Genetic Diversity and Tests of the Hybrid Origin of the Endangered Yellow Larkspur

Abstract: Delphinium luteum ( Ranunculaceae), an endangered larkspur, is restricted to two wild populations near Bodega Bay, California. The total number of individuals in these two populations is estimated to be <100. We used allozyme and random amplified polymorphic DNA ( RAPD) markers to (1) assess levels and patterns of genetic diversity in one wild population and two cultivated populations and (2) test the hypothesis that D. luteum is of hybrid origin between D. decorum and D. nudicaule . These data will be used to aid in developing a management plan to conserve the species. The wild population maintains high levels of genetic diversity. Genetic data indicate that both cultivated populations, especially the north Sonoma population, have several allozymes and RAPD markers not found in the wild population and could be used to establish new populations of D. luteum or to enhance the diversity and size of the wild population. The allozyme data did not reveal any fixed differences between D. decorum and D. nudicaule , although allele frequencies of the putative parental populations differed. At these loci, D. luteum resembled D. nudicaule more than D. decorum  . Many unique RAPD markers distinguish each of the three species. The diagnostic markers from populations of D. nudicaule and D. decorum were not additive in the putative hybrid, and these data indicate that D. luteum is not of recent hybrid origin. Conservation of the yellow larkspur should include strategies that use the cultivated populations of D. luteum , but hybridizing D. decorum and D. nudicaule to "recreate" D. luteum is not recommended.     

Demographic Stability Metrics for Conservation Prioritization of Isolated Populations

Abstract:  Systems of geographically isolated habitat patches house species that occur naturally as small, disjunct populations. Many of these species are of conservation concern, particularly under the interacting influences of isolation and rapid global change. One potential conservation strategy is to prioritize the populations most likely to persist through change and act as sources for future recolonization of less stable localities. We propose an approach to classify long-term population stability (and, presumably, future persistence potential) with composite demographic metrics derived from standard population-genetic data. Stability metrics can be related to simple habitat measures for a straightforward method of classifying localities to inform conservation management. We tested these ideas in a system of isolated desert headwater streams with mitochondrial sequence data from 16 populations of a flightless aquatic insect. Populations exhibited a wide range of stability scores, which were significantly predicted by dry-season aquatic habitat size. This preliminary test suggests strong potential for our proposed method of classifying isolated populations according to persistence potential. The approach is complementary to existing methods for prioritizing local habitats according to diversity patterns and should be tested further in other systems and with additional loci to inform composite demographic stability scores.     

Facilitation drives local abundance and regional distribution of a rare plant in a harsh environment

The importance of facilitation to local community dynamics is becoming increasingly recognized. However, the predictability of positive interactions in stressful environments, the balance of competition and facilitation along environmental gradients, and the scaling of local positive interactions to regional distributions are aspects of facilitation that remain unresolved. I explored these questions in a habitat specialist, Delphinium uliginosum, and a moss, Didymodon tophaceus, both found in small serpentine wetlands. I tested three hypotheses: (1) moss facilitates germination, growth, and/or fecundity of D. uliginosum; (2) facilitation is stronger at the harsher ends of gradients in soil moisture, toxicity, and/or biomass; and (3) facilitation is reflected in positive associations at the levels of local abundance and regional occurrence. Although considerable competitive interactions occurred in later life stages, moss strongly facilitated D. uliginosum seedling emergence. There was no evidence that this facilitative effect weakened, or switched to competition, in benign environments. D. uliginosum was more locally abundant and more frequently present, across a large portion of its range, with than without moss, indicating a net facilitative effect in the face of competitive influences. Facilitated recruitment, possibly by seed retention, was found to be an important control on abundance and distribution in this rare species.     

Evaluating the Effectiveness of Corridors: a Genetic Approach

Abstract: The effectiveness of corridors in maintaining dispersal in fragmented landscapes is a question of considerable conservation and ecological importance. We tested the efficacy of corridors as residual landscape structures in maintaining population structure in the red-backed vole ( Clethrionomys gapperi ), a closed-canopy specialist, and the deer mouse (   Peromyscus maniculatus ), a habitat generalist. In coniferous forests managed for timber production in northeastern Washington, we sampled pairs of populations in three landscape classes: (1) contiguous landscapes, in which sites were located completely within a matrix of closed-canopy forest; (2) corridor landscapes, in which sites were connected by a corridor of closed-canopy forest; and (3) isolated landscapes, in which sites were separated from one another by clearcut or young regeneration stands. For each species, we used four microsatellite loci to quantify genetic distance between population pairs. Nei's genetic distance (   D _s  ) increased from smallest to largest in the order of contiguous, corridor, and isolated landscapes for C. gapperi. For P. maniculatus, genetic distances across landscape configurations were not significantly different. The differences between the two species indicate that they respond differently to the presence of forest corridors. In managed forests, corridors between unlogged habitats appear to maintain higher population connectivity for C. gapperi than landscapes without corridors.     

