The clonal mat-forming fern, Dicranopteris linearis (N. L. Burm.) Underw., dominates vast areas o... more The clonal mat-forming fern, Dicranopteris linearis (N. L. Burm.) Underw., dominates vast areas of rainforests on the windward slopes of Mauna Loa Volcano on the island of Hawai'i. Because clone size has important ecological and evolutionary consequences in such a dominant species, we used isozyme analysis to investigate clone size and other aspects of genetic diversity and reproduction over a broad range of environmental conditions on primary successional sites (pahoehoe lava substrates).Isozyme analysis provided a measure of the upper limit of clonal size in this interdigitating clonal species. Each 0.5-ha primary successional site on Mauna Loa was comprised of a minimum of two to four clones. Genetic diversity in Dicranopteris was low; of 32 putative loci investigated, only 4 were polymorphic, with 2 or 3 alleles/locus. Over the 17 study locations on Mauna Loa and Kilauea Volcanoes, we identified nine multilocus genotypes based on unique combinations of allozymes. Seven of the nine genotypes were heterozygous for at least one locus, evidence of an intergametophytic mating system. Highly dispersible spores, coupled with intergametophytic mating should promote higher genetic diversity. We propose that the following factors contributed to low genetic diversity: founder effects; extreme isolation from mainland gene pools; high potential for mating among different gametophytes produced from the same sporophyte; relatively low numbers of safe sites for gametophyte establishment over space and time; and long-term reliance on vegetative growth. Leaf phenotypes were associated with genotype, but also with environmental conditions. Enough variability within a genotype existed to support the current treatment of Hawaiian Dicranopteris as one species.Vegetative growth was the primary means by which Dicranopteris covered the landscape. Nevertheless, spore production, gametophyte establishment, and sexual reproduction were absolutely essential for colonization of the few favorable microsites available on pahoehoe lava substrates of Mauna Loa following lava eruptions, dieback, and similar landscape-level disturbances.
The endemic Hawaiian genus Brighamia (Campanulaceae) comprises two federally endangered, morpholo... more The endemic Hawaiian genus Brighamia (Campanulaceae) comprises two federally endangered, morphologically similar species, B. insignis from Kaua'i and Ni'ihau and B. rockii from Moloka'i. To assist the design of conservation management programs for these taxa, isozyme analyses were performed to assess the levels of genetic diversity at the population and species levels, including comparisons within and among seven natural populations and one ex situ collection each of B. insignis and B. rockii. Our sampling (N ϭ 80) represents ϳ41% of all known individuals in the wild. Isozyme analyses revealed levels of genetic variation comparable to those reported for other Hawaiian flowering plant taxa but low levels of genetic variation at the population and species levels when compared to flowering plants in general. Ex situ individuals (N ϭ 61) were genetically representative of natural populations and hence may appropriately serve as stock for population augmentations. The two morphologically similar Brighamia species were highly distinct genetically. The combination of morphological and ecological similarity with allozymic dissimilarity observed in Brighamia is unique among the Hawaiian taxa studied to date.
Abstract.— The vascular-plant flora of the Hawaiian Islands is characterized by one of the highes... more Abstract.— The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single-island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single-island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single-island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single-island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago-level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed-mating system.
Proceedings of The Royal Society B: Biological Sciences, 2005
The enigmatic fern genus Diellia, endemic to the Hawaiian archipelago, consists of five extant an... more The enigmatic fern genus Diellia, endemic to the Hawaiian archipelago, consists of five extant and one recently extinct species. Diellia is morphologically highly variable, and a unique combination of characters has led to several contrasting hypotheses regarding the relationship of Diellia to other ferns. A phylogenetic analysis of four chloroplast loci places Diellia within 'black-stemmed' rock spleenworts of the species-rich genus Asplenium, as previously suggested by W. H. Wagner. Using an external calibration point, we estimate the divergence of the Diellia lineage from its nearest relatives to have occurred at ca. 24.3 Myr ago matching an independent estimate for the renewal of Hawaiian terrestrial life (ca. 23 Myr ago). We therefore suggest that the ancestor of the Diellia lineage may have been among the first successful colonists of the newly emerging islands in the archipelago. Disparity between morphological and nucleotide sequence variation within Diellia is consistent with a recent rapid radiation. Our estimated time of the Diellia radiation (ca. 2 Myr ago) is younger than the oldest island of Kaua'i (ca. 5.1 Myr ago) but older than the younger major islands of Maui (ca. 1.3 Myr ago), Lana'i (ca. 1.3 Myr ago) and Hawaii (ca. 0.43 Myr ago).
