THE ROLE OF DIATOMS IN STROMATOLITE GROWTH: TWO EXAMPLES FORM MODERN FRESHWATER SETTINGS1

Some modern laminated find calcified stromatolitic structures are partially or completely formed by eukaryotes. Diatom populations in freshwater environments with elevated ionic concentrations contribute to calcite precipitation, and the formation of distinctive mineral-rich stromatolitic laminae. Two types of stromatolite-forming diatom populations were observed. In the first example, in stromatolites growing on a quarry ledge near Laegerdorf, North Germany, calcite crystals with biogenic imprints form around polysaccharide stalks of the diatom Gomphonema olivaceum var. calcarea (Cleve) Cleve-Euler. These individually precipitated crystals eventually become cemented together in layers, forming rigid, laminated stromatolitic deposits which drape over the quarry ledge. In the second example, in stromatolites forming in a shallow stream near Cuatro Ciénegas, Coahuila, Mexico, diatomaceous laminae also form by the accumulation of carbonate particles in a matrix of diatoms and their extracellular polysaccharide products. These laminae become thick enough to drape over individual stromatolite heads. The diatoms responsible for these deposits are Amphora aff. A. katii Selva, Nitzschia denticula Grun., and six other species. At Cuatro Ciénegas, in addition to the diatomaceous laminae, carbonate-rich cyanobacterial layers, dominated by two cyanobacterial species with different fabrics and porosities, are also present and contribute substantially to the growth of the stromatolites. In both the Laegerdorf and Cuatro Ciénegas examples, entire stromatolites or thick laminations on stromatolites are built by a small number of diatom species which produce copious amounts of extracellular stalk, gel, and sheath material, a propertuy they share with cyanobacterial stromatolite builders.     

Phylogeography, genetic structure, and gene flow in the endemic freshwater shrimp Palaemonetes suttkusi from Cuatro Ci��negas, Mexico

The valley of Cuatro Ciénegas in Mexico has the highest degree of local endemicity of any habitat in North America. A large number of endemic aquatic species occur both in the Cuatro Ciénegas basin and in the upper parts of the Río Salado de los Nadadores drainage, located immediately to the east of the valley. No natural surface flow occurs between these basins but artificial canals connect both areas. It is not clear whether the sharing of endemics between the Cuatro Ciénegas basin and the Río Salado is due to migration through canals. We conducted a phylogeographic study of mitochondrial haplotypes of the endemic shrimp Palaemonetes suttkusi to determine the evolutionary distinctiveness of the population found in Río Salado. We discovered that P. suttkusi likely has occurred naturally in both basins well into the distant past. Based on molecular clock analyses of the COI gene, the populations in the Río Salado and much of the Cuatro Ciénegas basin likely began diverging from each other between 1.9 and 11.2 Myr ago. The general levels of divergence are substantial but our results suggest there has also likely been recent gene flow between these basins. This is consistent with migration through human-made canals, but also consistent with the occurrence of natural gene flow during intermittent wet periods in the past million years. We also found significant differentiation of the Pozas Azules area from the rest of the Cuatro Ciénegas basin, a finding that is concordant with several phylogeographic studies on other aquatic endemics in Cuatro Ciénegas. We recommend that the upper parts of the Río Salado, the Pozas Azules area, and the rest of the Cuatro Ciénegas basin should each be considered independent evolutionarily significant units for conservation, and that migration of species through human-made canals should be monitored and controlled.     

Role of Fish in Structuring Invertebrates on Stromatolites in Cuatro Ciénegas, México