Extirpation and Recolonization in a Metapopulation of an Endangered Fish, the Tidewater Goby

Abstract: The tidewater goby ( Eucyclogobius newberryi ), an endangered species in the United States, occurs in a series of isolated coastal wetlands in California. Using historical presence-absence data and our own surveys, we estimated annual rates of extirpation and recolonization for several populations of the goby in southern California. As predicted, large wetlands had lower rates of extirpation than small wetlands. There was a negative but statistically nonsignificant correlation between recolonization rate and distance to the nearest northerly source population. Populations at small sites were sensitive to drought, presumably because droughts can eliminate suitable habitat at small wetlands. Populations in small wetlands have declined over time, even after accounting for variation in stream flow, supporting the species' endangered status. Our study emphasizes the need to understand metapopulation dynamics for conserving species where the unit of conservation is a local population. It is also emphasizes the importance of not treating metapopulations as identical units. Finally, our results provide a means for describing the decline of a species that is complex in time and space and provide insight into how to target protection measures among metapopulations.     

Developing a Sampling Strategy for Baptisia arachnifera Based on Allozyme Diversity

We analyzed the amount and distribution of genetic variation in Baptisia arachnifera Duncan to develop a sampling strategy for ex situ research. Baptisia arachnifera is an endangered plant species endemic to the coastal plain of Georgia (U.S.) where all populations are within 16 km of each other. A reduction in numbers of individuals has been observed during the last 50 years. Baptisia arachnifera was polymorphic at 24% of the 37 loci examined with an average of 1.32 alleles per locus. The genetic diversity index was relatively low ( H_e = 0.097) as expected for endemic species. Populations were in Hardy-Weinberg equilibrium, suggesting that the species is outcrossing. Consistent with this conclusion is the observation that the majority (approximately 90%) of the genetic variation present in the species is found within individual populations. Indirect evidence of gene flow between populations was detected (   Nm = 2.35). The close proximity of the populations and the recent reduction in population sizes suggest that the populations surveyed may be fragments of a once more continuous gene pool. Based on the observed distribution of genetic diversity among populations (G_ST = 0.096), sampling two populations would capture 99% of the allozyme diversity surveyed. Allozyme data were used to determine which 2 of the 10 populations surveyed should be sampled to maximize the ex situ conservation of genetic diversity. Although the paper-producing companies that own most of the land where Baptisia arachnifera occurs are modifying their harvesting techniques, the species could become extinct without more effective management and preservation efforts.     

Conservation Value of Sites of Hybridization in Peripheral Populations of Rare Plant Species

Abstract:  Populations at the periphery of a species' range are of interest to conservation biologists because they can show marked genetic differentiation from populations at the center of a range and because of potential hybridization among rare and common species. We examined two closely related Cyclamen species. One is a narrow endemic, and the other is more geographically widespread (both protected by law in continental southern France). We used floral traits and genetic variability to test for hybridization among the species in peripheral populations of the rare species. The species co-occurred on Corsica in a disjunct, peripheral part of the distribution of the endemic species and in an ecologically marginal area for the widespread species. The two species have hybridized and the endemic species showed high levels of introgression with its widespread congener. Genetic and floral variability in sites with both species was markedly higher than in sites with a single species. Our results highlight the need for a conservation strategy that integrates hybrid populations because they represent a source of novel diversity that may have adaptive potential.     

