Partitioning a morphology among its controlling factors

In the first known attempt to use a sympatric, non-mimetic convergence (‘a naturally controlled experiment in evolutionary morphology’) to quantitatively partition a morphology among its controlling factors, shells of the polygyrid land snails Neohelix major (Binney) and Mesodon normalis (Pilsbry) were analysed in a factorially designed rearing experiment. The complete absence of character displacement or character release confirmed previous field studies. Growth rate, a determinant of adult shell size, was controlled 50% by direct environmental induction (with temperature twice as effective as humidity), 10% by proximate natural selection (with a shorter growing season selecting for faster growth), 10% by genotype-environment interaction (with moisture generally inducing a greater growth surge in a drier climate), and 30% by within-cohort variation (apparently heritable). Spire height confirmed predictions that flatter shells make better estivators (shelterers from drought) and that taller shells make better climbers and tall/flat niches are readily filled: high within-cohort variation (70% of total) provides the raw material for niche-filling (but is reduced in severe climates by stabilizing selection); environmental induction (10%) is strictly by humidity, which makes shells taller for foraging more widely under more favourable conditions; proximate natural selection (10%) produces taller shells in moister climates (with intensity of effect dependent on evolutionary time); and genotype-environment interaction (10%) makes the drought-induction of flat shells stronger in more drought-prone areas. Coiling tightness, allied to whorl-expansion rate, shows distinct evidence of adaptive neutrality: 60% of total variance is explained by phylogenetic constraints despite an estimated 120 million years of genetic isolation and despite the demonstrated ability of related clades to converge; the 10% due to genotype-environment interaction is generally random and nonadaptive in its response; and the remaining 30% appears random, non-induced, and non-selected. This study's protocol of experimentally partitioning a ‘closed universe’ of morphology among a full set of controlling factors is recommended as an alternative to the usual practice of analysing the effects of one or a subset of factors on an ‘open universe’ of morphology. One advantage of this new protocol is that it obviates the ‘nature-nurture debate’ by quantifying the relative contributions of each, which in this snail-shell case differ drastically among the various components of a single morpholocgyy.     

Morphological convergence and coexistence in three sympatric North American species of Microtus (Rodentia: Arvicolinae)

Aim  Niche theory predicts that ecologically identical species cannot stably coexist in local communities. My aim was to investigate morphological diversity as a possible factor enabling the coexistence of a species-rich Microtus (Rodentia: Arvicolinae) fauna in a hotspot of North American mammalian diversity, the Greater Yellowstone Ecosystem (GYE).
Location  The Greater Yellowstone Ecosystem, North America.
Methods  Using in-hand morphological measurements of size and shape, I compared the morphologies of three North American vole species ( Microtus spp.), in sympatry in the GYE and in allopatry across their ranges, in order to examine: (1) whether morphologies are fixed or plastic and (2) the degree of morphological character displacement or convergence in sympatric species.
Results  Support was found for plasticity of morphology for all three vole species: M. longicaudus , M. montanus and M. pennsylvanicus. However, Microtus individuals of all species from the GYE area of sympatry were more similar to each other than to allopatric individuals of the same species.
Main conclusions  Competition among these congeners is not manifested in morphological overdispersion. The response of these congeneric species to the same local ecological conditions is convergent. The relative strength of environmental conditions appears to be stronger than the strength of competitive interactions among the study species.     