Stromatolites, the dominant Precambrian life form, declined in the Phanerozoic to occur today in only a few sites. This decline has been attributed to evolution of metazoan grazers, but stromatolites in our study site, Cuatro Ciénegas, Coahuila, México, harbor diverse macroinvertebrates. Drawing on food chain theory, we hypothesized that fish predation on invertebrates controls invertebrate populations, allowing stromatolites to flourish in Cuatro Ciénegas. Our experiment used small mesh (1 mm) cages to exclude all but larval fishes, and larger (6.5 mm) cages to exclude all larger fishes (including the molluscivorous and omnivorous endemic polymorphic cichlid, Herichthys minckleyi), but allow access to all sizes of the abundant endemic pupfish, Cyprinodon bifasciatus. No effects of treatments on invertebrate densities were noted at 6 week, but significant effects were observed on specific taxonomic groups after 3 month. In absence of fishes, hydrobiidae snails and ceratopogonids increased 3- and 5-fold, respectively, and invertebrate assemblage composition varied among treatments. Algal biomass was not affected by treatments, but algal species composition appeared to change. Overall results suggest that fish assemblages structure invertebrate assemblages, and that fishes may also be factors in determining algal communities.     

A new species of Mimosa (Leguminosae) from Coahuila,Mexico

Mimosaunipinnata from the Sierra de San Marcos in the Cuatro Ciénegas Basin of central Coahuila, Mexico, is described. It is unique among mimosas in having oncepinnate leaves.     

Response of grazing snails to phosphorus enrichment of modern stromatolitic microbial communities

1. The effects of added phosphorus (P) on the growth, P and RNA : DNA contents, and survivorship of snails grazing on laminated microbial mats (living ‘stromatolites’) were examined in the Rio Mesquites at Cuatro Ciénegas, Mexico (total P, c. 0.60 μmol L^?1) to test the hypothesis that strong P‐limitation of microautotroph growth produces a stoichiometric constraint on herbivores because of mineral P‐limitation. 2. In a 3‐week experiment performed in summer 2001, addition of phosphorus (+15 μmol L^?1) resulted in a strong decline in stromatolite biomass C : P ratio from very high levels (c. 2300 : 1 by atoms) to moderate levels (c. 550 : 1). The endemic hydrobiid snail Mexithauma quadripaludium responded to P‐enrichment with elevated body P content and higher RNA : DNA ratios, especially for small animals likely to be actively growing. This positive response is consistent with the existence of a stoichiometric constraint on snail growth. 3. In a longer experiment (8 weeks) involving a more moderate P enrichment (+5 μmol L^?1) in summer 2002, P enrichment reduced stromatolite C : P ratio from moderate values in control treatments (c. 750) to very low values (<100 : 1). Snails responded to stromatolite P‐enrichment with increased body P content but, in contrast to the first experiment, with lower RNA : DNA ratio, lower growth rates, and higher mortality. 4. These contrasting results suggest that both very high and very low biomass C : P ratios in stromatolites are detrimental to M. quadripaludium performance, leading us to hypothesise that these herbivores live on a ‘stoichiometric knife edge’.     

Grain trapping by filamentous cyanobacterial and algal mats: implications for stromatolite microfabrics through time

Archean and Proterozoic stromatolites are sparry or fine‐grained and finely laminated; coarse‐grained stromatolites, such as many found in modern marine systems, do not appear until quite late in the fossil record. The cause of this textural change and its relevance to understanding the evolutionary history of stromatolites is unclear. Cyanobacteria are typically considered the dominant stromatolite builders through time, but studies demonstrating the trapping and binding abilities of cyanobacterial mats are limited. With this in mind, we conducted experiments to test the grain trapping and binding capabilities of filamentous cyanobacterial mats and trapping in larger filamentous algal mats in order to better understand grain size trends in stromatolites. Mats were cut into squares, inclined in saltwater tanks at angles from 0 to 75° (approximating the angle of lamina in typical stromatolites), and grains of various sizes (fine sand, coarse sand, and fine pebbles) were delivered to their surface. Trapping of grains by the cyanobacterial mats depended strongly on (i) how far filaments protruded from the sediment surface, (ii) grain size, and (iii) the mat's incline angle. The cyanobacterial mats were much more effective at trapping fine grains beyond the abiotic slide angle than larger grains. In addition, the cyanobacterial mats actively bound grains of all sizes over time. In contrast, the much larger algal mats trapped medium and coarse grains at all angles. Our experiments suggest that (i) the presence of detrital grains beyond the abiotic slide angle can be considered a biosignature in ancient stromatolites where biogenicity is in question, and, (ii) where coarse grains are present within stromatolite laminae at angles beyond the abiotic angle of slide (e.g., most modern marine stromatolites), typical cyanobacterial‐type mats are probably not solely responsible for the construction, giving insight into the evolution of stromatolite microfabrics through time.     

