). CUMBERLAND CO.: cliffs near Moose River, about 8 mi. east of Parrsboro, Scoggan 13820 (CAN). DIGBY CO.: Brier Island, Smith, Roland, Collins, Erskine §_Schofield 126 (DAO). GUYSBOROUGH CO.: Cansco, Fowler 25191 (WIS); Boyleston, Hamilton 18294 (CAN, US); Cook's Cover, Perry, Wetmore, Hicks §_Prince 10229 (GH); Guysborough, Rousseau 35324 (CAN, US). HALIFAX CO.: Halibut Cove, near Halifax, Dore, Senn §_Gorham 45.521 (DAO). INVERNESS CO.: near Pleasant Bay, Cape Breton Island, Pease 133 26634 (GH); Cheticamp River, Cape.Breton Island, Smith, Schofield, Taylor, Webster §_Slipp 7799 (DAO); near Margaree Forks, Cape Breton Island, Smith, Taylor, Webster.9181ipp.6228 (DAO); West Mabou Harbour, Smith, Schofield, Sampson §1Bent 4178 (DAO). KINGS CO.: Hall Harbor, Fassett 19049 (WIS); Scott Bay, Prince §_Atwood 1064 (DAO); Black River, Zinch 224 (DAO). LUNENBURG CO.: .Chester, Pease 26655 (GH); near Petite Riviere, Greene Bay, Zinch 700 (DAO). PICTOU CO.: Pictou, Robinson 209 (NY). SHELBURNE CO.: on the island, Barrington Passage, Macoun 80869 (CAN, MO). VICTORIA CO.: Aspy Bay, Cape Breton Island, Churchill s.n., 25 July 1909 (MIN); Baddeck, Cape Breton Island, Macoun 19031 (CAN, GH); Money Rocks, St. Paul Island, Perry 9_Roscoe 194 (CAN, GH, MO, NY, US); Ciboux Island, Smith, Schofield, Taylor, Webster, Slipp 9 Bentley 10932 (CAN, DAO). YARMOUTH CO.: Jassy Lake, Lake Annis, Bean, White §_Linden 21193 (GH, NY, US); Wedgeport, Klawe 1080 (GH); Yarmouth, Macoun 80868 (CAN). Sable Island: 43°59'N., 59°47'W., St. John 1226 (CAN, GH, NY, US). PRINCE EDWARD ISLAND:. Tracadie Beach, Churchill s.n., 23 July 1901 (GH, MO. Brackley Beach, Dore §_Gorham 45.245 (DAO). Near Bon- shaw, Erskine 9_Dore 1105 (DAO, NY). Rocky Point, Fernald 9_St. John 7438 (WS). Charlottetown, Groh s.n., 18 July 1932 (DAO). Souris, Malte s.n., 29 August 1924 (CAN). Near Park Corner, Smith 230 (DAO). QUEBEC: ANTICOSTI ISLAND CO.: Pointe de 1'est, Marie—Victorin, Rolland-Germain E'Louis-Marie 21 642 (GH). BONAVENTURE CO.: Port Daniel, Lepage 13542 (DAO); New Carlisle, Williams §_Fernald 69140 (CAN). BROME CO.: Bolton Pass, Marie-Victorin, Rolland-Germain, Raymond E_ Rouleau 56363 (GH). GASPE-EST CO.: Bonaventure Island, Fernald §_ 134 Collins 1020.(GH); cliffs between Cape.Rosier and Cape Gaspé, Johansen- s.n., 12 August 1922 (CAN); Perce, Marie—Victorin §_RollandeGermain 49 462 (DAO, MICH); Riviere.AuseeAux—Canards, Marie—Victorin, Rolland- Germain 9_Jacques 44 527 (DAO, UC);.Coin du Banc, Scoggan 515 (CAN). LEVIS CO.: Point Garneau, CingeMars.§_Cayouette s.n., 15 September 1954 (DAO). MAGDALINE ISLANDS CO.:..Grindstone, Grindstone Island, Fernald, Bartram, Long §_St. John 7439 (GH); Grand Ruisseau, Ile-aux—Meules, Grandtner, Lamieux §_Rousseau 8191 (DAO); Brion Island, St. John 1875 (GH, US). MATANE CO.: Sandy Bay, St. Lawrence River, Fernald §_Pease 25041 (GH); Riviere Blanche, Forbes 1494 (CAN, DS, GH). RIMOUSKI CO.: Cap a‘l'Original, Bic, Clausen §_Trapido 2827 (MIN, UC); St.-Fabien-sur— mer, Lepage 15843 (DAO). RIVIERE—DE-LOUP CO.: Trois-Pistoles, Lepage 15325 (DAO). SANGUENAY CO.: Blanc Sablon, Straits of Belle Isle, Fernald, Wiegand 9 Long 28156 (GH); Ile des Perroquets, Archipel de Blanca Sablon, St. John s.n., 28 July.1915 (CAN, GH). TEMISCOUATA CO.: Saint-Hongré, Blouin, Carrier, Lemiaux §_Richard 7373 (DAO); Ile Aux Basque, Marie-Victorin, Rolland—Germain §_Jacques 33 032 (GH, RM). WOLFE CO.: Notre-Dame de Ham, Canton d'Ham Nord, Hamel §_Brisson 15399 (CAN); Riviere Saumon, canton de Lingwick, Hamel §_Brisson 14790 (UC). ILES SAINT—PIERRE (France): .Savoyard, Le Hors s.n., 16 August 1950 (DAO); Vigie, Le Hors s.n., 21 September 1931 (DAO). MEXICO: MEXICO: Vertiente sw.del Ixtaccihuatl, La Joya, elev. 3850 m, Rzedowski 21578 (MICH); Vertiente nw del Ixtaccihuatl, en la regiOn de Penas Cuatas, La Ciénega, Rzedowski 21813 (MICH). 135 UNITED STATES: ALASKA: .Haines (Chilkoot Inlet, Lynn Canal, n. of Juneau), Anderson 6047 (CAN). Unalaska (Aleutian Islands), Eyerdam 2273 (CAN, CAS, DS, NY, UC, US).< Girdwood (Cook Inlet, Turnagain Arm, se. of Anchorage), Hultén, s.n., 1 July 1961 (US). Sand Point, Shumagin Islands, Riggs s.n., 31 July 1913 (US). Vicinity of Massacre Bay, Peaceful Valley, Attu Island (Aleutian Islands), VanSchaack 499a (US).. ARKANSAS: PULASKI CO.: without definite local— ity, Hasse s.n., May 1886 (MONTU). CALIFORNIA: AMANDA CO.: weed in University of California Botanical Garden, Berkeley, Crow 1179 (MSC). DEL NORTE CO.: 1 mi. south of Crescent City, Abrams §_Baciga1upi 8393 (DS); along road in redwoods north of Crescent City, Eastwood 12299 (CAS); Laguna Creek, 5 mi. north of Regna, Tracy 5872 (UC). HUMBOLDT CO.: Stone Lagoon, Humboldt Coast, Jepson 9337 (JEPS); Freshwater Creek near Wrangletown, Tracy 5338 (CAS, UC, WTU). MARIN CO.: Shell Beach, Howell 20929 (CAS); Inverness.Ridge, Pt. Reyes Peninsula, about 3 mi. northward from town of InverneSS, Thomas 10413 (DS). MENDOCINO CO.: Highway 1, Caspar, Nobs §_Smith 1149 (CAS). SAN FRANCISCO CO.: Golden Gate Park on cross park boulevard, San Francisco, Howell 93571 (CAS, MSC); crevices of sidewalk on 9th Ave. near Fulton Street, San Francisco, Rubtzoff 2449 (CAS, MSC). SAN MATEO CO.: ravine north of Seal Cove, Endley s.n., 16 March 1900 (DS). SANTA CLARA CO.: weed in Stanford Experimental Garden, Stanford University, Stanford, Thomas 8725 (DS). CONNECTICUT: FAIRFIELD CO.: Fairfield, Johnson s.n., 30 June 1890 (NY); Trumbull s.n., 10 June 1893 (MIN). HARTFORD CO.: Southington, Bissell S.n., 17 June 1900 (GH); crest of Avon Mt., 1/4 mi. south of Farmington 136 Ave., Farmington Twp., Churchill s.n., 10 July 1952 (MSC); South Glastonbury, Wilson Z§_(RM). LITCHFIELD CO.: Norfolk, Redfield 13062 (MO). NEW HAVEN CO.: North.Gui1ford,.Bart1ett s.n., 22 June 1906 (GH); Waterbury, DuBois s.n., August 1889.(UC); Milford, Eames 1494 (MIN); Mt. Carmel, Stafford.s.n., 16 June 1885.(US). NEW LONDON CO.: Mumford”s Point, Groton, Graves s.n., 6 June 1903 (GH); Taftville, Setchell s.n., 28 June 1884 (UC); Franklin, Woodward s.n., 30 August 1914 (GH). TOLLAND CO.: Hop River, Andover, Seymour 17643 (WIS). WINDHAM CO.: Connecticut River, Westminster, Blanchard Z9 (GH). DELAWARE: NEW CASTLE CO.: Wilmington, Tatnall s.n., 21 May 1930 (GH). IDAHO: KOOTENAI CO.: Hayden Lake,.Baker 14882 (WTU). MAINE: CUMBER— LAND CO.: Portland, Garber s.n., 29.August 1874 (F). FRANKLIN CO.: South Chesterville,.Eaton.17001 (WIS). HANCOCK CO.: vicinity of Blue Hill, Maxon 11036 (US); Seal Harbor, Mount Desert Island, Williams s.n., 15 July 1897 (GH). KENNEBEC CO.: Chesterville, Chamberlain, Knowlton 9 Eaton s.n., 18 July 1902 (GH); South.Litchfie1d, Fassett 18292 (WIS). KNOX CO.: Union, Cole 973 (US); Washington, Steyermark 4209 (F). LINCOLN CO.: Ocean Point, Fassett 15442 (WIS); Rams Island, Boothbay, Fassett 3580 (WIS); Monhegan Island, Jenney, Churchill §_Hi11 s.n., 3 July 1919 (M0); Medomak, Steyermark s.n., August 1928 (F). SAGADAHOC CO.: Bowdoinham, Fassett 2850 (WIS). YORK CO.: York, Blake s.n., 23 July 1863 (NY); Fryeburg, Harvey 140 (US); South Berwick, True 1141 (US). WASHINGTON CO.: Cutler, Kennedy,.Wi11iams, Collins 9 Fernald Son., 2 July 1902° MARYLAND: BALTIMORE CO.: Baltimore, Jones s.n., 25 May 1904 (F). MASSACHUSETTS: BARNSTABLE CO.: Woods Hole, Bacon 9§_ 137 (MSC); South Orleans,.Murdoch 2056 (F); Nashawena, Elizabeth Islands, Northrop s.n., July—August 1901 (NY). BERKSHIRE CO.: Stockbridge, Britton s.n., 28-31 July 1901 (NY); Great Barrington, Leavenworth s.n., in 1820 (MSC). BRISTOL CO.: New