Epidemiologic Analysis of Factors Associated with Local Disappearances of Native Ranid Frogs in Arizona

Abstract:  We examined factors that may independently or synergistically contribute to amphibian population declines. We used epidemiologic case–control methodology to sample and analyze a large database developed and maintained by the Arizona Game and Fish Department that describes historical and currently known ranid frog localities in Arizona, U.S.A. Sites with historical documentation of target ranid species ( n = 324) were evaluated to identify locations where frogs had disappeared during the study period (case sites) and locations where frog populations persisted (control sites). Between 1986 and 2003, 117 (36%) of the 324 sites became case sites, of which 105 were used in the analyses. An equal number of control sites were sampled to control for the effects of time. Risk factors, or predictor variables, were defined from environmental data summarized during site surveys and geographic information system data layers. We evaluated risk factors with univariate and multifactorial logistic-regression analyses to derive odds ratios (OR). Odds for local population disappearance were significantly related to 4 factors in the multifactorial model. Disappearance of frog populations increased with increasing elevation (OR = 2.7 for every 500 m, p < 0.01). Sites where disappearances occurred were 4.3 times more likely to have other nearby sites that also experienced disappearances (OR = 4.3, p < 0.01), whereas the odds of disappearance were 6.7 times less (OR = 0.15, p < 0.01) when there was a source population nearby. Sites with disappearances were 2.6 times more likely to have introduced crayfish than were control sites (OR = 2.6, p = 0.04). The identification of factors associated with frog disappearances increases understanding of declines occurring in natural populations and aids in conservation efforts to reestablish and protect native ranids by identifying and prioritizing implicated threats.     

Demographic Evidence of Illegal Harvesting of an Endangered Asian Turtle

Harvesting pressure on Asian freshwater turtles is severe, and dramatic population declines of these turtles are being driven by unsustainable collection for food markets, pet trade, and traditional Chinese medicine. Populations of big‐headed turtle (Platysternon megacephalum) have declined substantially across its distribution, particularly in China, because of overcollection. To understand the effects of chronic harvesting pressure on big‐headed turtle populations, we examined the effects of illegal harvesting on the demography of populations in Hong Kong, where some populations still exist. We used mark‐recapture methods to compare demographic characteristics between sites with harvesting histories and one site in a fully protected area. Sites with a history of illegal turtle harvesting were characterized by the absence of large adults and skewed ratios of juveniles to adults, which may have negative implications for the long‐term viability of populations. These sites also had lower densities of adults and smaller adult body sizes than the protected site. Given that populations throughout most of the species’ range are heavily harvested and individuals are increasingly difficult to find in mainland China, the illegal collection of turtles from populations in Hong Kong may increase over time. Long‐term monitoring of populations is essential to track effects of illegal collection, and increased patrolling is needed to help control illegal harvesting of populations, particularly in national parks. Because few, if any, other completely protected populations remain in the region, our data on an unharvested population of big‐headed turtles serve as an important reference for assessing the negative consequences of harvesting on populations of stream turtles. Evidencia Demográfica de la Captura Ilegal de una Tortuga Asiática en Peligro     

DNA Fingerprinting Reveals a Lack of Genetic Variation in Northern Populations of the Western Pond Turtle (Clemmys marmorata)

I used DNA fingerprinting to provide the first analysis of the genetic composition of western pond turtle ( Clemmys marmorata ) populations in Washington, Oregon, and California. Populations of the western pond turtle in Washington and northern Oregon are rapidly approaching extinction. Genetic similarity within the largest northern populations, which are located inland, is high. An analysis of population substructure (F_st) revealed significant genetic divergence between inland populations, indicating a lack of dispersal and gene flow between sites. In contrast, northern coastal sites are not genetically distinct, but there are few if any viable populations remaining in this region. Genetic variability within southern California populations is a great deal higher than in northern inland sites. Similarly, a low F_st value indicated a lack of genetic differentiation between southern sites. An inter-regional analysis of population substructure (F_st = 0.24) revealed a significant degree of genetic divergence between geographical regions throughout the range. In addition, an estimate of western pond turtle phylogeny showed a genetic break in the species between northern and southern populations. Both population subdivision and phylogenetic analyses suggest a lack of appreciable gene flow between geographical regions for a considerable period of time. Genetic analyses support traditional subdivision based solely on the morphological variation of Clemmys marmorata into two subspecies: northern Clemmys marmorata marmorata and southern Clemmys marmorata pallida . Recovery of dwindling northern populations must combine demographic and genetic considerations. A first step should be to preserve local gene pools while augmenting population numbers, with the goal of preventing the extinction of this genetically and morphologically distinct subspecies.     

