Mechanisms of rapid sympatric speciation by sex reversal and sexual selection in cichlid fish

Mechanisms of speciation in cichlid fish were investigated by analyzing population genetic models of sexual selection on sex-determining genes associated with color polymorphisms. The models are based on a combination of laboratory experiments and field observations on the ecology, male and female mating behavior, and inheritance of sex-determination and color polymorphisms. The models explain why sex-reversal genes that change males into females tend to be X-linked and associated with novel colors, using the hypothesis of restricted recombination on the sex chromosomes, as suggested by previous theory on the evolution of recombination. The models reveal multiple pathways for rapid sympatric speciation through the origin of novel color morphs with strong assortative mating that incorporate both sex-reversal and suppressor genes. Despite the lack of geographic isolation or ecological differentiation, the new species coexists with the ancestral species either temporarily or indefinitely. These results may help to explain different patterns and rates of speciation among groups of cichlids, in particular the explosive diversification of rock-dwelling haplochromine cichlids.

Explosive speciation rates and unusual species richness in haplochromine cichlid fishes: Effects of sexual selection

Ancient lakes are often unusually species rich, mostly as a result of radiation and species-flock formation having taken place in only one or a few of many taxa present. Understanding why some taxa radiate and others do not is at the heart of understanding biodiversity. In this chapter I discuss possible explanations for disproportionally large species numbers in some cichlid fish lineages in East African Great Lakes: the halochromine cichlid fishes in Lakes Victoria and Malawi. I show that speciation rates in this group are higher than in any other lacustrine fish radiation. Against this background, I review hypotheses put forward to explain diversity in cichlid species flocks. The evolution of species diversity requires three processes: speciation, ecological radiation and anatomical diversification, and it is wrong to consider hypotheses that are relevant to different processes as alternatives to each other. The African cichlid species flocks show unusually high ecological species packing in several phylogenetic groups and unusually high speciation rates in haplochromines. Therefore, it maybe concluded that at least two evolutionary models are required to explain the difference between cichlid diversity and other fish diversity in East African Lakes: one for speciation in haplochromines and one for coexistence. Subsequently I review work on speciation in haplochromines, and in particular studies aimed at testing the hypothesis of speciation by sexual selection. Haplochromines have a polygynous mating system, conducive to sexual selection, but other polygynous cichlids are not particularly species rich. This suggests that more than just strong sexual selection is required to explain haplochromine species richness. Recent palaeoecological evidence undermines the previously popular hypotheses that explained the species richness of Lake Victoria in terms of speciation under varying natural or sexual selection regimes in satellite lakes or in isolated lake basins. I summarize experimental and comparative studies, which provide evidence for two mechanisms of sympatric speciation by disruptive sexual selection on polymorphic coloration. Such modes of speciation may explain (i) the high speciation rates in colour polymorphic lineages of haplochromine cichlids under conditions where colour variation is visible in clear water, and (ii) in combination with factors that affect population survival, the unusual species richness in haplochromine species flocks. I argue that sexual selection, if disruptive, can accelerate the pace of adaptive radiation because the resultant genetic population fragmentation allows a much increased rate of differential response to disruptive natural selection. Hence, the ecological pattern of diversity resembles that produced by disruptive natural selection, with the difference that disruptive sexual selection continues to cause (gross) speciation even after niche space is saturated. This may explain the unusually high numbers of very closely related and ecologically similar species in haplochromine species flocks. The role of disruptive sexual selection is twofold: it not only causes speciation, but also maintains reproductive isolation in sympatry between species that have evolved in sympatry or allopatry. Therefore, the maintenance of diversity in species flocks that originated through sexual selection depends on the persistence of the selection regime within the environmental signal space under which that diversity evolved.

Color polymorphism and sex ratio distortion in a cichlid fish as an incipient stage in sympatric speciation by sexual selection

We investigated a Lake Victoria cichlid with a complex colour polymorphism that apparently represents one original species and two incipient species, all of which are sympatric. In laboratory breeding experiments we observed sex ratio distortion in certain matings between original and incipient species. Mate choice experiments show that males of the incipient species exhibit mating preferences against the original species, and males and females of the original species exhibit strong mating preferences against the incipient species. Mating preferences might evolve by sex ratio selection to avoid matings with distorted progeny sex ratios. Phenotype frequencies in nature suggest that mating preferences translate into mating frequencies, thus restricting gene flow and exerting disruptive sexual selection between the original and incipient species. The incipient species do not differ in morphology or ecology from the original species, implying that colour polymorphism, associated with sex ratio distortion, can be an incipient stage in sympatric speciation, and that disruption of gene flow can precede ecological differentiation

Can sympatric speciation by disruptive sexual selection explain rapid evolution of cichlid diversity in Lake Victoria?

