The ups and downs of trophic control in continental shelf ecosystems

Traditionally, marine ecosystem structure was thought to be determined by phytoplankton dynamics. However, an integrated view on the relative roles of top-down (consumer-driven) and bottom-up (resource-driven) forcing in large-scale, exploited marine ecosystems is emerging. Long time series of scientific survey data, underpinning the management of commercially exploited species such as cod, are being used to diagnose mechanisms that could affect the composition and relative abundance of species in marine food webs. By assembling published data from studies in exploited North Atlantic ecosystems, we found pronounced geographical variation in top-down and bottom-up trophic forcing. The data suggest that ecosystem susceptibility to top-down control and their resiliency to exploitation are related to species richness and oceanic temperature conditions. Such knowledge could be used to produce ecosystem guidelines to regulate and manage fisheries in a sustainable fashion.     

BATHYMETRIC PATTERNS OF BODY SIZE IN DEEP-SEA GASTROPODS

The shift to smaller body size in marine invertebrates at the deep-sea threshold and size-depth clines within the deep-sea ecosystem are global biogeographic phenomena that remain poorly understood. We present the first standardized measurements of larval and adult size among ecologically and phylogenetically similar species across a broad and continuous depth range, using the largest family of deep-sea gastropods (the Turridae). Size at all life stages increases significantly with depth from the upper bathyal region to the abyssal plain. These consistent clines may result from selection favoring larger size at greater depths because of its metabolic and competitive advantages. The unusually small size of deep-sea mollusks, in general, may represent an independent evolutionary process that favors invasion by inshore taxa composed of small organisms.     

Ecosystem productivity is associated with bacterial phylogenetic distance in surface marine waters

Understanding the link between community diversity and ecosystem function is a fundamental aspect of ecology. Systematic losses in biodiversity are widely acknowledged but the impact this may exert on ecosystem functioning remains ambiguous. There is growing evidence of a positive relationship between species richness and ecosystem productivity for terrestrial macro‐organisms, but similar links for marine micro‐organisms, which help drive global climate, are unclear. Community manipulation experiments show both positive and negative relationships for microbes. These previous studies rely, however, on artificial communities and any links between the full diversity of active bacterial communities in the environment, their phylogenetic relatedness and ecosystem function remain hitherto unexplored. Here, we test the hypothesis that productivity is associated with diversity in the metabolically active fraction of microbial communities. We show in natural assemblages of active bacteria that communities containing more distantly related members were associated with higher bacterial production. The positive phylogenetic diversity–productivity relationship was independent of community diversity calculated as the Shannon index. From our long‐term (7‐year) survey of surface marine bacterial communities, we also found that similarly, productive communities had greater phylogenetic similarity to each other, further suggesting that the traits of active bacteria are an important predictor of ecosystem productivity. Our findings demonstrate that the evolutionary history of the active fraction of a microbial community is critical for understanding their role in ecosystem functioning.     

Linking the Trophic Fingerprint of Groundfishes to Ecosystem Structure and Function in the California Current

Mean trophic level (MTL) is one of the most widely used indicators of marine ecosystem health. It usually represents the relative abundance of fished species across a spectrum of TLs. The reality, ubiquity, and causes of a general decline in the MTL of fisheries catch through time, and whether fisheries catch tracks ecosystem level changes, have engendered much attention. However, the consequences of such patterns for broader ecosystem structure and function remain virtually unexplored. Along the Pacific U.S. Coast, previous work has documented fluctuations and a slow increase in ecosystem MTL from 1977 to 2004. Here, we document a decline in the ecosystem MTL of groundfishes in the same ecosystem from 2003 to 2011, the proximate cause of which was a decrease in the biomass of higher TL groundfishes. Using a food web model, we illustrate how these shifts in ecosystem structure may have resulted in short term, positive responses by many lower TL species in the broader ecosystem. In the longer term, the model predicts that initial patterns of prey release may be tempered in part by lagged responses of other higher TL species, such as salmon and seabirds. Although ecosystem functions related to specific groups like piscivores (excluding high-TL groundfishes) changed, aggregate ecosystem functions altered little following the initial reorganization of biomass, probably due to functional redundancy within the predator guild. Efforts to manage and conserve marine ecosystems will benefit from a fuller consideration of the information content contained within, and implied by, fisheries-independent TL indicators.     