MACHAERANTHERA RESTIFORMIS (ASTERACEAE) A BIZARRE NEW GYPSOPHILE FROM NORTHCENTRAL MEXICO

A rayless, diploid (n = 4), shrubby, new species of Machaeranthera is described from gypsum dunes near Cuatro Ciénegas, Coahuila. It is most closely related to M. johnstonii, another rayless, shrubby, endemic gypsophile of this region. Machaeranthera restiformis often occurs with the wide-ranging, white-rayed species, M. gypsophila (n = 4) with which it occasionally hybridizes.     

Balance between microbial calcification and metazoan bioerosion in modern stromatolitic oncolites

Stromatolites date back some 3.5 billion years and constitute the most common and conspicuous fossils through the Proterozoic. These organosedimentary structures decreased dramatically in diversity and abundance by the late Neoproterozoic, a phenomenon often ascribed to destructive grazing by newly evolved metazoans. We investigated the concurrent processes of microbial calcification and metazoan bioerosion in one of the few locations (Rio Mesquites, Cuatro Ciénegas, Coahuila, Mexico) where living freshwater stromatolites, formed by cyanobacteria and diatoms, coexist with significant populations of metazoan grazers. We used microsensor chemical profiling and monitoring of bulk water Ca²⁺ concentrations to determine calcification rates and their dependence on microbial metabolism. The bioerosive impact resulting from grazing by endemic hydrobiid gastropods was assessed by gravimetric quantification of carbonaceous faecal pellet production. Calcification was clearly light‐dependent, reaching maximal rates (saturation) at low incident light intensity, and was surprisingly efficient, with O₂/Ca²⁺ exchange ratios well above unity, and with absolute rates similar to those found in corals. However, the erosive action of grazing snails removed most of these carbonate inputs from the oncolites. Thus, a precarious balance between constructive and destructive geobiological processes was at play in the system. The fact that accretion barely exceeded bioerosion in an environment highly conducive to calcification supports the potential impact of faunal grazing as causal agent in the demise of stromatolites in the late Proterozoic. Our findings indicate that a search for fossil evidence of bioerosive grazing in the form of carbonaceous faecal pellets associated with fossil stromatolites may provide a means to test that hypothesis directly.     

Relationship between soil P fractions and microbial biomass in an oligotrophic grassland-desert scrub system

Phosphorus (P) is an essential element of the biosphere, both as a constituent of living organisms and as a regulator of biological processes. The Cuatro Ciénegas Basin in the central Chihuahuan Desert of Mexico is characterized by extreme P oligotrophy. The aim of this study was to quantify P distribution in soil P fractions, P sorption capacity, and P in microbial biomass in a desert scrub and grassland soil system in the Churince area of the Cuatro Ciénegas Basin over summer and winter seasons. Our objective, as part of an exploration of ecosystem functioning, was to ascertain the relationship between soil P fractions and P in microbial biomass. Our results demonstrate a scarcity of P, mainly in grassland, and also a higher P sorption capacity in grassland soil than in desert scrub. Desert scrub soil retained more P (228 ± 5 μg g^?1 dry soil) than grassland soil (87 ± 10 μg g^?1 dry soil), mainly in inorganic forms, but grassland soil retained more P in accessible organic forms. We suggest that biotic controls regulated by access to water shape the dynamics of soil P availability in the Churince grassland-desert scrub system.     