The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of ... more The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single-island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single-island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single-island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single-island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago-level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed-mating system.
Few studies of cpDNA have provided evolutionary and/or phylogenetic information at the intraspeci... more Few studies of cpDNA have provided evolutionary and/or phylogenetic information at the intraspecific level. We analyzed restriction site variation using 19 endonucleases in 37 populations representing both diploid (2n = 14) and autotetraploid ( 2 n = 28) Tolmiea menziesii. Seven restriction site mutations and five length mutations were observed. Although diploid and tetraploid Tolmiea have been intensively studied using nuclear markers, cpDNA variation provided additional evolutionary insights not revealed previously. The chloroplast genomes of diploid and tetraploid Tolmiea are as distinct as those of many pairs of congeneric species of angiosperms. Based on outgroup comparisons, the primitive chloroplast genome is present in tetraploid rather than diploid Tolmiea. These findings suggest that either: (1) diploid and tetraploid Tolmiea may have diverged since the origin of the autotetraploid, (2) the original diploid donor of the cytoplasm present in the tetraploid subsequently became extinct, or (3) the diploid was actually derived from the tetraploid via polyhaploidy. cpDNA variation also revealed that despite their close geographic proximity, diploid and tetraploid Tolmiea do not experience cytoplasmic gene flow. Last, three cytoplasmically distinct groups of diploid populations exist, two of which occupy distinct geographic areas. These findings demonstrate that, at least in some plant species, restriction fragment analysis of cpDNA can provide important evolutionary and phylogenetic information at low taxonomic levels.
Maximum parsimony and maximum likelihood analyses of the combined data sets of two chloroplast g... more Maximum parsimony and maximum likelihood analyses of the combined data sets of two chloroplast genes, rbcL and rps4, demonstrate that nk;the monotypic genus Gymnogrammitis is part of the polygrammoid clade (Polypodiaceae + Grammitidaceae), and not the Davalliaceae as proposed in most studies. The genus forms a clade together with two Asiatic genera of the Polypodiaceae, Arthromeris and Selliguea. These last two genera have either simple or once-pinnate leaves, whereas Gymnogrammitis has highly divided (3- to 4-pinnate) blades. Two characters of this genus, the basic chromosome number of x=36 and the absence of indusia, support a relationship with the Polypodiaceae. Neither feature is found within Davalliaceae. Three morphological characters support the placement of Gymnogrammitis within the selligueoid lineage of Polypodiaceae: spores with a thick perine extending in microspines, sclerenchymatous strands in the rhizome, and non-clathrate rhizome scales. These results demonstrate that molecular and morphological data are phylogenetically congruent with the exception of blade dissection. Our study clearly shows the pitfalls of classifications based on single characters, and illustrates the importance of phylogenetic assessment of all taxonomic conclusions.