Rapid speciation can occur on ecological time scales and interfere with ecological processes, resulting in species distribution patterns that are difficult to reconcile with ecological theory. The haplochromine cichlids in East African lakes are an extreme example of rapid speciation. We analyse the causes of their high speciation rates. Various studies have identified disruptive sexual selection acting on colour polymorphisms that might cause sympatric speciation. Using data on geographical distribution, colouration and relatedness from 41 species endemic to Lake Victoria, we test predictions from this hypothesis. Plotting numbers of pairs of closely related species against the amount of distributional overlap between the species reveals a bimodal distribution with modes on allopatric and sympatric. The proportion of sister species pairs that are heteromorphic for the traits under disruptive selection is higher in sympatry than in allopatry. These data support the hypothesis that disruptive sexual selection on colour polymorphisms has caused sympatric speciation and help to explain the rapid radiation of haplochromine species flocks.

Coccolithophore sensitivities to changing carbonate chemistry - an ecological framework

Coccolithophores are a group of unicellular phytoplankton species whose ability to calcify has a profound influence on biogeochemical element cycling. Calcification rates are controlled by a large variety of biotic and abiotic factors. Among these factors, carbonate chemistry has gained considerable attention during the last years as coccolithophores have been identified to be particularly sensitive to ocean acidification. Despite intense research in this area, a general concept harmonizing the numerous and sometimes (seemingly) contradictory responses of coccolithophores to changing carbonate chemistry is still lacking to date. Here, we present the "substrate-inhibitor concept" which describes the dependence of calcification rates on carbonate chemistry speciation. It is based on observations that calcification rate scales positively with bicarbonate (HCO3-), the primary substrate for calcification, and carbon dioxide (CO2), which can limit cell growth, whereas it is inhibited by protons (H+). This concept was implemented in a model equation, tested against experimental data, and then applied to understand and reconcile the diverging responses of coccolithophorid calcification rates to ocean acidification obtained in culture experiments. Furthermore, we (i) discuss how other important calcification-influencing factors (e.g. temperature and light) could be implemented in our concept and (ii) embed it in Hutchinson's niche theory, thereby providing a framework for how carbonate chemistry-induced changes in calcification rates could be linked with changing coccolithophore abundance in the oceans. Our results suggest that the projected increase of H+ in the near future (next couple of thousand years), paralleled by only a minor increase of inorganic carbon substrate, could impede calcification rates if coccolithophores are unable to fully adapt. However, if calcium carbonate (CaCO3) sediment dissolution and terrestrial weathering begin to increase the oceans' HCO3- and decrease its H+ concentrations in the far future (10 -100 kyears), coccolithophores could find themselves in carbonate chemistry conditions which may be more favorable for calcification than they were before the Anthropocene.

A test of alternative models of diversification in tropical rainforests: Ecological gradients vs. rainforest refugia

Comparison of mitochondrial and morphological divergence in eight populations of a widespread leaf-litter skink is used to determine the relative importance of geographic isolation and natural selection in generating phenotypic diversity in the Wet Tropics Rainforest region of Australia. The populations occur in two geographically isolated regions, and within each region, in two different habitats (closed rainforest and tall open forest) that span a well characterized ecological gradient. Morphological differences among ancient geographic isolates (separated for several million years, judging by their mitochondrial DNA sequence divergence) were slight, but morphological and life history differences among habitats were large and occurred despite moderate to high levels of mitochondrial gene flow. A field experiment identified avian predation as one potential agent of natural selection. These results indicate that natural selection operating across ecological gradients can be more important than geographic isolation in similar habitats in generating phenotypic diversity. In addition, our results indicate that selection is sufficiently strong to overcome the homogenizing effects of gene flow, a necessary first step toward speciation in continuously distributed populations. Because ecological gradients may be a source of evolutionary novelty, and perhaps new species, their conservation warrants greater attention. This is particularly true in tropical regions, where most reserves do not include ecological gradients and transitional habitats.

Evidence for sympatric speciation by host shift in the sea

The genetic divergence and evolution of new species within the geographic range of a single population (sympatric speciation) contrasts with the well-established doctrine that speciation occurs when populations become geographically isolated (allopatric speciation). Although there is considerable theoretical support for sympatric speciation [1, 2], this mode of diversification remains controversial, at least in part because there are few well-supported examples [3]. We use a combination of molecular, ecological, and biogeographical data to build a case for sympatric speciation by host shift in a new species of coral-dwelling fish (genus Gobiodon). We propose that competition for preferred coral habitats drives host shifts in Gobiodon and that the high diversity of corals provides the source of novel, unoccupied habitats. Disruptive selection in conjunction with strong host fidelity could promote rapid reproductive isolation and ultimately lead to species divergence. Our hypothesis is analogous to sympatric speciation by host shift in phytophagous insects [4, 5] except that we propose a primary role for intraspecific competition in the process of speciation. The fundamental similarity between these fishes and insects is a specialized and intimate relationship with their hosts that makes them ideal candidates for speciation by host shift.