Bedford, Greene s.n., date unknown (WIS). DUKES CO.: Gay Head, Martha's Vineyard,.Seymour 1201 (GH, NY, US). ESSEX CO.: Newburyport, Fernald s.n., 2 October 1902 (GH); Forbes 17194 (MIN, WIS); Nahant, Russell-s.n., August 1863 (GH). FRANKLIN CO.: Mt. Toby,.Harris s.n., 30 June 1877 (MIN); Charlemont, Hunnewell 10659 (GH); Buckland, Seymour 3812 (WIS). HAMPDEN CO.: Granville, Seymour 121 (GH, MO); Mt..Shatterack, Montgomery, Seymour 8121 (DAO, WIS). HAMPSHIRE CO.: Middlefield Valley of Westfield River, Fernald §_Long 9480 (GH); Huntington, Robinson 776 (GH); MIDDLESEX CO.: Merrimac River, Lowell, Beattie s.n., 21 May 1930 (RM); Ashland, Morong s.n., 19 June 1878 (F, NY); Waltham, Seymour §_Seymour s.n., 4 July 1911 (CAS, MSC, WIS). NANTUCKET CO.: Brant Point road, Nantucket Island, Bicknell s.n., 20 June 1908 (NY); "Ram Pasture," Nantucket Island, MacKeever 417 (NY). PLYMOUTH CO.: Middleboro, Murdoch 596 (F). SUFFOLK CO.: Cambridge, Beal s.n., date unknown (MSC); Revere, Clark s.n., in 1873 (MSC). WORCESTER CO.: Lancaster, Seymour s.n., 5 July 1944 (WIS); Douglas, Weatherby D2197 (US). MICHIGAN: HOUGHTON CO.: Douglas Houghton Falls, Lake Linden, Farwell 6595 (MICH); 1.5 mi. east of Laurium, Hermann 7692 (MSC, NY); Otter.Lake, Hyypio 409 (MSC). KEWEENAW CO.: Allouez, Hermann 7797 (DS, F, MICH, MO, NY. US: WS); Agate Harbor, Hermann 7797 (NY, RM, UC). MARQUETTE CO.: Champion, Hill s.n., 10 July 1889 (GH, NY). WAYNE CO.: golf links, Detroit, 138 Piper s.n., 8 June 1922 (US). MINNESOTA: ST. LOUIS CO.: Duluth, Lakela 2561 (MIN, NY, UC, WS). MONTANA: LINCOLN CO.: Stanley Creek base of Stanely Mt., elev. 760 m, Harvey 5492 (MONTU). NEW HAMPSHIRE: CHESHIRE CO.: Alstead, Seymour §_Seymour s.n., 27 June 1913 (WIS). GRAFTON CO.: road near confluence of Batchelders Brook and Bakers River, Warren Twp., Churchill s.n., 11 July 1939 (MSC); Holdeness, Faxon s.n., 7 June 1886 (GH); Lebanon, Kennedy s.n., 25 September 1894 (GH). HILLSBOROUGH CO.: Peterborough, Batchelder s.n., 3 September 1928 (M0). ROCKINGHAM CO.: Isles.of Shoals, Canby s.n., August 1866 (WS); North Hampton, Eaton s.n., June 1896 (GH). STRATTFORD CO.: Dover, Hodgdon 4679 (WIS). NEW JERSEY: OCEAN CO.: Beach Haven Terrace, Long Beach Island, Long 3798 (GH). PASSAIC CO.: Hewitt, Mackenzie 2850 (M0). NEW YORK: ALBANY CO.: Albany, House 28498 (NY); Loudonville, House 30942 (NY). BRONX CO.: Bronx, Moldenke 20157 (CAN). FRANKLIN CO.: Adirondack Hatchery, Saranac Inn, Muenscher 9 Maguire 1113 (F, MO). FULTON CO.: Nick Stone Golf Course, Caroga Lake, Fassett 10374 (WIS). ONADAGA CO.: near Jamesville, House s.n., 1 August 1903 (NY). ONEIDA CO.: Utica, Harberer 129 (GH, NY). RENSSELAER CO.: Troy, Hall s.n., 1828—1834 (F). RICHMOND CO.: Richmond Valley, Staten Island, E. G. Britton s.n., 24 June 1894 (NY, GH); Giffords, Staten Island, N. L. Britton s.n., 21 May 1889 (NY). ROCKLAND CO.: Clarkes— tOWn Twp., Lehr 1020 (NY). SARATOGA CO.: Saratoga Springs, House 27928 (GH, NY). SUFFOLK CO.: Wading River, Long Island, Miller 327 (UC); Southampton, Long Island, Piper s.n., 12 August 1921 (US); Northville, Long Island, Young s.n., June 1873 (IRS). TOMPKINS CO.: Ithaca, 139 Eames 9870 (GH, MO). WESCHESTER CO.: Tarrytown, Barnhart 1446 (NY). OHIO: PICKAWAY CO.: Tarlton, Bartley 1526 (NY, US). OREGON: CLATSOP CO.: Cannon Beach, Thompson 19746 (MO, NY, WS, WTU). COOS CO.: Fossil Point, Coos Bay, Abrams §_Benson 10577 (DS). CURRY CO.: 4 mi. north of Brookings, VanDeventer s.n.,.23 April, 1963 (CAS); The Heads, Port Orford, Peck 9062 (GH, MO,.NY). .KLAMATH CO.: north end of Lake-of— the—Woods, Peck 16598 (DS). .LAND CO.: just south of Heceta Head lighthouse, Cronquist 6112 (GH, NY, UC, WTU); Cleawox Lake, Detling 2989 (UC). LINCOLN CO.: Yachats, Cooke 10717 (OSC); 1/4 mi. south of Yaquina Bay, Lawrence 1562 (DS, OSC, US). MULTNOMAH CO.: Portland, Piper s.n., June 1921 (US); Albina, Suksdorf 1744 (WS). TILLAMOOK CO.: Garibaldi, Erickson s.n., 15 July 1954 (OSC). Oregon, no specific locality, Elihu Hall s.n. in 1871 (CAN). PENNSYLVANIA: BERKS CO.: Reading, Fisher s.n., 23 August 1905 (MICH). CUMBERLAND CO.: Newville, Wahl, Wherry, Hammond, Stafford 9 Tanger 6624 (GH). LACKAWANA CO.: Scranton, Glowenke 6861 (GH). LEHIGH CO.: Bethlehem, Lochman, s.n., 6 August 1891; Allentown, Schaeffer 56239 (US). MONROE CO.: Pocono Manor, Wherry s.n., 18 June 1945 (DAO). MONTGOMERY CO.: Philadelphia, Parker s.n., 2 July 1865 (F); Philadelphia, Witte s.n., 7 July 1934 (RM, NY). RHODE ISLAND: NEWPORT CO.: Newport, Tweedy s.n., June 1881 (DS). PROVIDENCE CO.: Providence, Collins s.n., 30 May 1891. WASHINGTON CO.: Block Island, Watson s.n., June 1885 (GH); Middletown, Williams s.n., 4 July 1909 (GH). Rhode Island, no specific locality, George Thurber s.n., in 1846 (GH). UTAH: PIUTE CO.: Tate Mine, Marysvale, elev. 9000 ft., Jones 5855 (MO). VERMONT: CALDONIA CO.: Peacham, Blanchard 140 s.n., 31 July 1884 (UC, US); Groton, Seymour 18328 (WIS). ORANGE CO.: Newbury, Jesup 9 Sargent s.n., 29 July 1891 (GH). WASHINGTON CO.: Walden and Cabot, Burbank, Grout §_Egg1eston s.n., 4 July 1894 (NY). WINDHAM CO.: Newfane, Grout.s.n., 2 July 1895 (F). WINDSOR CO.: Rochester, Dutton s.n., 10 July 1914 (GH, MICH, MO). WASHINGTON: CLALLAM CO.: Elcoha River, Olympic Peninsula, Jones 3522. CHELAN CO.: head of Poison Creek, north side of Lake.Che1an, Ward 700 (CAS, WS, WTU). CLATSOP CO.: Cornet Bay, Whidbey Island, Smith 1720 (UC). KING CO.: Kirkland Lake, Eyerdam 1686 (DAO);.Redmond, Frye s.n., 24 October 1908 (WTU); Seattle, Rigg s.n., May 1908; near Renton, Thompson 8956 (M0, WTU). KITTITAS CO.: Table Mt., Wenatchee Mts., elev. 4500 ft., Thompson 9743 (WTU); Snoqualmie.Pass, Wiegand 840 (F). KITSAP CO.: Colby, Warren 276 (WS, WTU). KLICKITAT.CO.:. Bingen, Suksdorf 5013 (WS). PACIFIC CO.: .Ilwaco, Abrams 11272 (DS). SAN JUAN CO.: Long Island, Muenscher 9_Muenscher 15979 (MIN). SKAGIT CO.: Bear Creek, T36N, R8E, sec. 10, Mt. Baker Nationa1-Forest, Crow 1241 (MSC). SNOHOMISH CO.: Everett, Minch s.n., 23 July 1928 (WS). WAHKIAKUM CO.: Cathlamet, Foster s.n., 10 May 1907 (WS). WHATCOM CO.: Little Sandy Creek, Baker Lake, T37N, R8E, sec. 12, Crow 1238 (MSC); Birch Bay, Muencher 7786 (UC, WS); Kendall, Muenscher 10184 (DS, WTU); Bellingham, Thomas 9967 (DS). WISCONSIN: IRON CO.: Hurley, Fassett 9541 (WIS). A few specimens from the coast of Labrador and of the St. Lawrence seaway have been recognized as S, procumbens var. compacta Lange. These plants appear to be depauperate, environmentally induced growth forms. Dr. J. K. Morton (personal communication) has likewise 141 . _ ' h Figure 23~ Geographic distribution of Sagina procumbens in Nort America. n North 142 SCALE o mo too we coo soc coo voo coo coo mm m: i \ gk 4 L _L A A A A A._ _J 7 Y v v v T v Y r v 7" v—W 0 [W 000 I00 .00 '0“ IIOO I00 “LOU! VI l\ LANBERT‘S AZIMUTHAL EQUAL-AREA PQOJECYION ;.__:_._ : _ 120 no :06 .0 um macaw; Figure 23 143 questioned the validity of this taxon.. His field observations in a dune situation indicate that there seemed to be a cline from the very compact form ("good” var. compacta) growing away from the shore and becoming more normal toward the shore.: Morton collected seed and vouchers from both extremes and grew the plants under cool greenhouse conditions. Plants grown.from the compact form appear as normal S, procumbens. We agree that var. compacta should be considered no more than a growth form. 4. Sggjfla_subulata (Sw.) Presl §agina_subulata (Sw.) Presl, Fl. Sic. p. 158. 1826. Spergula subulata Sw., Sven, Vet, Acad. Handl. Stockh. p. 45. 