Evolution and history of hummingbirds (Aves: Trochilidae) from the Juan Fernandez Islands, Chile

The Juan Fernandez Firecrown Sephanoides fernandensis is an endangered endemic hummingbird that inhabits the Juan Fernandez Islands, 667 km off the coast of Chile. Its population has decreased from several thousand in the early part of this century to approximately 250–400 individuals at present. The reasons for its decline include habitat degradation by anthropogenic forest clearance and the introduction of grazing mammals and rodents. Another hummingbird, the Green-backed Firecrown Sephanoides sephaniodes, inhabits the Juan Fernandez Islands but is also found on the Chilean mainland. It currently numbers several thousand on the Juan Fernandez Islands but was considered rare during the late 19th and early 20th centuries. The sister relationship between the two species has not been critically tested, and so their evolutionary histories on the Juan Fernandez Islands remain uncertain. With the use of mtDNA cytochrome b and ND2 phylogenetic reconstructions, our study supports the two species as sister taxa. Moreover, the molecular data suggest that the genus Sephanoides is closely related to the higher altitude Andean hummingbirds typical of the paramo and puna habitats. The molecular divergence between the two species of Sephanoides indicates they may have become isolated from each other less than 1 million years ago, suggesting that S. fernandensis evolved in situ on the Juan Fernandez Islands. We find no evidence of genetic subdivision between populations of S. sephaniodes from the Juan Fernandez Islands and the mainland. In addition, high genetic variation of the Juan Fernandez Islands population does not indicate a long period of isolation of a limited number of S. sephaniodes but instead suggests a recent colonization event, perhaps from several mainland populations. As a result of molecular, morphological and apparent ecological similarities, we suggest that competition by S. sephaniodes may be an additional factor stifling the recovery of S. fernandensis. Possible conservation strategies include habitat restoration and the removal of introduced mammals; immediate implementation of such conservation management plans are necessary to save this species from extinction.     

RIBOSOMAL DNA AND RAPD VARIATION IN THE RARE PLANT FAMILY LACTORIDACEAE

Lactoris fernandeziana, endemic to the island of Masatierra in the Juan Fernandez Archipelago, is the only living member of the primitive angiosperm family, Lactoridaceae. The species was surveyed for ribosomal DNA (rDNA) and RAPD (Random Amplified Polymorphic DNA) variation. Previous analyses of allozymes had revealed no variation within the species. Variation was found for length in the intergenic spacer and for restriction sites in the 18S–25S genes of rDNA, and for the presence of amplified bands using 16 primers. Different rDNA repeat lengths and restriction site variants were detected within individuals as well as within and among populations. The level of variation in RAPDs is low relative to other Juan Fernandez endemic species surveyed, and nearly all variants were restricted to single populations. The rDNA length variants were distributed throughout the island, whereas the rDNA restriction site variants and RAPD markers indicated minor genetic differences among the populations.     

FLAVONOID EVOLUTION IN DENDROSERIS (COMPOSITAE,LACTUCEAE) FROM THE JUAN FERNANDEZ ISLANDS,CHILE

Leaf flavonoid chemistry was examined from the three subgenera and 11 species of the endemic genus Dendroseris (Compositae, Lactuceae) of the Juan Fernandez Islands, Chile. Eight of the species are restricted to the older island (Masatierra, ca. 4 million years old), which is also closer to the mainland. Three species, one from each subgenus, are restricted to Masafuera, which is younger geologically (1–2 million years old) and 145 km further west of Masatierra. A total of 16 compounds was identified, with the 7-0-glucosides of the flavones apigenin and luteolin accounting for 12 of the constituents. Two glucosides of the flavonol quercetin were detected. Despite considerable interpopulation variation within species, six of the taxa have distinctive flavonoid profiles. Although there are few absolute differences among the subgenera, they can be distinguished chemically. Subgenus Rea contains the greatest number of compounds, and a previous cladistic analysis based on morphological features suggested this subgenus as most primitive. Subgenus Phoenicoseris is considered highly derived morphologically, and it has a reduced flavonoid chemistry. Very little reduction in flavonoid diversity was seen in the morphologically specialized subg. Dendroseris as compared to subg. Rea. A trend in reduction of numbers of compounds was seen for two of the three species on the younger island of Masafuera when compared to their presumed ancestors on Masatierra. Flavonoids of selected species of Hieracium and Hypochaeris, presumptive mainland progenitors of Dendroseris, reveal a close chemical affinity with the former genus.     