Character displacement, frequency-dependent selection, and divergence of shell colour in land snails Mandarina (Pulmonata)

Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an exceptional example of habitat and character divergence among closely related species. In this study, microhabitat differences between sympatric ground-dwelling species were studied by distinguishing habitats on the basis of vegetation and types of litter. In all sites where two ground species coexisted, segregation occurred with each species showing preference for the microhabitat in which they were found. When they were in sympatry, one species was predominant in relatively wet and sheltered sites and the other in relatively dry and exposed sites. Although most species can live in both types of habitat, occupation by one species is inhibited by occupation by another. This suggests that competitive interaction between sympatric species caused segregation. Except for populations that have undergone interspecific hybridization, no examples were found of sympatric populations of two ground species sharing a similar shell colour. Species that were predominant in relatively wet and sheltered sites possessed shells with dark coloration and their colour patterns were mostly of one type. Species that were predominant in relatively dry and exposed sites possessed shells with bright coloration and their color patterns were polymorphic. Most populations from areas in which single species were distributed had shells with medium coloration. Microhabitat differentiation between sympatric species possibly caused diversification of shell colour, because bright shells are advantageous in sites where snails are largely exposed, and dark shells are advantageous in sites in where they are mostly sheltered from sunlight. In addition, frequency-dependent selection by predators hunting by sight may have operated to maintain colour polymorphism in the populations which are restricted to exposed habitats by competition with other sympatric species. This reveals the importance of interaction among closely related species as a cause of diversification in ecological and morphological traits.     

Genetic structure and differentiation of Plantago major reveals a pair of sympatric sister species

Seeds of the widespread weed Plantago major were collected from 10 European countries, as well as Trinidad and North America. The seed collections were from populations of two taxa which are ecologically rather than geographically separated and formally recognized as the subspecies Plantago major ssp. major and P.m. ssp. intermedia (also called P.m. ssp. pleiosperma). Eight polymorphic allozyme loci and 73 random-primed DNA fragments were scored, as well as 11 morphological characters. Complete concordance between morphological traits and genetic data provides evidence that these two taxa, although very similar, are distinct species. They are both widespread, they are broadly sympatric and capable of interbreeding. However, slight morphological and ecological differences coincide with genetic clustering of populations from widely separated locations. In addition, P. major and P. intermedia differ in their population structure: P. intermedia has greater genetic diversity among populations and less genetic variance within populations than P. major. We suggest that differences between the two species in their levels of selfing may explain the distinctive genetic structure of each species. We hypothesize a link between selfing rate and lifespan of the two taxa. P. major is characterized by lower genetic variation among populations, a higher rate of outcrossing, longer lifespan and production of fewer seeds per seed capsule. P. intermedia is more highly structured with much differentiation among populations, a higher rate of inbreeding and it often grows as an annual.     

Recent and rapid diversification of the small carpenter bees in eastern North America

Bees fulfil a critical ecological role as pollinators, significantly contributing to the reproductive success of myriad angiosperm species. Although increasingly appreciated for their agricultural contributions, relatively little is yet known about the natural history of the vast majority of the more than 20 000 wild bee species worldwide. The small carpenter bee genus Ceratina occurs globally, and is represented in North America by its most recently diverged subgenera, Ceratinula and Zadontomerus. Recent genetic analysis of eastern Ceratina (Zadontomerus) supports the existence of five closely related, yet genetically distinct species living in sympatry. This phylogeographical study employs molecular barcoding of the most comprehensive specimen collection yet assembled to confirm the identities of these recently diverged eastern North American Ceratina (Zadontomerus) species. Delineation of extant population structure, evolutionary history and known range of this emerging model native pollinator are greatly improved by this study. We consider ecological and behavioural factors potentially contributing to the maintenance of genetic identity among these sympatric species.     

The historical ecology of yellow perch (Perca flavescens[Mitchill]) and their parasites

