A link between host dispersal and parasite diversity in two sympatric cichlids of Lake Tanganyika

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Increased genetic divergence between two closely related fir species in areas of range overlap

Because of introgressive hybridization, closely related species can be more similar to each other in areas of range overlap (parapatry or sympatry) than in areas where they are geographically isolated from each other (allopatry). Here, we report the reverse situation based on nuclear genetic divergence between two fir species, Abies chensiensis and Abies fargesii, in China, at sites where they are parapatric relative to where they are allopatric. We examined genetic divergence across 126 amplified fragment length polymorphism (AFLP) markers in a set of 172 individuals sampled from both allopatric and parapatric populations of the two species. Our analyses demonstrated that AFLP divergence was much greater between the species when comparisons were made between parapatric populations than between allopatric populations. We suggest that selection in parapatry may have largely contributed to this increased divergence.     

Behavioral modifications lead to disparate demographic consequences in two sympatric species

Life‐history theory suggests species that typically have a large number of offspring and high adult mortality may make decisions that benefit offspring survival in exchange for increased adult risks. Such behavioral adaptations are essential to understanding how demographic performance is linked to habitat selection during this important life‐history stage. Though studies have illustrated negative fitness consequences to attendant adults or potential fitness benefits to associated offspring because of adaptive habitat selection during brood rearing, equivocal relationships could arise if both aspects of this reproductive trade‐off are not assessed simultaneously. To better understand how adaptive habitat selection during brood rearing influences demographics, we studied the brood survival, attendant parental survival, and space use of two sympatric ground‐nesting bird species, the northern bobwhite (hereafter: “bobwhite”; Colinus virgininanus) and scaled quail (Callipepla squamata). During the 2013–2014 breeding seasons, we estimated habitat suitability across two grains (2 m and 30 m) for both species and determined how adult space use of these areas influenced individual chick survival and parental risk. We found the proportion of a brood's home range containing highly suitable areas significantly increased bobwhite chick survival (β = 0.02, SE = 0.006). Additionally, adult weekly survival for bobwhite was greater for individuals not actively brooding offspring (0.9716, SE = 0.0054) as compared to brooding adults (0.8928, SE = 0.0006). Conversely, brood habitat suitability did not influence scaled quail chick survival during our study, nor did we detect a survival cost for adults that were actively brooding offspring. Our research illustrates the importance of understanding life‐history strategies and how they might influence relationships between adaptive habitat selection and demographic parameters.     

Niche divergence facilitated by fine‐scale ecological partitioning in a recent cichlid fish adaptive radiation

Ecomorphological differentiation is a key feature of adaptive radiations, with a general trend for specialization and niche expansion following divergence. Ecological opportunity afforded by invasion of a new habitat is thought to act as an ecological release, facilitating divergence, and speciation. Here, we investigate trophic adaptive morphology and ecology of an endemic clade of oreochromine cichlid fishes (Alcolapia) that radiated along a herbivorous trophic axis following colonization of an isolated lacustrine environment, and demonstrate phenotype‐environment correlation. Ecological and morphological divergence of the Alcolapia species flock are examined in a phylogenomic context, to infer ecological niche occupation within the radiation. Species divergence is observed in both ecology and morphology, supporting the importance of ecological speciation within the radiation. Comparison with an outgroup taxon reveals large‐scale ecomorphological divergence but shallow genomic differentiation within the Alcolapia adaptive radiation. Ancestral morphological reconstruction suggests lake colonization by a generalist oreochromine phenotype that diverged in Lake Natron to varied herbivorous morphologies akin to specialist herbivores in Lakes Tanganyika and Malawi.     