Recent studies of the phylogeny of several groups of native Hawaiian vascular plants have led to ... more Recent studies of the phylogeny of several groups of native Hawaiian vascular plants have led to significant insights into the origin and evolution of important elements of the Hawaiian flora. No groups of Hawaiian pteridophytes have been subjected previously to rigorous phylogenetic analysis. We conducted a molecular phylogenetic analysis of the endemic Hawaiian fern genus Adenophorus employing DNA sequence variation from three cpDNA fragments: rbcL, atpβ, and the trnL-trnF intergenic spacer (IGS). In the phylogenetic analyses we employed maximum parsimony and Bayesian inference. Bayesian phylogenetic inference often provided stronger support for hypothetical relationships than did nonparametric bootstrap analyses. Although phylogenetic analyses of individual DNA fragments resulted in different patterns of relationships among species and varying levels of support for various clades, a combined analysis of all three sets of sequences produced one, strongly supported phylogenetic hypothesis. The primary features of that hypothesis are: (1) Adenophorus is monophyletic; (2) subgenus Oligadenus is paraphyletic; (3) the enigmatic endemic Hawaiian species Grammitis tenella is strongly supported as the sister taxon to Adenophorus; (4) highly divided leaf blades are evolutionarily derived in the group and simple leaves are ancestral; and, (5) the biogeographical origin of the common ancestor of the Adenophorus–G. tenella clade remains unresolved, although a neotropical origin seems most likely.
The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic ... more The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic taxa and one non-endemic, native species. The goals of this study were to determine whether Dryopteris in Hawai’i is monophyletic and to infer the biogeographical origins of Hawaiian Dryopteris by determining the geographical distributions of their closest living relatives. We sequenced two chloroplast DNA fragments, rbcL and the trnL-F intergenic spacer (IGS), for 18 Hawaiian taxa, 45 non-Hawaiian taxa, and two outgroup species. For individual fragments, we estimated phylogenetic relationships using Bayesian inference and maximum parsimony. We performed a combined analysis of both cpDNA fragments employing Bayesian inference, maximum parsimony, and maximum likelihood. These analyses indicate that Hawaiian Dryopteris is not monophyletic, and that there were at least five separate colonizations of the Hawaiian Islands by different species of dryopteroid ferns, with most of the five groups having closest relatives in SE Asia. The results suggest that one colonizing ancestor, perhaps from SE Asia, gave rise to eight endemic taxa (the glabra group). Another colonizing ancestor, also possibly from SE Asia, gave rise to a group of five endemic taxa (the exindusiate group). Dryopteris fusco-atra and its two varieties, which are endemic to Hawai’i, most likely diversified from a SE Asian ancestor. The Hawaiian endemic Nothoperanema rubiginosum has its closest relatives in SE Asia, and while the remaining two species, D. wallichiana and D. subbipinnata, are sister species, their biogeographical origins could not be determined from these analyses due to the widespread distributions of D. wallichiana and its closest non-Hawaiian relative.
Genetic loads, estimated from sporophyte production by isolated gametophyte cultures, indicate ma... more Genetic loads, estimated from sporophyte production by isolated gametophyte cultures, indicate mating systems of intragametophytic selfing in Campyloneurum angustifolium (in part), C. phyllitidis, Phlebodium aureum and Phymatosorus scolopendria, and intergametophytic mating in C. angustifolium (in part), Microgramma heterophylla and Polypodium pellucidum. Polyploidy characterizes the intragametophytic-selfing species, whereas the intergametophyticmating taxa are diploid. The duplicated loci of polyploid taxa may mitigate the expression of recessive lethal alleles caused by intragametophytic selfing, whereas genetic load probably maintains the mating systems of the intergametophytically mating taxa. Enzyme electrophoretic patterns of fixed heterozygosity support allopolyploid origins of C. phyllitidis and P. aureum and confirm their intragametophytic mating systems. Antheridiogens, present in both groups, may promote intergametophytic mating in diploids through promotion of the early development of male plants in gametophyte populations and bisexuality in isolated gametophytes of polyploids if these gametophytes delay or do not attain insensitivity to their own antheridiogen. In the polyploids, antheridiogens may also alleviate low genetic variability through promotion of occasional outcrossing. The perennial, clone-forming habit of epiphytic Polypodiaceae increases the duration and the physical space occupied by derivatives of a single spore, thus expanding the chance of interaction with a later migrant. Genetic load, duplicated genes, and antheridiogens, together with a perennial and clone-forming gametophyte growth habit, interact to produce successful breeding strategies of these epiphytic species.