Assessing the Legacy of Red Mud Pollution in a Shallow Freshwater Lake: Arsenic Accumulation and Speciation in Macrophytes

Little is known about long-term ecological responses in lakes following red mud pollution. Among red mud contaminants, arsenic (As) is of considerable concern. Determination of the species of As accumulated in aquatic organisms provides important information about the biogeochemical cycling of the element and transfer through the aquatic food-web to higher organisms. We used coupled ion chromatography and inductively coupled plasma mass spectrometry (ICP-MS) to assess As speciation in tissues of five macrophyte taxa in Kinghorn Loch, UK, 30 years following the diversion of red mud pollution from the lake. Toxic inorganic As was the dominant species in the studied macrophytes, with As species concentrations varying with macrophyte taxon and tissue type. The highest As content measured in roots of Persicaria amphibia (L.) Gray (87.2 mg kg-1) greatly exceeded the 3 - 10 mg kg-1 range suggested as a potential phytotoxic level. Accumulation of toxic As species by plants suggested toxicological risk to higher organisms known to utilise macrophytes as a food source.

Experiences From The Brazilian Atlantic Forest: Ecological Findings And Conservation Initiatives.

The Brazilian Atlantic Forest hosts one of the world's most diverse and threatened tropical forest biota. In many ways, its history of degradation describes the fate experienced by tropical forests around the world. After five centuries of human expansion, most Atlantic Forest landscapes are archipelagos of small forest fragments surrounded by open-habitat matrices. This 'natural laboratory' has contributed to a better understanding of the evolutionary history and ecology of tropical forests and to determining the extent to which this irreplaceable biota is susceptible to major human disturbances. We share some of the major findings with respect to the responses of tropical forests to human disturbances across multiple biological levels and spatial scales and discuss some of the conservation initiatives adopted in the past decade. First, we provide a short description of the Atlantic Forest biota and its historical degradation. Secondly, we offer conceptual models describing major shifts experienced by tree assemblages at local scales and discuss landscape ecological processes that can help to maintain this biota at larger scales. We also examine potential plant responses to climate change. Finally, we propose a research agenda to improve the conservation value of human-modified landscapes and safeguard the biological heritage of tropical forests.

All Green, But Equal? Morphological Traits And Ecological Implications On Spores Of Three Species Of Mosses In The Brazilian Atlantic Forest.

Spores of the tropical mosses Pyrrhobryum spiniforme, Neckeropsis undulata and N. disticha were characterized regarding size, number per capsule and viability. Chemical substances were analyzed for P. spiniforme and N. undulata spores. Length of sporophyte seta (spore dispersal ability) was analyzed for P. spiniforme. Four to six colonies per species in each site (lowland and highland areas of an Atlantic Forest; Serra do Mar State Park, Brazil) were visited for the collection of capsules (2008 - 2009). Neckeropsis undulata in the highland area produced the largest spores (ca. 19 µm) with the highest viability. The smallest spores were found in N. disticha in the lowland (ca. 13 µm). Pyrrhobryum spiniforme produced more spores per capsule in the highland (ca. 150,000) than in lowland (ca. 40,000); longer sporophytic setae in the lowland (ca. 64 mm) than in the highland (ca. 43 mm); and similar sized spores in both areas (ca. 16 µm). Spores of N. undulata and P. spiniforme contained lipids and proteins in the cytoplasm, and acid/neutral lipids and pectins in the wall. Lipid bodies were larger in N. undulata than in P. spiniforme. No starch was recorded for spores. Pyrrhobryum spiniforme in the highland area, different from lowland, was characterized by low reproductive effort, but presented many spores per capsule.

Ecological Literacy And Beyond: Problem-based Learning For Future Professionals.

Ecological science contributes to solving a broad range of environmental problems. However, lack of ecological literacy in practice often limits application of this knowledge. In this paper, we highlight a critical but often overlooked demand on ecological literacy: to enable professionals of various careers to apply scientific knowledge when faced with environmental problems. Current university courses on ecology often fail to persuade students that ecological science provides important tools for environmental problem solving. We propose problem-based learning to improve the understanding of ecological science and its usefulness for real-world environmental issues that professionals in careers as diverse as engineering, public health, architecture, social sciences, or management will address. Courses should set clear learning objectives for cognitive skills they expect students to acquire. Thus, professionals in different fields will be enabled to improve environmental decision-making processes and to participate effectively in multidisciplinary work groups charged with tackling environmental issues.