Hot spring siliceous stromatolites from Yellowstone National Park: assessing growth rate and laminae formation

Stromatolites are commonly interpreted as evidence of ancient microbial life, yet stromatolite morphogenesis is poorly understood. We apply radiometric tracer and dating techniques, molecular analyses and growth experiments to investigate siliceous stromatolite morphogenesis in Obsidian Pool Prime (OPP), a hot spring in Yellowstone National Park. We examine rates of stromatolite growth and the environmental and/or biologic conditions that affect lamination formation and preservation, both difficult features to constrain in ancient examples. The "main body" of the stromatolite is composed of finely laminated, porous, light-dark couplets of erect (surface normal) and reclining (surface parallel) silicified filamentous bacteria, interrupted by a less-distinct, well-cemented "drape" lamination. Results from dating studies indicate a growth rate of 1-5 cm year(-1) ; however, growth is punctuated. (14)C as a tracer demonstrates that stromatolite cyanobacterial communities fix CO(2) derived from two sources, vent water (radiocarbon dead) and the atmosphere (modern (14)C). The drape facies contained a greater proportion of atmospheric CO(2) and more robust silica cementation (vs. the main body facies), which we interpret as formation when spring level was lower. Systematic changes in lamination style are likely related to environmental forcing and larger scale features (tectonic, climatic). Although the OPP stromatolites are composed of silica and most ancient forms are carbonate, their fine lamination texture requires early lithification. Without early lithification, whether silica or carbonate, it is unlikely that a finely laminated structure representing an ancient microbial mat would be preserved. In OPP, lithification on the nearly diurnal time scale is likely related to temperature control on silica solubility.     

Cyanobacterial construction of hot spring siliceous stromatolites in Yellowstone National Park

Living stromatolites growing in a hot spring in Yellowstone National Park are composed of silica-encrusted cyanobacterial mats. Two cyanobacterial mat types grow on the stromatolite surfaces and are preserved as two distinct lithofacies. One mat is present when the stromatolites are submerged or at the water-atmosphere interface and the other when stromatolites protrude from the hot spring. The lithofacies created by the encrustation of submerged mats constitutes the bulk of the stromatolites, is comprised of silica-encrusted filaments, and is distinctly laminated. To better understand the cyanobacterial membership and community structure differences between the mats, we collected mat samples from each type. Molecular methods revealed that submerged mat cyanobacteria were predominantly one novel phylotype while the exposed mats were predominantly heterocystous phylotypes (Chlorogloeopsis HTF and Fischerella). The cyanobacterium dominating the submerged mat type does not belong in any of the subphylum groups of cyanobacteria recognized by the Ribosomal Database Project and has also been found in association with travertine stromatolites in a Southwest Japan hot spring. Cyanobacterial membership profiles indicate that the heterocystous phylotypes are 'rare biosphere' members of the submerged mats. The heterocystous phylotypes likely emerge when the water level of the hot spring drops. Environmental pressures tied to water level such as sulfide exposure and possibly oxygen tension may inhibit the heterocystous types in submerged mats. These living stromatolites are finely laminated and therefore, in texture, may better represent similarly laminated ancient forms compared with more coarsely laminated living marine examples.     

JET‐SUSPENDED,CALCITE‐BALLASTED CYANOBACTERIAL WATERWARTS IN A DESERT SPRING1

We describe a population of colonial cyanobacteria (waterwarts) that develops as the dominant primary producer in a bottom‐fed, O₂‐poor, warm spring in the Cuatro Ciénegas karstic region of the Mexican Chihuahuan Desert. The centimeter‐sized waterwarts were suspended within a central, conically shaped, 6‐m deep well by upwelling waters. Waterwarts were built by an Aphanothece‐like unicellular cyanobacterium and supported a community of epiphytic filamentous cyanobacteria and diatoms but were free of heterotrophic bacteria inside. Sequence analysis of 16S rRNA genes revealed that this cyanobacterium is only distantly related to several strains of other unicellular cyanobacteria (Merismopedia, Cyanothece, Microcystis). Waterwarts contained orderly arrangements of mineral crystallites, made up of microcrystalline low‐magnesium calcite with high levels of strontium and sulfur. Waterwarts were 95.9% (v/v) glycan, 2.8% cells, and 1.3% mineral grains and had a buoyant density of 1.034 kg·L^?1. An analysis of the hydrological properties of the spring well and the waterwarts demonstrated that both large colony size and the presence of controlled amounts of mineral ballast are required to prevent the population from being washed out of the well. The unique hydrological characteristics of the spring have likely selected for both traits. The mechanisms by which controlled nucleation of extracellular calcite is achieved remain to be explored.     