1789. Ehalge subulata (Sw.) Dumort., F1. Belgica p. 110. 1827. Spergella subulata (Sw.) Reichb., F1. Germ. Excurs. p. 794. 1832. §ggjga_subulata (Sw.) Wimm., Fl. Schles. p. 76. 1841. Aléiflg subulata Krause, in Sturms, Fl. Deutschl. 2 ed. 5: 34. 1901. Type: not seen. Original material: near Alingsas, Sweden. Sagjfla_Revelieria Jord. & Flourr., Brev. P1. Nov. fasc. i. p. 11. 1866. Type: not seen. Original material: Corsica Mountains, Quenza, Corsica, France. Perennial. Plants tufted, caespitose, frequently forming dense mats. Horizontal stems becoming slightly woody with extensive mat formation. Branches short, with short internodes, ascending or decum- bent, often not exceeding basal leaves. Stems densely glandular pubescent or less frequently glabrous. Leaves densely glandular 144 pubescent or with glandular hairs restricted to the margins, and then often minutely glandular ciliate, rarely glabrous. Leaves with long aristae, 0.5-0.75 mm long. Basal tufts of leaves linear, 3.0-12 mm long, curled inward. Cauline leaves linear-subulate, 3.0-10 mm long. Connate leaf bases scarious, forming a conspicuous cup. Flowers axillary or terminal, usually solitary. Pedicels long, filiform, mean length 22.5 mm, erect during capsular development. Pedicels densely to weakly pubescent. Flowers S-merous, rarely 4- and 5-merous. Sepals elliptical, 1.5-2.0 mm long, the hyaline margins white. Petals elliptical, 1.5-2.0 mm long, shorter than or equaling sepals. Stamens 10, filaments (1.0-) 1.5 mm long, anthers 0.25 mm long. Capsules slightly exceeding sepals. Capsule valves thin, 2.0-3.0 (-3.5) mm long, dehiscing to base. Sepals remaining appressed following capsule dehiscence. Seeds brown, obliquely triangular, with dorsal groove, a distinct notch at hilum, surface smooth, 0.4 (-0.5) mm long. Chromosome number: an=l8, 22. Figure 24. Ecology and distribution: In wet gravelly sands of stream margins. Introduced from Eurasia and known only from Harney Co., Oregon, Bedford Co., Virginia and La Laguna, Baja California. Native to Eurasia. Flowering June to August. Representative specimens: MEXICO: BAJA CALIFORNIA: Sierra de la Laguna, Brandegee s.n., 22 January 1890 (UC). Sierra de la Laguna, Brandegee s.n., 27 March 1892 (GH, UC). Sierra de la Laguna, Brandegee s.n., 4 October 1899 (GH, NY, UC). Cape Region, La Laguna, elev. 6000 ft., Thomas 7885 (CAS, DS, GH, MICH, US)° Figure 24. 145 Photographs of Sagina subulata. (a) Habit, Sweden (Sarnquist s.n., 20 June 1948, DAO). (b) Close-up of aristae, photographed under epi-illumination, Sweden (Bagge 919,, 16 June 1890, UC). 146 Figure 24 147 UNITED STATES: OREGON: BARNEY CO.: Steens Mt., Faegri s.n., 25 July 1965 (OSC); Dino Creek, Steens Mt., Train s.n., 30 July 1935 (US) and 31 July 1935 (MIN). VIRGINIA: BEDFORD CO.: specific locality unknown, Curtiss s.n., 20 May 1872 (M0). Sagina subulata is an extremely variable, wide ranging species of Eurasia. Sagina glabra and S, pilifera, two closely allied European montane taxa of restricted distribution, overlap in numerous character- istics with S, subulata and appear to be divergent expressions of this variable species. Further study of these taxa may reveal that this complex should be considered a single species with three infraspecific taxa. I have seen but a single specimen from eastern North America which is referable to Sagina subulata. Plants treated by Torrey and Gray as S. subulata belong to S, decumbens subsp. decumbens (see dis- cussion under that taxon). In the Northwest, three specimens (IE919.999., July 30, 1935 and July 31. 1935; fgggri_§;n., July 25, 1965) have been collected in the alpine zone of Steens Mt., Harney Co., Oregon which exhibit glandular pubescence. The Train specimens are extremely pubescent while the Faegri specimen is weakly glandular, with foliar pubescence restricted to leaf margins. Introduction of this population of S, subulata into this remote area is without explanation. Another isolated population of 9. Egbgla:g_occurs at the tip of Baja California. Specimens collected by T. S. Brandegee during the period 1890-1899 and by J. H. Thomas in 1959 in the mountains at l48 La Laguna (ca. 5,500 ft. elevation) occur in the region of an abandoned ranch which had been operated in the l800‘s (Goldman, l95l). Introduc- tion of this population took place during the period that the ranch was actively worked. A mat-producing form of this taxon is widely used as a ground cover and is available from Nurserymen under the names ”Scottish Moss" and "Corsican Pearlwort.“ Living plants observed in the w. J. Beal Botanical Garden, Michigan State University, were noted to flower profusely but with no subsequent capsular development. Herbarium specimens from several localities in the western United States likewise show little capsular development. The cultivar propagates easily by vegetative means, but is not found readily escaping cultivation. The cultivar differs from the native European mat-producing form by being glabrous, except for the minutely glandular-ciliate leaf margins. The cultivated form originated in the Corse Mountains, Corsica, France (Vilmorin, l894). 5. Sagina nivalis (Lindbl.) Fries Sagina nivalis (Lindbl.) Fries, Nov. Fl. Suec. Mant. 3: 3l. l842. Spergula saginoides var. niyalis Lindbl., in Physiogr. sallsk. Tidskr. p. 328. l838. Type: not seen. Original material: region of Kongsvold near Doores, Norway, 24 September l837. Sagina intermedia Fenzl, in Ledeb., Fl. Ross. l: 339. l842. Type: not seen. Original material: region of Tschuktschorum along bays of St. Laurent, Russia. l49 Perennial. Caespitose, forming low cushions. Basal rosette of succulent, subulate leaves, tips apiculate. Flowering stems numerous, radiating from axils of basal rosette leaves. Stems slender, sometimes purple tinged. Cauline leaves subulate,.with connate leaf bases forming shallow scarious cup, often purplish, becoming shorter toward stem apex. Pedicels long, filiform, glabrous.. Flowers 4-merous or 4- and S—merous. Sepals l.5-2.0 mm long, nearly orbicular to elliptical, rounded at tip, glabrous, frequently purplish, hyaline margins nearly always purple, sometimes only at tip. Petals equaling to slightly shorter than sepals, l.5—2.0 mm long. Stamens 8 or l0, filaments l.5 mm long, anthers 0.25 mm long. Capsules 4- or 5-valved, 2.0—3.0 mm long, dehiscing to base. Capsule valves thick. Sepals remaining appressed following capsule dehiscence. Seeds brown, obliquely.triangular, with dorsal groove, distinctly notched at hilum, lateral surfaces frequently with elongate ridges, dorsal surface appearing smooth to pebbled, 0.5 mm long. Chromosome number: 23;=84, 88. Figure 25. Ecology and distribution: Growing on sandy or gravelly beaches, coastal rocks, alluvial plains, fresh glacial moraines and low, swampy tundra and in alpine areas. Widely distributed in the Arctic Archipel— ago, Alaska, Hudsoanames Bay region and the coast of Labrador. Disjunct population in Alberta. Flowering July and August. Figure 26. figpresentative specimens: CANADA: ALBERTA: 'Medicine Lake, Jasper National Park, Scamman 2527 (GH). Mt. Edith Cavell, Angel Glacier, Jasper National Park, Scamman 2446 (CH), mixed collection. LABRADOR: Chateau, 51°49“N. Allen s.n., 8 August 1882 (F, NY). 