Aim To determine the origins of the host–parasite association between among yellow perch (Perca flavescens[Mitchill]) and the parasites Crepidostomum cooperi Hopkins, Proteocephalus pearsei La Rue and Urocleidus adspectus Beverly Burton. Of secondary interest are the parasites Bunodera luciopercae (Muller) and Proteocephalus percae (Muller) predictably associated with the Eurasian perch. Location The areas considered are the Holarctic, since the upper‐Cretaceous, and contemporary North America. Methods Published and new information from host and parasite phylogenies, palaeontology, palaeogeography and plate tectonics and host biology is incorporated to assess the origins of yellow perch and several of its parasites. This information is used to determine the origins for these host–parasite associations. Results Cladistic analysis suggests a Laurasian origin for Percidae and Perca, and that Perca is sister to the other genera in the family. Parasite phylogenies support a North American origin for the three species associated with yellow perch and a Laurasian origin for B. luciopercae. Proteocephalus pearsei and P. percae are not sister taxa. The fossil record for Perca dates to the Miocene in Europe and the Pleistocene in North America. North America and Europe were connected across the North Atlantic since at least the upper Cretaceous with separation complete by the Miocene. Europe was separated from Asia by the Obik Sea from the late Cretaceous until the Oligocene. Western cordillera orogeny and its accompanying high rates of water flow and Pleistocene glaciation represent barriers to Perca dispersal. Main conclusions The origin of Perca in North America dates at least to the late Oligocene when North America and Europe were connected across the North Atlantic and Europe and Asia were separate landmasses, and does not result from Pleistocene dispersal across Beringia from Asia. The present disjunction of Perca species in North America and Europe is due to the vicariant separation of North America and Europe. Based on the available information, yellow perch and its parasites have a North America origin. The association between yellow perch and the parasites in all cases is a consequence of host switching from other sympatric host species in North America and is not explained by co‐speciation. Even the association between the host‐specific Urocleidus adspectus and yellow perch originated with a host switch and is not due to co‐speciation. The basis for this host switching is geographical and ecological sympatry, especially shared feeding habits, with other North American fish hosts.     

Sympatric relationship between redband trout and non-native brook trout in the Southeastern Oregon Great Basin

Brook trout (Salvelinus fontinalis) and rainbow trout (Oncorhynchus mykiss) have been widely introduced outside their respective ranges within North America causing declines and displacement of native trout. Yet, successful coexistence of native and non-native trout has received little attention. Here we evaluated the effect of introduced brook trout on the size and density of native redband trout in two invaded sub-basins in southeastern Oregon. In a multi-year study, we investigated whether habitat and fish communities differed between streams and stream reaches where redband trout were allopatric versus where redband trout were sympatric with brook trout. We hypothesized that redband trout would be less dense and have smaller total length in sympatry with brook trout than in allopatry, but that total trout density would not differ. We investigated whether differences in habitat existed between sympatric and allopatric locations that would indicate differentiation in site level habitat preferences for each trout species. We found that sympatric locations had more wood but similar fish community structure. Mean length and densities of redband trout were higher at allopatric locations. However, in most years at sympatric locations total trout density was twice that of allopatric redband trout sites. Using comparable data from an eastern United States system where brook trout are native, sympatric sites had lower densities of brook trout; however, total trout density did not differ. We conclude that invading trout negatively impact native trout densities; but in southeastern Oregon system the negative impact is minimized.     

THE RATE TEST OF SPECIATION: ESTIMATING THE LIKELIHOOD OF NON‐ALLOPATRIC SPECIATION FROM REPRODUCTIVE ISOLATION RATES IN DROSOPHILA

Among the most debated subjects in speciation is the question of its mode. Although allopatric (geographical) speciation is assumed the null model, the importance of parapatric and sympatric speciation is extremely difficult to assess and remains controversial. Here I develop a novel approach to distinguish these modes of speciation by studying the evolution of reproductive isolation (RI) among taxa. I focus on the Drosophila genus, for which measures of RI are known. First, I incorporate RI into age‐range correlations. Plots show that almost all cases of weak RI are between allopatric taxa whereas sympatric taxa have strong RI. This either implies that most reproductive isolation (RI) was initiated in allopatry or that RI evolves too rapidly in sympatry to be captured at incipient stages. To distinguish between these explanations, I develop a new “rate test of speciation” that estimates the likelihood of non‐allopatric speciation given the distribution of RI rates in allopatry versus sympatry. Most sympatric taxa were found to have likely initiated RI in allopatry. However, two putative candidate species pairs for non‐allopatric speciation were identified (5% of known Drosophila). In total, this study shows how using RI measures can greatly inform us about the geographical mode of speciation in nature.     

