AUTOGAMOUS ALLOHOMOPLOIDY IN ALSOPHILA AND NEPHELEA (CYATHEACEAE): A NEW HYPOTHESIS FOR SPECIATION IN HOMOPLOID HOMOSPOROUS FERNS

Morphological and chemosystematic studies demonstrate that three endemic Puerto Rican tree ferns, Alsophila bryophila, A. dryopteroides and Nephelea portoricensis have been hybridizing and have produced a hybrid swarm. Three distinct hybrids are present, one of which is strikingly uniform in its morphology. The population ecology of species and hybrids indicates that the most uniform hybrid is more abundant and has a narrower ecological adaptation than its parents. The hybrids are fertile diploids and are producing F₂ segregates which resemble species from other islands in the Greater Antilles. An hypothesis for the stabilization of hybrid reproduction at the homoploid level, autogamous allohomoploidy, is proposed. The hypothesis explains the sharing of unique characters between sympatric species pairs of Alsophila and Nephelea in the Greater Antilles and in Central and South America. It is suggested that other genera of homosporous ferns have also speciated by autogamous allohomoploidy.     

ELECTROPHORESIS IS MODIFYING OUR CONCEPTS OF EVOLUTION IN HOMOSPOROUS PTERIDOPHYTES

Analyses of electrophoretically detectable enzyme variants in homosporous pteridophytes are facilitating the development of new insights into their genetics and evolution. The number of isozymes per enzyme indicates that homosporous pteridophytes are genetic diploids, in spite of the fact that they have high chromosome numbers. High levels of heterozygosity and genetic variability in sporophytic populations indicate that many diploid species are outcrossing with inbreeding representing a derived character state. Because the congeneric homosporous pteridophyte species analyzed to date have low genetic identities, allozymic characters are also proving to be useful as genomic markers for elucidating patterns of reticulate evolution. The accumulated data suggest that the genetic system of homosporous pteridophytes differs fundamentally from that of seed plants. The present genomic constitution of extant taxa may be the result of repeated cycles of allopolyploidy followed by gene silencing and extinction of progenitor taxa. Alternatively, the original homosporous pteridophytes may have had high chromosome numbers. Although current species probably evolved recently, their phylogenetic roots may be difficult to trace because even closely related pteridophytes are genetically distant and extinction has obliterated the ancestral intermediates between lineages. These hypotheses can and should be tested using a combination of molecular, phylogenetic, and population biology methods.     

MYCORRHIZAE IN HAWAIIAN PTERIDOPHYTES: OCCURRENCE AND EVOLUTIONARY SIGNIFICANCE

Forty-five percent of the Hawaiian pteridophyte flora were examined for mycorrhizae. Vesicular-arbuscular mycorrhizae were present in 66 of the 89 species examined. Nonmycotrophy was significantly higher in Hawaiian pteridophytes than in non-Hawaiian pteridophytes. The highest mycorrhizal colonization occurred in the leptosporangiate families Dicksoniaceae, Dryopteridaceae, and Lindsaeaceae. Mycotrophy was lower in the eusporangiate families. Substrate strongly influenced the presence of mycotrophy. Mycorrhizae were present in 83% of terricolous species, in 86% of epilithic species, in 55% of epiphytic species, and absent from aquatic species. A phylogram of pteridophyte families is constructed incorporating the intensity of mycorrhizal colonization as one criterion for classification.     

A NEW KIND OF SEXUAL DIMORPHISM IN ORDOVICIAN OSTRACODES

Abstract:  In Ordovician ostracodes (genus Incisua ) a new kind of sexual dimorphism is described, which is the first example among this group with males being the heteromorphs. The valves of males are larger and less frequent than the females, and are characterized by having a furrow in the ventral part. The furrow may have functioned as a device to enable the two sexes to cling together during mating.     

GENETIC LOAD AND THE MATING SYSTEM IN HOMOSPOROUS FERNS

The mutational genetic load was calculated assuming mutation-selection-inbreeding equilibrium and applied to homosporous ferns. Diploid species with past inbreeding should have a low genetic load while outcrossers should have a high genetic load. These predictions are consistent with the bimodal pattern of genetic load found in 18 diploid homosporous fern species. The prediction that tetraploids should have a low genetic load is also consistent with estimates of genetic load in several species.     

EVIDENCE FOR GENETIC HETEROZYGOSITY IN A HOMOSPOROUS FERN

Homosporous fern sporophytes from natural populations exhibited heterozygous electrophoretic patterns for several enzyme systems. Genetic tests utilizing individual gametophytes demonstrate that the observed heterozygosity is coded by alleles at single loci. This simple procedure makes it possible to distinguish segregating from fixed or phenotypic heterozygosity, previously a problem in homosporous vascular plants.     