Involvement of cyclodipeptides in the competition of bacterial communities in the oligotrophic Churince aquatic system of Cuatro Ciénegas Basin dominated by Gammaproteobacteria

The Cuatro Ciénegas Basin (CCB) within the Chihuahuan Desert in México is an extremely oligotrophic oasis with negligible phosphorous levels, described as a hot spot of biodiversity, not only in stromatolites and microbial mats, but also in living forms in general. The microorganisms possess the capability to produce a wide variety of virulence factors, antibiotics, and quorum-sensing (QS) crosstalk signals such as non-ribosomal cyclodipeptides (CDPs) which enables them to colonize diverse ecological niches. In the aquatic system of CCB known as Churince, a bacterial population was isolated from the Lagunita pond dominated by Gammaproteobacteria. In this work, we determined the relationships between the antagonism and CDPs production in this bacterial population. Results indicate that 68% of isolates showed antagonistic effects over other isolates, correlating with production of CDPs and the antibiotic 2,4-diacetylphloroglucinol (DAPG). Although a minority of the isolates were capable of inducing a QS biosensor strain, bacterial QS interference was not the main mechanism in the antagonism observed. Thus, our results indicate that CDPs primarily, and DAPG to a lesser degree, are involved with the growth-inhibition competition mechanisms of bacterial communities in the Lagunita pond and was associated with a Gammaproteobacteria dominancy phenomena.

     

Predation drives morphological convergence in the Gambusia panuco species group among lotic and lentic habitats

Fish morphology is often constrained by a trade‐off between optimizing steady vs. unsteady swimming performance due to opposing effects of caudal peduncle size. Lotic environments tend to select for steady swimming performance, leading to smaller caudal peduncles, whereas predators tend to select for unsteady swimming performance, leading to larger caudal peduncles. However, it is unclear which aspect of performance should be optimized across heterogeneous flow and predation environments and how this heterogeneity may affect parallel phenotypic evolution. We investigated this question among four Gambusia species in north‐eastern Mexico, specifically the riverine G. panuco, the spring endemics G. alvarezi and G. hurtadoi, and a fourth species, G. marshi, found in a variety of habitats with varying predation pressure in the Cuatro Ciénegas Basin and Río Salado de Nadadores. We employed a geometric morphometric analysis to examine how body shapes of both male and female fish differ among species and habitats and with piscivore presence. We found that high‐predation and low‐predation species diverged morphologically, with G. marshi exhibiting a variable, intermediate body shape. Within G. marshi, body morphology converged in high‐predation environments regardless of flow velocity, and fish from high‐predation sites had larger relative caudal peduncle areas. However, we found that G. marshi from low‐predation environments diverged in morphology between sub‐basins of Cuatro Ciénegas, indicating other differences among these basins that merit further study. Our results suggest that a morphological trade‐off promotes parallel evolution of body shape in fishes colonizing high‐predation environments and that changing predation pressure can strongly impact morphological evolution in these species.     

Modern marine stromatolites in the Exuma Cays,Bahamas: Uncommonly common

Summary Modern stromatolites in open marine environments, unknown until recently, are common throughout the Exuma Cays, Bahamas. They occur in three distinct settings: subtidal tidal passes, subtidal sandy embayments and intertidal beaches. These stromatolites have a relief of up to 2.5 m and occur in water depths ranging from intertidal to 10 m. Surfaces near the sediment-water interface are typically colonized by cyanobacterial mats, whereas high relief surfaces are commonly colonized by algal turf and other macroalgae such asBatophora, Acetabularia, andSargassum. The internal structure of the stromatolites is characterized by millimeter-scale lamination defined by differential lithification of agglutinated sediment. In thin section, the lithified laminae appear as micritic horizons with distinct microstructures: they consist of thin micritic crusts (20–40 μm thick) overlying layers of micritized sediment grains (200–1000 μm thick); the micritized grains are cemented at point-contacts and are trucated along a surface of intense microboring. The Exuma stromatolites are built by cyanobacterial-dominated communities. These laminated prokaryotic structures grade to unlayered thrombolites built by eukaryotic algae. The variety of sites, settings and shapes of stromatolites in the Exuma Cays present excellent opportunities for future studies of stromatolite morphogenesis.     