Effects of Widespread Fish Introductions on Paedomorphic Newts in Europe

Abstract:  As a result of factors such as global warming, habitat destruction, and species introduction, amphibians are declining worldwide. No one, however, has analyzed the status of polymorphic amphibian species at a national or continental scale, although some local reports exist. Our aim was to report on the loss of intraspecific heterochrony as a loss to diversity in determining the consequences of fish stocking on European populations of paedomorphic newts. Paedomorphosis is a polymorphism in which larval traits are retained in the adult stage. We surveyed 39 paedomorphic populations of the alpine (  Triturus alpestris ) and palmate ( T. helveticus ) newts, all but one of which initially occupied fishless ponds and lakes in France, Italy, Slovenia, Bosnia, Montenegro, and Greece. Exotic fishes were found in 44% of the studied aquatic habitats, with a 100% presence in Montenegro. At all sites paedomorphs disappeared and metamorphs declined. Only fish explained these population changes because alternative factors such as drying were not significant. More catastrophically, fish introductions occurred in habitats known to support the largest populations of newts and even some endemic subspecies. If management and legislative measures are not taken to stop fish stocking, protect paedomorphs as conservation units at national and international levels, and restore natural habitats, all the largest paedomorphic populations may disappear in the near future. Their disappearance would represent a loss of one of the rare, fascinating examples of intraspecific heterochrony.     

Microsatellite Analysis of Population Structure in the Endangered Hawaiian Monk Seal

Abstract: The endangered Hawaiian monk seal breeds at six locations in the northwestern Hawaiian Islands. To determine whether significant genetic differentiation exists among these sites, we used microsatellite loci to examine the monk seal population structure at the five largest breeding colonies. Of 27 loci isolated from other seal species, only 3 were polymorphic in an initial screening of one individual from each breeding site. Only two alleles were found at each of these 3 loci in samples of 46–108 individuals. This extremely low variation is consistent with other measures of genetic variability in this species and is probably the result of a recent severe population bottleneck, combined with a long-term history of small population sizes. Although the smallest monk seal subpopulation in this study ( Kure Atoll) showed some evidence of heterozygote deficit, possibly indicative of inbreeding, the next smallest ( Pearl and Hermes Reef) had an apparent excess of heterozygous individuals. Genetic differentiation was detected between the two subpopulations at extreme ends of the range ( Kure and French Frigate Shoals). This trend was significant only at the microsatellite locus for which we had the largest sample size ( Hg6.3: R _ST = 0.206, p = 0.002; allelic goodness of fit G _h = 15.412, p < 0.005). French Frigate Shoals is the source population for translocated animals that have been released primarily at Kure Atoll. Differentiation between these sites consisted of allele frequency differences (with the same allele predominant in each location at all three loci), rather than the preservation of alternative alleles. Although the translocations have had positive demographic effects, we recommend continued genetic monitoring of both the source and recipient populations because translocated individuals are now entering the breeding population.     

Genetic Variability in mtDNA of the Silvery Gibbon: Implications for the Conservation of a Critically Endangered Species

Abstract: The silvery gibbon (   Hylobates moloch ), endemic to the island of Java, relies on closed-canopy, lowland evergreen forest for its survival. Because Java has lost over 91% of its original forest, silvery gibbons currently occupy small, isolated forest fragments and are threatened with extinction. To contribute to a comprehensive conservation strategy for this species, we analyzed the mtDNA control region of 31 silvery gibbons representing most remaining populations. Our results suggest the presence of at least two genetically differentiated lineages: a "western" lineage, represented by the largest remaining natural population in Gunung Halimun National Park and a "central" lineage, consisting of smaller, more isolated populations in and around the Gunung Masigit/Simpang/Tilu complex, Gunung Gede/Pangrango, and Gunung Slamet. These two lineages, at a minimum, represent different management units that should, except in the most dire circumstances, be managed separately.     

Population Structure and Cryptic Evolutionary Units in the Alligator Snapping Turtle

The alligator snapping turtle (  Macroclemys temminckii ) is a long-lived, slow-growing chelydrid turtle found in Gulf of Mexico drainages from Florida to Texas (U.S.A.). Populations are thought to be depleted throughout the range due in part to an increased harvest in the 1960s through 1980s. To identify population and evolutionary units, 420 base pairs were sequenced within the mitochondrial DNA control region of 158 specimens from 12 drainages. Results indicate substantial phylogeographic structuring and strong population-level separations among river drainages. Eight of 11 haplotypes were observed to be river-specific, providing diagnostic markers for most drainages. Three partitions are resolved in the mtDNA genealogy, corresponding to the eastern, central, and western portion of the species' range. These separations coincide with recognized biogeographic provinces. The population structure by river system indicates that many drainages are distinct management units, with the Suwannee River lineage possibly deserving special attention, based on the criterion of genetic distinctiveness. The partitioning of M. temminckii into river-specific populations illustrates the management framework and conservation challenges that apply to a broad array of riverine species. Drainage-specific molecular markers may be used to identify the geographic origin of turtle products in the marketplace.     

