Interspecific Studies Research Articles

Compensatory traits and the evolution of male ornaments

How ornaments that are used during mate choice and rival assessment remain reliable has been a source of contention for many years. Such signals are hypothesized to be costly (i.e., 'handicaps'), but empirical studies testing for costs of sexually selected ornaments are equivocal at best, contradictory at worst. We review recent studies finding compensation for sexually selected ornaments, in both intra- and inter-specific studies, suggesting that other traits evolve to mitigate costs of ornaments. We synthesize these studies to elucidate the role of compensatory traits in the evolution of reliable ornaments and explain how selection to reduce ornament costs may influence aspects of the phenotype that are not subjected to direct sexual selection and may obscure our ability to directly measure ornament costs. Both intraspecific studies and comparative studies in a phylogenetic framework are important for our understanding of how the costs of signals may be reduced by compensation, but each approach answers different questions about ornament evolution. We also elaborate on a general theoretical model that can be useful when testing for costs of ornaments in correlational and experimental studies. We recommend that future investigators should consider compensatory traits when testing for ornament costs, especially when manipulating ornamentation.

On the intraspecific variability in basal metabolism and the food habits hypothesis in birds

Abstract The food habits hypothesis (FHH) stands as one of the most striking and often-cited interspecific patterns to emerge from comparative studies of endothermic energetics. The FHH identifies three components of diet that potentially produce variability in mass-independent BMR, i.e. food quality, food availability, and food predictability or environmental productivity. The hypothesis predicts that species with diets of low energy content and/or low digestibility should evolve low mass-independent BMRs. The effects of food habits on BMR have been widely investigated at the interspecific level, but the variation between individuals and populations has been largely ignored. Our focus is to compare predictions derived from interspecific studies with data collected from within-species studies to explore the mechanisms and functional significance of adaptive responses predicted by the food-habits hypothesis among birds. We conclude that if BMR is correlated with daily energy expenditure, then organisms that can lower BMR will reduce daily energy expenditure and hence, food requirements. Birds that lower BMR in stressful environments may increase survival. Nevertheless, the mechanism (s) by which birds eating a low quality diet reduce BMR and whether lower BMR affects fitness remain to be determined.

Intraspecific trait variation and covariation in a widespread tree species (Nothofagus pumilio) in southern Chile

• The focus of the trait-based approach to study community ecology has mostly been on trait comparisons at the interspecific level. Here we quantified intraspecific variation and covariation of leaf mass per area (LMA) and wood density (WD) in monospecific forests of the widespread tree species Nothofagus pumilio to determine its magnitude and whether it is related to environmental conditions and ontogeny. We also discuss probable mechanisms controlling the trait variation found. • We collected leaf and stem woody tissues from 30-50 trees of different ages (ontogeny) from each of four populations at differing elevations (i.e. temperatures) and placed at each of three locations differing in soil moisture. • The total variation in LMA (coefficient of variation (CV) = 21.14%) was twice that of WD (CV = 10.52%). The total variation in traits was never less than 23% when compared with interspecific studies. Differences in elevation (temperature) for the most part explained variation in LMA, while differences in soil moisture and ontogeny explained the variation in WD. Traits covaried similarly in the altitudinal gradient only. • Functional traits of N. pumilio exhibited nonnegligible variation; LMA varied for the most part with temperature, while WD mostly varied with moisture and ontogeny. We demonstrate that environmental variation can cause important trait variation without species turnover.

Is intraspecific variation in diet and morphology related to environmental gradients? Exploring Liem's paradox in a cichlid fish

Interspecific studies have demonstrated that trophic morphology and ecology are not always tightly matched: a phenomenon rarely reported at the intraspecific level. In the present study, we explored relationships among diet, morphology and the environment in a widespread cichlid fish, Astatoreochromis alluaudi (Pellegrin 1904), from 6 sites in southern Uganda to test for evidence of eco-morphological matching at the interdemic level. Previous studies of Astatoreochromis alluaudi have demonstrated developmental plasticity in trophic morphology in response to diet: a mollusk diet produces specimens with large pharyngeal jaws and muscles, whereas a soft-food diet produces smaller pharyngeal jaws and corresponding changes in musculature. Sites were chosen to maximize variability in environmental variables that might directly or indirectly affect trophic morphology. We found significant differences in pharyngeal jaw and muscle morphology among populations. Similarly, we found differences in diets among sites: mollusks were found in the stomachs of fish from only 2 populations sampled, despite the presence of mollusks in 5 of the 6 sites. Although trophic morphology did match the observed diet in 2 sites, diet did not correlate with either morphology or environmental variables across sites, nor were environmental variables correlated with morphological variation among sites. These results suggest that mismatch can occur among different populations of a single species for reasons such as seasonality in resources, developmental plasticity and/or complex indirect interactions. Intraspecific mechanisms should be further studied in order to better understand the complex relationships between morphological specialization and ecological generalization.

