Genetic analysis of the endemic island loggerhead shrike, <Emphasis Type="Italic">Lanius ludovicianus anthonyi</Emphasis>

Based on limited research, the island loggerhead shrike, Lanius ludovicianus anthonyi has been considered a distinct subspecies endemic to the northern California Channel Islands. We used mtDNA control region sequences and microsatellite genotyping to compare loggerhead shrikes from the southern California mainland (L. l. gambeli), San Clemente Island (L. l. mearnsi), and the northern islands (L. l. anthonyi). Habitats on the islands are recovering due to the removal of non-native ungulates on the islands, but may be transitioning to habitats less supportive of loggerhead shrikes, so this evaluation comes at a critical time. We utilized 96 museum specimens that were collected over a century to evaluate both spatial and temporal genetic patterns. Analysis of multilocus microsatellite genotypes indicated that historical specimens of loggerhead shrikes (collected between 1897 and 1986) from the two northern islands of Santa Rosa and Santa Cruz are genetically distinct from adjacent mainland and island shrikes. Birds from Santa Catalina Island showed mixed ancestry and did not cluster with the northern island birds. Historical specimens of L. l. mearnsi from San Clemente Island also showed mixed ancestry. Our study provides evidence that a genetically distinct form of loggerhead shrikes, L. l. anthonyi, occurred on the islands of Santa Rosa and Santa Cruz.     

Population structure of loggerhead shrikes in the California Channel Islands

The loggerhead shrike (Lanius ludovicianus), a songbird that hunts like a small raptor, maintains breeding populations on seven of the eight California Channel Islands. One of the two subspecies, L. l. anthonyi, was described as having breeding populations on six of the islands while a second subspecies, L. l. mearnsi, was described as being endemic to San Clemente Island. Previous genetic studies have demonstrated that the San Clemente Island loggerhead shrike is well differentiated genetically from both L. l. anthonyi and mainland populations, despite the fact that birds from outside the population are regular visitors to the island. Those studies, however, did not include a comparison between San Clemente Island shrikes and the breeding population on Santa Catalina Island, the closest island to San Clemente. Here we use mitochondrial control region sequences and nuclear microsatellites to investigate the population structure of loggerhead shrikes in the Channel Islands. We confirm the genetic distinctiveness of the San Clemente Island loggerhead shrike and, using Bayesian clustering analysis, demonstrate the presence and infer the source of the nonbreeding visitors. Our results indicate that Channel Island loggerhead shrikes comprise three distinct genetic clusters that inhabit: (i) San Clemente Island, (ii) Santa Catalina Island and (iii) the Northern Channel Islands and nearby mainland; they do not support a recent suggestion that all Channel Island loggerhead shrikes should be managed as a single entity.     

Using GIS mapping of the extent of nearshore rocky reefs to estimate the abundance and reproductive output of important fishery species

Kelp Bass (Paralabrax clathratus) and California Sheephead (Semicossyphus pulcher) are economically and ecologically valuable rocky reef fishes in southern California, making them likely indicator species for evaluating resource management actions. Multiple spatial datasets, aerial and satellite photography, underwater observations and expert judgment were used to produce a comprehensive map of nearshore natural rocky reef habitat for the Santa Monica Bay region (California, USA). It was then used to examine the relative contribution of individual reefs to a regional estimate of abundance and reproductive potential of the focal species. For the reefs surveyed for fishes (i.e. 18 out of the 22 in the region, comprising 82% the natural rocky reef habitat <30 m depth, with a total area of 1850 ha), total abundance and annual egg production of California Sheephead were 451 thousand fish (95% CI: 369 to 533 thousand) and 203 billion eggs (95% CI: 135 to 272 billion). For Kelp Bass, estimates were 805 thousand fish (95% CI: 669 to 941 thousand) and 512 billion eggs (95% CI: 414 to 610 billion). Size structure and reef area were key factors in reef-specific contributions to the regional egg production. The size structures of both species illustrated impacts from fishing, and results demonstrate the potential that relatively small increases in the proportion of large females on larger reefs could have on regional egg production. For California Sheephead, a substantial proportion of the regional egg production estimate (>30%) was produced from a relatively small proportion of the regional reef area (c. 10%). Natural nearshore rocky reefs make up only 11% of the area in the newly designated MPAs in this region, but results provide some optimism that regional fisheries could benefit through an increase in overall reproductive output, if adequate increases in size structure of targeted species are realized.     