Management reference points to account for direct and indirect impacts of fishing on marine mammals

Reference points can help implement an ecosystem approach to fisheries management (EAF), by establishing precautionary removal limits for nontarget species and target species of ecological importance. PBR (Potential Biological Removal), developed under the U.S. Marine Mammal Protection Act (MMPA), is a limit for direct mortality for marine mammals, but it does not account for indirect effects of fishing due to prey depletion. I propose a generalization of PBR (called PBR*) to account for plausible changes in marine mammal carrying capacity (ΔK) from prey biomass decline relative to two example benchmarks: SSB_MSY (maximum sustainable yield biomass for all known prey species) or SSB_K (unfished prey biomass). PBR* can help identify when indirect fishing effects (alone, or combination with direct mortality estimates) may stymie MMPA objectives, and could inform catch limit estimates for target species that are also important as marine mammal prey. As a case study, I applied PBR* estimates to evaluate the possible combined direct + indirect effects of fishing on cetaceans in northeastern U.S. waters. Estimated distributions for ΔK were based on fish stock assessments and meta‐analysis of predator‐prey relationships from the mammalian literature. Based on this analysis, increased risk of marine mammal depletion due to indirect fishing effects was not evident, although this result must be interpreted cautiously given our limited understanding of cetacean diets and marine trophic dynamics. This study is intended to illustrate a possible practical approach for incorporating indirect fisheries impacts on marine mammals into a comprehensive management framework, and it raises several scientific and policy issues that merit further investigation.     

Phylogeography of the Chionodraco genus (Perciformes,Channichthydae) in the Southern Ocean

Antarctic fish of the suborder Notothenioidei represent one of the most notable examples of adaptive radiation in the marine environment. The evolutionary relationships between and within the eight families of this suborder have been well established by numerous studies, whereas the microevolutionary processes of notothenioid species remain largely unexplored. In the present paper we investigated the evolutionary relationships between three closely related species of the genus Chionodraco (family Channichthyidae), namely Chionodraco hamatus, Chionodraco rastrospinosus, and Chionodraco myersi by analysing portions of the mitochondrial genome (D-loop and 16S rRNA). The taxonomic status of C. hamatus and C. rastrospinosus as separate species has been questioned because of the limited number of key morphological characters that distinguish these two taxa. Our results, based on the analysis of several specimens belonging to both morphological groups revealed a small genetic differentiation among haplotypes, however, a clear separation between the two nominal species emerged since all individuals of each of the two taxa clustered together in distinct monophyletic groups. C. myersi appeared more distantly related in the phylogenetic analysis. For one species, C. hamatus, sampling was carried out at three different geographic locations in the area of the Ross Sea and Weddell Sea. The results showed that the partition of the genetic variation within this species is not compatible with the hypothesis of panmixia as gene flow between populations was significantly reduced.     

The biodiversity of macrofaunal organisms in marine sediments

Marine sediments cover most of the ocean bottom, and the organisms that reside in these sediments therefore constitute the largest faunal assemblage on Earth in areal coverage. The biomass in these sediments is dominated by macrofauna, a grouping of invertebrate polychaetes, molluscs, crustaceans and other phyla based on size. Globally, only a small portion of marine habitats have been sampled for macrofauna, but sampled areas have led to global estimates of macrofaunal species number ranging from 500,000 to 10,000,000. Most of these species are undescribed, and global syntheses of patterns of individual taxa and biodiversity are few and based on limited samples. The significance of biodiversity in marine sediments to ecosystem processes is poorly understood, but individual species and functional groups are known to carry out activities that have global importance. Macrofaunal activity impacts global carbon, nitrogen and sulphur cycling, transport, burial and metabolism of pollutants, secondary production including commercial species, and transport of sediments. Documented extinctions of marine macrofauna are few, but the ramifications of species loss through habitat shrinkage and undocumented extinctions are unknown. Limited data suggest there is substantial functional redundancy in macrofauna within trophic groups but whether this redundancy is sufficient to allow species loss without significantly altering ecosystem processes is unknown. Sorely needed are experiments that test specific hypotheses on biodiversity, redundancy, and ecosystem processes as they relate to marine macrofauna.     