Cryptic diversity within and amongst spring‐associated Stygobromus amphipods (Amphipoda: Crangonyctidae)

Multiple species of troglomorphic, spring‐associated Stygobromus amphipods, including the endangered, narrow‐range endemic Stygobromus pecki, occupy sites in the Edwards Plateau region of North America. Given the prevalence of cryptic diversity observed in disparate subterranean, animal taxa, we evaluated geographical genetic variation and tested whether Stygobromus contained undetected biodiversity. Nominal Stygobromus taxa were treated as hypotheses and tested with mitochondrial sequence cytochrome oxidase C subunit 1, nuclear sequence (internal transcribed spacer region 1), and AFLP data. Stygobromus pecki population structure and diversity was characterized and compared with congeners. For several Stygobromus species, the nominal taxonomy conflicted with molecular genetic data and there was strong evidence of significant cryptic diversity. Whereas S. pecki genetic diversity was similar to that of congeners, mitochondrial data identified two significantly diverged but sympatric clades. AFLP data for S. pecki indicated relatively recent and ongoing gene flow in the nuclear genome. These data for S. pecki suggest either a substantial history of isolation followed by current sympatry and ongoing admixture, or a protracted period of extremely large effective population size. This study demonstrates that Edwards Plateau Stygobromus are a complex, genetically diverse group with substantially more diversity than currently recognized. © 2013 The Linnean Society of London     

Floral traits and mating systems in sister species of <Emphasis Type="Italic">Nicotiana</Emphasis>: interpopulational variability and sympatry effects

Mating systems of Angiosperms are important determinants of population genetic structure and evolutionary potential. Nicotiana longiflora and N. plumbaginifolia are self-compatible, sister species, with contrasting floral morphology and can be found in allopatry and sympatry in North Argentina. In two sympatric and 10 nearby allopatric populations we studied their natural interpopulational variability and sympatry effects on corolla length and anther–stigma distance. We also estimated seed set by selfing and via pollinators using pollination treatments. Both corolla length and anther–stigma distance varied significantly among N. longiflora, but not among N. plumbaginifolia populations. We did not detect an effect of sympatry in either species for any of the floral traits studied. Pollination treatments suggest that N. longiflora is mainly an outcrosser, although selfing occurs at some extent. Seed set attributed to pollinators was significantly higher in N. longiflora whereas most seeds in N. plumbaginifolia were sired through self-pollination. In N. plumbaginifolia, selfing seems to assure reproduction in sympatric populations, where floral visitors have a strong preference for N. longiflora. Corolla length was significantly negatively correlated with an increase in the percentage of self-seeds estimated by pollination treatments. Within N. longiflora, anther–stigma distance showed a positive correlation with selfing.     

Canid hybridization: contemporary evolution in human‐modified landscapes

Contemporary evolution through human‐induced hybridization occurs throughout the taxonomic range. Formerly allopatric species appear especially susceptible to hybridization. Consequently, hybridization is expected to be more common in regions with recent sympatry owing to human activity than in areas of historical range overlap. Coyotes ( Canis latrans) and gray wolves ( C. lupus) are historically sympatric in western North America. Following European settlement gray wolf range contracted, whereas coyote range expanded to include eastern North America. Furthermore, wolves with New World (NW) mitochondrial DNA (mtDNA) haplotypes now extend from Manitoba to Québec in Canada and hybridize with gray wolves and coyotes. Using mtDNA and 12 microsatellite markers, we evaluated levels of wolf‐coyote hybridization in regions where coyotes were present (the Canadian Prairies, n = 109 samples) and absent historically (Québec, n = 154). Wolves with NW mtDNA extended from central Saskatchewan (51°N, 69°W) to northeastern Québec (54°N, 108°W). On the Prairies, 6.3% of coyotes and 9.2% of wolves had genetic profiles suggesting wolf‐coyote hybridization. In contrast, 12.6% of coyotes and 37.4% of wolves in Québec had profiles indicating hybrid origin. Wolves with NW and Old World ( C. lupus) mtDNA appear to form integrated populations in both regions. Our results suggest that hybridization is more frequent in historically allopatric populations. Range shifts, now expected across taxa following climate change and other human influence on the environment, might therefore promote contemporary evolution by hybridization.     