Cryptic intraspecific variation in sex determination in Caenorhabditis elegans revealed by mutations

Sex determination mechanisms (SDMs) show striking diversity and appear to evolve rapidly. Although interspecific comparisons and studies of ongoing major transitions in sex determination (such as the establishment of new sex chromosomes) have shed light on how SDMs evolve, comparatively little attention has been paid to intraspecific variation with less drastic effects. In this study, I used mutant strains carrying a temperature-sensitive sex determination mutation, along with a second null mutation, in different wild genetic backgrounds to uncover hidden variation in the SDM of the model nematode Caenorhabditis elegans. I then used quantitative trait locus (QTL) mapping to begin to investigate its genetic basis. I identified several QTLs, and although this variation apparently involved genotype-by-temperature interactions, QTL effects were generally consistent across temperatures. These QTLs collectively and individually explained a relatively large fraction of the variance in tail morphology (a sexually dimorphic trait), and two QTLs contained no genes known to be involved in somatic sex determination. These results show the existence of within-species variation in sex determination in this species, and underscore the potential for microevolutionary change in this important developmental pathway.

Leaf traits and water relations of 12 evergreen species in Costa Rican wet and dry forests: patterns of intra-specific variation across forests and seasons

This study examined variation in leaf traits and water relations in 12 evergreen and semideciduous woody species that occur in both seasonal wet and dry forests in Costa Rica and compared intra-specific leaf–trait correlations to those found in inter-specific global studies. The following traits were measured in both forests across seasons for 2 years: leaf nitrogen (N), leaf carbon (C), specific leaf area (SLA), toughness, cuticle thickness, leaf thickness, and leaf lifespan (LLS). Leaf water potential (LWP) and water content (LWC) were measured as indices of plant available water. Canopy openness, soil moisture, and herbivory were also measured to compare environmental variation across sites. Although species contributed the greatest amount to variation in traits, season, forest, and their interaction had a large influence on patterns of intra-specific leaf–trait variation. Leaf traits that contributed most to variation across sites were C, LWP, leaf thickness, and SLA. Traits that contributed most to variation across seasons were leaf toughness, LWP, and LWC. Furthermore, leaf traits were more correlated (i.e., number and strength of correlations) in the dry than in the wet forest. In contrast to results from global literature syntheses, there was no correlation between LLS and N, or LLS and SLA. Both light and water availability vary seasonally and may be causing variation in a number of leaf traits, specifically those that relate to water relations and leaf economics. Strong seasonality may cause leaf–trait relationships at the local scale to differ from those documented in continental and global-scale studies.

Scaling and mechanics of carnivoran footpads reveal the principles of footpad design

In most mammals, footpads are what first strike ground with each stride. Their mechanical properties therefore inevitably affect functioning of the legs; yet interspecific studies of the scaling of locomotor mechanics have all but neglected the feet and their soft tissues. Here we determine how contact area and stiffness of footpads in digitigrade carnivorans scale with body mass in order to show how footpads' mechanical properties and size covary to maintain their functional integrity. As body mass increases across several orders of magnitude, we find the following: (i) foot contact area does not keep pace with increasing body mass; therefore pressure increases, placing footpad tissue of larger animals potentially at greater risk of damage; (ii) but stiffness of the pads also increases, so the tissues of larger animals must experience less strain; and (iii) total energy stored in hindpads increases slightly more than that in the forepads, allowing additional elastic energy to be returned for greater propulsive efficiency. Moreover, pad stiffness appears to be tuned across the size range to maintain loading regimes in the limbs that are favourable for long-bone remodelling. Thus, the structural properties of footpads, unlike other biological support-structures, scale interspecifically through changes in both geometry and material properties, rather than geometric proportions alone, and do so with consequences for both maintenance and operation of other components of the locomotor system.