150 . IS, Figure 25. Photographs of Sagina nivalis (a) Habit, Axel Heiberg N. w . (He and Bes__chel 10857, GH) (b) Habit Attu 15’ Aleutians Jordal and Miller 3077- A, MICH) hm!HIdunlmiluulnllllmlufihmlufil METRIC 1 MUFIC I Figure 25 l52 Nunaksuk Island, Bishop 285b (GH). Crater Lake vicinity, ca. 52 mi. west southwest of Hebron, 58°02’N., 64°02”W., Gillett 8938 (DAO, US). Makkovik, 55°N., Hustich 68 (CAN). Cutthroat Harbour, south of Cape Mugford, 57°30'N., 62°W., Porsild 189 (CAN). Hebron, Sornborger 207 (GH). Battle Harbour, Waghorne 4901 (CAN). Okkak, near Cutthroat Tickle, 57°40'N., 62°W., Wynn-Edwards 7476 (CAN). NORTHWEST TERRITORIES: DISTRICT OF FRANKLIN: Northwest Middle Fiord, Axel Heiberg Island, Beschel 13118 (CAN). Mould Bay, Prince Patrick Island, 76°14'N., 118°59”W., Bruggemann 361 (DAO, NY, UC). Frobisher Bay, Baffin Island, 63°45°N., 68°32“W., Calder 2085 (DAO, US). Erik Harbour, Baffin Island, 72°40"N., 76°30"W., Coombs 31;(DAO). Head of Inugsuin Fiord, Baffin Island, Hainault 3634 (CAN). Ferguson Lake, Victoria Island, 69°25'N., 105°15'W., Jones 29a (DAO). Clyde, Baffin Island, Martin 23 (DAO). Beekman Peninsula, southeast Baffin Island, ca. 63°25'N., 64°75'W., McLaren 103 (CAN). Pangnirtung, Baffin Island, Polunin 1168 (US). Arctic Bay, Baffin Island, Polunin 2555 (CAN). Banks Island, near northeast corner, ca. 73°24"N., 117°W., Porsild 17677 (CAN). Bernard Island on northwest coast of Banks Island, Porsild 17747 (CAN). Resolute Bay, Cornwallis Island, Porsild.2l672 (CAN). Head of Minto Inlet, Victoria Island, Porsild 17388 (CAN). Botany Bay, Kangerdluak Fiord, Ekalugad Fiord region, Baffin Island, Webber 1302 (CAN). Isortoq Fiord, Baffin Island, Webber 382 (CAN). Isortoq River at Lewis Glacier, Webber 140 (CAN). Cumberland Sound, opposite Pangnirtung, Baffin Island, WXflnerEdwards 9364 (CAN). Cape Searle, Padloping, ca. 67°10'N., 62°30'W., W nne-Edwards 9147 (CAN). DISTRICT OF KEEWATIN: Cape Jones, Baldwin, M 153 Hustich, Kucyniak §_Tuomikoski.664 (CAN). Smith Island, east coast of Hudson Bay, Baldwin 1824.(CAN,.GH).. Coral Harbour, South Hampton Island, Calder, Savile §_Kukkonen.24228 (DAO).. Whale Point, northwest coast Hudson Bay, Comer s.n.,.August 1894 (GH). Fullerton, Hudson Bay, 63°57'N., Macoun 79091 (CAN, GH). Port Burwell, Hudson Strait, 60°22'N., 64°50'W., Malte s.n., 25-28 July 1928 (CAN, GH). Mistake Bay, 62°05'N., 93°06'W., Porsild 5664 (CAN). Yathkyed or Hikolikdjuak Lake on the Kazan River, 62°30'-63°N., 97°-98°30”W., Porsild 5795 (CAN). Chester- field Inlet, 1 mi. north of settlement, 63°21'N., 90°42'W., Saville g Watts 1266 (DAO, MO, WTU) . DISTRICT OF MACKENZIE: O'Grady Lake, Mackenzie Mts., 63°00'N., 129°02'W., Cody 16416 (DAO). Bathurst Inlet, west side, Keisall & McEwen 190 (CAN). Atkinson Point, Arctic Coast, ca. 70°N., 131°20'W., Porsild §_Porsild 2601 (CAN). Cape Dalhousie, Arctic Coast, ca. 70°20'N., 129°55'W., Porsild §_Porsi1d 2747 (CAN). 6 mi. east of Kittigazuit, Arctic Coast, ca. 69°20'N., 133°W., Porsild §_Porsild 2482 (CAN). Shingle Point, Arctic Coast, ca. 69°N., 137°30'W., Egrsild 7099 (CAN). QUEBEC. Richmond Gulf, Cairn Island, east coast of Hudson Bay, ca. 56°15'N., 76°30'W., Abbe, Abbe fi_fl3££_fi§22_(MINlu Stromness Harbor, Fort George, James Bay, 53°56“N., Dutilly E Lepage L239 (GH) . YUKON TERRITORY: Mayo Landing, WEE. §_ (DAO) . Canol Rd., Mile 95, upper Rose River valley, Porsild §_Breitung 10368 (CAN). Wain- UNITED STATES: ALASKA: Mendenhall, Anderson 434 (NY). Wright, Anderson 4364 (UC). Lowell Creek Canyon, Seward, Kenai Penin— sula, Calder 5898 (DAO). Snow River delta, Kenai Lake, Kenai Peninsula, 60°19'Nor 149°21'W., Calder 6573 (DAO). Head of Katmai River, Katmai l54 National Monument, Cahalane 518 (US). Ca. 40 mi. east of Cape Lisburne, 4 mi. inland along Pitmegea River, Cantlon §_Gillis 57-2453 (MSC). Okpilak River, 69°23“N., 144904'W., Cantlon E Gillis 57—2287 (MSC, US). Glacier Bay, Cooper 222 (US). Columbia Bay, Prince William Sound, Cooper 315 (F, MIN). Muir Inlet,.G1acier Bay, Coville §_Kearney 630 (US). Port Vita, Raspberry Island, Kodiak group, Eyerdam 5137 (CAS, DAO, MIN). Sable Pass, Mt. McKinley National Park, Frohne 54-256 (DS). Tangle Lakes area, east of Landmark Gap, Alaska Range, Gjaerevoll 1324 (CAN). Mead River village, Northern Coastal Plain, Hultén s.n., 5-8 Aug— ust 1960 (US). Uyak, Kodiak Island, Jepson 391 (US, UC). Olga Bay, Kodiak Island, Looff §_Looff 1027 (GH). Dexter Creek, Nome, Seward Peninsula, Porsild §_Porsild.l340 (CAN). Pastolik, Norton Sound, Porsild & Porsild 985 (GH). Unalaklet, Norton Sound, Porsild §_Porsild 1148 (CAN). Port Clarence, Teller, Seward Peninsula, Porsild §_Porsi1d 1432 (CAN). Paxon, Alaska Range, Porsild_§_Porsild 574 (CAN). Point Hope, Scamman 6390 (CH). Meade River, 1 mi. southeast of Atkasuk, E:22:,199 (DS, RM). Iko Bay, vicinity of Point Barrow, Thompson 1395 (D57 US). Columbia Glacier, Heather Island, 60°N., l47°W., Xiereck g Viereck 2312 (COLO). Nanivak, 7 mi. southwest of Point Barrow, Wiggins 13293 (0s, GH, UC, us, WTU). Point Barrow, Wiggins 12946 (DS, GH, UC, WTU). Near Walakpa triangulation target, southwest of Barrow Village, Wiggins 12511A (DS). Cape Thompson, 68°05“N., 165°40"W., Wood §_Wood Ell (CAN). ALLEUTIAN ISLANDS: Amchitka Island, Erdman 578 (COLO). Amlia Island, Eyerdam 1244 (CAS, DS, GH). Nunivak Island, Haley s.n., 1 July 1927 (CAS). Attu Island, Hardy 385 (GH, MIN, WTU). Attu Island, Howard 30 (US). Umnak Island, Nikolski, Hultén 7041 (CAS). Unalaska, 155 Figure 26. Geographic distribution of Sagina nivalis in North America. 156 _L.—__ -~. J I ‘,.'J“‘- -f /’\{/'\f I I \ r/l I \l/J ‘. NM”, \ a—___..-(.- r /, J‘K/"I K (K; I/\}-——-.-‘ mint 1 ,' I, ‘ - -)_ \y (' .\ \ . l ., \ - \. m .n”_ ‘ i .’ w 1 f \‘ -Alt'-‘_/',’ /‘ -_r_‘._1_-_.1—--—.—"‘;. / é ""‘ \1'7’KI‘Q'H“ \. ' .J/\fW\ : . x 5 A. ( ‘ : ‘\ .. . _ _ 1' - _ 1. - _\ u(\ 7 . . — - 4’ l | r _ _ n: 4 \_\‘ . ,~ \ \\ ' - -'-~l._ w‘. -' .* _ ‘ \— \ \ . I. ._r' I n \ , 1 I. \ ' \ '\ . I ‘-' ‘ . 1 l ‘ ,_ 1 ,' x- ___ , I \‘ q I ‘~ I I -. . . .7‘_ \ . { \.«". ' ' c -. e I ' r I ao- ‘~«-. . ~ " i ’ ~\ » . ~ , \ __ -‘~. ' - —___ n ' — _7 _____ \‘I\ _ \ -’r~ .4.-.1.,_ _. _ --— x‘ - 1 1 .. \p I .‘ . I I - a - v-0 » n 1'!,\_.. ‘ L‘_ \ ‘ ,1 \ ‘r . \’_. ‘->“':-, ° ¢ 1 -l\ , __ -‘ . t s, . r v < \’ he... ~-, A v\"‘ 1 , . .. r " ' 1‘ “ .z‘ ‘I ’3 , " a 7 a I, u- . .: I‘ ‘Llla ‘-,_ —-.' I .,‘_ -/‘.“ /_ \f", __. g . . . , - . ._ (I - f I\ I _ , . \a‘.- I,_ I .l' . ‘ , . 0 I‘..-\. 0-‘ o |O '. P - I SCALE 0 00 no ran 4W 500 000 '00 .00 .00 ‘00) U~Ll$ . 0 too v.00 fooo 00-0 '000 v '50 400 I|LOU['l-3 LMBCRT'S AZIMUTHAL EQUAL-ARCA PROJECTION o \,,\ O I K __ __ _ __ __ ___ _ ———-—————°-- — - A — ,wv, g_ “a _ _ # _ . \ yo '---— E5 :10 _ _ no __ _ _ _ _ ‘0 .0 u" Lawn... Figure 26 l57 Hultén 6708b (US). Ananuliak.Island,.Umnak,.5 mi. of Nikolski, Johnson 290 (WIS). Adak Island, Jordal 262l-A (MICH, US). Atka Island, Oliver g Oliver 55 (MICH, US). BERING.SEA: Pribilof Islands, St. George Island, Johnston s.n., 5 May.l920 (CAS). Pribilof Islands, St. Paul Island, Macoun s.n., 28 July 1891.(GH, MO, NY, US). St. Lawrence Island, Gambell, Anderson 5185 (CAN). Sagina nivalis occurs disjunction in Alberta. I have seen only two specimens from Alberta which representfgood" S, nivalis (Scamman 2446 and 2527). This population is mixed with S, saginoides. There are several specimens from Banff and Jasper National Parks which appear intermediate between S, nivalis and S, saginoides. There are several specimens from alpine habitats in Colorado, Utah and Wyoming which are suggestive of S, nivalis. These specimens are densely caespitose and the sepal margins are distinctly purple. These 5~merous specimens, however, belong to S, saginoides. 