Genomics detects population structure within and between ocean basins in a circumpolar seabird: The white‐chinned petrel

The Southern Ocean represents a continuous stretch of circumpolar marine habitat, but the potential physical and ecological drivers of evolutionary genetic differentiation across this vast ecosystem remain unclear. We tested for genetic structure across the full circumpolar range of the white‐chinned petrel (Procellaria aequinoctialis) to unravel the potential drivers of population differentiation and test alternative population differentiation hypotheses. Following range‐wide comprehensive sampling, we applied genomic (genotyping‐by‐sequencing or GBS; 60,709 loci) and standard mitochondrial‐marker approaches (cytochrome b and first domain of control region) to quantify genetic diversity within and among island populations, test for isolation by distance, and quantify the number of genetic clusters using neutral and outlier (non‐neutral) loci. Our results supported the multi‐region hypothesis, with a range of analyses showing clear three‐region genetic population structure, split by ocean basin, within two evolutionary units. The most significant differentiation between these regions confirmed previous work distinguishing New Zealand and nominate subspecies. Although there was little evidence of structure within the island groups of the Indian or Atlantic oceans, a small set of highly‐discriminatory outlier loci could assign petrels to ocean basin and potentially to island group, though the latter needs further verification. Genomic data hold the key to revealing substantial regional genetic structure within wide‐ranging circumpolar species previously assumed to be panmictic.     

Phylogeny as a Proxy for Ecology in Seagrass Amphipods: Which Traits Are Most Conserved?

Increasingly, studies of community assembly and ecosystem function combine trait data and phylogenetic relationships to gain novel insight into the ecological and evolutionary constraints on community dynamics. However, the key to interpreting these two types of information is an understanding of the extent to which traits are phylogenetically conserved. In this study, we develop the necessary framework for community phylogenetics approaches in a system of marine crustacean herbivores that play an important role in the ecosystem functioning of seagrass systems worldwide. For 16 species of amphipods and isopods, we (1) reconstructed phylogenetic relationships using COI, 16S, and 18S sequences and Bayesian analyses, (2) measured traits that are potentially important for assembling species between and within habitats, and (3) compared the degree to which each of these traits are evolutionarily conserved. Despite poor phylogenetic resolution for the order Amphipoda as a whole, we resolved almost all of the topology for the species in our system, and used a sampling of ultrametric trees from the posterior distribution to account for remaining uncertainty in topology and branch lengths. We found that traits varied widely in their degree of phylogenetic signal. Body mass, fecundity, and tube building showed very strong phylogenetic signal, and temperature tolerance and feeding traits showed much less. As such, the degree of signal was not predictable based on whether the trait is related to environmental filtering or to resource partitioning. Further, we found that even with strong phylogenetic signal in body size, (which may have large impacts on ecosystem function), the predictive relationship between phylogenetic diversity and ecosystem function is not straightforward. We show that patterns of phylogenetic diversity in communities of seagrass mesograzers could lead to a variety of interpretations and predictions, and that detailed study of trait similarities and differences will be necessary to interpret these patterns.     

Resilience and stability of a pelagic marine ecosystem

The accelerating loss of biodiversity and ecosystem services worldwide has accentuated a long-standing debate on the role of diversity in stabilizing ecological communities and has given rise to a field of research on biodiversity and ecosystem functioning (BEF). Although broad consensus has been reached regarding the positive BEF relationship, a number of important challenges remain unanswered. These primarily concern the underlying mechanisms by which diversity increases resilience and community stability, particularly the relative importance of statistical averaging and functional complementarity. Our understanding of these mechanisms relies heavily on theoretical and experimental studies, yet the degree to which theory adequately explains the dynamics and stability of natural ecosystems is largely unknown, especially in marine ecosystems. Using modelling and a unique 60-year dataset covering multiple trophic levels, we show that the pronounced multi-decadal variability of the Southern California Current System (SCCS) does not represent fundamental changes in ecosystem functioning, but a linear response to key environmental drivers channelled through bottom-up and physical control. Furthermore, we show strong temporal asynchrony between key species or functional groups within multiple trophic levels caused by opposite responses to these drivers. We argue that functional complementarity is the primary mechanism reducing community variability and promoting resilience and stability in the SCCS.     