Habitat Utilization of Three Sympatric Cheirogaleid Lemur Species in a Littoral Rain Forest of Southeastern Madagascar

I compared the habitat utilization in 3 sympatric species of Cheirogaleidae (Microcebus murinus [81 g], Cheirogaleus medius [183 g] and Cheirogaleus major [362 g]) in a littoral rain forest in southeastern Madagascar during 3 rainy seasons. Females of promiscuous Microcebus murinus had small home ranges and the males had large overlapping home ranges. Home ranges of family groups of monogamous Cheirogaleus medius and C. major overlapped extensively. Home ranges of all 3 species overlapped completely in the study area but home range sizes differed among species and correlate positively with body masses. Male Microcebus murinus slept in open vegetation (79%) and alone (71%), whereas female M. murinus and family group members of Cheirogaleus spp. preferred communal sleeping in tree holes. There are significant interspecific differences in the choice of sleeping sites: smaller lemurs chose smaller trees and used more sleeping sites than larger lemurs did. Species also differed significantly in the vertical dimension of forest utilization: Cheirogaleus major used the upper part of the trees, C. medius used the middle parts, and Microcebus murinus used the understory during nocturnal activities. The 3 species differed mainly in vertical habitat utilization and showed vertical stratification.     

CONVERGENT EVOLUTION OF COURTSHIP SONGS AMONG CRYPTIC SPECIES OF THE CARNEA GROUP OF GREEN LACEWINGS (NEUROPTERA: CHRYSOPIDAE: CHRYSOPERLA)

Although traits of related species are likely to be similar due to common ancestry, mating signals are an exception. In singing insects, for example, song similarity has been documented only for allopatric or allochronic species pairs, and even then, not often. Where song similarity does occur, it has been logically attributed to the inheritance of ancestral traits rather than convergence. It is quite common for related, sympatric insect species to differ dramatically in calling song, which is predicted by evolutionary theory to maximize intraspecific mating success. Given that there are a limited number of ways to make sounds on anatomically similar organs and given that there would be no selective pressure for songs to differ in widely separated geographic areas, convergence in songs among related species living on different continents might be expected. Here we present the first well-documented case of such convergence, in a group of sibling, cryptic species characterized by substrate-borne vibrational mating songs. In this example from green lacewings of the carnea group of the genus Chrysoperla, a variety of statistical tests shows that one species in North America and another in Asia possess songs that are strikingly similar to each other. DNA data demonstrate that the species involved belong to divergent speciose lineages, and behavioral data demonstrate that the convergent songs are readily accepted by members of both species.     

THE ORIGIN AND EVOLUTIONARY RELATIONSHIPS OF ‘HUAUZONTLE’ (CHENOPODIUM NUTTALLIAE SAFFORD), DOMESTICATED CHENOPOD OF MEXICO

A New World assemblage of tetraploid Chenopodium species (section Chenopodium, subsection Cellulata) includes two domesticates, C. quinoa of Andean South America and C. nuttalliae of Mexico. Both have been combined into a single species and the Mexican form has been considered as a possible derivative of C. quinoa. The domesticates and related, sympatric weed forms, C. berlandieri of North America and C. hircinum of the Andes, were examined for variation in morphological and biochemical characteristics and also were included in a program of artificial hybridization. Results indicate that the domesticated forms are more closely related to their sympatric weeds than to each other. The Mexican cultigen is placed as a subspecies of C. berlandieri, the taxon from which it most likely evolved under human selection in North America. Possible origins for the Andean weed-crop complex are considered. Southward migration of a North American tetraploid appears to be more likely than independent allotetraploidy in South America. Of the North American tetraploids examined, C. berlandieri var. zschackei of the western U.S. shows closest affinities to the Andean complex.     

