GENETIC AND MORPHOLOGICAL DIVERGENCE AMONG MORPHOTYPES OF THE ISOPOD EXCIROLANA ON THE TWO SIDES OF THE ISTHMUS OF PANAMA

Excirolana braziliensis is a dioecious marine isopod that lives in the high intertidal zone of sandy beaches on both sides of Central and South America. It possesses no larval stage and has only limited means of adult dispersal. Indirect estimates of gene flow have indicated that populations from each beach exchange less than one propagule per generation. Multivariate morphometrics have discovered three morphs of this species in Panama, two of them closely related and found on opposite sides of Central America (“C morph” in the Caribbean and “C′ morph” in the eastern Pacific), the third found predominantly in the eastern Pacific (“P morph”). Though the P and C′ morphs are seldom found on the same beach, they have overlapping latitudinal ranges in the eastern Pacific. A related species, Excirolana chamensis, has been described from the Pacific coast of Panama. Each beach contains populations that remain morphologically and genetically stable, but a single drastic change in both isozymes and morphology has been documented. We studied isozymes and multivariate morphology of 10 populations of E. braziliensis and of one population of E. chamensis. Our objective was to assess the degree of genetic and morphological variation, the correlation of divergence on these two levels of integration, the phylogenetic relationships between morphs, and the possible contributions of low vagility, low gene flow, and occasional extinction and recolonization to the genetic structuring of populations. Genetic distance between the P morph, on one hand, and the other two morphotypes of E. braziliensis, on the other, was as high as the distance between E. braziliensis and E. chamensis. Several lines of evidence agree that E. chamensis and the P morph had diverged from other morphs of E. braziliensis before the rise of the Panama Isthmus separated the C and C′ forms, and that the P morph constitutes a different species. A high degree of genetic differentiation also exists between populations of the same morph. On the isozyme level, every population can be differentiated from every other on the basis of at least one diagnostically different locus, regardless of geographical distance or morphological affiliation. Morphological and genetic distances between populations are highly correlated. However, despite the high degree of local variation, evolution of E. braziliensis as a whole has not been particularly rapid; divergence between the C and C′ morphs isolated for 3 million yr by the Isthmus of Panama is not high by the standard of within-morph differentiation or by comparison with other organisms similarly separated. Alleles that are common in one population may be absent from another of the same morph, yet they appear in a different morph in a separate ocean. The high degree of local differentiation, the exclusive occupation of a beach by one genotype with rare arrival of foreign individuals that cannot interbreed freely with the residents, the genetic stability of populations with infrequent complete replacement by another genetic population, and the sharing by morphs of polymorphisms that are not shared by local populations, all suggest a mode of evolution concentrated in rare episodes of extinction and recolonization, possibly coupled with exceptional events of gene flow that help preserve ancestral variability in both oceans.     

TWO SNOW SPECIES OF THE QUADRIFLAGELLATE GREEN ALGA CHLAINOMONAS (CHLOROPHYTA,VOLVOCALES): ULTRASTRUCTURE AND PHYLOGENETIC POSITION WITHIN THE CHLOROMONAS CLADE1

The quadriflagellate snow alga Chlainomonas Christen, distributed in New Zealand and North America, has several unusual structural attributes. A process assumed to be cytokinesis involves extrusion of protoplasm from the parent through a narrow canal, C. kolii (J. T. Hardy et Curl) Hoham produces a net‐like outer envelope rather than a cell wall, and the flagellar basal apparatus of Chlainomonas consists of two semi‐independent pairs of basal bodies. Structural connections between basal body pairs appear minimal, but a connecting system different from that observed in other genera exists within each pair. Phylogenetic analysis using rbcL sequences places Chlainomonas in the Chloromonas clade, other known members of which are all biflagellate. Chlainomonas is split into two robust lineages, with New Zealand collections sharing an origin with northern North American collections. Although the quadriflagellate condition is regarded as ancestral in the Chlorophyceae, we speculate—based on ultrastructural and molecular data presented here—that Chlainomonas represents a derived form that has arisen from fusion of two ancestral biflagellate cells. Other explanations (for example, that Chlainomonas represents a diploid form of a biflagellate species) are remotely possible but are presently at odds with extensive observations of field material. Improvements in techniques for experimental manipulation of these sensitive cryophiles will be required to fully characterize their structure and progress our understanding of their biology.     