FLAVONOID EVOLUTION IN ROBINSONIA (COMPOSITAE) OF THE JUAN FERNANDEZ ISLANDS

Leaf flavonoids were isolated and identified from 54 populations representing all seven species of Robinsonia, a genus of dioecious rosette trees endemic to the Juan Fernandez Islands. Fourteen compounds were detected consisting of flavonols, flavones, flavanones and dihydroflavonols. The distribution of these compounds in Robinsonia largely corresponds to specific and sectional limits based on morphological data. The morphologically similar species, R. gayana and R. thurifera, have identical flavonol profiles (derivatives of quercetin). Likewise, the closely related R. evenia and R. masafuerae are unique in the genus by possessing flavones. The inclusion of Rhetinodendron (i.e., R. berteroi) in Robinsonia is supported by its strong flavonoid similarity with species in two other sections of the genus. The morphologically diverse section Eleutherolepis exhibits the greatest flavonoid variation of any section, and only here are found flavones, flavanones and dihydroflavonols. The direction of flavonoid evolution in Robinsonia is hypothesized to be from fewer to more classes of compounds. Biosynthetic considerations suggest that this gain in compounds is due both to a gain of an additional enzymatic step and to the sequestering of precursors. This interpretation of direction of flavonoid evolution is in agreement with several lines of evidence including the flavonoid chemistry of the hypothesized outgroup (i.e., species of Senecio on mainland Chile), the ages of the two islands, and morphological trends.     

Notes on the Poaceae of the Robinson Crusoe (Juan Fernández) Islands, Chile

Notes on the Poaceae of the Robinson Crusoe (Juan Fernández) Islands, Chile, Brittonia 54: 154–163. 2001.—Poaceae in the Robinson Crusoe (=Juan Fernández) Islands number 53 species in 32 genera, of which 9% of the species are endemic, 9% indigenous, and 81% adventitious. The endemic taxa (and their conservation status) are:Agrostis masafuerana (rare),Chusquea fernandeziana (not endangered),Megalachne berteroana (not endangered),M. masafuerana (not endangered), andPodophorus bromoides (extinct).Megalachne andPodophorus are endemic genera. Comparisons with Poaceae in the Bonin and Volcano Islands, Canary Islands, Galápagos Islands, and Hawaiian Islands show different levels of endemism: number of endemic taxa, respectively, 5, 10, 12, 40; percent specific endemism, 8, 6, 21, 19. No endemic genera occur.Anthoxanthum odoratum, Avena barbata andHordeum murinum are noxious weeds in the Robinson Crusoe Islands. Many adventives are shared among floras of the archipelagos, with the highest ties of Robinson Crusoe being to the Canaries (53% of total Poaceae known in Juan Fernández) and the Hawaiian Islands (47%). Low levels of adventives occur within the Bonin (5%) and Galápagos (7%) Islands. In contrast, there are many endemic genera of Asteraceae in these same archipelagos: Bonin and Volcano Islands (1), Canary Islands (8), Galápagos Islands (5), and Hawaiian Islands (6); percent of specific endemism is also higher (20, 53, 54 and 56, respectively). Hypotheses for greater levels of endemics among oceanic island Asteraceae include more efficient dispersal mechanisms by wind and birds, animal pollination systems that result in greater degrees of geographic populational genetic isolation, and a vascular cambium that offers enhanced growth-form evolutionary opportunities.     