Habitat-associated morphological divergence in two Neotropical fish species

We examined intraspecific morphological diversification between river channel and lagoon habitats for two Neotropical fish ( Bryconops caudomaculatus , Characidae; Biotodoma wavrini , Cichlidae). We hypothesized that differences between habitats (e.g. flow regime, foraging opportunities) might create selective pressures resulting in morphological divergence between conspecific populations. We collected fish from four channel-lagoon habitat pairs in the Río Cinaruco, Venezuela, and compared body morphology using geometric morphometrics. There were two aspects of divergence in both species: (1) placement of maximum body depth and (2) orientation of the mouth. For both species, maximum body depth was positioned more anteriorly (i.e. fusiform) in the river channel than in lagoons. Both species exhibited a relatively terminal mouth in lagoons compared to the channel. The mouth of B. caudomaculatus was relatively upturned, whereas the mouth of B. wavrini was relatively subterminal, in channel habitats. Observed morphological patterns are consistent with functional morphological principles suggesting adaptive divergence. We also show that spatial distance between habitats, presumably reflecting rates of population mixing, appears to have constrained diversification. For both species, morphological divergence increased with distance between habitats. Thus morphological divergence between channel and lagoon habitats apparently reflects a balance between diversification driven by natural selection, and homogenization driven by population mixing.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 689–698.     

Trophic dynamics within a hybrid zone – interactions between an abundant cyprinid hybrid and sympatric parental species

1. Recent proliferation of hybridisation in response to anthropogenic ecosystem change, coupled with increasing evidence of the importance of ancient hybridisation events in the formation of many species, has moved hybridisation to the forefront of evolutionary theory. 2. In spite of this, the mechanisms (e.g. differences in trophic ecology) by which hybrids co‐exist with parental taxa are poorly understood. A unique hybrid zone exists in Irish freshwater systems, whereby hybrid offspring off two non‐native cyprinid fishes often outnumber both parental species. 3. Using stable isotope and gut content analyses, we determined the trophic interactions between sympatric populations of roach (Rutilus rutilus), bream (Abramis brama) and their hybrid in lacustrine habitats. 4. The diet of all three groups displayed little variation across the study systems, and dietary overlap was observed between both parental species and hybrids. Hybrids displayed diet, niche breadth and trophic position that were intermediate between the two parental species while also exhibiting greater flexibility in diet across systems.     

Deep sympatric mtDNA divergence in the autumnal moth (Epirrita autumnata)

Deep sympatric intraspecific divergence in mtDNA may reflect cryptic species or formerly distinct lineages in the process of remerging. Preliminary results from DNA barcoding of Scandinavian butterflies and moths showed high intraspecific sequence variation in the autumnal moth, Epirrita autumnata. In this study, specimens from different localities in Norway and some samples from Finland and Scotland, with two congeneric species as outgroups, were sequenced with mitochondrial and nuclear markers to resolve the discrepancy found between mtDNA divergence and present species‐level taxonomy. We found five COI sub‐clades within the E. autumnata complex, most of which were sympatric and with little geographic structure. Nuclear markers (ITS2 and Wingless) showed little variation and gave no indications that E. autumnata comprises more than one species. The samples were screened with primers for Wolbachia outer surface gene (wsp) and 12% of the samples tested positive. Two Wolbachia strains were associated with different mtDNA sub‐clades within E. autumnata, which may indicate indirect selection/selective sweeps on haplotypes. Our results demonstrate that deep mtDNA divergences are not synonymous with cryptic speciation and this has important implications for the use of mtDNA in species delimitation, like in DNA barcoding.     

Shared dietary niche between sexes in Magellanic Penguins

Prey use by male and female Magellanic Penguins (Spheniscus magellanicus) was quantified during chick rearing in northern Golfo San Jorge, Argentina and the dietary niches and diet composition of each sex were assessed and compared. A total of 67 males and 45 females were flushed during the chick stages of 2011 and 2012. Argentine Anchovy (Engraulis anchoita) was the main prey in terms of importance by mass in males and females in all stages and years (52.4–86.6% and 83.8–88.7%, respectively), except during the old chick stage in 2011 when the main prey in males and females was the Shortfin Squid (Illex argentinus) (63.1% and 44.4%, respectively). Analyses of similarity showed that diet composition in terms of contribution by mass of the main prey species was similar between sexes. Based on the isotopic values of δ¹³C and δ¹⁵N corresponding to 38 males and 21 females, the Bayesian mixing model outputs showed that Argentine Anchovy was also the main prey. In both years and chick stages, the isotopic niche of males and females was similar. The similarity in the dietary niche between male and female Magellanic Penguins raising chicks may be due to the high availability of Argentine Anchovy during the study period and the consequent dominance of this forage species in the diet of both sexes.     