Analysis of DNA sequence polymorphism at the cMWG699 locus reveals phylogenetic relationships and allopolyploidy withinHordeum murinumsubspecies

Hordeum murinum L. is one of the most widely distributed species in the genus Hordeum. This species is composed of three subspecies with three ploidy levels, namely subsp. glaucum (2x=14), subsp. murinum (4x=28) and subsp. leporinum (4x=28, 6x=42). These three subspecies are morphologically similar and are frequently referred to as the 'murinum complex'. Although many cytological studies suggest that the murinum complex is allopolyploid, one inter-specific hybridization study suggested that it is autopolyploid. The goals of the present study are to identify nucleotide variation in the cMWG699 locus in the polyploid genomes of the murinum complex, to conduct molecular phylogenetic analysis of this locus, and to clarify the allo- versus auto-polyploidy status of the murinum complex. For this purpose, PCR-RFLP analysis was conducted with HhaI and SspI restriction enzymes on 80 H. murinum accessions. Single enzyme digestion data revealed polymorphism between diploid and polyploids, and double-digestion revealed polymorphism between tetra- and hexaploids. The nucleotide sequences of clones clearly show that polyploid murinum species are allopolyploid. In addition, DNA sequence analysis indicated that one donor of the tetraploid was subsp. glaucum (2x), as has been suggested previously by cytological studies. The other diploid donors were not identified, but at least one group of sequences common to 4x and 6x genomes (namely clonetype B) was highly diverged from 2x subsp. glaucum. The two tetraploid subspecies, 4x subsp. murinum and 4x subsp. leporinum, had identical DNA sequences, suggesting that these two subspecies are not differentiated at the cMWG699 locus.

An interspecific QTL study of Drosophila wing size and shape variation to investigate the genetic basis of morphological differences

The Drosophila wing has been used as a model in studies of morphogenesis and evolution; the use of such models can contribute to our understanding of mechanisms that promote morphological divergence among populations and species. We mapped quantitative trait loci (QTL) affecting wing size and shape traits using highly inbred introgression lines between D. simulans and D. sechellia, two sibling species of the melanogaster subgroup. Eighteen QTL peaks that are associated with 12 wing traits were identified, including two principal components. The wings of D. simulans and D. sechellia significantly diverged in size; two of the QTL peaks could account for part of this interspecific divergence. Both of these putative QTLs were mapped at the same cytological regions as other QTLs for intraspecific wing size variation identified in D. melanogaster studies. In these regions, one or more loci could account for intra- and interspecific variation in the size of Drosophila wings. Three other QTL peaks were related to a pattern of interspecific variation in wing size and shape traits that is summarized by one principal component. In addition, we observed that female wings are significantly larger and longer than male wings and the second, fourth and fifth longitudinal veins are closer together at the distal wing area. This pattern was summarized by another principal component, for which one QTL was mapped.

Studies on Heterosis in Abelmoschus esculentus (L.) Moench and A. callei (A. Chev) Stevels Cultivars During Shorter Day Photoperiods in South Eastern Nigeria

Better parent heterosis was studied in direct and reciprocal crosses using 9 early and late okra cultivars which belonged to the Early-early, Early-late, Late-early and Late-late okra flowering types. The ANOVA for length of pod, circumference of pod, number of locules per pod, number of seeds per pod, Number of pods per plant, weight of 1000 seeds, density of seeds, weight of pods per plant, days to flower opening, plant height and circumference of stem at 10 cm above soil level, showed almost very highly significant differences-an indication that the cultivars are genetically diverse. Very highly significant, narrow and intermediate, heterosis was recorded for most of the direct and reciprocal crosses, showing that selections could be made from the hybrids to meet desired local okra qualities. A cross between an Early-late and Late-early parents, using early okra as the maternal parent, gave rise to a stable viable bridge hybrid which outperformed the better parent in many respects thereby overcoming the hitherto strong barrier to gene flow in interspecific hybridization studies involving the two okra types and also indicating the existence of maternal effects. Consequently with this development, the window has been opened for possible accelerated transfer of several desirable genes from late okra types to many promising but vulnerable early okra types. This could result to minimizing the further erosion of such early okra germplasm still in the custody of the local farmers.