Ecological roles of zoosporic parasites in blue carbon ecosystems

Pathosystems describe the relationships between parasites, hosts and the environment. Generally these systems remain in a dynamic equilibrium over time. In this review we examine some of the evidence for the potential impacts of change in dynamic equilibrium in blue carbon ecosystems and the relationships to the amount of stored carbon. Blue carbon ecosystems are marine and estuarine ecosystems along the coasts. Virulent pathogens can be introduced into ecosystems along with non-native hosts. Alteration of environmental conditions, such as temperature, pH and salinity, may cause parasites to dominate the pathosystems resulting in significant decreases in productivity and population sizes of producer hosts and in changes in the overall species composition and function in these ecosystems. Such changes in blue carbon ecosystems may result in accelerated release of carbon dioxide back into the ocean and atmosphere, which could then drive further changes in the global climate. The resiliency of these ecosystems is not known. However, recent evidence suggests that significant proportions of blue carbon ecosystems have already disappeared.     

Diversification of unicellular eukaryotes: cryptomonad colonizations of marine and fresh waters inferred from revised 18S rRNA phylogeny

The cryptomonads is a well-defined lineage of unicellular eukaryotes, composed of several marine and freshwater groups. However, the evolutionary relationships among these groups are unclear due to conflicting inferences between morphological and molecular phylogenies. Here, we have inferred the evolutionary relationships among marine and freshwater species in order to better understand the importance of the marine-freshwater boundary on the historical diversification patterns of cryptomonads. We have constructed improved molecular phylogenies by taking into account rate variation both across sites and across sequences (covarion substitutions), and by analysing the vast majority of publicly available cryptomonad 18S rRNA sequences and related environmental phylotypes. The resulting phylogenies included 55 sequences, and revealed two novel freshwater cryptomonad clades (CRY1 and CRY2) and a large hidden diversity of cryptomonads. CRY1 was placed deeply within the cryptomonad phylogeny together with all the major freshwater lineages (i.e. Goniomonas and Cryptomonas), while CRY2 was placed within a lineage of marine species identified as Plagioselmis-like with the aid of a new sequence generated from a cultured species. The inferred phylogenies suggest only few successful marine-freshwater transitions over the history of cryptomonads. Most of the transitions seem to have occurred from marine to fresh waters, but re-colonizations of marine habitats have also taken place. This implies that the differences in the biogeophysical conditions between marine and fresh waters constitute a substantial barrier for the cross-colonization of these environments by cryptomonads.     

Are kelp holdfasts islands on the ocean floor? – indication for temporarily closed aggregations of peracarid crustaceans

During the colonisation process of islands, newly immigrating species often arrive as single individuals. Islands that have received single colonisers may subsequently harbour large populations of a species, while other islands may completely lack this species. Exchange between islands is limited, thereby strongly affecting evolutionary processes. While this concept is widely used in the context of oceanic islands or habitat patches on the mainland, it is rarely used to explain and examine the distribution patterns of marine invertebrates. Benthic marine organisms inhabiting patches with island-like features may also be restricted in their movements between patches. Once established in a patch, it may be more favourable to remain there rather than moving to another patch. Juveniles of species with direct development may recruit to the island patch of their parents. Herein, we examined the peracarid fauna in patches that have island-like features, i.e. kelp holdfasts. The number of peracarid species within an individual holdfast increased with its size. Similarly, the number of individuals per holdfast increased with holdfast size. However, several peracarid species showed a strongly aggregated distribution pattern, being highly abundant in some holdfasts and almost completely absent in others. Our results suggest that these aggregations of conspecifics may be a consequence of the peracarid reproductive biology: fully developed juveniles emerge from the female's marsupium and recruit to the immediate vicinity of their mother, showing little or no tendency to emigrate towards other patches. At present, while it is not known how long peracarid aggregations within kelp holdfasts persist, our data suggest that some juveniles may remain with the natal holdfast and possibly reproduce therein. It is concluded that, during certain time periods, reproduction rates of peracarids in a holdfast may exceed their migration rates between holdfasts.     