Reproductive character displacement of female,but not male song discrimination in an avian hybrid zone

Divergence of male sexual signals and female preferences for those signals often maintains reproductive boundaries between closely related, co‐occurring species. However, contrasting sources of selection, such as interspecific competition, can lead to weak divergence or even convergence of sexual signals in sympatry. When signals converge, assortative mating can be maintained if the mating preferences of females diverge in sympatry (reproductive character displacement; RCD), but there are few explicit examples. Pied flycatchers (Ficedula hypoleuca) are sympatric with collared flycatchers (F. albicollis) on the Baltic island of Öland, where males from both species compete over nestboxes, their songs converge, and the two species occasionally hybridize. We compare song discrimination of male and female pied flycatchers on Öland and in an allopatric population on the Swedish mainland. Using field choice trials, we show that male pied flycatchers respond similarly to the songs of both species in sympatry and allopatry, while female pied flycatchers express stronger discrimination against heterospecific songs in sympatry than in allopatry. These results are consistent with RCD of song discrimination of female pied flycatchers where they co‐occur with collared flycatchers, which should maintain species assortative mating despite convergence of male sexual signals.     

Cryptic, genetically extremely divergent, polytypic, convergent, and polymorphic taxa in Madagascan Tropidophora (Gastropoda: Pomatiasidae)

Madagascar's magnificent and environmentally threatened endemic radiation of the land-snail genus Tropidophora Troschel has recently been classified into three subgenera, 95 species and 142 varieties, based on often subtle conchological variation among small samples; it seems best to ignore temporarily this confusing plethora of names until true biological species and their relationships are better understood. The author's field work in 1990 succeeded in obtaining live Tropidophora from 40 populations distributed throughout the island. Allozyme analysis (108 snails, 15 loci, 117 alleles) yielded a cladogram suggesting nine genetically coherent taxa (T. taxa A-I), each represented in the collections by 1–10 populations. Comparisons among shells (total 1634) and penes (total 31 from 20 populations representing eight of the nine taxa) revealed: (1) two conchologically indistinguishable taxa (H and I) fixed for alternative alleles at 13 of 15 loci, with only a very subtle difference in penes, and with mosaic and overlapping geographical distributions in the Northeast; (2) two extremely polytypic taxa (C in the Southwest, F in the Southeast) with parallel trends toward depressed, broadly umbilicate, heavily sculptured shells with apertural lips widely reflected at the umbilicus at inland, more arid sites, resulting in sympatric convergence; (3) one southeastern taxon (G) in which the penis apparently doubles in length but the shell does not change 6.0 km to the northwest, but in which the shell shifts dramatically in sculpture, colour, and lip reflection 0.5 km to the westnorthwest; and (4) generally such extreme intra- and inter-population variation in shell and male-genital characters as to render many of them dubious at best for systematics. Thus the Madagascan Tropidophora present a fascinatingly complex problem in evolutionary morphology/ecology, the solution of which will require even more extensive collecting, followed by molecular comparisons or detailed anatomical comparisons, or both. The total number of biological species is still likely to be quite large, despite irrelevance of much of the current taxonomy, because many smaller species remain to be discovered.     