6. Sagina caespitosa (J. Vahl) Lange Sagina caespitosa (J. Vahl).Lange, Consp. Fl. Groenl., Meddel. Groenl. l: 22. l880. Arenaria caespitosa J. Vahl, Icon. Fl. Danica. l8: 4. l840. Sagina nivalis var. caespitosa (J. Vahl) Boiv., Nat. Can. 93: 583-646. Type: not seen. Original material: growing in wet clayey and sandy gravel filled places, ca. 60 ft. above sea level, ESpecially in bays of Greenland from Godthaab to Upernavik, and occurs from 64° to 72° 48'. Perenn rosette of lea linear to line radiating, fre shorter towarc Turning shallr glandular pub: Flowers 5-mer ovate to lanc usually purpl pubescent or equaling, sen 0.25-0.3 nm ' to base. Cal callsule dehi- lrunve, dist e”Nate l‘ld lung, Chrom E01 lines and Si Welly hil Bay and non Hlure 28. 3e, 1 ‘kordlearsu l58 Perennial. Caespitose, forming small mats or cushions. Basal rosette of leaves lacking; secondary rosettes usually present, leaves linear to linear-subulate. Flowering stems numerous, ascending to radiating, frequently purple tinged. Cauline leaves subulate, becoming shorter toward apex, midvein frequently.conspicuous; connate leaf bases forming shallow scarious cup, often purplish. Pedicels long, filiform, glandular pubescent, rarely glabrous (in North American plants). Flowers 5-merous or 5- and 4-merous. Sepals 2.0-2.5 mm long, broadly ovate to lanceolate, tips obtuse to somewhat acute, hyaline margins usually purple tinged, at least at the tip. Calyx base glandular pubescent or glabrous. Petals 2.5-3.0 mm long, exceeding, seldom equaling, sepals. Stamens l0 or 8, filaments l.5-2.0 mm long, anthers 0.25-0.3 mm long. Capsules 5- or 4-valved, 3.0-3.5 mm long, dehiscing to base. Capsule valves thick. Sepals remaining appressed following capsule dehiscence. Seeds brown, obliquely triangular, with dorsal groove, distinctly notched at hilum, lateral surfaces frequently with elongate ridges, dorsal surface appearing smooth to pebbled, 0.5 mm long. Chromosome number: 2§f=88. Figure 27. Ecology and distribution: In wet sands and gravels of shore- lines and stream margins, wet mossy places, and dry rocky barrens and gravelly hillocks. Northeast Arctic, south to northern Manitoba, James Bay and northern Labrador. Amphi-Atlantic. Flowering July and August. Figure 28. Representative specimens: CANADA: LABRADOR: Cape Mugford Peninsula, Kaumajet Mts., 57°50'N., 62°50'W., Abbe 269 (GH). East Bay, Ikordlearsuk, Torngate region, 59°57‘N., 64°24'W., Abbe §_Ode11 270 (GH). 159 Figure 27. Photographs of Sagina caespitosa. (a) Habit. (b) Close-up showing pubescence on calyx base and pedicel. Both Baralzon Lake, Manitoba (Scoggan and Baldwin 8206, CAN). 160 Figure 27 161 MANITOBA: -Baralzon Lake, 60°00'N., 98°10'W., Scoggan §_Ba1dwin 8206 (CAN, MIN). NORTHWEST TERRITORIES: DISTRICT OF FRANKLIN: Inugsuin Fiord, Baffin Island, ca. 70°N., 68°30'W., Hainault 3836 (CAN). Beekman Peninsula, southeast Baffin Island, ca. 63°20'N., 64°50'W., McLaren 142 (CAN). Pangnirtung, Baffin Island, Polunin 1568 (WIS). Point Brewster, Frobisher Bay, Baffin Island, Potter 8237 (GH).- Resolution Island, Frobisher Bay, Baffin Island, Potter 8238 (GH). Tolnes Road, Baffin Island, 66°27'N., Seidenfaden 1281 (NY). Isortoq Fiord, Baffin Island, ca. 70°N., 77°W., Webber 407 (CAN). DISTRICT OF KEEWATIN: Cape Jones, Baldwin, Hustich, Kucyniak §_Tuomikoski 681a (CAN). Kaminak Lake, 62°N., 95°W., Gfissow 114a (DAO). Port Burwell, Hudson Strait, 60°22'N., 64°50'W., Malte s.n., 25—28 July 1928 (CAN, GH). Baker Lake, north shore, 64°30'N., 97°W., Porsild 6094 (CAN). Kazan River, 62°30'-63°N., 97°—98°30'W., Porsild 5798.(CAN). QUEBEC: Fort Chimo area, 58°07‘N., 68°23'W., Calder 2662 (DAO). Fort George, James Bay, 53°53'N., Dutilly §_Ernest 12500 (GH). Korok River, east side of Ungava Bay, 42 mi. in— land from Korok Bay, 58°35“N., 64°15“-66°W., Rousseau 1111 (DAO). The material of North America and western Greenland differs from European plants of this taxon in having glandular trichomes on the upper portion of the pedicels and on the calyx bases. The European plants are totally glabrous. Amount of glandular pubescence in the North American material is variable. Weakly glandular forms are fre- quent and occasionally glabrous specimens are encountered. Plants of the glandular pubescent form examined possessed an average of 70% glandular flowers while 30% of the flowers were glabrous. Separate 162 Figure 28. Geographical distribution of Sagina caespitosa in North America. am arsaei 1.111 /. 163 14.1111I1( -1 11.1 . Cu 4,11 164 nomenclatural recognition of European and North American material seems unnecessary . 7. Sagina decumbens (Ell.) T. & G. Key to the subspecies: Seeds light tan, with delicate reticulate ridge pattern; surface smooth or tuberculate . . . 7a. subsp. decumbens Seed light brown, never with reticulate ridge pattern; surface smooth to slightly pebbled . . . 7b. subsp. occidentalis 7a. Sggiflg decumbens (Ell.) T. & G. subsp. decumbens Sggjpg_decumbens (Ell.) T. & G., Fl. N. Am. 1: 77. 1838. Spergula decumbens Ell., Sketch l: 523. 1821. Type: unknown. Sggjp§_subulata var. Smithii Gray, Manual 5 ed. p. 95. 1867. Type: S, S, Sgfl§b_§,fl,, June 1865, sandy road in the pine woods, at the mouth of Great Egg Harbor, New Jersey (GH, lectotypel). Annual with slender taproot. Branches slender, ascending or decumbent. Basal rosette of leaves lacking or early deciduous. Lower cauline leaves linear, 4-22 mm long, leaf bases connate, typically with conspicuous hyaline margins, connate portion not generally appearing inflated, apices apiculate. Anthocyanins frequently giving stems and connate leaf bases a purplish tinge. Upper cauline leaves becoming subulate toward apex, 1.5-5 mm long at tip, apiculate. Pedicels fili— form, glabrous or glandular pubescent. Flowers 5-merous, rarely 4- merous, Calyx base glabrOus or glandular pubescent, often sparsely so. 165 Sepals ovate, hyaline margin conspicuous, margins or apex frequently purple. Sepals (1.0-) 1.5-2.0 (-3.0) mm long. Petals elliptical, slightly exceeding sepals at anthesis, equal or shorter than sepals during capsule development, (O.75-) 1.0-2.0 (—2.25) mm long. Stamens 10 or fewer, filaments (1.0-) 1.5 mm long, anthers 0.25 mm long, dehiscing less than half the length of the capsule. Sepals remaining appressed after capsule dehiscence. Seeds light tan, obliquely tri- angular, with dorsal groove, surface smooth or tuberculate, with delicate reticulate ridge pattern (sometimes obscure), (0.25-) 0.3-1.4 um long. Chromosome number 2pg=36. Figure 29. Ecology and distribution: In moist or dryish sandy places, frequently at field margins, open pine woods, paths, roadsides, sidewalk cracks and lawns. Southeastern half of the United States, from Connect- icut and southern New Jersey, west to southern Ohio, southern Indiana, Illinois, Missouri and southeastern Kansas, south to northern Florida and eastern Texas. Disjunct populations in Arizona, Alberta and New Brunswick. Sagina decumbens subsp. decumbens is chiefly a Coastal Plain and Piedmont taxon. It appears to have extended its range westward with civilization, particularly into Kansas, Oklahoma, northeast Texas, Arizona, Alberta and Saskatchewan. Flowering April to June. Figure 30. Representative specimens: CANADA: ALBERTA: east of Hand Hills, Macoun s.n., 8 August 1879 (CAN, DAO). Buffalo Plains, Macoun s.n., 10 August 1879 (US). SASKATCHEWAN: hillsides, Farewell Creek, Cypress Hills, Macoun s.n., 27 June 1895 (CAN). Figure 29. 