Dioecy derived from distyly and pollination in Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands,Japan

Floral morphology and breeding system were examined in natural populations of Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands in southwest Japan. This species is morphologically dimorphic with long‐ and short‐styled morphs; however, these morphs are functionally dioecious. All long‐styled morphs set fruits, but their short anthers completely lack pollen grains; thus, they function as pistillate flowers. Alternately, short‐styled morphs usually have fertile pollen and never set fruit, whether after open or obligate pollinations; thus, they function as staminate flowers. However, some short‐styled morphs in Mounts Katsuu‐dake and Oppa‐dake populations have no fertile pollens. No fruit was obtained from bagging experiments, indicating that P. rubra requires pollinators for its reproduction. In our pollinator observations, flies and short‐tongued wasps were the main visitors of the flowers of P. rubra. Thus, these insects are probably effective pollinators for P. rubra in the Ryukyu Islands. Considering the morphological features, the dioecy found in P. rubra is probably derived from distyly; however, no sufficient evidence is available showing that dioecy has evolved “directly” from distyly.     

Synchronized dichogamy and dioecy in neotropicalLauraceae

In hermaphrodite neotropicalLauraceae a highly evolved dichogamous system is present which represents a kind of temporal dioecy. This system involves the existence of two flower morphs which are characterized by reciprocal phases of receptivity of the stigmas and pollen release. In some genera (Persea, Cinnamomum), nectar is produced as a reward for the flower visitors, while in other genera (Aniba, Clinostemon, Licaria), nectar is absent and pollen seems to be the only reward. This implies that in this case the flowers in the female stage must be deceptive flowers. In dioecious species of the generaOcotea andNectandra, both the male and female flowers attract the visitors with nectar. The pollen-ovule ratio of theLauraceae is comparatively low. — The type of reproductive system that characterizes theLauraceae, comprising functional dioecy, small, inconspicuously coloured flowers, pollination by small bees, and large, one-seeded fruits dispersed by birds, is quite prominent among trees of various families in the tropical lowland forest. The relationship between the different modes of flowering within theLauraceae and the causes for the correlation of their reproductive traits are discussed.     

Molecular comparisons of native range collections of Gadirtha fusca,a potential biological control agent of Chinese tallowtree

Chinese tallowtree, Triadica sebifera (L.) Small (Euphorbiaceae), is one of the worst invasive weeds of the southeastern USA impacting coastal wetlands, forests, and natural areas. A proposed biological control candidate, the defoliating moth Gadirtha fusca Pogue (Lepidoptera: Nolidae) shows high specificity for the target weed Chinese tallowtree. A total of 13 field sites were sampled in the native range of this herbivore species. To determine if all individuals were the same taxon, molecular DNA analyses were conducted of these collections. These included collections from 2012, 2015 and 2016, from three regions and two adult color morphs. Molecular COI analysis was performed on thirty colonized individuals from these collections. The results of this analysis arranged all taxa into a single clade with average genetic divergence values of 0.3%. Comparison of these G. fusca sequences and those from other Gadirtha spp. all had divergence values that were equal to or exceeded 5%. These results indicate that all the G. fusca collections were a single species and they were distinct from other known members of this genus.     

CHROMOSOME NUMBERS IN UMBELLIFERAE. III

Chromosome numbers are reported for 156 collections representing 100 taxa of Umbelliferae. Approximately two thirds of the collections are from Mexico, Central and South America and indicate a high percentage of polyploid species in certain genera found in this area. Chromosome numbers for plants belonging to 78 taxa are published here for the first time, previously published chromosome numbers are verified for 18 taxa and chromosome numbers differing from those previously published are reported in seven instances. No chromosome counts have been previously published for nine of the genera included here. Further aneuploidy and polyploidy were found in Eryngium, and Lomatium columbianum has been found to be a high polyploid with 2n = 14x. Every chromosome count is referable to a cited herbarium specimen.     