Dictyota cyanoloma (Dictyotales,Phaeophyceae), a Newly Introduced Brown Algal Species in California

Here, we report for the first time the presence of Dictyota cyanoloma in southern California. Dictyota cyanoloma is conspicuous in harbors and bays by its distinctive bright blue‐iridescent margins. This species was originally described from Europe, but subsequent studies have revealed that it represented an introduction from Australia. The current distribution of D. cyanoloma comprises southern Australia and the North East Atlantic, including the Mediterranean Sea and the Macaronesian islands. The presence of D. cyanoloma in southern California is supported by molecular cox1 and psbA gene sequences. A reconstruction of the invasive history based on nine polymorphic microsatellite markers reveals a close affinity of the Californian specimens with European populations. Dictyota cyanoloma in the United States appears to be (so far) restricted to the Californian coast from San Diego Bay in the south to Santa Catalina Island and Long Beach Harbor in the north. A correlative species distribution model suggests gradually declining habitat suitability north of the Southern Californian Bight and high suitability in Baja California, including the Gulf of California. Finally, its widespread abundance in bays and harbors suggests shipping is a likely transport mechanism.     

Population genetic structure of the round stingray Urobatis halleri (Elasmobranchii: Rajiformes) in southern California and the Gulf of California

The round stingray, Urobatis halleri, is a viviparous elasmobranch that inhabits inshore, benthic habitats ranging from the western U.S.A. to Panama. The population genetic structure of this species was inferred with seven polymorphic microsatellite loci in samples collected at three sites in coastal southern California, one near Santa Catalina Island, California and one in the eastern Gulf of California. Urobatis halleri is relatively common, but little is known of its movement patterns or population structure. Small F_ST values (?0·0017 to 0·0005) suggested little structure among coastal populations of southern and Baja California. The population sampled at Santa Catalina Island, which is separated by a deep‐water channel from the coastal sites, however, was significantly divergent (large F_ST, 0·0251) from the other populations, suggesting low connectivity with coastal populations. The Santa Catalina Island population also had the lowest allele richness and lowest average heterozygosity, suggesting recent population bottlenecks in size.     

The foraging ecology of two sympatric gobiid fishes: importance of behavior in prey type selection

Synopsis The foraging ecology of two temperate marine gobies (Pisces: Gobiidae) was studied in rocky subtidal habitats off Santa Catalina Island, California. The bluebanded goby, Lythrypnus dalli, foraged from exposed ledges and fed on planktonic and benthic prey, although planktonic prey were more important in diets by number and weight. The more cryptic zebra goby, Lythrypnus zebra, remained hidden under rocks and in crevices feeding on benthic prey almost exclusively. The active selection of particular prey taxa from the two prey sources (water column and substratum), mediated by species-specific differences in foraging behavior, resulted in interspecific differences in type, number, size and weight of prey consumed. Interspecific differences in foraging ecology reflect the selection of prey most readily available to these fishes that occupy specific and fixed microhabitats within rocky reefs.     

Biogeography and the structure of coral reef fish communities on isolated islands

Aim To determine the applicability of biogeographical and ecological theory to marine species at two remote island locations. This study examines how biogeography, isolation and species geographic range size influence patterns of species richness, endemism, species composition and the abundance of coral reef fishes. Location Christmas Island and the Cocos (Keeling) Islands in the tropical eastern Indian Ocean. Methods Published species lists and underwater visual surveys were used to determine species richness, endemism, species composition and abundance of reef fishes at the islands. These data were statistically compared with patterns of species composition and abundance from the neighbouring ‘mainland’ Indonesian region. Results The two isolated reef fish communities were species‐poor and contained a distinct taxonomic composition with an overrepresentation of species with high dispersal potential. Despite low species richness, we found no evidence of density compensation, with population densities on the islands similar to those of species‐rich mainland assemblages. The mix of Indian and Pacific Ocean species and the proportional representations of the various regional faunas in the assemblages were not influenced by the relative proximity of the islands to different biogeographical provinces. Moreover, species at the edge of their range did not have a lower abundance than species at the centre of their range, and endemic species had substantially higher abundances than widespread species. At both locations, endemism was low (less than 1.2% of the community); this may be because the locations are not sufficiently isolated or old enough to promote the evolution of endemic species. Main conclusions The patterns observed generally conform to terrestrial biogeographical theory, suggesting that similar processes may be influencing species richness and community composition in reef fish communities at these remote islands. However, species abundances differed from typical terrestrial patterns, and this may be because of the life history of reef fishes and the processes maintaining isolated populations.     