Growth of modern branched columnar stromatolites in Lake Joyce,Antarctica

Modern decimeter‐scale columnar stromatolites from Lake Joyce, Antarctica, show a change in branching pattern during a period of lake level rise. Branching patterns correspond to a change in cyanobacterial community composition as preserved in authigenic calcite crystals. The transition in stromatolite morphology is preserved by mineralized layers that contain microfossils and cylindrical molds of cyanobacterial filaments. The molds are composed of two populations with different diameters. Large diameter molds (>2.8 μm) are abundant in calcite forming the oldest stromatolite layers, but are absent from younger layers. In contrast, <2.3 μm diameter molds are common in all stromatolites layers. Loss of large diameter molds corresponds to the transition from smooth‐sided stromatolitic columns to branched and irregular columns. Mold diameters are similar to trichome diameters of the four most abundant living cyanobacteria morphotypes in Lake Joyce: Phormidium autumnale morphotypes have trichome diameters >3.5   μm, whereas Leptolyngbya antarctica, L. fragilis, and Pseudanabaena frigida morphotypes have diameters <2.3   μm. P. autumnale morphotypes were only common in mats at <12 m depth. Mats containing abundant P. autumnale morphotypes were smooth, whereas mats with few P. autumnale morphotypes contained small peaks and protruding bundles of filaments, suggesting that the absence of P. autumnale morphotypes allowed small‐scale topography to develop on mats. Comparisons of living filaments and mold diameters suggest that P. autumnale morphotypes were present early in stromatolite growth, but disappeared from the community through time. We hypothesize that the mat‐smoothing behavior of P. autumnale morphotypes inhibited nucleation of stromatolite branches. When P. autumnale morphotypes were excluded from the community, potentially reflecting a rise in lake level, short‐wavelength roughness provided nuclei for stromatolite branches. This growth history provides a conceptual model for initiation of branched stromatolite growth resulting from a change in microbial community composition.     

Diversity of culturable thermo-resistant aquatic bacteria along an environmental gradient in Cuatro Ciénegas,Coahuila, México

At the desert oasis of Cuatro Ciénegas in Coahuila, México, more than 300 oligotrophic pools can be found and a large number of endemic species of plants and animals. The most divergent taxa of diatoms, snail and fishes are located in the Churince hydrological system, where we analyzed the local diversification of cultivable Firmicutes and Actinobacteria. The Churince hydrological system is surrounded by gypsum dunes and has a strong gradient for salinity, temperature, pH and dissolved oxygen. In August 2003, surface water samples were taken in 10 sites along the Churince system together with the respective environmental measurements. 417 thermo-resistant bacteria were isolated and DNA was extracted to obtain their BOX-PCR fingerprints, revealing 55 different patterns. In order to identify similarities and differences in the diversity of the various sampling sites, an Ordination Analysis was applied using Principal Component Analysis. This analysis showed that conductivity is the environmental factor that explains the distribution of most of the microbial diversity. Phylogenetic reconstruction from their 16S rRNA sequences was performed for a sample of 150 isolates. Only 17 sequences had a 100% match in the Gene Bank (NCBI), representing 10 well known cosmopolitan taxa. The rest of the sequences cluster in 22 clades for Firmicutes and another 22 clades for Actinobacteria, supporting the idea of high diversity and differentiation for this site.     