New Records of Lycophytes and Ferns from Moorea, French Polynesia.-We report here 11 new taxon re... more New Records of Lycophytes and Ferns from Moorea, French Polynesia.-We report here 11 new taxon records for the island of Moorea, French Polynesia, records that were not listed in the floristic study by Murdock and Smith (Pacific Sci. 57: 253-265. 2003.). These additions bring the total number of pteridophyte species known from the island to 83. These new records, seven of them collected during an ascent of one of the highest peaks on Moorea (Mt. Mouaputa), were found at the following three localities.
, occurring only in Boulder County, CO, Eldon Mountain, AZ, Zion National Park, UT, and the state... more , occurring only in Boulder County, CO, Eldon Mountain, AZ, Zion National Park, UT, and the states of Chihuahua and Tamaulipas, Mexico. The species also occurs in the Hawaiian Archipelago, Eurasia, and Africa where it is more common. The present study was undertaken to determine 1) the mode of origin of the North American populations (i.e., remnants of a once more widely dispersed population or established via long-distance dispersal) and, 2) the phylogenetic distinctness of the Boulder population which has occasionally been treated as a separate species, (A. andrewsii). Our results are more consistent with the hypothesis that North American populations arose from at least two independent long-distance dispersal events with subsequent intracontinental dispersal establishing new populations.
Pleistocene glaciations have had a profound influence on the genetic structure of plant species t... more Pleistocene glaciations have had a profound influence on the genetic structure of plant species throughout the Northern Hemisphere because of range contractions, fragmentations, and expansions. Phylogeographic studies have contributed to our knowledge of this influence in several geographic regions of North America, however, very few phylogeographic studies have examined plant species in the Sonoran, Mojave, and Peninsular deserts. In this study, we used sequence data from the chloroplast DNA psbA–trnH intergenic spacer to obtain information on phylogeographic patterns among 310 individuals from 21 populations of Encelia farinosa (“brittlebush”; Asteraceae) across its range. We applied several population and spatial genetic analyses that allowed us to interpret our data with respect to Pleistocene climate change. These analyses indicate that E. farinosa displays patterns of genetic differentiation and geographic structuring consistent with postglacial range expansion. Populations of E. farinosa are characterized by distinct haplotype lineages significantly associated with geography. Centers of genetic diversity for the species occur in southwestern Arizona, the plains of Sonora, and Baja California Sur, all of which are putative sites of glacial refugia as predicted by analyses of macrofossil and pollen data. Nested clade analysis suggests that genetic structure in E. farinosa has been affected by past fragmentation followed by range expansion. Range expansion in several locations is further supported by significant departures from neutrality for values of Fu’s FS and Tajima’s D, and mismatch analyses.
Encelia, Enceliopsis, and Geraea are three closely related genera of shrubs and herbs distributed... more Encelia, Enceliopsis, and Geraea are three closely related genera of shrubs and herbs distributed in the arid lands of western North and South America. Resolution of relationships within Encelia has traditionally been difficult because there is some morphological overlap among species, and species hybridize when sympatric. In this study, we used DNA sequence data from two nuclear regions (ITS and ETS) and two chloroplast regions (psbA-trnH and trnS-trnfM) to infer phylogenetic relationships among 19 species, subspecies, and varieties of Encelia. Eight species of Enceliopsis, Geraea, Simsia, and Bahiopsis were also included as outgroups. These data support previous hypotheses suggesting that Encelia is monophyletic and sister to Enceliopsis + Geraea. In addition, our data provide evidence for two major subclades within Encelia that are supported by morphological synapomorphies. However, little differentiation was observed among species of Encelia within subclades. This lack of differentiation may be the result of a recent diversification of the genus including recent radiation in the Peninsular Desert.