The effect of biogeographic and phylogeographic barriers on gene flow in the brown smoothhound shark,Mustelus henlei,in the northeastern Pacific

We assessed the effects of the prominent biogeographic (Point Conception and the Peninsula of Baja California) and phylogeographic barriers (Los Angeles Region) of the northeastern Pacific on the population connectivity of the brown smoothhound shark, Mustelus henlei (Triakidae). Data from the mitochondrial control region and six nuclear microsatellite loci revealed significant population structure among three populations: northern (San Francisco), central (Santa Barbara, Santa Catalina, Punta Lobos, and San Felipe), and southern (Costa Rica). Patterns of long‐term and contemporary migration were incongruent, with long‐term migration being asymmetric and occurring in a north to south direction and a lack of significant contemporary migration observed between localities with the exception of Punta Lobos that contributed migrants to all localities within the central population. Our findings indicate that Point Conception may be restricting gene flow between the northern and central populations whereas barriers to gene flow within the central population would seem to be ineffective; additionally, a contemporary expansion of tropical M. henlei into subtropical and temperate waters may have been observed.     

Length–weight relationships of cryptic reef fishes from the southwestern Gulf of California, Mèxico

Length–weight relationships of 33 cryptic reef fishes from the southwestern Gulf of California were estimated. The most common families were Gobiidae (seven species), Labrisomidae (four), Pomacentridae (three), and Tripterygiidae (three). These are the first length–weight parameters reported for 32 of these species. Specimens were sampled during a comparative ecological study of community structure of fish assemblages associated with coral heads, rocky walls, and artificial reefs in Bahía de La Paz, Baja California Sur. The b values varied between 2.63 and 3.61. New maximum length records for five of the species were established in the study.     

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs

The previously sub‐dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub‐tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co‐occurring habitat‐forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef‐associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon‐Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.     

Geography and island geomorphology shape fish assemblage structure on isolated coral reef systems

We quantify the relative importance of multi‐scale drivers of reef fish assemblage structure on isolated coral reefs at the intersection of the Indian and Indo‐Pacific biogeographical provinces. Large (>30 cm), functionally‐important and commonly targeted species of fish, were surveyed on the outer reef crest/front at 38 coral reef sites spread across three oceanic coral reef systems (i.e. Christmas Island, Cocos (Keeling) Islands and the Rowley Shoals), in the tropical Indian Ocean (c. 1.126 x 106 km²). The effects of coral cover, exposure, fishing pressure, lagoon size and geographical context, on observed patterns of fish assemblage structure were modelled using Multivariate Regression Trees. Reef fish assemblages were clearly separated in space with geographical location explaining ~53 % of the observed variation. Lagoon size, within each isolated reef system was an equally effective proxy for explaining fish assemblage structure. Among local‐scale variables, ‘distance from port’, a proxy for the influence of fishing, explained 5.2% of total variation and separated the four most isolated reefs from Cocos (Keeling) Island, from reefs with closer boating access. Other factors were not significant. Major divisions in assemblage structure were driven by sister taxa that displayed little geographical overlap between reef systems and low abundances of several species on Christmas Island corresponding to small lagoon habitats. Exclusion of geographical context from the analysis resulted in local processes explaining 47.3% of the variation, highlighting the importance of controlling for spatial correlation to understand the drivers of fish assemblage structure. Our results suggest reef fish assemblage structure on remote coral reef systems in the tropical eastern Indian Ocean reflects a biogeographical legacy of isolation between Indian and Pacific fish faunas and geomorphological variation within the region, more than local fishing pressure or reef condition. Our findings re‐emphasise the importance that historical processes play in structuring contemporary biotic communities.     