Relationships between spatio-temporal environmental and genetic variation reveal an important influence of exogenous selection in a pupfish hybrid zone

The importance of exogenous selection in a natural hybrid zone between the pupfishes Cyprinodon atrorus and Cyprinodon bifasciatus was tested via spatio‐temporal analyses of environmental and genetic change over winter, spring and summer for three consecutive years. A critical influence of exogenous selection on hybrid zone regulation was demonstrated by a significant relationship between environmental (salinity and temperature) and genetic (three diagnostic nuDNA loci) variation over space and time (seasons) in the Rio Churince system, Cuatro Ciénegas, Mexico. At sites environmentally more similar to parental habitats, the genetic composition of hybrids was stable and similar to the resident parental species, whereas complex admixtures of parental and hybrid genotypic classes characterized intermediate environments, as did the greatest change in allelic and genotypic frequencies across seasons. Within hybrids across the entire Rio Churince system, seasonal changes in allelic and genotypic frequencies were consistent with results from previous reciprocal transplant experiments, which showed C. bifasciatus to suffer high mortality (75%) when exposed to the habitat of C. atrorus in winter (extreme temperature lows and variability) and summer (abrupt salinity change and extreme temperature highs and variability). Although unconfirmed, the distributional limits of C. atrorus and C. atrorus‐like hybrids appear to be governed by similar constraints (predation or competition). The argument favouring evolutionary significance of hybridization in animals is bolstered by the results of this study, which links the importance of exogenous selection in a contemporary hybrid zone between C. atrorus and C. bifasciatus to previous demonstration of the long‐term evolutionary significance of environmental variation and introgression on the phenotypic diversification Cuatro Ciénegas Cyprinodon.     

Introgressive hybridization in a trophically polymorphic cichlid

Trophically polymorphic species could represent lineages that are rapidly diverging along an ecological axis or could phenotypically mark the collapse of species through introgressive hybridization. We investigated patterns of introgression between the trophically polymorphic cichlid fish Herichthys minckleyi and its relative H. cyanoguttatus using a combination of population genetics and species tree analyses. We first examined the distribution of mitochondrial haplotypes within the alternative H. minckleyi pharyngeal jaw morphotypes that are endemic to the small desert valley of Cuatro Ciénegas. We recovered two clusters of mitochondrial haplotypes. The first contained a number of slightly differentiated cytochrome b (cytb) haplotypes that showed some phylogeographic signal and were present in both jaw morphotypes. The other haplotype was monomorphic, highly differentiated from the other cluster, present in equal frequencies in the morphotypes, and identical to H. cyanoguttatus haplotypes found outside Cuatro Ciénegas. Then, we investigated whether H. minckleyi individuals with the H. cyanoguttatus cytb were more evolutionarily similar to H. cyanoguttatus or other H. minckleyi using a species tree analysis of 84 nuclear loci. Both H. minckleyi pharyngeal morphotypes, regardless of their cytb haplotype, were quite distinct from H. cyanoguttatus. However, hybridization could be blurring subdivision within H. minckleyi as the alternative jaw morphotypes were not genetically distinct from one another. Accounting for introgression from H. cyanoguttatus will be essential to understand the evolution of the trophically polymorphic cichlid H. minckleyi.     

Environmental variation,hybridization, and phenotypic diversification in Cuatro Ciénegas pupfishes

Hybridization can generate novel phenotypes, and in combination with divergent selection along environmental gradients, can play a driving role in phenotypic diversification. This study examined the influence of introgressive hybridization and environmental variation on the phenotypic diversity of two pupfish species (Cyprinodon atrorus and Cyprinodon bifasciatus) endemic to the Cuatro Ciénegas basin, Mexico. These species occupy opposite environmental extremes and are comprised of multiple, intraspecifically isolated populations. However, interspecific hybridization occurs to various degrees within connecting, intermediate environments. Using geometric morphometric analysis, extensive variation of body shape was observed between and within species, and phenotypic variation was strongly correlated with environmental conditions. Furthermore, some introgressed populations exhibited unique phenotypes not found in either of the parents, and overall morphospace occupation was significantly higher in introgressed populations when compared to the parentals. Overall, we find environmental variation and transgressive segregation both appear to have been important in shaping phenotypic variation in this system.