The clonal mat-forming fern, Dicranopteris linearis (N. L. Burm.) Underw., dominates vast areas o... more The clonal mat-forming fern, Dicranopteris linearis (N. L. Burm.) Underw., dominates vast areas of rainforests on the windward slopes of Mauna Loa Volcano on the island of Hawai'i. Because clone size has important ecological and evolutionary consequences in such a dominant species, we used isozyme analysis to investigate clone size and other aspects of genetic diversity and reproduction over a broad range of environmental conditions on primary successional sites (pahoehoe lava substrates).Isozyme analysis provided a measure of the upper limit of clonal size in this interdigitating clonal species. Each 0.5-ha primary successional site on Mauna Loa was comprised of a minimum of two to four clones. Genetic diversity in Dicranopteris was low; of 32 putative loci investigated, only 4 were polymorphic, with 2 or 3 alleles/locus. Over the 17 study locations on Mauna Loa and Kilauea Volcanoes, we identified nine multilocus genotypes based on unique combinations of allozymes. Seven of the nine genotypes were heterozygous for at least one locus, evidence of an intergametophytic mating system. Highly dispersible spores, coupled with intergametophytic mating should promote higher genetic diversity. We propose that the following factors contributed to low genetic diversity: founder effects; extreme isolation from mainland gene pools; high potential for mating among different gametophytes produced from the same sporophyte; relatively low numbers of safe sites for gametophyte establishment over space and time; and long-term reliance on vegetative growth. Leaf phenotypes were associated with genotype, but also with environmental conditions. Enough variability within a genotype existed to support the current treatment of Hawaiian Dicranopteris as one species.Vegetative growth was the primary means by which Dicranopteris covered the landscape. Nevertheless, spore production, gametophyte establishment, and sexual reproduction were absolutely essential for colonization of the few favorable microsites available on pahoehoe lava substrates of Mauna Loa following lava eruptions, dieback, and similar landscape-level disturbances.
The endemic Hawaiian genus Brighamia (Campanulaceae) comprises two federally endangered, morpholo... more The endemic Hawaiian genus Brighamia (Campanulaceae) comprises two federally endangered, morphologically similar species, B. insignis from Kaua'i and Ni'ihau and B. rockii from Moloka'i. To assist the design of conservation management programs for these taxa, isozyme analyses were performed to assess the levels of genetic diversity at the population and species levels, including comparisons within and among seven natural populations and one ex situ collection each of B. insignis and B. rockii. Our sampling (N ϭ 80) represents ϳ41% of all known individuals in the wild. Isozyme analyses revealed levels of genetic variation comparable to those reported for other Hawaiian flowering plant taxa but low levels of genetic variation at the population and species levels when compared to flowering plants in general. Ex situ individuals (N ϭ 61) were genetically representative of natural populations and hence may appropriately serve as stock for population augmentations. The two morphologically similar Brighamia species were highly distinct genetically. The combination of morphological and ecological similarity with allozymic dissimilarity observed in Brighamia is unique among the Hawaiian taxa studied to date.
Abstract.— The vascular-plant flora of the Hawaiian Islands is characterized by one of the highes... more Abstract.— The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single-island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single-island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single-island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single-island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago-level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed-mating system.
Proceedings of The Royal Society B: Biological Sciences, 2005
The enigmatic fern genus Diellia, endemic to the Hawaiian archipelago, consists of five extant an... more The enigmatic fern genus Diellia, endemic to the Hawaiian archipelago, consists of five extant and one recently extinct species. Diellia is morphologically highly variable, and a unique combination of characters has led to several contrasting hypotheses regarding the relationship of Diellia to other ferns. A phylogenetic analysis of four chloroplast loci places Diellia within 'black-stemmed' rock spleenworts of the species-rich genus Asplenium, as previously suggested by W. H. Wagner. Using an external calibration point, we estimate the divergence of the Diellia lineage from its nearest relatives to have occurred at ca. 24.3 Myr ago matching an independent estimate for the renewal of Hawaiian terrestrial life (ca. 23 Myr ago). We therefore suggest that the ancestor of the Diellia lineage may have been among the first successful colonists of the newly emerging islands in the archipelago. Disparity between morphological and nucleotide sequence variation within Diellia is consistent with a recent rapid radiation. Our estimated time of the Diellia radiation (ca. 2 Myr ago) is younger than the oldest island of Kaua'i (ca. 5.1 Myr ago) but older than the younger major islands of Maui (ca. 1.3 Myr ago), Lana'i (ca. 1.3 Myr ago) and Hawaii (ca. 0.43 Myr ago).