Delimitation of sympatric Palaemon (Decapoda,Palaemonidae) species of the Laguna Madre,Mexico

Grass shrimps Palaemon mundusnovus, P. pugio and P. vulgaris sympatrically inhabit in the Laguna Madre, Mexico. They exhibit very close morphological similarity and overlap in their diagnostic characteristics, which has hindered certainty in their identification and has raised doubt regarding their taxonomic validity. In this work, we analyse intra‐ and interspecific morphometric and meristic variability through a multivariate analysis, and we determine the molecular variation using mitochondrial sequences of the 12S, 16S and COX1 genes to confirm the validity of the three taxa as having distinct lineages or to recognize a smaller number of species with phenotypic plasticity. Our results corroborate the taxonomic validity of the three species; however, there is intraspecific plasticity, interspecific overlap of characteristics and greater morphological and molecular similarity between the species P. mundusnovus and P. pugio, whereas P. vulgaris was better delimited. These species form a monophyletic group but the phylogenetic relationship obtained is discussed. Telson length was the primary variable in the principal components analysis, whereas the length of the propodus of the second pereiopod was the best discriminant. The range of variation reported in the characteristics linked to the rostrum is extended, and its influence on the separation of these species is recognized. The joint application of multivariate analysis from morphological variables and molecular tools is recommended to clarify the taxonomic status of species featuring close morphological similarity and sympatric distribution.     

Pollen competition between two sympatric Orchis species (Orchidaceae): the overtaking of conspecific of heterospecific pollen as a reproductive barrier

The frequency of hybrid formation in angiosperms depends on how and when heterospecific pollen is transferred to the stigma, and on the success of that heterospecific pollen at fertilising ovules. We applied pollen mixtures to stigmas to determine how pollen interactions affect siring success and the frequency of hybrid formation between two species of Mediterranean deceptive orchid. Plants of Orchis italica and O. anthropophora were pollinated with conspecific and heterospecific pollen (first conspecific pollen then heterospecific pollen and vice versa) and molecular analysis was used to check the paternity of the seeds produced. In this pair of Mediterranean orchids, competition between conspecific and heterospecific pollen functions as a post‐pollination pre‐zygotic barrier limiting the frequency of the formation of hybrids in nature. Flowers pollinated with heterospecific pollen can remain receptive for the arrival of conspecific pollen for a long time. There is always an advantage of conspecific pollen for fruit formation, whether it comes before or after heterospecific pollen, because it overtakes the heterospecific pollen. The conspecific pollen advantage exhibited in O. italica and O. anthropophora is likely to result from the reduced germination of heterospecific pollen or retarded growth of heterospecific pollen tubes in the stigma and ovary. Overall, the results indicate that our hybrid zone represents a phenomenon of little evolutionary consequence, and the conspecific pollen advantage maintains the genetic integrity of the parental species.     

Aging alters interspecific competition between two sympatric insect–parasitic nematode species

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Trophic relationships between the larvae of two freshwater mussels and their fish hosts

Freshwater mussels of the order Unionoida have life cycles that include larval attachment to and later metamorphosis on suitable host fishes. Information on the trophic relationship between unionoid larvae and their host fishes is scarce. We investigated the trophic interaction between fish hosts and encysted larvae of two species of freshwater mussels, Margaritifera margaritifera and Unio crassus, using stable isotope analyses of larvae and juvenile mussels as well as of host fish gill and muscle tissues before and after infestation. Due to different life histories and durations of host‐encystment, mass and size increase in M. margaritifera during the host‐dependent phase were greater than those of U. crassus. δ¹³C and δ¹⁵N signatures of juvenile mussels approached isotopic signatures of fish tissues, indicating a parasitic relationship between mussels and their hosts. Shifts were more pronounced for M. margaritifera, which had a five‐fold longer host‐dependent phase than U. crassus. The results of this study suggest that stable isotope analyses are a valuable tool for characterizing trophic relationships and life history strategies in host–parasite systems. In the case of unionoid mussels, stable isotopic shifts of the larvae are indicative of the nutritional versus phoretic importance of the host.     