Evolution ofPeperomia (Piperaceae) in the Juan Fernandez Islands,Chile

Four species ofPeperomia (Piperaceae) occur in the Juan Fernandez Islands, Chile:P. berteroana, P. margaritifera, P. skottsbergii, andP. fernandeziana. The last species is found also in continental Chile, whereas the other three are endemic to the archipelago.Peperomia margaritifera is found only on the older island of Masatierra, whereasP. skottsbergii is confined to the younger island of Masafuera, andP. berteroana occurs on both islands. Phenetic analyses of mainland taxa suggest thatP. fernandeziana belongs to subg.Sphaerocarpidium whereas the endemic taxa form their own subg.Tildenidium connecting to subg.Tildenia. Cladistic analyses indicate thatP. margaritifera is the most primitive species in the archipelago and thatP. berteroana is the most derived, especially patristically. Chromosomally, the four species are all n = 22, which may be tetraploid on a base of x = 11. Sulfated flavones occur only inP. berteroana andP. skottsbergii, which are otherwise unknown for the family. Dispersal of propagules to the islands from the continent and between islands is believed to have been accomplished by birds.     

Flavonoid chemistry of the endemic species of Myrceugenia (Myrtaceae) of the Juan Fernandez Islands and relatives in continental South America

Twenty-one flavonoids were isolated from the leaves of two endemic species ofMyrceugenia on the Juan Fernandez Islands,M. fernandeziana andM. schulzei, from two related species from Brazil,M. campestris andM. rufescens, and from five species from continental Chile,M. colchaguensis, M. exsucca, M. lanceolata, M. pinifolia, andM. rufa. A phenetic analysis was used to evaluate chemical similarities.Myrceugenia campestris andM. rufescens appear most closely related to each other based on flavonoid profiles, and they are also different from the other seven species. The two endemic species in the Juan Fernandez Islands,M. fernandeziana andM. schulzei, group with the continental Chilean species. The former is most closely related toM. lanceolata, and the latter clusters withM. exsucca, although somewhat distantly. The results suggest that the two Juan Fernandez endemics are derived from two introductions from the Chilean continent and not from immigrants from the eastern side of the Andes.     

CHROMOSOME NUMBERS OF THE EAST AFRICAN GIANT SENECIOS AND GIANT LOBELIAS AND THEIR EVOLUTIONARY SIGNIFICANCE

The gametophytic chromosome number for the giant senecios (Asteraceae, Senecioneae, Dendrosenecio) is n = 50, and for the giant lobelias (Lobeliaceae, Lobelia subgenus Tupa section Rhynchopetalum) it is n = 14. Previous sporophytic counts are generally verified, but earlier reports for the giant senecios of 2n = 20 and ca. 80, the bases for claims of intraspecific polyploidy, are unsubstantiated. The 14 new counts for the giant senecios and the ten new counts for the giant lobelias are the first gametophytic records for these plants and include the first reports for six and four taxa, respectively, for the two groups. Only five of the 11 species of giant senecio and three of the 21 species of giant lobelia from eastern Africa remain uncounted. Although both groups are polyploid, the former presumably decaploid and the latter more certainly tetraploid, their adaptive radiations involved no further change in chromosome number. The cytological uniformity within each group, while providing circumstantial evidence of monophyly and simplifying interpretations of cladistic analyses, provides neither positive nor negative support for a possible role of polyploidy in evolving the giant-rosette growth-form.     

Variation in distribution and abundance of the endemic flora of Juan Fernández Islands, Chile. Pteridophyta

The Juan Fernández Archipelago is located 667 km west of central Chile (33°S) in the Pacific Ocean. It is composed of three islands: Robinson Crusoe (ex-Masatierra), Santa Clara and Marinero Alejandro Selkirk (ex-Masafuera).The first botanical collections were made 250 years after its discovery and colonization. These studies showed the great importance of ferns in the vascular flora of the islands. Of the 53 species described up to the present, 23 are endemic (45%), with one endemic monotypic genus, Thyrsopteris, whose only species is T. elegans Kunze (Dicksoniaceae).During a period of 4 years, I have studied the flora of the islands. There has been surprisingly little change in the distribution of the ferns, considering the devastation of the angiosperm flora. Some species have even increased their abundance, especially the tree ferns. Nevertheless, the more delicate species and those with a more restricted habitat have been reduced in abundance.Ten of the endemic species (43%) are in danger of extinction, such as Asplenium macrosorum and Ophioglossum fernandezianum, four are vulnerable (17%), including Blechnum schottii and Polypodium intermedium, and nine (40%), including Pteris berteroana and Blechnum cycadifolium are not endangered.     