The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of ... more The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single-island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single-island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single-island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single-island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago-level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed-mating system.
Few studies of cpDNA have provided evolutionary and/or phylogenetic information at the intraspeci... more Few studies of cpDNA have provided evolutionary and/or phylogenetic information at the intraspecific level. We analyzed restriction site variation using 19 endonucleases in 37 populations representing both diploid (2n = 14) and autotetraploid ( 2 n = 28) Tolmiea menziesii. Seven restriction site mutations and five length mutations were observed. Although diploid and tetraploid Tolmiea have been intensively studied using nuclear markers, cpDNA variation provided additional evolutionary insights not revealed previously. The chloroplast genomes of diploid and tetraploid Tolmiea are as distinct as those of many pairs of congeneric species of angiosperms. Based on outgroup comparisons, the primitive chloroplast genome is present in tetraploid rather than diploid Tolmiea. These findings suggest that either: (1) diploid and tetraploid Tolmiea may have diverged since the origin of the autotetraploid, (2) the original diploid donor of the cytoplasm present in the tetraploid subsequently became extinct, or (3) the diploid was actually derived from the tetraploid via polyhaploidy. cpDNA variation also revealed that despite their close geographic proximity, diploid and tetraploid Tolmiea do not experience cytoplasmic gene flow. Last, three cytoplasmically distinct groups of diploid populations exist, two of which occupy distinct geographic areas. These findings demonstrate that, at least in some plant species, restriction fragment analysis of cpDNA can provide important evolutionary and phylogenetic information at low taxonomic levels.
Maximum parsimony and maximum likelihood analyses of the combined data sets of two chloroplast g... more Maximum parsimony and maximum likelihood analyses of the combined data sets of two chloroplast genes, rbcL and rps4, demonstrate that nk;the monotypic genus Gymnogrammitis is part of the polygrammoid clade (Polypodiaceae + Grammitidaceae), and not the Davalliaceae as proposed in most studies. The genus forms a clade together with two Asiatic genera of the Polypodiaceae, Arthromeris and Selliguea. These last two genera have either simple or once-pinnate leaves, whereas Gymnogrammitis has highly divided (3- to 4-pinnate) blades. Two characters of this genus, the basic chromosome number of x=36 and the absence of indusia, support a relationship with the Polypodiaceae. Neither feature is found within Davalliaceae. Three morphological characters support the placement of Gymnogrammitis within the selligueoid lineage of Polypodiaceae: spores with a thick perine extending in microspines, sclerenchymatous strands in the rhizome, and non-clathrate rhizome scales. These results demonstrate that molecular and morphological data are phylogenetically congruent with the exception of blade dissection. Our study clearly shows the pitfalls of classifications based on single characters, and illustrates the importance of phylogenetic assessment of all taxonomic conclusions.