Gene flow,divergent selection and resistance to introgression in two species of morning glories (Ipomoea)

Gene flow is thought to impede genetic divergence and speciation by homogenizing genomes. Recent theory and research suggest that sufficiently strong divergent selection can overpower gene flow, leading to loci that are highly differentiated compared to others. However, there are also alternative explanations for this pattern. Independent evidence that loci in highly differentiated regions are under divergent selection would allow these explanations to be distinguished, but such evidence is scarce. Here, we present multiple lines of evidence that many of the highly divergent SNPs in a pair of sister morning glory species, Ipomoea cordatotriloba and I. lacunosa, are the result of divergent selection in the face of gene flow. We analysed a SNP data set across the genome to assess the amount of gene flow, resistance to introgression and patterns of selection on loci resistant to introgression. We show that differentiation between the two species is much lower in sympatry than in allopatry, consistent with interspecific gene flow in sympatry. Gene flow appears to be substantially greater from I. lacunosa to I. cordatotriloba than in the reverse direction, resulting in sympatric and allopatric I. cordatotriloba being substantially more different than sympatric and allopatric I. lacunosa. Many SNPs highly differentiated in allopatry have experienced divergent selection, and, despite gene flow in sympatry, resist homogenization in sympatry. Finally, five out of eight floral and inflorescence characteristics measured exhibit asymmetric convergence in sympatry. Consistent with the pattern of gene flow, I. cordatotriloba traits become much more like those of I. lacunosa than the reverse. Our investigation reveals the complex interplay between selection and gene flow that can occur during the early stages of speciation.     

Character displacement in Hydrobia

Summary Fenchel's study of size variation in Hydrobiid snails in the Limfjord, Denmark, provides one of the most convincing cases of ecological character displacement available. In order to assess the generality of the phenomena within the Hydrobia genus, allopatric and sympatric Hydrobia ventrosa, H. neglecta and H. ulvae were collected from 24 coastal sites around Eastern England in July and October, 1982. Shell heights of 5,850 snails from 55 samples were recorded. These data were analysed for intraspecific allopatric-sympatric, and interspecific height differences. Such differences were not significant for H. ventrosa/H. neglecta, but the species exhibited parallel variation at sympatric sites. H. ventrosa/H. ulvae showed only significant interspecific differences, but in both allopatry and sympatry. The pattern of mean size variation for these species resembles that in the Limfjord. However the statistical analyses fail to provide support for character displacement. Differences in character states attributed to the process of character displacement may result from a number of other causes. Environmental conditions at sympatric and allopatric sites may act differentially on the heights of H. ulvae and H. ventrosa. The conditions that prevail at sympatric H. ulvae sites appear to lead to increased size in this species irrespective of the presence of H. ventrosa.     

Variation in genetic structure among populations of the caddisfly Helicopsyche borealis from three streams in northern California, U.S.A.

1. Allozyme electrophoresis was used to describe the genetic structure of Helicopsyche borealis caddisflies collected from three sites in each of three streams in northern California, U.S.A.: the Rice Fork of the Eel River; Big Sulphur Creek; and Alameda Creek. Between 7 and 11km separated adjacent sites within these three streams. Helicopsyche borealis from three additional streams in eastern North America (Christiana Creek, Indiana; Byrd's Mill Creek, Oklahoma; Susquehanna River, Pennsylvania) were also analysed electrophoretically to address taxonomic questions that arose during the study. 2. Four genetically distinct groups of individuals were identified (i.e. Helicopsyche types A, B, C, and D). Lack of interbreeding between sympatric groups (as evidenced by fixed allelic differences) and large genetic differences (mean Nei's genetic distances = 0.396–0.693) indicate that these four groups of Helicopsyche were actually reproductively isolated species rather than genetic variants of a single species. 3. Occurrence of Helicopsyche type A at multiple sites permitted an analysis of spatial variation in genetic structure. Within a drainage basin, small differences in allele frequencies were observed among sites in the Rice Fork and Big Sulphur Creek, but not in Alameda Creek. Larger genetic differences were found among sites in separate drainage basins. Genetic distances (Nei's) between Helicopsyche type A from California and from eastern North America sites (mean = 0.236) were greater than interpopulation differences commonly observed for insects, which suggests that Helicopsyche type A from California may represent a different species than Helicopsyche type A from eastern North America. 4. Geographical and taxonomic differences observed in this study underscore the importance of understanding both population structure and genetic relationships among populations in the design and interpretation of stream faunal studies.