The community composition of Californian coastal sage scrub

Summary Coastal sage scrub is a community found extensively throughout cismontane California south of San Francisco, but has been surprisingly little studied. In the study area, which extends from Santa Barbara to the San Gorgonio Pass, two major floristic groupings can be found. In the basin bounded coastwards by a line drawn along the axis of the Santa Ana Mountains a large number of native and introduced annual herbs and a few shrubs (e.g.Encelia farinosa), rare or absent in the remainder of the study area, characterize one floristic group. In the coastal region the variety of shrub species increases, and the herbs are predominantly native and more restricted in number. Eleven groups defined by physiognomy, structure and species dominance, and arbitrarily called associations, are recognized. These associations can be grouped into four physiognomic-structural types which transgress the boundaries of the floristic groups. The results of this study and the limited previous literature suggest that Californian coastal sage scrub could be divided, mainly on floristic criteria, into Venturan, San Diegan and Riversidian sage.Plant nomenclature follows Munz & Keek (1968).We gratefully acknowledge the financial help provided by the Department of Earth Sciences, University of California, Riverside, and the aid in plant identification provided by Mr. Oscar Clarke, Museum Scientist, Department of Biology, University of California, Riverside.     

A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014–2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.     

Typological thinking and the conservation of subspecies: the case of the San Clemente Island loggerhead shrike

Abstract.  Hybridization with closely related taxa poses a significant threat to endangered subspecies (e.g. outbreeding depression, inbreeding) and confounds efforts to manage and conserve these taxa through a loss of taxonomic identity, in part because of the practical necessity of defining subspecies in a typological manner. We examined nine morphological characters in 167 post-juvenile museum specimens to determine if loggerhead shrikes Lanius ludovicianus Linnaeus 1766 on San Clemente Island (off the coast of California) remain diagnosable as L. l. mearnsi Ridgway (1903); an island endemic listed as endangered by the United States Fish and Wildlife Service. Four recent shrike specimens from the island were compared to historical specimens using a bivariate scatter plot and a discriminant function (the latter was used to classify recent specimens post hoc). The few recent specimens were not diagnosable as L. l. mearnsi , but instead appear to be intergrades between L. l. mearnsi and L. l. anthonyi Mearns 1898 (the subspecies endemic to Santa Cruz, Santa Catalina, Santa Rosa and Anacapa islands), and are perhaps closer to pure anthonyi . Our data and the species' natural history and distribution suggest that shrikes currently on San Clemente Island are the result of genetic 'swamping' of mearnsi by anthonyi . Under a necessarily typological definition of a subspecies, it is evident that mearnsi is probably no longer diagnosable. However, we conclude that protection of the entire Channel Islands population of the loggerhead shrike would be the best management strategy, as the species has declined drastically throughout the islands.     

Island biogeography of Caribbean coral reef fish

Aim The goal of our study was to test fundamental predictions of biogeographical theories in tropical reef fish assemblages, in particular relationships between fish species richness and island area, isolation and oceanographic variables (temperature and productivity) in the insular Caribbean. These analyses complement an analogous and more voluminous body of work from the tropical Indo‐Pacific. The Caribbean is more limited in area with smaller inter‐island distances than the Indo‐Pacific, providing a unique context to consider fundamental processes likely to affect richness patterns of reef fish. Location Caribbean Sea. Methods We compiled a set of data describing reef‐associated fish assemblages from 24 island nations across the Caribbean Sea, representing a wide range of isolation and varying in land area from 53 to 110,860 km². Regression‐based analyses compared the univariate and combined effects of island‐specific physical predictors on fish species richness. Results We found that diversity of reef‐associated fishes increases strongly with increasing island area and with decreasing isolation. Richness also increases with increasing nearshore productivity. Analyses of various subsets of the entire data set reveal the robustness of the richness data and biogeographical patterns. Main conclusions Within the relatively small and densely packed Caribbean basin, fish species richness fits the classical species–area relationship. Richness also was related negatively to isolation, suggesting direct effects of dispersal limitation in community assembly. Because oceanic productivity was correlated with isolation, however, the related effects of system‐wide productivity on richness cannot be disentangled. These results highlight fundamental mechanisms that underlie spatial patterns of biodiversity among Caribbean coral reefs, and which are probably also are functioning in the more widespread and heterogeneous reefs of the Indo‐Pacific.     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Changes in rocky reef fish assemblages throughout an estuary with a restricted inlet