Recent studies of the phylogeny of several groups of native Hawaiian vascular plants have led to ... more Recent studies of the phylogeny of several groups of native Hawaiian vascular plants have led to significant insights into the origin and evolution of important elements of the Hawaiian flora. No groups of Hawaiian pteridophytes have been subjected previously to rigorous phylogenetic analysis. We conducted a molecular phylogenetic analysis of the endemic Hawaiian fern genus Adenophorus employing DNA sequence variation from three cpDNA fragments: rbcL, atpβ, and the trnL-trnF intergenic spacer (IGS). In the phylogenetic analyses we employed maximum parsimony and Bayesian inference. Bayesian phylogenetic inference often provided stronger support for hypothetical relationships than did nonparametric bootstrap analyses. Although phylogenetic analyses of individual DNA fragments resulted in different patterns of relationships among species and varying levels of support for various clades, a combined analysis of all three sets of sequences produced one, strongly supported phylogenetic hypothesis. The primary features of that hypothesis are: (1) Adenophorus is monophyletic; (2) subgenus Oligadenus is paraphyletic; (3) the enigmatic endemic Hawaiian species Grammitis tenella is strongly supported as the sister taxon to Adenophorus; (4) highly divided leaf blades are evolutionarily derived in the group and simple leaves are ancestral; and, (5) the biogeographical origin of the common ancestor of the Adenophorus–G. tenella clade remains unresolved, although a neotropical origin seems most likely.
The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic ... more The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic taxa and one non-endemic, native species. The goals of this study were to determine whether Dryopteris in Hawai’i is monophyletic and to infer the biogeographical origins of Hawaiian Dryopteris by determining the geographical distributions of their closest living relatives. We sequenced two chloroplast DNA fragments, rbcL and the trnL-F intergenic spacer (IGS), for 18 Hawaiian taxa, 45 non-Hawaiian taxa, and two outgroup species. For individual fragments, we estimated phylogenetic relationships using Bayesian inference and maximum parsimony. We performed a combined analysis of both cpDNA fragments employing Bayesian inference, maximum parsimony, and maximum likelihood. These analyses indicate that Hawaiian Dryopteris is not monophyletic, and that there were at least five separate colonizations of the Hawaiian Islands by different species of dryopteroid ferns, with most of the five groups having closest relatives in SE Asia. The results suggest that one colonizing ancestor, perhaps from SE Asia, gave rise to eight endemic taxa (the glabra group). Another colonizing ancestor, also possibly from SE Asia, gave rise to a group of five endemic taxa (the exindusiate group). Dryopteris fusco-atra and its two varieties, which are endemic to Hawai’i, most likely diversified from a SE Asian ancestor. The Hawaiian endemic Nothoperanema rubiginosum has its closest relatives in SE Asia, and while the remaining two species, D. wallichiana and D. subbipinnata, are sister species, their biogeographical origins could not be determined from these analyses due to the widespread distributions of D. wallichiana and its closest non-Hawaiian relative.
Genetic loads, estimated from sporophyte production by isolated gametophyte cultures, indicate ma... more Genetic loads, estimated from sporophyte production by isolated gametophyte cultures, indicate mating systems of intragametophytic selfing in Campyloneurum angustifolium (in part), C. phyllitidis, Phlebodium aureum and Phymatosorus scolopendria, and intergametophytic mating in C. angustifolium (in part), Microgramma heterophylla and Polypodium pellucidum. Polyploidy characterizes the intragametophytic-selfing species, whereas the intergametophyticmating taxa are diploid. The duplicated loci of polyploid taxa may mitigate the expression of recessive lethal alleles caused by intragametophytic selfing, whereas genetic load probably maintains the mating systems of the intergametophytically mating taxa. Enzyme electrophoretic patterns of fixed heterozygosity support allopolyploid origins of C. phyllitidis and P. aureum and confirm their intragametophytic mating systems. Antheridiogens, present in both groups, may promote intergametophytic mating in diploids through promotion of the early development of male plants in gametophyte populations and bisexuality in isolated gametophytes of polyploids if these gametophytes delay or do not attain insensitivity to their own antheridiogen. In the polyploids, antheridiogens may also alleviate low genetic variability through promotion of occasional outcrossing. The perennial, clone-forming habit of epiphytic Polypodiaceae increases the duration and the physical space occupied by derivatives of a single spore, thus expanding the chance of interaction with a later migrant. Genetic load, duplicated genes, and antheridiogens, together with a perennial and clone-forming gametophyte growth habit, interact to produce successful breeding strategies of these epiphytic species.