166 Photographs of Sagina decumbens subsp. decumbens. (a) Habit. (b) Close-up showing glandular pubescence. Horry Co., South Carolina (Weatherby §_Griscom l6524, US). Figure 29 168 UNITED STATES: ALABAMA: CULLMAN CO.: Cullman, Mohr s.n., 1 April 1884 (US). ESCAMBIA CO.: Atmore, Blanton 201 (DAO, F, GH, MICH). LEE CO.: Auburn, Earle §_Baker s.n., 18 April 1898 (F, NY, RM, US). MOBILE CO.: Mobile, Mohr s.n., 24 March 1861 (US). ARIZONA: mountains between Miami and Superior (border of Pinal and Gila Counties), Nelson §_Nelson 1907 (CAS, GH, MO, NY, RM, UC, US). PIMA CO.: Rincon Range Station, Darrow s.n.,.ll April 1937 (CAS). ARKANSAS: BRENTON CO.: Decatur, Plank s.n., April 1899 (NY). BRADLEY CO.: Warren, Demaree 21484 (GH). CLAY CO.: Corning, Letterman s.n., May 1884 (M0). FRANKLIN CO.: Mulberry River, Cass, Fassett 17439 (WIS). GREENE CO.: specific locality unknown, Eggert s.n., 29 April 1893 (M0). HEMPSTEAD CO.: Fulton, Bush 2434 (M0). INDEPENDENCE CO.: Newark, Eggert s.n., 23 April 1896 (M0). NEVADA CO.: Prescott, Bush 533 (MO). PULASKI CO.: Little Rock, Demaree 22725 (MO). WASHINGTON CO.: Savoy, Fassett 17440 (WIS); Brentwood, Palmer 8206 (NY). CONNECTICUT: FAIRFIELD CO.: Fair— field, Eames s.n.. 19 June 1898 (GB): NEW HAVEN CO.: Milford, Eames s.n., 7 June 1898 (GH). DELAWARE: KENT CO.: Choptank Mills, Tatnall 2130 (GH). SUSSEX CO.: Millsboro, Commons s.n., 23 May 1876 (GH); Bethany Beach, Tatnall 3321 (GH). DISTRICT OF COLUMBIA: Washington, Morong s.n., 22 May 1877 (NY). FLORIDA: ALACHUA CO.: Gainsville, Wiggins 19394 (NY). BAY CO.: Lynn Haven, Banker 3671 (NY). DUVAL CO.: near Jacksonville, Curtiss 6353 (DS, GH, MIN, M0: NY, UC, US): GIL- CHRIST co.: 9 mi. south of Bell, Wiggins 19493 (US). HILLSBOROUGH CO.: no specific locality, Fredholm 6314 (GH, MIN). JACKSON CO.: Marianna Caverns State Park, Godfrey 55331 (GH, NY). LEON CO.: near Tallahassee, 169 Rugel s.n., April 1843 (M0). LIBERTY CO.: Apalachicola River swamp south of Bristol, Small, DeWinkelerfi Mosier 11263 (NY). ST. JOHNS CO.: St. Augustine, Leeds s.n., 4 March 1893 (F). GEORGIA: BARTOW CO.: 2 1/4 mi. northwest of Acworth, 7 1/2 mi. southeast of Centerville, Duncan 8041 (M0). CALHOUN CO.: Edison, Collom s.n., 18 April 1953 (M0). CHATHAM CO.: Tybee Island, Harper 2175 (GH, MO, US). CLARKE CO.: 3 mi. west of Winterville, Cronquist 4237 (GH, NY, US); Athens, Harper s.n., 20 June 1900 (NY). DEKALB CO.: Decatur, grgw_123§_(MSC); Stone Moun— tain State Park, grow 1212 (MSC). GLYNN CO.: 5 mi. west of Brunswick, Cronquist 4913 (US). GWINNETT CO.: Yellow River near McGuire's Mill, §g§$l_§;£,, 7 May 1895 (F, NY). MACON CO.: near Macon, Mgh£_§;2,, April 1915 (US). RABUN CO.: canyon at Tallulah Falls, Small_§;n,, 20 April 1893 (F). RICHMOND CO.: Augusta, Ergw_122§_(MSC). WILKES CO.: just east of Dry Fork of Long Creek, between Washington and Lexington, Cronquist 4229 (GH, MICH, MO, NY; UC, US). ILLINOIS: CHAMPAIGN CO.: Urbana, Jones 19605 (DAO). JACKSON CO.: Giant City State Park, Fassett 21467 (WIS). .JOHNSON CO.: Round Bluff, south of Goresville, Bailey 2194 (MIN); Vienna, Winterringer 6281 (F). PEORIA CO.: Peoria, Chase 10374 (F). PULASKI CO.: Wetang, Vasey s.n., no date (F, NY) . UNION CO.: Cobden, wgg (NY). INDIANA: BROWN CO.: Nashville, L192_§;23, 22 May 1930 (MICH). CLARK CO.: Charlestown, §§i£§_§;§,, 2 June 1877 (MICH). JEFFERSON CO.: Hanover, Barnes 12_ (WIS). POSEY CO.: Mt. Vernon, 22§§_§§;§§_(GH). SPENCER CO.: Rockport, Collector and date unknown (GH). KANSAS: CHEROKEE CO.: specific local— ity unknown, Hitchcock 628 (GH, MO, NY, RM. US); 2 mi. northwest Of I70 Baster Springs, McGregor 15337 (US). KENTUCKY: LYON CO.: Kuttawa, Eggleston 4641 (MIN, NY). LOUISIANA: EAST BATON ROUGE PARISH: Baton Rouge, Brown 860 (NY). IBERIA PARISH: Weeks Island, Thieret 16999 (US). IBERVILLE PARISH: Plaquemine, Barnhart 2822 (NY). LAFAYETTE PARISH: Lafayette, Thieret 10333 (DAO). LA SALLE PARISH: Catahoula Lake, 3 mi. southeast of Nebo, Ewan 19066 (GH). PLAQUEMINES PARISH: Point 1a Hache, Langlois s.n., April 1883 (NY, UC). RAPIDES PARISH: Bluffs of Red River, vicinity of Alexandria, Ball 411 (MO, NY, US). ST. MARTIN PARISH: St. Martinville, Langlois s.n., 15 March 1892 (MICH, MIN). ST. TAMMANY PARISH: vicinity of Covington, Arséne 11978 (US). MARYLAND: TALBOT CO.: 2 1/4 mi. southwest of Longwoods, Earle 4062 (GH). WORCES— TER CO.: 10 mi. southeast of Salisbury, Tatnall 1773 (WS). MISSISSIPPI: CARROLL CO.: North Carrollton, Clute 3§_(F, NY)° FORREST CO.: south— east of Hattiesburg, Cooley, Pease §_Ray 3243 (GH). JACKSON CO.: Biloxi, Tracy 5040 (F,.M1CH, MO, MSC, NY). LEE CO.: Natchez Trace -Parkway, McDougall 1804 (US)° PEARL RIVER CO.: 3 mi. north of Picayune, Rose 8046 (CAS). WARREN CO.: Snyder's Bluff, Cooley 3336 (GH). MISSOURI: BARRY CO.: Eagle Rock, BuSh 507 (MO) . BOLLINGER CO.:: 5 mi. west of Grassy, Steyermark 18982 (MO). BOONE CO.: 4 mi. south~ east of Ashland, Drones 1910 (CAS, GH). BUTLER CO.: Neelyville, Bush §§_(MIN); 5 mi. southwest of Qulin, Steyermark 26655 (F). CARTER CO.: Grandin, Bush 340 (MO). CEDAR CO.: 3 mi. north of Stockton, Steyermark 18655 (MO). CHRISTIAN CO.: Chadwick, Bush 4441A (MO). DALLAS CO.: between Plad and Buffalo, Steyermark 18691 (MO). DOUGLAS CO.: along north fork of White River between Roosevelt and Richville, Steyermark 19156 (MO). DUNKIN CO.: about 5 mi. northwest of Campbell, Steyermark I71 398 (MO). FRANKLIN CO.: Pacific, Eggert s.n., 23 May 1882 (MO, NY, US). GREENE CO.: specific locality unknown, Blankinship s.n., 23 April 1888 (MO). HENRY CO.: 3 mi. northeast of Finey, Steyermark 18774 (MO). JEFFERSON CO.: Hasse s.n., 24 May 1887 (M0). LACLEDE CO.: north of Hazel Green, Steyermark 8075 (MIN). LAWRENCE CO.: east of Chesapeak, Steyermark 4519 (M0). MAC DONALD CO.: specific locality unknown, Bush s.n., 24 April 1891 (M0). MILLER CO.: 4 mi. south of Bogville Dam, Steyermark 18798 (MO). OREGON CO.: 4 mi. south of Kosh- konong, Steyermark 18970 (MO). OSAGE CO.: east of Linn, Steyermark 18709 (MO). PHELPS CO.: 4 mi. southeast of St. James, Steyermark 22188 (F). POLK CO.: north of Burns, Steyermark 18665 (MO). PULASKI CO.: 1 mi. west of Jerome, Steyermark 4600 (M0). REYNOLDS CO.: south of Ellington, Steyermark 7944 (M0). ST. FRANCOIS CO.: Bismark, Russell s.n., 15 April 1898 (M0). ST. LOUIS CO.: Forest Park (in St. Louis), Steyermark 1728 (F). SCOTT CO.: 2 mi. south of Benton, Steyermark 10256 (MO). SHANNON CO.: Montier, Bush s3 (MO). TEXAS CO.: along Jack”s Fork of Current River, 3 mi. south of Arroll, Steyermark 18588 (MO). WAYNE CO.: south of Greenville, Anderson s.n., 31 May 1939 (M0). NEW JERSEY: ATLANTIC CO.: Pleasant Mills, Gross s.n., 12 May 1884 (NY); Atlantic City, Redfield 4180 (M0); Somer's Pt., Smith s.n., June 1865 (MO, NY). BERGEN CO.: near Hewitts, Britton s.n., 29 June 1886 (NY). BURLINGTON CO.: Atsion, Canby s.n., August 1863 (NY). CAMDEN CO.: Camden, Parker, s.n., 4 July 1866 (M0). CAPE MAY CO.: Cold Springs, Brown 5236 (GH). OCEAN CO.: Forked River, Churchill s.n., 27 May 1891 (GH); Surf City, Long 3821 (GH). NEW YORK: NASSAU CO.: Rockaway, I72 Schrenk s.n., 30 May 1879 (MICH). SUFFOLK CO.: Stony Brook, Miller s.n., 25 May 1877 (GH); Wading River, Mill§£_s.n., 22 May 1874 (F). NORTH CAROLINA: BUNCOMBE CO.: Asheville, Hayne 2766 (F). CARTERET CO.: Beaufort, Morton 2198 (US). BRUNSWICK CO.: Smith Island, Morton 2118 (US). NEW HANOVER CO.: near Wilmington, Canby s.n., May 1867 (MICH, NY). NORTHAMPTON CO.:. Garysburg, Ahles 38345 (DAO). ORANGE CO.: Chapel Hill, Carlton 3§_(MIN). PITT CO.: Gardnerville, Radford 32534 (NY). ROBESON CO.: Lumberton, Knowlton s.n., 9 April 1924 (GH). ROWAN CO.: vicinity of Heilio's Mill, Small §_He11er s.n., 4—9 June 1891 (WTU). WAKE