亚洲产Clavicorona的形态学及亲和性研究

中国产的Clavicorona pyxidata与北美的种样本间配对实验结果显示该种在不同的地理分布区内的种群之间完全性亲和或具有相同的交配型等位基因,表明中国产与北美产的该形态学种同属于一个生物种。日本产的标本与C. pyxidata在子实体外部形态及孢子尺度上有差异,虽然在日本被鉴定为该种。日本产的菌株与C. pyxidata菌株间配对实验证明二者之间完全不亲和,即属于不同的生物种。生殖隔离拌随着形态学变异。     

Phylogeography of the parthenogenic ant Platythyrea punctata: highly successful colonization of the West Indies by a poor disperser

Aim We investigated the spatio‐temporal patterns of genetic diversity in West Indian and mainland populations of a widespread parthenogenic ant (Platythyrea punctata F. Smith) to infer source populations and subsequent colonizations across its geographic range. Location Central America, Texas and the West Indies (Florida, the Bahamas, Greater and Lesser Antilles). Methods We employed phylogeographic reconstruction based on 1451 bp of mitochondrial DNA (cytochrome c oxidase subunits I and II) sequenced from 91 individuals of P. punctata. We employed standard population genetic analyses, Bayesian phylogenetic analyses, haplotype networks and molecular dating methods as performed by beast . We also employed phylogenetic analysis using two nuclear markers (970 bp) to understand the placement of P. punctata in the globally distributed genus Platythyrea. Results Based on highly reduced haplotypic variation and temporal estimates, rapid expansion and dispersal from Central America best explains the observed distribution of haplotypes. Platythyrea punctata successfully invaded the West Indies very few times. One haplotype occurred on every island surveyed from the Bahamas and Florida in the north to Barbados at the southern edge of its range. Haplotype diversity in the West Indies is quite low, despite a larger sample size relative to the mainland. Most mainland colonies collected each possessed a unique haplotype, whereas only Florida and the larger islands (the Dominican Republic, Puerto Rico and Guadeloupe) contained more than one haplotype. Island haplotypes were most similar to haplotypes collected in northern Mexico and southern Texas, but genetic distances were nevertheless high. The putative sister species of P. punctata appears to be an endemic of Hispaniola (P. strenua Wheeler & Mann), even though older, mainland populations of P. punctata are sympatric with at least two other congenerics. Main conclusions Dispersal seems very limited on the mainland, with well‐defined clades corresponding to geographical regions. Colonization of the islands from the mainland was extremely rare, but once successful there were very few barriers to expansion to nearly every island in the West Indies. We hypothesize that this invasion occurred during the late Pleistocene as the climate became warmer and less arid.     

Redescription of two genera of blastoids from China

In this paper we redescribe and reassign two genera of Chinese blastoids previously described by Mu (1955) and Chi (1943) based on our own new collections and those of other Chinese geologists. Mesoblastus tushanensis is reassigned to the Schizoblastidae from the Granatocrinidae based on the presence of ten spiracles and a separate anus and to the new genus Pinguiblastus. Previously Mississippian schizoblastids have been restricted to North America. Pinguiblastus provides a paleogeographic and evolutionary transitional taxon between the North American schizoblastid genera and the very abundant Permian faunas from Timor. Devonoblastus heitaiensis is reassigned to the new genus Sinoblastus based on the presence of large exposed cryptodeltoids, a taxonomically significant character not found in Devonoblastus. This removes known occurrences of Devonoblastus from Asia.     

Distribution and synonymy of Plagiochila punctata (Taylor) Taylor, with hypotheses on the evolutionary history of Plagiochila sect. Arrectae (Plagiochilaceae, Hepaticae)

For a long time, Plagiochila (sect. Arrectae) punctata (Taylor) Taylor has been treated as an endemic of Atlantic Europe. Studies of larger specimen sets demonstrated that the species is widespread in mountainous regions of the Neotropics where it is known under several names including P. stolonifera Lindenb. & Gottsche, P. choachina Gottsche and P. patzschkei Steph. In tropical Africa P. punctata is established as Plagiochila subalpina Steph., nom. illeg. The sporophyte of P. punctata is described for the first time, based on material from Costa Rica. A maximum likelihood analysis of a nrITS sequence alignment with sequences of P. sects. Fuscoluteae (outgroup), Arrectae, and Rutilantes as well as of P. rubescens (Lehm. & Lindenb.) Lindenb. results in a tree with well bootstrap supported Arrectae and Rutilantes clades. ITS sequences of Plagiochila punctata from the Comoros and Zaire are placed in an unsupported monophyletic lineage together with P. punctata sequences from the British Isles and Ecuador. The species is nested in a robust clade with P. spinulosa (Dicks.) Dumort. and P. stricta Lindenb. The weak genetic separation of these three species indicates a recent diversification; the disjunct ranges may be the result of long range dispersal events. Plagiochila rubescens from southern South America is placed sister to P. sect. Arrectae. The latter section possibly originated from southern South American ancestors, diversified in tropical America after the uplift of the Andes and reached the Holarctics and tropical Africa by long range dispersal of diaspores.We are grateful to the curators and owners of the herbaria cited in the text for the loan of specimens, especially to Tamás Pócs, Volker Buchbender and Eberhard Fischer for their continuous support with recent collections from tropical Africa. David Rycroft kindly sent a duplicate of P. rubescens. Financial support of the Deutsche Forschungsgemeinschaft (grant HE 3584/1) is gratefully acknowledged.     