New Records of Lycophytes and Ferns from Moorea, French Polynesia.-We report here 11 new taxon re... more New Records of Lycophytes and Ferns from Moorea, French Polynesia.-We report here 11 new taxon records for the island of Moorea, French Polynesia, records that were not listed in the floristic study by Murdock and Smith (Pacific Sci. 57: 253-265. 2003.). These additions bring the total number of pteridophyte species known from the island to 83. These new records, seven of them collected during an ascent of one of the highest peaks on Moorea (Mt. Mouaputa), were found at the following three localities.
, occurring only in Boulder County, CO, Eldon Mountain, AZ, Zion National Park, UT, and the state... more , occurring only in Boulder County, CO, Eldon Mountain, AZ, Zion National Park, UT, and the states of Chihuahua and Tamaulipas, Mexico. The species also occurs in the Hawaiian Archipelago, Eurasia, and Africa where it is more common. The present study was undertaken to determine 1) the mode of origin of the North American populations (i.e., remnants of a once more widely dispersed population or established via long-distance dispersal) and, 2) the phylogenetic distinctness of the Boulder population which has occasionally been treated as a separate species, (A. andrewsii). Our results are more consistent with the hypothesis that North American populations arose from at least two independent long-distance dispersal events with subsequent intracontinental dispersal establishing new populations.
Pleistocene glaciations have had a profound influence on the genetic structure of plant species t... more Pleistocene glaciations have had a profound influence on the genetic structure of plant species throughout the Northern Hemisphere because of range contractions, fragmentations, and expansions. Phylogeographic studies have contributed to our knowledge of this influence in several geographic regions of North America, however, very few phylogeographic studies have examined plant species in the Sonoran, Mojave, and Peninsular deserts. In this study, we used sequence data from the chloroplast DNA psbA–trnH intergenic spacer to obtain information on phylogeographic patterns among 310 individuals from 21 populations of Encelia farinosa (“brittlebush”; Asteraceae) across its range. We applied several population and spatial genetic analyses that allowed us to interpret our data with respect to Pleistocene climate change. These analyses indicate that E. farinosa displays patterns of genetic differentiation and geographic structuring consistent with postglacial range expansion. Populations of E. farinosa are characterized by distinct haplotype lineages significantly associated with geography. Centers of genetic diversity for the species occur in southwestern Arizona, the plains of Sonora, and Baja California Sur, all of which are putative sites of glacial refugia as predicted by analyses of macrofossil and pollen data. Nested clade analysis suggests that genetic structure in E. farinosa has been affected by past fragmentation followed by range expansion. Range expansion in several locations is further supported by significant departures from neutrality for values of Fu’s FS and Tajima’s D, and mismatch analyses.
Encelia, Enceliopsis, and Geraea are three closely related genera of shrubs and herbs distributed... more Encelia, Enceliopsis, and Geraea are three closely related genera of shrubs and herbs distributed in the arid lands of western North and South America. Resolution of relationships within Encelia has traditionally been difficult because there is some morphological overlap among species, and species hybridize when sympatric. In this study, we used DNA sequence data from two nuclear regions (ITS and ETS) and two chloroplast regions (psbA-trnH and trnS-trnfM) to infer phylogenetic relationships among 19 species, subspecies, and varieties of Encelia. Eight species of Enceliopsis, Geraea, Simsia, and Bahiopsis were also included as outgroups. These data support previous hypotheses suggesting that Encelia is monophyletic and sister to Enceliopsis + Geraea. In addition, our data provide evidence for two major subclades within Encelia that are supported by morphological synapomorphies. However, little differentiation was observed among species of Encelia within subclades. This lack of differentiation may be the result of a recent diversification of the genus including recent radiation in the Peninsular Desert.
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Papers by Tom Ranker