Habitat predictors of genetic diversity for two sympatric wetland‐breeding amphibian species

Population genetic diversity is widely accepted as important to the conservation and management of wildlife. However, habitat features may differentially affect evolutionary processes that facilitate population genetic diversity among sympatric species. We measured genetic diversity for two pond‐breeding amphibian species (Dwarf salamanders, Eurycea quadridigitata; and Southern Leopard frogs, Lithobates sphenocephalus) to understand how habitat characteristics and spatial scale affect genetic diversity across a landscape. Samples were collected from wetlands on a longleaf pine reserve in Georgia. We genotyped microsatellite loci for both species to assess population structures and determine which habitat features were most closely associated with observed heterozygosity and rarefied allelic richness. Both species exhibited significant population genetic structure; however, structure in Southern Leopard frogs was driven primarily by one outlier site. Dwarf salamander allelic richness was greater at sites with less surrounding road area within 0.5 km and more wetland area within 1.0 and 2.5 km, and heterozygosity was greater at sites with more wetland area within 0.5 km. In contrast, neither measure of Southern Leopard frog genetic diversity was associated with any habitat features at any scale we evaluated. Genetic diversity in the Dwarf salamander was strongly associated with land cover variables up to 2.5 km away from breeding wetlands, and/or results suggest that minimizing roads in wetland buffers may be beneficial to the maintenance of population genetic diversity. This study suggests that patterns of genetic differentiation and genetic diversity have associations with different habitat features across different spatial scales for two syntopic pond‐breeding amphibian species.     

Population genetics and sympatric divergence of the freshwater gudgeon,Gobiobotia filifer,in the Yangtze River inferred from mitochondrial DNA

The ecosystem and Pleistocene glaciations play important roles in population demography. The freshwater gudgeon, Gobiobotia filifer, is an endemic benthic fish in the Yangtze River and is a good model for ecological and evolutionary studies. This study aimed to decode the population structure of G. filifer in the Yangtze River and reveal whether divergence occurred before or after population radiation. A total of 292 specimens from eight locations in the upper and middle reaches of the Yangtze River were collected from 2014 to 2016 and analyzed via mitochondrial DNA Cyt b gene sequencing. A moderately high level of genetic diversity was found without structures among the population. However, phylogenetic and network topology showed two distinct haplotype groups, and each group contained a similar proportion of individuals from all sampled sites. This suggested the existence of two genetically divergent source populations in G. filifer. We deduced that a secondary contact of distinct glacial refugia was the main factor creating sympatric populations of G. filifer, and climate improvement promoted population expansion and colonization.     

Differences in seasonal survival suggest species‐specific reactions to climate change in two sympatric bat species

Long‐lived animals with a low annual reproductive output need a long time to recover from population crashes and are, thus, likely to face high extinction risk, if the current global environmental change will increase mortality rates. To aid conservation of those species, knowledge on the variability of mortality rates is essential. Unfortunately, however, individual‐based multiyear data sets that are required for that have only rarely been collected for free‐ranging long‐lived mammals. Here, we used a five‐year data set comprising activity data of 1,445 RFID‐tagged individuals of two long‐lived temperate zone bat species, Natterer's bats (Myotis nattereri) and Daubenton's bats (Myotis daubentonii), at their joint hibernaculum. Both species are listed as being of high conservation interest by the European Habitats Directive. Applying mixed‐effects logistic regression, we explored seasonal survival differences in these two species which differ in foraging strategy and phenology. In both species, survival over the first winter of an individual's life was much lower than survival over subsequent winters. Focussing on adults only, seasonal survival patterns were largely consistent with higher winter and lower summer survival but varied in its level across years in both species. Our analyses, furthermore, highlight the importance of species‐specific time periods for survival. Daubenton's bats showed a much stronger difference in survival between the two seasons than Natterer's bats. In one exceptional winter, the population of Natterer's bats crashed, while the survival of Daubenton's bats declined only moderately. While our results confirm the general seasonal survival pattern typical for hibernating mammals with higher winter than summer survival, they also show that this pattern can be reversed under particular conditions. Overall, our study points toward a high importance of specific time periods for population dynamics and suggests species‐, population‐, and age class‐specific responses to global climate change.     

Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Trophic level and overlap of sea lions (Zalophus californianus) in the Gulf of California, Mexico

Stable isotope and scat analyses were used in concert to determine trophic level and dietary overlap among California sea lions from different rookeries in the Gulf of California. Isotopic analysis of the fur of sea lion pups revealed differences in δ¹⁵N and δ¹³C values among rookeries during the breeding season. Mean δ¹⁵N and δ¹³C values varied from 20.2‰ to 22.4‰ and from −15.4‰ to −14.0‰, respectively. The pattern of differences among rookeries was similar between years in most cases. Isotopic variations among rookeries were associated with differences in prey consumption. There was a significant correlation between δ¹⁵N value and trophic level, as determined by scat analysis. Joint application of isotopic and scat analyses allowed us to identify how the feeding habits of sea lions vary with location. Our results suggest the presence of spatial structure in available prey as well as the localized use of prey by sea lions across the Gulf of California.     

Genetic divergence between the sympatric queen morphs of the ant Myrmica rubra

Pairs of obligate social parasites and their hosts, where some of the parasites have recently diverged from their host through intraspecific social parasitism, provide intriguing systems for studying the modes and processes of speciation. Such speciation, probably in sympatry, has also been propounded in the ant Myrmica rubra and its intraspecific social parasite. In this species, parasitism is associated with queen size dimorphism, and the small microgyne has become a social parasite of the large macrogyne. Here, we investigated the genetic divergence of the host and the parasite queen morphs in 11 localities in southern Finland, using nuclear and mitochondrial markers of queens and workers. We formulated and tested four speciation‐related hypotheses that differed in the degree of genetic divergence between the morphs. The queen morphs were genetically distinct from each other with little hybridization. In the nuclear data, when localities were nested within queen morphs in the hierarchical amova , 39% of the genetic variation was explained by the queen morph (standardized F'_CT = 0.63, uncorrected F_CT = 0.39), whereas 18% was explained by the locality (F'_SC = 0.39, F_SC = 0.29). This result corroborated the hypothesis of advanced sympatric speciation. In contrast, the mitochondrial DNA could not settle between the hierarchical levels of locality and queen morph, thus substantiating equally the hypotheses of incipient and advanced sympatric speciation. Together, our results support the view that the microgynous parasite has genetically diverged from its macrogynous host to the level of a nascent species.     

Dietary specialization and the effects of plant species on potential multitrophic interactions of three species of nymphaline caterpillars

The diet breadth of insect herbivores influences their response to variation in plant quality, and these bitrophic interactions have implications for the higher‐level trophic interactions between herbivores and their natural enemies. In this comparative study, we examined the role of host plant species and plant secondary chemistry on the potential interactions between three species of nymphaline caterpillars and their natural enemies. The caterpillar species (all Lepidoptera: Nymphalidae) varied in their degree of specialization: the buckeye, Junonia coenia Hübner, is a specialist on plants that contain iridoid glycosides (IGs); the white peacock, Anartia jatrophae L., feeds on plants in five families, some of which contain IGs and some of which do not; and the painted lady, Vanessa cardui L., is a generalist, feeding on plants in at least 15 families. Each species was reared on leaves of an introduced host plant, Plantago lanceolata L. (Plantaginaceae), which produces two IGs, aucubin and catalpol, and on another plant species that is a common host plant. These alternate host plants were Plantago major L. (Plantaginaceae) for J. coenia, Bacopa monnieri (L.) Pennell (Plantaginaceae) for A. jatrophae, and Malva parviflora L. (Malvaceae) for V. cardui. We examined growth, sequestration, and immune response of these caterpillars on the different host plant species. Junonia coenia developed more rapidly and sequestered higher IG concentrations when reared on P. lanceolata, whereas both other species grew more slowly on P. lanceolata. Host plant did not influence immune response of J. coenia or A. jatrophae, whereas V. cardui immune response was weaker when reared on P. lanceolata. Junonia coenia was most efficient at IG sequestration and A. jatrophae was least efficient, when all three species were reared on P. lanceolata. These results indicate that diet breadth may play an important role in structuring tritrophic interactions, and this role should be further explored.