A novel nuclear marker, Pm-int9, for phylogenetic studies of Opisthorchis felineus, Opisthorchis viverrini, and Clonorchis sinensis (Opisthorchiidae, Trematoda)

Opisthorchis felineus, O. viverrini, and Clonorchis sinensis, the trematodes of the family Opisthorchiidae, are important human parasites. Two previous studies (Kang et al. Parasitol Int 57:191-197, 2008; Katokhin et al. Dokl Biochem Biophys 421:214-217, 2008) have provided evidence using ribosomal and mitochondrial sequences that O. viverrini, O. felineus, and C. sinensis are closely related. We developed a novel nuclear marker, Pm-int9, which included the ninth intron of the paramyosin gene and flanking exon sequences. Samples of O. felineus from four localities of West Siberia, C. sinensis from the Russian Far East, and O. viverrini from Thailand were genotyped by Pm-int9. Little variation was detected in exon sequences, however, intron sequences turned out to be more variable than ribosomal internal transcribed spacers. We can conclude that Pm-int9 is valuable for interspecific variation studies. Phylogenetic analysis based on Pm-int9 revealed that O. viverrini and C. sinensis were closer to each other than either of them to O. felineus, supporting the opinion that C. sinensis should be considered the sister species of Opisthorchis spp.

Geographic body size gradients in tropical regions: water deficit and anuran body size in the Brazilian Cerrado

A recent interspecific study found Bergmann's size clines for Holarctic anurans and proposed an explanation based on heat balance to account for the pattern. However, this analysis was limited to cold temperate regions, and exploring the patterns in warmer tropical climates may reveal other factors that also influence anuran body size variation. We address this using a Cerrado anuran database. We examine the relationship between mean body size in a grid of 1° cells and environmental predictors and test the relative support for four hypotheses using an AIC‐based model selection approach. Also, we considered three different amphibian phylogenies to partition the phylogenetic and specific components of the interspecific variation in body size using a method analogous to phylogenetic eigen vector regression (PVR). To consider the potential effects of spatial autocorrelation we use eigenvector‐based spatial filters. We found the largest species inhabiting high water deficit areas in the northeast and the smallest in the wet southwest. Our results are consistent with the water availability hypothesis which, coupled with previous findings, suggests that the major determinant of interspecific body size variation in anurans switches from energy to water towards the equator. We propose that anuran body size gradients reflect effects of reduced surface to volume ratios in larger species to control both heat and water balance.

Interspecific cetacean interactions during the breeding season of humpback whale (Megaptera novaeangliae) on the north coast of Bahia State, Brazil

The large majority of cetacean interspecific studies report only on dolphin interactions, while studies on interactions between odontocete and mysticete are less common. The present work aims to report on sightings of cetacean interactions, during the breeding season of humpback whales(Megaptera novaeangliae), along 370 km of the Bahia State, north-eastern Brazil, addressing aspects of cetacean distribution and behaviour. During 7 seasons (2000–2006), a total of 230 research cruises were performed, in which 38 events of interactions among humpback whales and other cetaceans(Tursiops truncatus, Steno bredanensis, Peponocephala electraandBalaenoptera acutorostrata)were observed, plus another 5 encounters without the whale's presence, includingT. truncatus, S. bredanensis, P. electra, Stenella clymeneandStenella attenuata. Our results confirm the occurrence of multiple cetacean species in the Bahia State, being the first study in the world to report on a large range of interactions involving another 4 cetacean species, grouped with up to 3 mixed species per sighting, with humpback whales in their annual breeding ground.

Isolation and characterization of microsatellite markers for Brachiaria brizantha (Hochst. ex A. Rich.) Stap

The first set of nuclear simple sequence repeat (SSR) loci for Brachiaria brizantha (Hochst. ex A. Rich.) Stap is described. A microsatellite-enriched library was constructed and 19 loci were characterized. About 13 SSR loci were found to be polymorphic and across-taxa amplification tests showed that six of them can be transferred to four other species of Brachiaria. This new SSR resource will be a powerful tool for population genetic studies of B. brizantha, for interspecific genetic studies within the genus Brachiaria, for mapping and for marker assisted selection in breeding.