Rocky reef habitat is common in many estuaries, yet its role as a habitat for fishes is poorly understood. There is also limited understanding of how access of coastal species into estuaries and habitat quality can affect the distribution of rocky reef fishes within estuaries. This study used baited remote underwater video stations to determine spatial patterns in fish assemblages associated with rocky reef habitat throughout a barrier estuary with a permanently open but restricted inlet. Estuarine rocky reefs provided habitat for a diverse assemblage of fishes, many of which were large juveniles and subadults. In the absence of a pronounced salinity or temperature gradient, a clear transition in fish assemblages occurred from coastal waters, through the inlet channel, to the central estuary, and into the inner estuary. The inlet channel, notably its narrowness and length, limits tidal input into this estuary, which acts as a significant impediment to the dispersal of many coastal fishes, and insufficient habitat excludes many coastal rocky reef species from the inner estuary. This study highlights the need to recognise estuarine rocky reefs as providing habitat for diverse fish assemblages and the role inlets play in restricting access of coastal species.     

Site fidelity and homing behaviour of some kelp-bed fishes

Common species of kelp-bed fishes including adult blacksmith Chromis punctipinnis , adult senorita Oxyjulis californica , and sub-adult kelp bass Paralabrax clathratus were collected from two separate and isolated rocky reefs at Santa Catalina Island, California, U.S.A. where giant kelp Macrocystis pyrifera was present. After tagging, groups of fishes were either returned to their initial collection site or displaced to an alternative kelp-bed site to measure site fidelity and homing behaviour. No tagged fishes that were returned to their initial collection sites and subsequently resighted, exhibited any movement to another reef suggesting that these fishes have a limited home range, and handling during capture and tagging had no effect on site fidelity. Of fishes translocated to alternative kelp-bed sites, up to 80% of tagged senorita and 100% of tagged blacksmith that were resighted had returned to the site of their initial collection, indicating that these species are motivated and able to relocate previously utilized areas when displaced. However, when young-of-the-year and juvenile kelp bass were displaced, none were ever seen again suggesting that disruption of the relationship between a young kelp bass and its home site may influence its survival.     

Seri Indian traditional knowledge and molecular biology agree: no express train for island‐hopping spiny‐tailed iguanas in the Sea of Cortés

Aim The role of human activities in species biogeography can be difficult to identify, but in some cases molecular techniques can be used to test hypotheses of human‐mediated dispersal. A currently accepted hypothesis states that humans mediated the divergence of two species of spiny‐tailed iguanas in the Ctenosaura hemilopha species complex, namely C. conspicuosa and C. nolascensis, which occupy islands in the Sea of Cortés between the peninsula of Baja California and mainland Mexico. We test an alternative hypothesis that follows the traditional knowledge of the Seri Indians and states that the divergence of these species was not mediated by humans. Location Mexico, including Baja California, Sonoran and Sinaloan coastal regions, and Isla San Esteban and Isla San Pedro Nolasco in the Sea of Cortés. Methods We analysed mitochondrial (cytochrome b and cytochrome c oxidase subunit III) DNA sequences from four species in the C. hemilopha species complex. Maximum parsimony and Bayesian inference were used to infer matriarchal genealogical relationships between the species and several outgroup taxa. Bayesian methods were used to estimate divergence times for the major nodes on the trees based on previously published, fossil‐calibrated priors. Results Our analysis indicated that lineages within the C. hemilopha species complex diverged long before human colonization of the Americas. The divergence of C. nolascensis and C. conspicuosa could not be attributed to Seri translocations. The matriarchal genealogy of the species complex currently defies a simple biogeographical interpretation. Main conclusions We conclude that humans did not mediate the divergence of C. nolascensis and C. conspicuosa. This conclusion is consistent with the traditional knowledge of the Seri people. These results demonstrate the utility of molecular techniques in investigating potential cases of human‐mediated dispersal of plants and animals, and reinforce the importance of considering traditional knowledge in the formation of scientific hypotheses and the interpretation of results.