CHROMOSOME NUMBERS AND TAXONOMIC NOTES FOR ROCKY MOUNTAIN UMBELLIFERAE

Chromosome numbers are reported for 111 collections of Umbelliferae, which represent 38 species belonging to 23 genera. Three of these genera, Aletes, Neoparrya, and Musineon, have not been counted previously. In addition, first counts are presented for Angelica grayi, A, roseana, Cymopterus acaulis, C. bulbosus, C. longipes, C. montanus, Lomatium foeniculaceum subsp. macdougalii, L. megarrhizum, L. nuttallii, L. orientate, and L. simplex. Our reports of polyploidy in Ligusticum ported (hexaploid, 2n = ca. 66) and in Pteryxia terebinthina var. calcarea (tetraploid, 2n = 44) are the first for these taxa. The counts of 2n = 22 for Harbouria trachypleura differ from the previous report of 2n = 20.     

Agouticarpa, a new neotropical genus of Tribe Gardenieae (Rubiaceae)

Agouticarpa, a new genus of the informalAlibertia group (Rubiaceae), is described and illustrated. It comprises six species, and occurs from Costa Rica to Bolivia. Three species are here described as new:Agouticarpa grandistipula, A. hirsuta, andA. velutina. Additionally, three new combinations are made:A. curvifolia andA. williamsii (previously inGenipa) andA. isernii (previously inAlibertia). Agouticarpa is characterized by being dioecious, having elliptic to abovate, membranaceous stipules, male flowers in a branched dichasial or thyrse-like inflorescence, a poorly developed cup-shaped calyx, pollen grains with 3–7 apertures, and large globose fruits.     

Is Phillyrea angustifolia L. (Oleaceae) an androdioecious species?

LEPART, J. & DOMMEE B., 1992. Is Phillyrea angustifolia L. (Oleaceae) an androdioecious species? Observations during two or three years on three natural populations of Phillyrea angustifolia growing in contrasting environments in southern France show that two distinct phenotypes occur in approximately equal proportions: hermaphrodites, which produce pollen and seeds, and males, which produce pollen only. The populations studied are thus morphologically androdioecious. Biometric investigation of the two morphs shows that they are clearly distinguished by (1) the shape of the stigma and (2) the ovary which is well developed in hermaphrodites and aborted in males. Neither size of anthers and corolla, nor pollen germination, differ between the two sexes. In the Camargue, where P. angustifolia is presently an invader, controlled crosses between hermaphrodites (selfing and inter-individual) are partially successful. In contrast, such crosses are sterile in the well-established population of the La Gardiole. Hence the Camargue population appears to be functionally androdioecious and La Gardiole functionally dioecious. This breeding system is particularly puzzling, since the occurrence of anemogamy and possible selfing do not fit classical explanations of androdioecy. The ability of hermaphrodites to be functionally male and female could be particularly adaptive in founding populations. Since functional hermaphroditism occurs in expanding populations of the Camargue population, we consider that the androdioecy of P. angustifolia could be a kind of leaky dioecism.     

Dimorphisms and fluctuating asymmetry in the forceps of male earwigs

Male dimorphisms are particularly conspicuous examples of the alternative reproductive strategies employed within some species. Such dimorphisms are thought to exist as genetic polymorphisms under ESS conditions, or to be conditional strategies where exogenous conditions determine the adult body plan. Fluctuating asymmetry (FA) is currently considered to be a fitness correlate of significant use in interpreting the functional significance of secondary sexual characteristics. In particular, negative slopes of FA on trait size are thought to arise in traits whose expression is dependent on condition. We measured forceps lengths and asymmetries in 2 island populations of the European earwig Forficula auricularia and Museum specimens of 5 other earwig (Dermaptera, Forficulidae) species from different genera, that appeared to be dimorphic. In a detailed study of Forficula auricularia we found a significant fit to a statistical model for the identification of dimorphisms and, for all species examined, morphs differed in the slope and/or elevation of the allometric relationship between body size and forcep length. Possible determinates of male dimorphisms are suggested from the data. Contrary to expectation, FA was not found to be greater in the minor morphs. Negative relationships between FA and forceps length were absent in both morphs of species examined from museum collections. Of the two island populations of Forficula auricularia, the smaller and more isolated population had higher FA and a negative relationship between FA and forceps length in the major morph. We discuss these patterns in the light of recent theories of FA and honest signalling.     