ITS에 의한 한국 내 Pelvetia속 분류군의 계통학적 연구

갈조류 모자반과 Pelvetia속은 다세포 조류로 많은 해조류를 포함하고 있으며 북반구 태평양과 대서양에 분포되어 있다. Pelvetia속에 속하는 우리나라의 동종 및 근연한 같은 속내 종에 대해 유전자은행을 통해 밝혀진 ITS에 의한 서열을 이용하여 계통관계를 조사하였다. 한국의 P. babingtonii은 북미의 P. babingtonii AF102957과 유사한 핵산서열로 계통도 분석에서 같은 분지군을 형성하였다. 한국의 P. siliquosa은 역시 북미의 P. siliquosa AF102958과 유사한 핵산서열로 계통도 분석에서 같은 분지군을 형성하였다. Pelvetia 속 내 여러 종간은 결실이나 삽입에 의한 indel이 많은 반면 같은 종내 계통은 치환에 의한 차이가 현저하였다. 이 속은 NJ분석에서 크게 두 분지군으로 나뉘는데 한 그룹은 P. canaliculata와 P. limitata이며 나머지 그룹은 P. siliquosa, P. compressa, P. babingtonii을 포함하고 있었다. ITS 서열로 한국 내 분류군과 북미 간 분류군이 잘 구분되었다. MP 분석에서도 높은 지지도로 잘 구분되었다. 따라서 ITS 서열로 종 동정에 이용할 수 있었으며, 종의 보전이나 생식질 보전에 기초로 이용될 수 있을 것으로 사료된다. The brown algae, or phaeophytes, are a large group of multicellular algae, including many notable types of seaweed. We analysed intra- and interspecific phylogenic studies within the genus Pelvetia in Korea and compared them with results of both same and different species in GenBank. The sequences for P. babingtonii in Korea were generally similar to those for P. babingtonii AF102957, and the sequences of P. siliquosa in Korea were also similar to those of P. siliquosa AF102958. Sequence variation within the Pelvetia is mostly due to nucleotide substitutions, although several small indels and some large indels can be found. Another source of sequence divergence is length variation due to stretches of short repeats that occur at the ITS1 or ITS2 in all the Pelvetia. NJ analysis consists mainly of two clades. One of them contains P. canaliculata and P. limitata, and is sister to the rest of the genus (P. siliquosa, P. compressa, and P. babingtonii). P. babingtonii is not grouped one clade. In the MP analysis, ten accessions or populations were fully resolved and all accessions from the same species formed with 99% or 100% bootstrap supports.

An intra- and interspecific study of body size and autotomy as a defense in Orthoptera

Autotomy, the ‘voluntary’ shedding of a limb or other body part, is a highly effective escape mechanism to avoid predation or other forms of entrapment. Autotomy, however, comes with costs to locomotion, reproductive behavior, regeneration etc. It has been suggested that increasing body size and ‘robustness’ may allow for less reliance on this extreme form of predator defense, but this theory has never been tested. Here we present behavioral observations (‘willingness’ or time taken to lose an entrapped limb) of Orthoptera of a range of body size and mass. These data strongly suggest that body size and mass may be an important determinant of the use of autotomy as an escape mechanism within the Order, possibly due to the effects of body size upon the efficacy of autotomy, as well as other defense mechanisms. An ontogenetic study for Gryllus bimaculatus, however, shows no clear trend in willingness to autotomize a limb with body mass, suggesting that this defense tactic may be less affected by body size per se, but rather by the tactics developed by each individual species.

Suitability of EST-PCR Markers Developed in Highbush Blueberry for Genetic Fingerprinting and Relationship Studies in Lowbush Blueberry and Related Species

Little is known of the genetic structure and variability of wild fields, or of the dramatic differences in yield among clones (genetic individuals), of lowbush blueberry (Vaccinium angustifolium Ait.), Maine's most economically important fruit crop. Express sequence tag-polymerase chain reaction (EST-PCR) markers that were originally developed for genetic mapping purposes in highbush blueberry (Vaccinium corymbosum L.) are shown here to be valuable for genetic fingerprinting and relationship studies in the related species, V. angustifolium. As part of an interspecific genetic relationship study, 14 genotypes, including at least two specimens of each of four closely related Vaccinium L. species (V. pallidum Ait., V. corymbosum, V. boreale Hall & Aald., and V. myrtilloides Michx.) and the only four pedigreed cultivars of V. angustifolium, grouped out as expected in a genetic similarity dendrogram (matrix “r” correlation = 0.91). This work is ultimately aimed at using these markers in exploring how genetic relationship affects yield among proximal and distant breeding individuals via controlled field hand crosses. To help address this issue, a separate group of six individuals of V. angustifolium from two managed fields were also genotyped using the EST-PCR markers. The markers were very effective at intraspecific discrimination of individuals within the same field.