Conservation of Species in Dynamic Landscapes: Divergent Fates of Silene tatarica Populations in Riparian Habitats

Abstract:  In transient environments, where local extinctions occur as a result of destruction or deterioration of the local habitat, the long-term persistence of a species requires successful colonizations at new, suitable sites. This kind of habitat tracking should be associated with the asynchronous dynamics of local populations, and it can be especially important for the conservation of rare plant species in riparian habitats. We determined spatiotemporal variation in the demography of the perennial Silene tatarica (L.) Pers. in 15 populations (1998–2003) located in periodically disturbed riparian habitats. The habitats differed according to their morphology (flat shores, slopes) and the amount of bare ground (open, intermediate, closed) along a successional gradient. We used elasticity and life-table response analyses and stochastic simulations to study the variation in population demography. Finite population growth rate was higher in intermediate habitats than in open and closed habitats. In stochastic simulations population size increased in most cases, but four populations were projected to become extinct within 12–70 years. The viability of local populations depended most on the survival and growth of juvenile individuals and on the fecundity of large fertile individuals. On a regional scale, the persistence of this species will require a viable network of local populations as protection against local extinctions caused by natural disturbances and succession. Accordingly, the long-term persistence of riparian species may depend on habitat changes; thus, their conservation requires maintenance of natural disturbance dynamics. Along regulated rivers, management activities such as the creation of open habitats for new colonization should be implemented. Similarly, these activities can be rather general requirements for the conservation of endangered species dependent on transient habitats along successional gradients.     

Unprecedented Low Levels of Genetic Variation and Inbreeding Depression in an Island Population of the Black-Footed Rock-Wallaby

Abstract: It has been argued that demographic and environmental factors will cause small, isolated populations to become extinct before genetic factors have a significant negative impact. Islands provide an ideal opportunity to test this hypothesis because they often support small, isolated populations that are highly vulnerable to extinction. To assess the potential negative impact of isolation and small population size, we compared levels of genetic variation and fitness in island and mainland populations of the black-footed rock-wallaby ( Petrogale lateralis [Marsupialia: Macropodidae]). Our results indicate that the Barrow Island population of P. lateralis has unprecedented low levels of genetic variation (  H _e = 0.053, from 10 microsatellite loci) and suffers from inbreeding depression (reduced female fecundity, skewed sex ratio, increased levels of fluctuating asymmetry). Despite a long period of isolation ( ∼ 1600 generations) and small effective population size (  N _e ∼ 15), demographic and environmental factors have not yet driven this population to extinction. Nevertheless, it has been affected significantly by genetic factors. It has lost most of its genetic variation and become highly inbred (  F _e = 0.91), and it exhibits reduced fitness. Because several other island populations of P. lateralis also exhibit exceptionally low levels of genetic variation, this phenomenon may be widespread. Inbreeding in these populations is at a level associated with high rates of extinction in populations of domestic and laboratory species. Genetic factors cannot then be excluded as contributing to the extinction proneness of small, isolated populations.     

Genetic differentiation and reproductive incompatibility among Baja California populations of the copepod Tigriopus californicus

Populations of the harpacticoid copepod Tigriopus californicus were sampled from five sites from San Diego, California, to Playa Altamira, Baja California, Mexico. Allozyme analyses revealed that all the study populations are sharply differentiated genetically. At the extreme, two populations, Punta Baja and Playa Altamira, have no alleles in common at the seven allozyme loci studied. All pairwise interpopulation crosses successfully produced F₂ hybrids except those involving the Playa Altamira population. All crosses using Playa Altamira females failed to produce F₁ hybrids, while Playa Altamira males successfully produced F₁ progeny with females from all other sites. These F₁ offspring, however, were completely sterile (with San Diego and Punta Banda females) or only occasionally produced F₂ offspring (with Punta Morro and Punta Baja females). These results suggest that allopatric differentiation among Baja populations has resulted in exceptionally high levels of genetic divergence and nearly complete reproductive isolation of the Playa Altamira population, which should now be recognized at the semispecies (or perhaps sibling species) level.     