南海岛屿种子植物区系地理的研究

本文通过实地考察,广泛收集前人的研究资料,概述了南海岛屿地区的自然条件和植被,对南海岛屿种子植物的区系组成、特点、分布区类型、特有现象和替代现象等进行了较详细的分析,并与邻近植物区系进行了比较研究。同时,根据区内植物分布的特点和自然条件特征划分为５个植物区系小区,最后对南海岛屿地区植物区系的起源与演化进行了讨论。     

USE OF RAPD MARKERS TO DOCUMENT THE ORIGIN OF THE INTERGENERIC HYBRID × MARGYRACAENA SKOTTSBERGII (ROSACEAE) ON THE JUAN FERNANDEZ ISLANDS

Bands of random amplified polymorphic DNA (RAPDs) were used as markers to test the hypothesis that the species Margyracaena skottsbergii, which is endemic to the island of Masatierra in the Juan Fernandez archipelago, represents an intergeneric hybrid between Acaena argentea and Margyricarpus digynus. Thirteen 10-mer primers produced 18 consistent species-specific bands for A. argentea and 27 for M. digynus, with all 45 bands present in the presumed hybrid Margyracaena. A second species of Acaena on Masatierra, A. ovalifolia, has 23 unique amplified bands in all plants examined, and it shares none of these bands with Margyracaena. The data from RAPDs are concordant with morphology in implicating Acaena argentea rather than A. ovalifolia as one parent of Margyracaena. RAPDs can provide numerous genetic markers while requiring minimal quantities of DNA, thereby making them attractive for the study of hybridization, particularly in rare plants where DNA amounts may be limiting.     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Phylogenetics of Darwiniothamnus (Asteraceae: Astereae) – molecular evidence for multiple origins in the endemic flora of the Galápagos Islands

Aim The aims of this study were (1) to investigate whether the two growth forms of Darwiniothamnus Harling (Asteraceae) originated from the colonization of a single ancestor, (2) to identify the closest relative(s) of Darwiniothamnus, and (3) to review molecular phylogenies from other plant groups to infer the origin of Galápagos endemics. Location Darwiniothamnus is endemic to the Galápagos Islands. Methods All putative relatives of Darwiniothamnus plus 38 additional species were included. Nucleotide sequences of the internal transcribed spacers of the nuclear ribosomal DNA were used for Bayesian and parsimony analyses. Results Darwiniothamnus is polyphyletic. Two species (D. lancifolius (Hook. f.) Harling and D. tenuifolius (Hook. f.) Harling) are woody shrubs that usually grow to 1–2 m in height; they belong to a clade composed of species otherwise restricted to the Caribbean. These two species are sister to Erigeron bellidiastroides Griseb., a herbaceous species endemic to Cuba. The third species (D. alternifolius Lawesson & Adsersen) is a perennial herbaceous plant, woody at the base and reaching only up to 50 cm in height. It is sister to two Chilean (Coquimbo–Valparaiso region) species that also have a perennial herbaceous habit: E. fasciculatus Colla and E. luxurians (Skottsb.) Solbrig. They are placed in an assemblage restricted to South America. The review of previous molecular phylogenetic studies revealed that two of the endemic genera and endemic species of three non‐endemic genera have their closest relatives in South America. Endemic species belonging to three non‐endemic genera have sister species in North America or the West Indies. One endemic genus and endemic species in three non‐endemic genera have sister taxa with a widespread continental distribution, or their molecular phylogenies yielded equivocal results. Main conclusions The flora of Galápagos has affinities with both North America (including the Antilles) and South America. Darwiniothamnus exhibits both patterns: two species of this genus are sister to a taxon endemic to Cuba, supporting a connection between the Cocos plate and the West Indies; the third species, D. alternifolius, provides a link with the Coquimbo–Valparaiso region, suggesting a biogeographical connection between the Nazca plate and southern South America.     