ABNORMAL REDUCTIONAL AND NONREDUCTIONAL MEIOSIS IN CERATOPTERIS: ALTERNATIVES TO HOMOZYGOSITY AND HYBRID STERILITY IN HOMOSPOROUS FERNS

Natural and synthesized hybrids of Ceratopteris were investigated to determine the effect of hybridization on the genetic system. Studies indicated that the hybrids exhibited massive spore abortion and pairing abnormalities at meiotic prophase, characteristic of “sterile diploids and triploids” reported in hybridization studies of other fern genera. However, a small percentage of viable spores also was produced by the hybrids. Cytological investigations indicated the presence of previously unreported meiotic adaptations that allowed the production of unreduced spores and reduced spores exhibiting chromatid heterozygosity. The reduced spores allow haploid gametophytes to form heterozygous zygotes in spite of intragametophytic selfing. The unreduced spores were shown to be responsible for the fertility of the “sterile” hybrid and allowed the subsequent production of up to three generations of sporophytes. The literature suggests that these meiotic adaptations are present in other fern genera and may play a significant role in evolution through hybridization.     

SEXUAL DIMORPHISM,MATING SYSTEM AND BODY SIZE IN NEW WORLD BLACKBIRDS (ICTERINAE)

Although sexual selection is widely accepted as a primary functional cause of sexual size dimorphism in birds and mammals, results from some comparative studies have cast doubt on this conclusion. Chief among these contradictory results is the widespread association between body size and size dimorphism—large species tend to be more dimorphic than small species. This correlation is not directly predicted by the normal sexual selection scenario, and many hypotheses have been advanced to explain it. This paper reviews these hypotheses and evaluates them using data for the New World blackbirds (Icterinae). In this avian subfamily, (1) body size correlates with the intensity of sexual selection (as measured by mean harem size), and (2) size does not correlate with dimorphism if the effects of mating system are removed. Similar results are obtained when controlling for the confounding influence of phylogeny. Further, body size and mating system are associated with nesting dispersion. These results strongly argue that sexual dimorphism is a product of sexual selection in this subfamily, and suggest that either: (1) large body size itself, or the ecology of large species, promotes the development of coloniality and a polygynous mating system; or (2) polygyny and/or coloniality lead to the evolution of large size in both males and females. None of the other hypotheses examined predict an association between size and mating system, and all predict that size will correlate with dimorphism after the effects of mating system are removed. Thus, none of the other hypotheses seem applicable in this case. These results are compared to those obtained for other avian and mammalian taxa. Difficulties of analysis present in previous studies are discussed. I argue that it is inappropriate to assume that associations between a trait and body size or phylogeny are evidence of nonadaptive evolutionary “constraints.”     

A NEW MEASURE OF GENETIC IDENTITY BETWEEN POPULATIONS OF SEXUAL AND ASEXUAL SPECIES

We define a new genetic identity measure that is especially well suited for asexual polyploid species as it circumvents errors in the estimation of gene frequencies. It also can be applied to sexuals allowing the study of phylogenetic relationships in species complexes consisting of sexuals and asexuals of different ploidy levels. The measure groups genotypes into classes dependent on homozygosity vs heterozygosity and the number of ancestral allele types vs the number of presumed new mutations. Its value is related to evolutionary time since divergence. The application of the method is illustrated by using electrophoretic data on the species group Solenobia triquetrella (Lepidoptera: Psychidae). A high similarity of estimated relationships among the proposed as well as other genetic identity measures is shown in the case of diploid sexual and asexual races of this species group. The phylogenetic relationships within the group are reanalyzed and monophyletic vs polyphyletic origin of parthenogenesis in this species complex is discussed. The genetic identity values found by the proposed procedure are explained by a polyphyletic origin of parthenogenesis, though a monophyletic origin of parthenogenesis in a broader sense cannot be excluded. The explanation of the phylogenetic relationships is based on the assumption of hybridization between related species and the extinction of one ancestral species. Furthermore, the genetic diversity is compared among sexual and parthenogenetic races of the species.     