Phylogeography of the Chionodraco genus (Perciformes, Channichthydae) in the Southern Ocean

Antarctic fish of the suborder Notothenioidei represent one of the most notable examples of adaptive radiation in the marine environment. The evolutionary relationships between and within the eight families of this suborder have been well established by numerous studies, whereas the microevolutionary processes of notothenioid species remain largely unexplored. In the present paper we investigated the evolutionary relationships between three closely related species of the genus Chionodraco (family Channichthyidae), namely Chionodraco hamatus, Chionodraco rastrospinosus, and Chionodraco myersi by analysing portions of the mitochondrial genome (D-loop and 16S rRNA). The taxonomic status of C. hamatus and C. rastrospinosus as separate species has been questioned because of the limited number of key morphological characters that distinguish these two taxa. Our results, based on the analysis of several specimens belonging to both morphological groups revealed a small genetic differentiation among haplotypes, however, a clear separation between the two nominal species emerged since all individuals of each of the two taxa clustered together in distinct monophyletic groups. C. myersi appeared more distantly related in the phylogenetic analysis. For one species, C. hamatus, sampling was carried out at three different geographic locations in the area of the Ross Sea and Weddell Sea. The results showed that the partition of the genetic variation within this species is not compatible with the hypothesis of panmixia as gene flow between populations was significantly reduced.     

基于16S rDNA部分序列探讨中国近海30种石斑鱼类的分子系统进化关系

石斑鱼因其种类繁多、分布广泛及缺乏显著的形体特征,使其系统分类的研究颇为困难。为探讨中国近海石斑鱼类的系统进化关系,通过PCR扩增获得了石斑鱼亚科(Epinephelinae)6属30个种类的线粒体16SrDNA基因片段序列。采用多个生物软件对序列变异和碱基组成进行分析,计算了Kimura2parameter遗传距离、转换/颠换比等遗传信息指数,并结合GenBank石斑鱼属的同源序列,以多纹长尾(Pronotogrammusmultifasciatus)和皮氏叫姑鱼(Johniusbelengerii)为外群构建NJ、MP和ML系统树。根据所得分子依据并结合形态学特征,推论如下:1)在本研究的30种石斑鱼中,鳃棘鲈属(Plectropomus)最先分化,并呈明显单系性;九棘鲈属(Cephalopholis)是一个单系群,并且较石斑鱼属(Epinephelus)原始;侧牙鲈属(Variola)的进化地位介于鳃棘鲈属与九棘鲈属之间;2)宽额鲈(Promicropslanceolatus)可以归入石斑鱼属,而驼背鲈(Cromileptesaltivelis)也与石斑鱼属有很近的亲缘关系,甚至可能是石斑鱼属内的特化类群;3)石斑鱼属内部存在两个平行进化的姐妹分支,分支内部的种类组成与地理分布无关,暗示了石斑鱼属早期的分化模式。     

High-resolution SAR11 ecotype dynamics at the Bermuda Atlantic Time-series Study site by phylogenetic placement of pyrosequences

Advances in next-generation sequencing technologies are providing longer nucleotide sequence reads that contain more information about phylogenetic relationships. We sought to use this information to understand the evolution and ecology of bacterioplankton at our long-term study site in the Western Sargasso Sea. A bioinformatics pipeline called PhyloAssigner was developed to align pyrosequencing reads to a reference multiple sequence alignment of 16S ribosomal RNA (rRNA) genes and assign them phylogenetic positions in a reference tree using a maximum likelihood algorithm. Here, we used this pipeline to investigate the ecologically important SAR11 clade of Alphaproteobacteria. A combined set of 2.7 million pyrosequencing reads from the 16S rRNA V1–V2 regions, representing 9 years at the Bermuda Atlantic Time-series Study (BATS) site, was quality checked and parsed into a comprehensive bacterial tree, yielding 929 036 Alphaproteobacteria reads. Phylogenetic structure within the SAR11 clade was linked to seasonally recurring spatiotemporal patterns. This analysis resolved four new SAR11 ecotypes in addition to five others that had been described previously at BATS. The data support a conclusion reached previously that the SAR11 clade diversified by subdivision of niche space in the ocean water column, but the new data reveal a more complex pattern in which deep branches of the clade diversified repeatedly across depth strata and seasonal regimes. The new data also revealed the presence of an unrecognized clade of Alphaproteobacteria, here named SMA-1 (Sargasso Mesopelagic Alphaproteobacteria, group 1), in the upper mesopelagic zone. The high-resolution phylogenetic analyses performed herein highlight significant, previously unknown, patterns of evolutionary diversification, within perhaps the most widely distributed heterotrophic marine bacterial clade, and strongly links to ecosystem regimes.     