Gondwanan Gymnochilini (Coleoptera: Trogossitidae): generic concepts,review of New Zealand species and long‐range Pacific dispersal

Members of Leperina Erichson (Trogossitidae: Gymnochilini) from New Zealand, New Caledonia and Lord Howe Island are morphologically similar to members of the endemic Juan Fernandez Island genus Phanodesta Reitter, sharing at least one obvious character, elytral carinae that are beaded and contain well‐defined punctures. To test the monophyly of Leperina and Phanodesta, we reconstructed phylogenetic relationships of the genera of the tribe Gymnochilini by a cladistic analysis of 22 terminals and 47 adult characters rooted with one genus of trogossitine. Leperina is rendered paraphyletic by the placements of Seidlitzella Jakobson and Phanodesta. Kolibacia n.gen. (type species Leperina tibialis Reitter) is described for east Palaearctic species included formerly in Leperina (two new combinations); New Zealand Leperina and other species from New Caledonia and Lord Howe Island (Ostoma pudicum Olliff) are transferred to Phanodesta (six new combinations); and the remaining species are retained in Leperina. The following species are described as new: Phanodesta carinata n.sp., P. manawatawhi n.sp., P. oculata n.sp. and P. tepaki n.sp. Leperina ambiguum Broun is transferred to Grynoma Sharp resulting in a new combination and three new synonymies for New Zealand trogossitines: Leperina interrupta Brookes n.syn. and Leperina sobrina (White) n.syn. [= Phanodesta farinosa (Sharp)], and Trogosita affinis White n.syn. (= Tenebroides mauritanicus Linnaeus). A key to the New Zealand species and a checklist for the species of the Kolibacia, Leperina and Phanodesta are provided. The derived placement of Juan Fernandez Phanodesta in the phylogeny is evidence for long‐distance dispersal from Australasia. A tally of all Juan Fernandez Islands Coleoptera shows derivation mostly from Chile and South America, with few from the southern Pacific region, rarely from Australasia.     

SYSTEMATIC SIGNIFICANCE OF CHROMOSOME NUMBERS IN ACMELLA (ASTERACEAE)

Chromosome counts are reported for 372 individuals from 202 populations in 26 taxa of Acmella (Asteraceae: Heliantheae). Chromosome numbers for 15 taxa are first reports. A review of previous counts and the new reports supports a basic chromosome number of 13 for the genus. The results show that polyploidy, sometimes accompanied by hybridization and asexual reproduction, is widespread in Acmella and has contributed to the taxonomic difficulties in the genus. These factors have produced a variable polyploid pillar complex in sect. Acmella. In one taxon in this complex, A. oppositifolia var. oppositifolia, intrataxon and even intrapopulational chromosomal variation has been detected. Morphological studies in conjunction with observations of meiotic pairing suggest that most polyploids are alloploid in origin. The occurrence of polyploidy in 16 of the 27 taxa known chromosomally emphasizes the important role this process has had in speciation within Acmella. Although intrataxon chromosomal variation has limited the taxonomic utility of chromosome numbers, a few examples are presented in which these data have been valuable for separating some pairs of closely related taxa (A. decumbens var. affinis from var. decumbens and A. poliolepidica from A. oppositifolia).     

Floral resources and risk of exposure to pesticides for Melipona quadrifasciata anthidioides Lepeletier 1836 in a Cerrado of São Paulo (Brazil)

Honey and bee bread samples from storage pots of Melipona quadrifasciata anthidioides were collected monthly from April 2015 to May 2016 in the Mogi Guaçu Biological Reserve (22º 10? S, 47º 11? W). The flora in the site is characteristic of the Atlantic Forest with preserved areas of savanna-like vegetation surrounded by commercial forests, orchards and various crops of exotic and native plants. Samples were analysed with the use of melissopalynological methodology and 46 pollen types from 38 genera and 30 families were identified in 25 honey samples. Fabaceae, Asteraceae, Myrtaceae, Sapindaceae showed the greatest pollen richness in honey. Predominant nectariferous pollen types were Anadenanthera, Cordia, Eucalyptus, Mimosa scabrella, Schefflera, Sida, Serjania and Vernonia. Twenty-eight types of pollen from 21 genera and 19 families were identified in 22 bee bread samples. Fabaceae, Asteraceae and Myrtaceae showed the highest pollen richness. Anadenanthera, Cecropia, Eucalyptus, Melastomataceae, Mimosa scabrella, Mimosa verrucosa and Myrcia were the most frequent polliniferous pollen types. Principal component analysis (PCA) demonstrated that honey and pollen samples formed two main groups of similarity, mainly due to Eucalyptus’ nectar and pollen of Melastomataceae, respectively. Melipona quadrifasciata anthidioides collected nectar and pollen from the preserved areas as well as in the secondary and ‘ruderal’ vegetation and in cultivated forests/fields, suggesting their importance as pollinators both of native flora and exotic species. The use of trophic resources of plants grown with pesticides is a concern for the conservation of these species of bee and should be better studied.     