EQUIVALENCE OF COMPETITORS IN PLANT COMMUNITIES: A NULL HYPOTHESIS AND A FIELD EXPERIMENTAL APPROACH

One of the underlying assumptions of both theoretical and empirical community ecology is that the processes determining community composition and abundance of species are interactions specific to particular pairs of species. However, we argue that, in sessile plants at least, competitive interactions are not usually species-specific and that there exists a large degree of equivalence of the effect of species of similar growth form on the ability of any particular species to establish within a community. This null hypothesis of equivalence of competitive effects is based on three characteristics of plants: homogeneity of resource requirements among autotrophs; low encounter probabilities between individuals of any particular species pair; and the predominance of size asymmetries between competing individuals (e.g., seedling-adult interactions.) We present an experimental design to quantify competitive interactions among plant species under field conditions and therefore enable statistical comparisons of competitive abilities among species. The competitive effect of one “neighbor” species on one “target” species is measured as the slope of a regression of performance of target individuals on biomass (or other measure of amount) of its immediate neighbors. Use of the design to test for equivalence of competitive effects and other advantages are described.     

ESTIMATES OF INTRAGAMETOPHYTIC SELFING AND INTERPOPULATIONAL GENE FLOW IN HOMOSPOROUS FERNS

Relatively little information is available on mating systems and interpopulational gene flow in species of homosporous pteridophytes. Because of the proximity of antheridia and archegonia on the same thallus, it has long been maintained that intragametophytic selling is the predominant mode of reproduction in natural populations of homosporous ferns and other homosporous plants. Furthermore, quantitative estimates of interpopulational gene flow via spore dispersal are lacking. In this paper, we examine five species of homosporous ferns (Botrychium virginianum, Polystichum munitum, P. imbricans, Blechnum spicant, and Dryopteris expansa) and present estimates of 1) rates of intragametophytic selling, 2) levels of interpopulational gene flow, and 3) interpopulational genetic differentiation (F_ST). Our data demonstrate that mating systems vary among species of ferns, just as they do among species of seed plants. The data also suggest that levels of interpopulational gene flow are generally high. The F_ST values indicate little genetic divergence among populations for all species except Dryopteris expansa, which exhibits significant levels of interpopulational genetic differentiation. Patterns of genetic diversity in the five species examined are related to the mating system and rate of interpopulational gene flow in each species. The F_ST values for all species except Botrychium virginianum are in close agreement with those predicted for an island model of population structure.     

OBLIGATE OUTCROSSING IN A HOMOSPOROUS FERN: FIELD CONFIRMATION OF A LABORATORY PREDICTION

While homosporous ferns are potentially capable of producing totally homozygous sporophytes in one generation via selfing of their bisexual gametophytes, laboratory analyses indicate that a variety of mechanisms promote gametophytic outcrossing. The operation of these mechanisms in natural sporophyte populations, however, has not been previously demonstrated. Laboratory analyses of gametophyte ontogeny show that Bommeria hispida is obligately outcrossing. Electrophoretic data presented here indicate that individuals from natural sporophyte populations of this species are highly heterozygous. Electrophoretic data, therefore, corroborate evidence from the in vitro analysis of gametophyte development and demonstrate that sporophytes of B. hispida in nature typically are products of outcrossing between genetically different gametophytes. Extrapolations from the literature, together with our findings, indicate that outcrossing mechanisms may operate frequently in ferns, thereby maintaining genetic variability between individuals within populations. This evidence questions whether most ferns are highly inbred and therefore predominantly homozygous.     

BIOGEOGRAPHY OF ASEXUAL AND SEXUAL REPRODUCTION IN CALOGLOSSA (DELESSERIACEAE,RHODOPHYTA) FROM AUSTRALIA AND NEW ZEALAND

Caloglossa species are widely distributed in mangroves and salt marshes around the world and their life history patterns are being investigated in laboratory culture. In Australia all isolates of C. monosticha, C. postiae and C. ogasawaraensis have Polysiphonia‐type (P‐type) sexual life histories. Among the 70 C. leprieurii isolates from Australia and New Zealand P‐type sexual reproduction also is dominant. However, ten isolates of C. leprieurii from the Spencer Gulf and the Gulf of St. Vincent in South Australia give rise to successive tetrasporphyte generations without gametophytes. Moreover, one isolate from Queensland is asexual. Only one South Australia isolate, obtained from Lake Alexandrina at the mouth of the Murray River, is sexual. South Australia and Pacific Mexico are two regions in which asexual reproduction is dominant. In another mangrove dwelling red alga Bostrychia moritiziana (Rhodomelaceae) non‐sexual reproduction also is frequent in Australia, New Caledonia and Bali (Indonesia). This asexual reproductive pattern of tetrasporophytic recycling appears to have arisen independently among individual populations of various red algal species in different regions. Investigations are underway on the molecular phylogeny of the Caloglossa leprieurii isolates.