CO.: Raleigh, Godfrey, s.n., 8 April 1937 (GH). OHIO: LAWRENCE CO.: Ironton, Werner s.n., 27 May 1892. OKLAHOMA: CARTER CO.: 6 mi. north- west of Ardmore, Nelson, Nelson, Goodman §_Waterfa11 5696 (RM). COMANCHE CO.: Vicinity of Fort Sill, Clemens 11576 (MO). CRAIG CO.: Vinita, Indian Territory, Carleton 12_(NY, US). CLEVELAND CO.: 2 mi. east of Norman, Bruner s.n., 15 April 1924 (M0, RM). MARSHALL CO.: near Lake Texoma, Goodman 5806 (GH, US). MURRAY CO.: Crusher Spur, Stevens 52 (DS, MIN, MO, NY, US). PONTOTOC CO.: 4—5 mi. east of Ada, Robbins 2339 (UC). PENNSYLVANIA: BUCKS CO.: Doylestown, Pond s.n., 27 May 1885 (US). LANCASTER CO.: Safe Harbor, Porter s.n., 15 May 1861 (NY). PHILADELPHIA CO.: ballast ground, Philadelphia, Parker s.n., 30 May 1865 (NY). SOUTH CAROLINA: ALLENDALE CO.: Fairfax, AhleS 10616 (COLO). BEAUFORT CO.: Beaufort, Churchill 379 (MO). CHARLESTON CO.: Charleston, Smith 5:2.,.April 1865 (GH). DORCHESTER CO.: Summer- Ville, Hunnewell 8139 (MIN). EDGEFIELD CO.: north of Edgefield, §teyermark 63373 (F). HORRY CO.: 2 mi. south of Myrtle Beach, Clausen 173 & Trapigg_3748 (NY); Longwood.Landing, Weatherby §_Griscom 16524 (NY, US). KERSHAW CO.: Lynches River, 7 mi. south of Jefferson, Redford 9068 (WTU). TENNESSEE: DECATUR CO.: specific locality unknown, Ames s.n., July 1865 (MICH). FRANKLIN CO.: north of Estill Springs, Svenson 9993 (GH). GRUNDY CO.: Goose Pond, near Pelham, Svenson 7618 (GH). GRUNDY CO.: Goose Pond, near Pelham, Svenson 7618 (GH). KNOX CO.: Knoxville, Ruth 4310 (M0). OBION CO.: near Samburg, Eyles 7801 (GH). ROAN CO.: Harriman, McMorine.s.n.,.22 April 1893 (DAO). SHELBY CO.: near Memphis, Palmer 17447 (MO). TEXAS: BASTROP CO.: Alum Creek, south of Bastrop State Pines Park, Tharp, Warnock g Barkley 16T011 (F, UC). BELL CO.: near Belton, Wolff 374 (US). BURNET CO.: 3 mi. south of Bertram, Johnson 6158 (DS). COLORADO CO.: Columbia, Bush 96 (MIN). DALLAS CO.: Dallas, Reverchon s.n., April 1876 (M0, NY). DENTON CO.: 4 1/2 mi. north of Grapevine, Whitehouse 17970.(MICH). FAYETTE CO.: Specific locality unknown,.Wurzlow s.n., in 1891 (F). FANNIN CO.: Bonham, Milligan s.n., April 1892 (US). GALVESTON CO.: Galveston, Lindheimer s.n., April 1863 (M0). GONZALES CO.: Ottine Swamp, Cory 18133 (GH). HARRIS CO.: Houston, Bush 22 (MO). KLEBERG CO.: Kingsville, Rees §Z_(NY). LIBERTY CO.: Liberty, Palmer 7733 (MIN)° MATAGORDA CO.: Matagorda, Palmer 4253 (M0)» MONTGOMERY C003 near Conroe, Palmer 33339 (GH). TARRANT CO.: near Handley, Ruth 452 (F, NY, US, WIS). TRAVIS CO.:, Austin, Tharp gin” 19 March 1932 (CAS, GH, MICH, NY, UC, WTU). VAN ZANDT CO.: west of Canton, Correll §_Correll 35679 (UC). WALKER CO.: specific locality unknown, Warner 64 (GH, US). WOODS CO.: Mineola, Reverchon, s.n., 22 April, no year (MO). VERMONT: I74 WINDHAM CO.: Brattleboro,.Grout.s.n., 25 July 1895 (GH). VIRGINIA: BEDFORD CO.: specific locality unknown, Curtiss s.n., 20 May 1872 (F, MO). DINWIDDE CO.: near Burgess.Station, Fernald §_Long 9917 (GH). GREENSVILLE CO.: 1 mi. south of Emporia,.Ferna1d §_Long 7016 (NY, US). HAMPTON CO.: Hampton, Chickering s.n., 15 May 1877 (NY). NANSEMOND CO.: Norfolk, Earll s.n., 14 May.1880 (US). PRINCES ANNE CO.: False Cape, Fernald, Griscom.§;Long.4636.(NY). .PITTSYLVANIA CO.: Dansville, Small §_He11er 230 (DAO, F, GH, MIN, MO, US). HENRICO CO.: Richmond, Churchill s.n., 11 May 1894 (GH).. SMYTH CO.: south fork of Holston River near Add.Wolf, Sma11.s.n., 15 June 1902 (F). YORK CO.: Yorktown, Thomas 2694 (DS). SOUTHAMPTON CO.: Franklin, collector unknown, May 1867 (GH). There is considerable.variation in pubescence in this taxon. Pubescent forms predominate over glabrous forms 3:l with no geographical segregation of the character. .Within the pubescent forms there is a complete range from just a few flowers with glandular hairs to all flowers on a single plant bearing glandular trichomes. Presence of the tuberculate seed character is likewise variable and without geographical segregation. While only smooth seeds or tuber— culate seeds may be present in a single population, frequent occurrences of mixtures of the two seed types are encountered. The frequency of the tuberculate seed type is ca. 60 percent. The nomenclature has been somewhat confusing in subsp. decumbens. In.A Flora 9f North America Torrey and Gray (l938) correctly transferred Spergula decumbens Ell. to Sagina. In the same work they included 175 Figure 30. Geographical distribution of Sagina decumbens subsp° We 176 m r. .'—r‘,. ..;’1;.;' ’ 2.. W... LAMOERY AIIMUYHAL EOUAL'AREA PROJECTION \j \ ~ ‘ O ‘ ‘ . -~ 1 7 7 I: ' ' ’ ” ' 1’5 3...}? ) Figure 30 I77 Sggiga subulata based on a collection cited as "Rocky Mountains, Drummond" (p. l78) and simultaneously made the transfer of Spergula subulata Swartz to Sagina, The description provided for this taxon more nearly applies to Sagina.decumbens.subsp. decumbens and is not descriptive of any of the Saginas.native to the Rocky Mountains. In Gray's Manual ed. 2 (l856) the.name Sagina decumbens is replaced by the name Sagifla Elliottii Fenzl with Spergula decumbens Ell. indicated as the synonym. The binomial Sggjgg_Elliottii was never validly published, either by Fenzl or by Gray. In the 5th edition.of.Gray's Mangal (I867) §§gjng_decumbens is treated as S, subulata, Nimmer.being recognized as the author of the transfer, and §agjn§_Elliottii is cited in synonymy. Gray also de- scribed a new variety of the taxon, Sagina subulata var. Smithii in that edition. In Gray's Mangal ed. 6, revised by Watson and Coulter (l899), the binomial Sagina_decumbens is correctly used for the taxon. Although var. Smithii is included, a nomenclatural transfer to §Efllfli decumbens was never actually made. This slender, nearly apetalous variety described by Gray does not warrant recognition as a distinct taxon. Gray's variety represents an extreme in the range of variability of subsp. decumbens exhibiting a tendency toward a habit which is more slender, with much branched fili- form stems and a greater frequency of 4-merous flowers which produce fewer-seeded capsules. The range of variability is continuous and it seems best to consider the material a single taxon. 178 No single specimen was cited with.the original description to typify var. Smithii. Of the.four collections studied by Gray, all glued to a single sheet, only one specimen bears the notation “no petals” in Gray's handwriting. I therefore designate this specimen, C, E. Smith §;Q., June l865, sandy road in the pine woods, at the mouth of Great Egg Harbor, New Jersey (GH), as the lectotype of Sagina_subulata var. Smithii. 7b. Sagina decumbens.subsp..occidentalis (Wats.).Crow, comb..nov. Sagina occidentalis.Wats., Proc. Am. Acad. lO: 344. l875. Alsinella occidentalis.(wats.) Greene, Fl. Franc. p. l25. l89l. Type: Bolander 3891, in the streets of.Ukiah, Mendocino Co., California, 1864 (GH, holotypeI; UC, MO, isotypesl). Annual with slender taproot. Branches slender, ascending or sometimes decumbent. Basal rosette of leaves lacking. Lower cauline leaves linear, 5.0—23 mm long. Upper cauline leaves becoming subulate toward tip, l.0-4.5 mm long at apex. Cauline leaves apiculate. Pedi- cels filiform, weakly glandular pubescent or glabrous. Sepals ovate to orbicular, tips frequently purple, occasionally the entire hyaline margin purple tinged. Sepals (l.5-) l.75—2.0 (-2.5) mm long. Petals elliptical, nearly equaling the sepals, (l.25—) l.5-2.0 mm long. Stamens 5 or l0, filaments (l.O—) l.5 mm long, anthers 0.25 mm long. Capsules globose prior to dehiscence, valves thin, dehiscing to ca. half the capsule length. Valves (2.0-) 2.5-3.0 (—3.5) mm long. Sepals remaining appressed following dehiscence. Seeds light brown, obliquely I79 triangular, with dorsal groove, surface smooth to slightly pebbled, rarely with elongate ridges on the lateral surfaces, 0.4 mm long. Figure 3l. EcoZogy and distribution: .On dryish hillsides, margins of vernal pools, along streams, open spots in redwood and pine woods, along roadsides and around dwellings. Ranging northward from southern California along the Great Valley and.Coastal Range to the southern border of British Columbia. Flowering April to June. Figure 32. Representative.specimens: CANADA: .BRITISH COLUMBIA: Gordon Head, Vancouver Island, Macoun.