Atherinella blackburni (Schultz, 1949) at Itamambuca Beach, Ubatuba, SP: ecological characterization and distribution on the Brazilian coast (Teleostei: Atheriniformes: Atherinopsidae)

Atherinella blackburni is a silverside species whose occurrence on the Brazilian coast was not properly registered until recently. So far, records of its distribution along the Brazilian shore were limited to Itaparica Island, Bahia State, and Porto Inhaúma, Rio de Janeiro State. In a recent survey of the ichthyofauna of Itamambuca Beach, Ubatuba, São Paulo State, 100 specimens of this species were collected, yielding a considerable source of information regarding its ecology and a new southern limit of its distribution. A detailed survey of an ichthyological collection revealed lots of this species from regions of the Brazilian Northeast, resulting in a northward expansion of the occurrence of A. blackburni in Brazilian waters. Besides the populations found on the Brazilian coast, the species also occurs discontinuously in the Atlantic coasts of Costa Rica, Panama and Venezuela. Meristic and morphometric analysis showed overlapped values between Brazilian populations and the ones closer to the Caribbean. An ecological study of A. blackburni at Itamambuca Beach revealed that it is a predominantly diurnal species with preference for warm and salty waters, often occurring in sandy-bottom environments. It was more abundant in Summer and Winter, when individuals with smaller values of length and weight also occurred.

Ecological aspects of the surf-zone ichthyofauna of Itamambuca Beach, Ubatuba, SP

Sandy beaches are among the most extensive environments in Brazil and are generally threatened by disorderly urban growth due to their location along the coast, where human populations tend to concentrate. This is especially true in southeastern Brazil, where the largest urban areas are located. Thus, better knowledge regarding these natural resources is of considerable importance. The surf zone of sandy beaches is inhabited by a number of fish species, including juveniles seeking both food and shelter from predators. An ecological survey was conducted in the surf zone of Itamambuca beach in the city of Ubatuba, São Paulo State, Brazil, in order to evaluate the composition, structure and dynamics of the fish fauna using community descriptors. Diurnal and nocturnal samples were taken with a beach seine at five oceanographic stations over a 12-month period. Water temperature and salinity were also recorded. A total of 2,147 individuals representing 34 species and 18 families were sampled. The main variation in fish fauna was detected on the diel level, followed by a spatial repartition possibly related to salinity. Seasonal variation was also observed. The five most important species in the overall context were Atherinella brasiliensis, Eucinostomus melanopterus, Mugil sp., Trachinotus goodei and Atherinella blackburni.

Culicidae (Diptera, Culicomorpha) from the western Brazilian Amazon: Juami-Japurá Ecological Station

With 312 trap-hours of sampling effort, 1554 specimens of Culicidae (Diptera) were collected, using CDC and Malaise traps, in nine different locations along the Juami River, within the Juami-Japurá Ecological Station, Amazonas State, Brazil. A list of mosquito species with 54 taxa is presented, which includes three new distributional records for the state of Amazonas. The species found belong to the genera Anopheles, Aedeomyia, Aedes, Psorophora, Culex, Coquillettidia, Sabethes, Wyeomyia and Uranotaenia.

American cutaneous leishmaniasis in the Pontal of Paranapanema - SP, Brazil: ecological and entomological aspects

American cutaneous leishmaniasis (ACL) occurs in epidemic outbreaks and in sporadic cases with small annual variation in the Pontal of Paranapanema, SP. There is little research on the sandfly fauna of this region. The last outbreaks were related to the Movement of the Landless Workers (MST) and with the ecological tourism in preserved forest of the Parque Estadual do Morro do Diabo (PEMD). AIM: identification of the sandfly fauna within the PEMD, mainly anthropophilic species already incriminated as vectors of ACL, as well as their seasonality, hourly frequency and data of the behavior. M&M: The captures were undertaken with CDC light and Shannon traps from 6:00 pm to 10:00 pm, monthly from May 2000 to December 2001. The temperature and relative humidity data were registered at hourly intervals. RESULTS: The captured species were: Brumptomyia brumpti, Nyssomyia neivai, Nyssomyia whitmani, Pintomyia fischeri and Pintomyia pessoai. The P. pessoai predominated (34.39%) and N. neivai was less found (0.74%), only being captured in CDC traps. Shannon trap captured more sandflies (63.01%) than the CDC traps (36.99%). Despite the environmental degradation anthropophilic species, indicates favorable bioecological conditions for persistence of vectors and potential transmission of leishmaniasis.