ATTEMPTED HYBRIDIZATION BETWEEN MONOECIOUS AND DIOECIOUS CLONES OF CHARA

The suggestion has been made that within the genus Chara the condition of monoecism or dioecism is of limited taxonomic significance, and that this character reflects only minor genetic variation, rarely sufficient to prevent hybridization between two otherwise similar clones. Objections have been raised to this suggestion. The present investigation was undertaken to determine whether the eggs of dioecious C. rusbyana could be fertilized by sperm from monoecious C sejuncta, monecious C. zeylanica, or dioecious C. kenoyeri; and in turn whether the eggs of C. sejuncta and C. zeylanica could be fertilized by sperm from C. rusbyana. On the basis of 83 crosses involving 28 clones of the four taxa, we conclude that C. rusbyana is genetically isolated from the other three.     

GERMINATION AND SEEDLING EMERGENCE IN SPERGULA ARVENSIS

Spergula arvensis L. (Caryophyllaceae) is a cosmopolitan weedy annual that exhibits a genetically based seed coat polymorphism in which two primary seed types, papillate (P) and nonpapillate (NP), are produced. This study examined germination response of the two morphs and seedling emergence in Pacific coast populations. Germination behavior was complex and variable among populations and between seed morphs. Seeds from some populations exhibited greater temperature sensitivity and/or differentiation between morphs. Freshly produced seeds from grassland and ruderal populations were strongly dormant at the time of dispersal, in contrast to seeds from cultivated or summer-wet populations. In San Francisco Bay area populations, germination and seedling emergence was largely simultaneous, following the first substantial fall rains.     

Does habitat-specific variation in trematode infection risks influence habitat distribution of two closely related freshwater snails?

Parasitism may be an important factor determining the geographic distribution of closely related species. A habitat-specific risk of parasitism may lead to exclusion of susceptible host types from parasite-rich environments, and promote speciation if it leads to reproductive isolation between susceptible and resistant types. We surveyed populations of the freshwater snail Lymnaea peregra for differences in habitat distribution and trematode parasitism between its two distinct shell morphs, L. ovata and L. peregra. We surveyed 58 populations (43 L. ovata, 15 L. peregra). At each location we recorded an array of habitat characteristics that were summarized using a nonlinear principal components analysis. This yielded two orthogonal habitat score variables. Discriminant analysis with these habitat dimensions indicated that the snail morphs differed in their habitat distribution. L. ovata preferred larger, more permanent natural habitats surrounded by forests, while L. peregra was found more often at a higher altitude, in nonpermanent habitats, often surrounded by meadows. The snails were parasitized by four cercarial types of castrating trematodes. The morphs had a similar prevalence of infection by each of the parasite types, with one exception: monostomid cercariae were found at a higher prevalence in L. ovata than in L. peregra. However, monostomes were rare parasites, and the difference in prevalence of infection was not significant when only populations with monostomes were compared. Our results indicate that variation in the overall prevalence of infection seems to be independent of snail morph, and do not support the idea that a difference in the rate of parasitism might explain differences in the habitat distribution of these snail morphs. Received: 4 January 1999 / Accepted: 30 June 1999     

British Bryological Society Expedition to Mulanje Mountain,Malawi. 17 Sematophyllaceae (Bryopsida): Part 1

Abstract

The family Sematophyllaceae is a difficult family to treat in Africa because of the large number of species described without any subsequent revision or monography and, despite the many collections mentioned in this paper, only two species had previously been identified from Malawi. A total of four species in four genera (Gammiella, Mρacrohymenium, Radulina and Wijkia) are recorded in this paper, all but one new to Malawi. The two remaining genera (Sematophyllum and Trichosteleum) present additional difficulties of identification, and will be dealt with separately.