Uncoupling ecological innovation and speciation in sea snakes (Elapidae, Hydrophiinae, Hydrophiini)

The viviparous sea snakes (Hydrophiini) are by far the most successful living marine reptiles, with ～ 60 species that comprise a prominent component of shallow-water marine ecosystems throughout the Indo-West Pacific. Phylogenetically nested within the ～ 100 species of terrestrial Australo-Melanesian elapids (Hydrophiinae), molecular timescales suggest that the Hydrophiini are also very young, perhaps only ～ 8-13 Myr old. Here, we use likelihood-based analyses of combined phylogenetic and taxonomic data for Hydrophiinae to show that the initial invasion of marine habitats was not accompanied by elevated diversification rates. Rather, a dramatic three to six-fold increase in diversification rates occurred at least 3-5 Myr after this transition, in a single nested clade: the Hydrophis group accounts for ～ 80% of species richness in Hydrophiini and ～ 35% of species richness in (terrestrial and marine) Hydrophiinae. Furthermore, other co-distributed lineages of viviparous sea snakes (and marine Laticauda, Acrochordus and homalopsid snakes) are not especially species rich. Invasion of the oceans has not (by itself) accelerated diversification in Hydrophiini; novelties characterizing the Hydrophis group alone must have contributed to its evolutionary and ecological success.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Diversification of sympatric broadcast-spawning limpets (Cellana spp.) within the Hawaiian archipelago

Speciation remains a central enigma in biology, and nowhere is this more apparent than in shallow tropical seas where biodiversity rivals that of tropical rainforests. Obvious barriers to gene flow are few and most marine species have a highly dispersive larval stage, which should greatly decrease opportunities for speciation via geographic isolation. The disparity in the level of geographic isolation for terrestrial and marine species is exemplified in Hawai'i where opportunities for allopatric speciation abound in the terrestrial realm. In contrast, marine colonizers of Hawai'i are believed to produce only a single endemic species or population, due to the lack of isolating barriers. To test the assertion that marine species do not diversify within Hawai'i, we examine the evolutionary origin of three endemic limpets (Cellana exarata, C. sandwicensis and C. talcosa) that are vertically segregated across a steep ecocline on rocky shores. Analyses of three mtDNA loci (12S, 16S, COI; 1565bp) and two nDNA loci (ATPSβ, H3; 709bp) in 26 Indo-Pacific Cellana species (N=414) indicates that Hawai'i was colonized once ～3.4-7.2Ma from the vicinity of Japan. Trait mapping demonstrates that high-shore residence is the ancestral character state, such that mid- and low-shore species are the product of subsequent diversification. The Hawaiian Cellana are the first broadcast-spawners demonstrated to have speciated within any archipelago. The habitat stratification, extensive sympatry, and evolutionary history of these limpets collectively indicate a strong ecological component to speciation and support the growing body of evidence for non-allopatric speciation in the ocean.     

SYSTEMATIC EVALUATION OF THE GENUS HILDENBRANDIA (RHODOPHYTA): A SYNTHESIS OF TECHNIQUES

The evolutionary relationships among members of the red algal genus Hildenbrandia have not been well understood for several reasons. For example, the genus contains both marine and freshwater representatives, all of which are non-calcified and crustose, and few have definitive morphological characters for classification. Hildenbrandia is also assumed to be completely asexual (reproduction by tetrasporangia in marine forms and by gemmae in freshwater populations), and characters of the female gametangial system and post-fertilization structures are not available for comparative studies. Currently there are 14 marine and five freshwater species and infraspecific taxa recognized within the genus. We used phylogenetic analyses (parsimony, distance and maximum likelihood) of DNA sequences of commonly employed genes ( rbc L and 18S rRNA) to examine the evolutionary relationships among representatives of many of these taxa. In addition, we employed morphometrics (principal co-ordinates and cluster analyses) of several measured characters of these same representatives, as well as all available type specimens, to determine the number of morphologically-delimited entities within the genus. Thus far our results indicate that some characters traditionally used to distinguish species of Hildenbrandia , such as tetrasporangial division pattern, may not be useful in some cases, and a revision of the taxonomy of the genus will be necessary. Although the marine and freshwater species of Hildenbrandia appear to be well separated in our molecular analyses of European specimens, this trend was not observed for North American specimens. High sequence divergence values were calculated for both the rbc L and 18S rRNA genes of Hildenbrandia , compared to other red algal genera.