Genetic variation in three species of freshwater crayfish; Astacus astacus L., Astacus leptodactylus Aesch., and Pacifastacus leniusculus (Dana), revealed by isozyme electrophoresis

Agerberg, A. 1990. Genetic variation in three species of freshwater crayfish; Astacus astacus L., Astacus leptodactylus Aesch., and Pacifastacus leniusculus (Dana), revealed by isozyme electrophoresis. —Hereditas 113: 101–108. Lund, Sweden. ISSN 0018–0661. Received March 15, 1990. Accepted June 20, 1990 Isozyme electrophoresis was applied to three species of freshwater crayfish, Astacus astacus L., Asfacus leptodactylus Esch., and Pacifastacus leniusculus (Dana), in order to detect genetic variation and to find the most suitable enzyme systems for intra- and interspecific studies of these species. To obtain maximum information a number of different tissues and organs were used. lnitjally 21 enzymes and general proteins were surveyed and 15 of them, with a putative number of 29 loci, were found to be suitable. No genetically interpretable variation was observed in any of the A. astacus populations. However, in A. leptodactylus and P. leniusculus, polymorphisms were detected in one and three loci, respectively. The genetic identity (nei 1972, Am. Nat. 106: 283–292), calculated for enzymatic loci only, was estimated at 0.7243 within genus Astacus and 0.5983+0.019 betweeen the genera Astacus and Pacifastacus.

The degree and pattern of phylogenetic signal in primate long-bone structure

Interspecific scaling is a fundamental tool for comparative studies of primate long-bone structure and adaptation. However, scaling analyses based on conventional statistical methods can lead to false positives regarding adaptive relationships when traits exhibit strong phylogenetic signal. This problem can be addressed through the use of phylogenetic comparative methods (PCMs). To date, PCMs have not been incorporated into comparative studies of primate long-bone structure because it has been assumed that long-bone structure is free of phylogenetic signal once appropriately scaled. To test this assumption, we evaluated the degree of phylogenetic signal in three types of long-bone structural traits (bone length, articular surface areas, and cross-sectional geometric properties) from 17 quadrupedal primate species. We compared the pattern of phylogenetic signal in raw trait values and residual trait values after regression against body mass, bone length, and the product of body mass × bone length. Our results show that significant phylogenetic signal is present in all traits before scaling, due in part to their strong covariance with body mass. After scaling, bone length still exhibits strong phylogenetic signal, but articular surface areas do not, and cross-sectional properties exhibit different levels of signal depending on the variable used to scale the data. These results suggest that PCMs should be incorporated into interspecific studies of bone length and perhaps cross-sectional geometric properties. Our results also demonstrate that tests for phylogenetic signal prior to implementing a PCM should focus on residual variance, not individual traits.

Ontogenetic scaling of fish metabolism in the mouse-to-elephant mass magnitude range

Intraspecific or ontogenetic analyses of mass-metabolism relationships do not often conform to the same allometric correlations as those seen in interspecific analyses. A commonly cited reason for this discrepancy is that ontogenetic studies examine smaller mass ranges than interspecific studies, and are therefore not statistically comparable. In this study the metabolic rate of yellowtail kingfish was measured from 0.6 mg–2.2 kg, a mass range comparable to that between a mouse and an elephant. Linear regression of the log transformed data resulted in a scaling exponent of 0.90 and high correlation coefficient. Statistical and information theory comparisons of three other models showed that a segmented linear regression and curvilinear quadratic function were an improvement over a simple linear regression. This confirmed previous observations that the metabolic scaling exponent of fish changes during ontogeny. Ammonia excretion rates were also measured and scaled linearly with an exponent of 0.87. The data showed that the metabolism of yellowtail kingfish during ontogeny did not scale with the commonly cited 2/3 or 3/4 mass exponent. This demonstrates that differences between interspecific and ontogenetic allometries are not necessarily statistical artefacts.