Plant Invasions on an Oceanic Archipelago

Plant invasions are particularly noticeable on oceanic islands. For many ecological or evolutionary phenomena, oceanic islands offer advantages in comparison to continental regions, because they are often simpler systems. The Juan Fernández (Robinson Crusoe) Islands, located 667 km west of continental Chile, provide an especially favorable case study of plant invasions on an oceanic archipelago. They have little flora, no influence from aboriginal peoples, and good historical and recent documentation of flora, vegetation and human impacts. The total flora of the archipelago consists of 441 vascular plants, of which 209 are native (125 of them endemic) and 232 are aliens. Many alien species exist on the Juan Fernández Archipelago, but three shrubs are particularly invasive: Aristotelia chilensis, Rubus ulmifolius, and Ugni molinae. About 15% of the total area of the island has been impacted by at least one of these shrubs. In addition, the herbaceous Acaena argentea, now occurs at high abundance in 12% of the total area of the island. Comparisons of earlier and recent surveys of vegetation reveal that the area impacted by Aristotelia, Rubus, and Ugni molinae has increased tremendously. Among the most frequent aliens are Euro-Mediterranean taxa, also present on other archipelagos. A few aliens that are serious invasives on other archipelagos have been recently observed near San Juan Bautisata, the only village on the island. Effective measures involving stronger monitoring and sanitation of incoming materials (foodstuffs, building materials, etc.), cutting or poisoning of invasives, and reseeding with native species, are necessary to help preserve the native and endemic flora (and biota) of these islands.     

Ribosomal and chloroplast DNA restriction site mutations and the radiation ofRobinsonia (Asteraceae: Senecioneae) on the Juan Fernandez Islands

Restriction site mutations in the chloroplast (cpDNA) and ribosomal DNA (rDNA) were examined in 41 populations representing five of the seven recognized species of the genusRobinsonia, which is endemic to the Juan Fernandez Islands. No intraspecific variation was detected for cpDNA but one population of one of the species (R. evenia) had a restriction site mutation in rDNA not detected elsewhere. No restriction site mutations were unique to all species ofRobinsonia relative to the species ofSenecio used as outgroups. All 13 mutations (eight from cpDNA and five from rDNA) are restricted to single species, and thus provide no cladistically useful information within the genus. The distribution of mutations is concordant with the hypothesis of a rapid adaptive radiation ofRobinsonia subsequent to the dispersal of its ancestor to Masatierra.     

ON THE METHODS OF EXPERIMENTAL TAXONOMY

By studying seedling progenies from individual plants it may be decided whether the material under investigation is allogamous, autogamous or apomictic. Chromosome counts disclose whether the material is cytologically stable or variable. If variation occurs it may be a question of polyploidy or aneuploidy. Aneuploidy may either be an occasional consequence of polyploidy or it may represent dysploidy and the formation of new basic numbers. Chromosome counts combined with measurements of chromosome size may reveal cases of pseudopolyploidy of the kind first observed in Luzula. If chromosome morphology is different between two taxa this indicates reproductive isolation. Different species usually differ with regard to their chromosome structure. Studies of meiosis are desirable in polyploids in order to distinguish between auto- and alloploidy. Cases of genic control of meiosis in polyploids of Phleum and Triticum demonstrate, however, that partial or complete homology between genomes does not always result in multivalent formation. Apparent alloploids may in reality be more or less autoploid. Incompatibility is not a reliable criterion of species differentiation. Strong or absolute barriers between diploids and autotetraploids may have purely quantitative causes. Hybrid sterility is not always a reliable measure of the degree of genetic difference between the parents, but may be caused by heterozygosity for chromosomal rearrangements. Cytological and experimental methods are, nevertheless, indispensable tools in plant taxonomy.