Experimental interbreeding between geographically separated populations of the marine wood-boring isopod Limnoria tripunctata with preliminary indications of hybrid vigor

Populations of the marine wood-boring isopod Limnoria tripunctata Menzies were cultivated and interbred in the laboratory in order to determine whether geographically separated populations would interbreed. Two series of interbreeding studies were conducted. In the first, field populations from 11 Atlantic and Caribbean sites and 3 Pacific sites were interbred with a North Western Atlantic population from Beaufort, North Carolina (USA). Viable crossings, to at least F ₁, occurred in all but one case—that of a BeaufortxChatham (Massachusetts, USA) cross. In the second series of experiments, field populations from 9 Atlantic and Caribbean sites were crossed with populations from St. Teresa, Florida, Gulf of Mexico. In this case also, all crosses except one produced viable offspring, not only to the F ₁, but to the F ₂ generations as well. The St. Teresa x Chatham cross was not successful. Geographic distance was not a factor in the success of crossbreeding between disjunct populations. Chatham populations (mainly) are now believed to represent a valid but previously obscure species, Limnoria tuberculata Sowinski, and not a variant of L. tripunctata Menzies. Enhanced viality or vigor, as determined from rate of wood destruction, population growth rate and survival, was indicated in several crossings, but this was not necessarily correlated with distance between populations. Observations were made on the manner in which the species has achieved a worldwide distribution, and a hypothesis is put forward to explain the evolution of species related to L. tripunctata.Contribution from Zoology Department, Duke University, Durham, North Carolina, and Duke University Marine Laboratory, Beaufort, North Carolina, and the Department of Oceanography, No. 289, Florida State University, Tallahassee, Florida. Support in part from NSF Grant No. GB-6898.     

<Emphasis Type="Italic">Acropora austera</Emphasis> connectivity in the south-western Indian Ocean assessed using nuclear intron sequence data

The hypervariable carbonic anhydrase 3/550 intron marker was sequenced in order to ascertain the levels of genetic variability and connectivity within and between reefal populations of the hard coral, Acropora austera, on the south-east African coastline. Populations were sampled from (north to south) Bazaruto and Inhaca islands (Mozambique), Rabbit Rock, Two-mile and Red Sands Reefs and Leadsman Shoal (Maputaland, South Africa). Populations at Inhaca Island contained two private alleles, part of the only monophyletic clade with fixed differences between populations in this study. Haplotype and nucleotide diversity were higher in the north of the study area. Indices of migration and haplotype sharing supported significant connectivity between populations in South Africa and Mozambique, which may be important in sustaining genetic diversity in the down-current South African A. austera populations. Measures of population subdivision indicated a significant amount of fixation of allele frequencies amongst populations. Although fine, such differentiation in a marker from the nuclear genome of a hard coral is consistent with some demographic isolation between A. austera populations in southern Mozambique and South Africa. Populations at Rabbit Rock and Inhaca Island were found to be significantly isolated from, and thus less connected to, A. austera populations at other reefs.     

Population structure of the widely dispersing marine bryozoan Membranipora membranacea (Cheilostomata): implications for population history, biogeography, and taxonomy

Morphologically plastic, cryptic, or geographically widespread species pose similar challenges to the evolutionary biologist: their taxonomic status is often unclear yet must be known to study almost any aspect of their biology, ecology, evolution, or biogeography. The marine bryozoan Membranipora membranacea (L.) is morphologically plastic and geographically widespread in temperate oceans of the Northern and Southern Hemispheres, and its taxonomy is unclear. This study examined genetic relationships among allopatric populations and sympatric morphs of this species, or species complex. Colonies were collected from 1992 to 1995. Allozymes were used to elucidate the relationships among four widely separated populations, two in the North Atlantic and two in the North Pacific Ocean. Allozymes and mtDNA sequencing were used to clarify the genetic relationships among three sympatric morphs that might correspond to the species M. villosa Hincks and M. membranacea in the northeastern Pacific (Washington State). Populations in the North Atlantic and North Pacific had no fixed allelic differences at the loci tested but were separated by an average Nei's genetic distance of 0.581, suggesting their near-sibling species status. Populations from Friday Harbor (Washington) and Catalina Island (California) were not significantly differentiated, which was attributed to high gene flow. Populations on either side of the North Atlantic were genetically indistinguishable, which is most likely due to the recent establishment of the West Atlantic populations from European founders. At Friday Harbor, sympatric morphs varying in their spination and spine inducibility were genetically indistinguishable, supporting the hypothesis that M. villosa is an induced phenotype of M. membranacea and not a distinct species in the northeastern Pacific. Since such phenotypic plasticity is common in cheilostome bryozoans, the morphospecies concept must be used with caution. Received: 31 August 1998 / Accepted: 10 August 1999