§;n., 30 May 1887 (US). Vicinity of Victoria, Vancouver Island, Macoun §;n., 23 May 1893 (F, MICH, MIN). UNITED STATES: CALIFORNIA: .ALAMEDA CO.: near summit on south side of Redwood Ridge, Constance 592 (UC); vicinity of Oakland, Holder 2523 (US); Berkeley Hills, Mulliken 29 (UC). AMADOR CO.: New York Falls, elev. 2000 ft.,.Hansen.§§Z_(DS, GH). CALAVERAS CO.: Mokelumne Hill, Blaisdell £42., no date (CAS); Big Meadow, elev. 6600 ft., Jepson Igggg (JEPS). CONTRA COSTA CO.: Rock City Camp, Mont Diablo, Bowerman 2938 (UC). DEL NORTE CO.: Gasquet to Patricks, Bacigalupi 8542 (DS). HUMBOLDT CO.: at Eureka, gzagy_21§1 (DS, UC); Garberville, I£322.l§21£ (UC). LOS ANGELES CO.: Pasadena, §£§23_§;n., 28 March 1898 (DS); Los Angeles, Ha§§3_§;n., May 1889 (DS); grassy hillsides, specific locality unknown, H§§§§_§;fl,, April 1890 (M0); Avalon, Santa Catalina Island, IEEEE §;2., February 1898 (US). MADERA CO.: along Fresno—Yosemite road (Calif. 41), 3 mi. north.of crossing with Madera Lateral, elev. 750 ft., Bacigalupi 4876 A (JEPS). MARIN CO.: McClure Beach, Pt. Reyes National h 180 Figure 3l. Photographs of Sagina decumbens subsp. occidentalis. (a) Habit. (b) Close-up. Santa Cruz Is., California (Howell 6356, CAS). Nil Nil HH I Ii; 1 IV IHiuiliillllullillilinm I81 Figure 31 182 Seashore, CE w 11892 (MSC); Inverness Ridge, Howell 12299 (CAS, UC). MARIPOSA CO.: Mariposa, Congdon §1n., 11 April 1897 (MIN). MENDOCINO CO.: Ukiah, Bolander 1821 (GH, MO, UC, US); redwoods east of Mendocino, Eastwood §_Howe11 g§§§_(CAS, NY, WTU). Albion Ridge, McMurphy 11'(DS). MERCED CO.: 5 1/2 mi. southeast of Planada, north of LeGrand, elev. 245 ft., Bacigalupi 2319 (JEPS, RM, WTU). MONTEREY CO.: Monterey, Congdon s12}, 12 May 1901 (MIN); Pacific Grove, Heller 8501 (DS, F, GH, MIN, MO, NY, WTU); Carmel, Hardam 19§4§_(CAS); about 5 mi. north of Jolon, Howell 11146 (CAS). NAPA CO.: Napa River basin, Oakville, Jepson §1n1, 26 April 1893 (JEPS); Howell Mountain, 3—4 mi. east of Angwin's, 1539y_1§91 (UC).. RIVERSIDE CO.: Lake Suprise, San Jacinto Mts., Reg§_2441 (UC). SACRAMENTO CO.: Elk Grove, Congdon Ein., 31 March 1894 (MIN). SAN DIEGO CO.: Spencer Valley, near Julian, Abrams 3131 (DS, NY); La Jolla, Jepson 11889 (JEPS). SAN FRANCISCO CO.: Point Richmond, Ha11_1§§1 (GH, MIN); ocean bluffs near Point Lobos, San Fran— cisco, Razen_§121, 9 May 1954 (CAS); serpentine slope above Baker's Beach, San Francisco, Razen_§2§1_(WTU). SAN LUIS OBISPO CO.: Coast west of Cambria, Hoover 19123_(CAS); Price Canyon, HOOVer 6221 (CAS, UC). SAN MATEO CO.: road from La Honda to Pescadero Creek, Mason 2685 (Uc). SANTA BARBARA CO.: Santa Rosa Island,.Brandegee E22}? June 1888 (UC); Mission La Purisima, Jepson 11211 (JEPS); Kinevan Canyon, Marcos Pass, Santa Barbara, Pollard §1n., 2 June 1958 (DAO); Lady's Cove, East Canyon, Santa Cruz Island, elev. 400 ft., W911 §1n:, 27 March 1932 (DS). SANTA CLARA CO.: Stanford University, Dudley sinf, 25 April 1905 (DS); ISabel Creek, east base of Mt. Hamilton, elev. 2100 ft., Sharsmith & 183 Sharsmith 11§§.(UC). SANTA CRUZ CO.: .near Jamison Creek, H§§§g_111§ (DS); near Jamison Creek Road and Big Basin, Boulder Creek Highway, elev. ca. 800 ft., Thomas 1g1§.(DS). SHASTA CO.: Olinda, Blankenship E (JEPS, WS). SOLANO.CO.: Montezuma Hills, Jepson E121, 14 May 1892; Violet Station, near Vacaville, Jepson 11923 (JEPS). SONOMA CO.: Analy Twp., Congdon E13,, 16 May 1880 (MIN); Santa Rosa, Eastwood 10112'(CAS); southern half of Lower Marsh, Rubtzoff 418 (CAS). TRINITY CO.: Junc- tion City, EEEEX.Z§§9.(UC)° TUOLUMNE CO.: Mather, elev. 1400 m, Clausen 1§42 (DS). VENTURA CO.: Kennedy Canyon, Ventura River basin, Pollard §;E:' 5 May 1946 (CAS). OREGON: CLACKAMAS CO.: Oregon City, Thompson 681 (WTU). CLATSOP CO.: beach.near Seaside, Morrill 82 (WTU). COLUMBIA CO.: St. Helens, Suksdorf fill” 28 May 1895 (ws) . CURRY CO.: Port Orford, EESE.§£§£ (GH, MO, NY). JACKSON CO.: Wimir, Hammond §1n., 21 May 1892 (M0); Evans Creek, Hammond £13., 3 June 1893 (M0). JOSEPHINE CO.: Grants Pass,.P1pegyé191.(WS). LANE CO.: 4 mi. above Takilma on the East Fork of the Illinois River, Henderson 5891 (CAS, Ds, MO, RM); near Oakridge, EEEE.E£§2§ (CAS). LINCOLN CO.: Waldo, Howell E121, June 1887 (MIN). LINN CO.: Santiam slough near Lebanon, Gilkey §.P£EEE §1§., June 1934 (OSC). MARION CO.: Jefferson, Nelson 111_(DS); Salem, Nelson §§_(DS). MULTNOMAH CO.: Lower Albina, Portland, Sheldon 8.10328 (F, GH, MIN, MO, NY, Us, WS); southwest of Sylvan, Suksdorf Q1 (ws) . POLK CO.: Nesbit, Nelson 2911 (GH). TILLAMOOK CO.: Sand Lake south of Tillamook, Thompson 111_(WTU). WASCO CO.: Eight Mile Creek, Mt. Hood National Forest, ggn§§_4911_(CAS). WASHINGTON CO.: Forest Grove, 112y§ S.n., 20 April 1893 (GH, NY); Millsboro, Lloyd s.n., 12 May 1894 (NY). 184 YAMHILL CO.: specific locality unknown, Summers s.n., June 1880 (MONTU). WASHINGTON: KING CO.: Seattle, Piper s.n., 29 May 1889 (WTU); Alki Point near Seattle, Shumway s.n., May 1894 (MICH). KLICKITAT CO.: Bingen, Suksdorf 5014 (WS); western portion, Specific locality unknown, Suksdorf s.n., May 1883 (CAN, F, MO, NY, US, WS). PIERCE CO.: prairies, Tacoma, Flett s.n., 2 October 1897 (WTU). SAN JUAN CO.: Cattle Point, San Juan Islands, Peck 12944 (WS); Friday Harbor, Pope s.n., 25 June 1904 (WTU). As in subsp. decumbens there is considerable variation in the glandular pubescence character. Although in the original description Watson indicates the taxon as being glabrous, both glabrous plants and plants with sparsely pubescent flowers are present on the type sheet. However, few of the flowers of the pubescent form in this collection are glandular. The glandular form, however, is predominant in the taxon and relatively few herbarium sheets consist entirely of glabrous specimens. In the northern portion of its range more robust plants aLPpear very similar to more slender growth forms of S. m subsp. crassicaulis. This observation led Piper (l906) to state that species lines in Sgging_are not well defined and he doubted that the two taxa were really distinct. The saginoid seed type readily distinguishes this taxon from S, maxima subsp. crassicaulis. Althoughrunzpreviously recognized nomenclaturally, subsp. Qgpidentalis has long been considered to be the western equivalent of S, decumbens. Watson (l875, p. 344) makes note to this effect in the 185 Figure 32. Geographical distribution of Sagina decumbens subsp. gggidentalis. . ___. 186 a‘. l--—-;—~.,p——v‘-‘--y—_— mm orgasm \._J t _ j. . .- \ — f‘l ..»/ \w P . . . . . . . . (\l _ _ x... l I. I” lslll‘fll IJ .. _ \ _ 7. . _ ,, l, . . y y. _ (oh. H k.\ I \. _ Q . ... .. l. _ .\ .1. ,_... __ arm illnitllllnr, llllll vaIVI N. ... _ I H a .. .I.\ 1.11 llJllilulllllll-tl. lOiUUfiOCt 030 00. 00‘ och 0.. h r r UJ- 11111 .Arux EU 1],Tilliiiillilliiiif