Dealing with uncertain absences in habitat modelling: a case study of a rare ground-dwelling parrot

In the development of a species distribution model based on regression techniques such as generalized linear or additive modelling (GLM/GAM), a basic assumption is that records of species presence and absence are real. However, a common concern in many studies examining species distributions is that absences cannot be inferred with certainty. This is particularly the case where the species is rare, difficult to detect and/or does not occupy all available habitat considered suitable. The western ground parrot ( Pezoporus wallicus flaviventris ) of southern Western Australia, Australia, is a case in point, as not only is it rare and difficult to detect, but it is also unlikely to occupy all available suitable habitat. A recent survey of ground parrots provided the opportunity to develop a predictive distribution model. As the data were susceptible to false absences, these were replaced with randomly selected 'pseudo' absences and modelled using GLM. As a comparison, presence-only information was modelled using a relatively new approach, MAXENT, a machine-learning technique that has been shown to perform comparatively well. The predictive performance of both models, as assessed by the receiver operating characteristic plot (ROC) was high (AUC > 0.8), with MAXENT performing only marginally better than the GLM. These approaches both indicated that the ground parrot prefers areas relatively high in altitude, distant from rivers, gently sloping to level habitat, with an intermediate cover of vegetation and where there is a mosaic of vegetation ages. In this case, the use of presence-only information resulted in the identification of important environmental attributes defining the occurrence of the ground parrot, but additional factors that account for the inability of the bird to occupy all suitable habitat should be a component of model refinement.     

Ancient Divergence in the Trans-Oceanic Deep-Sea Shark Centroscymnus crepidater

Unravelling the genetic structure and phylogeographic patterns of deep-sea sharks is particularly challenging given the inherent difficulty in obtaining samples. The deep-sea shark Centroscymnus crepidater is a medium-sized benthopelagic species that exhibits a circumglobal distribution occurring both in the Atlantic and Indo-Pacific Oceans. Contrary to the wealth of phylogeographic studies focused on coastal sharks, the genetic structure of bathyal species remains largely unexplored. We used a fragment of the mitochondrial DNA control region, and microsatellite data, to examine genetic structure in C. crepidater collected from the Atlantic Ocean, Tasman Sea, and southern Pacific Ocean (Chatham Rise). Two deeply divergent (3.1%) mtDNA clades were recovered, with one clade including both Atlantic and Pacific specimens, and the other composed of Atlantic samples with a single specimen from the Pacific (Chatham Rise). Bayesian analyses estimated this splitting in the Miocene at about 15 million years ago. The ancestral C. crepidater lineage was probably widely distributed in the Atlantic and Indo-Pacific Oceans. The oceanic cooling observed during the Miocene due to an Antarctic glaciation and the Tethys closure caused changes in environmental conditions that presumably restricted gene flow between basins. Fluctuations in food resources in the Southern Ocean might have promoted the dispersal of C. crepidater throughout the northern Atlantic where habitat conditions were more suitable during the Miocene. The significant genetic structure revealed by microsatellite data suggests the existence of present-day barriers to gene flow between the Atlantic and Pacific populations most likely due to the influence of the Agulhas Current retroflection on prey movements.     

Drivers for genetic structure at different geographic scales for Pacific red snapper (Lutjanus peru) and yellow snapper (Lutjanus argentiventris) in the tropical eastern Pacific

The tropical eastern Pacific (TEP) is a highly dynamic region and a model system to study how habitat discontinuities affect the distribution of shorefishes, particularly for species that display ontogenetic habitat shifts, including snappers (Lutjanidae). To evaluate the genetic structure of the Pacific red snapper (Lutjanus peru) and the yellow snapper (Lutjanus argentiventris) throughout their distribution range along the TEP, 13 and 11 microsatellite loci were analysed, respectively. The genetic diversity of L. peru (N = 446) and L. argentiventris (N = 170) was evaluated in 10 and 5 localities, respectively, showing slightly higher but non-significant values in the Gulf of California for both species. The genetic structure analysis identified the presence of significant genetic structure in both species, but the locations of the identified barriers for the gene flow differed between species. The principal driver for the genetic structure at large scales >2500 km was isolation by distance. At smaller scales (<250 km), the habitat discontinuity for juveniles and adults and the environmental differences throughout the distribution range represented potential barriers to gene flow between populations for both species.     

Spatial and environmental determinants of the distribution of Striped Marlin (Kajikia audax) in the western and central North Pacific Ocean

Striped Marlin is a highly migratory species distributed throughout tropical and temperate waters in the North Pacific Ocean. The habitat characteristics of Striped Marlin in the western and central North Pacific Ocean were examined using generalized additive models by modelling fishery catch-rates as a function of remotely-sensed environmental covariates, including sea surface temperature (SST), chlorophyll-a concentration (CHL), mixed layer depth (MLD) and sea height anomalies (SHA). SST explained the largest proportion of the deviance, and is therefore considered the best predictor for the habitat of Striped Marlin. Spatial distributions of the relative density of Striped Marlin indicated that there is a seasonal north–south migration, and that the highest densities occur in the central North Pacific Ocean. The preferred habitat characteristics of Striped Marlin in high density areas were identified as SST between 23 and 26 °C, MLD within 30 m depth, and CHL around 0.08 mg m^?3, while no preferred range was found for SHA. The results of this study could improve our understanding of Striped Marlin spatial distributions and habitat characteristics.     

Utilization of seagrass habitats by juvenile groupers and snappers in Banten Bay,Banten Province,Indonesia

Coastal development in Banten Bay, Indonesia, decreased seagrass coverage to only 1.5% of its surface area. We investigated the importance of seagrass as habitat for juvenile groupers (Serranidae) and snappers (Lutjanidae), by performing beam trawl hauls on a weekly basis in two seagrass locations and one mudflat area, and monthly trawl hauls in three different microhabitats (dense, mixed and patchy seagrass) in one of the seagrass locations. We studied the effects of location and microhabitat, as well as temporal patterns (diel, weekly and monthly) on the probability of occurrence and abundance of the most abundant grouper (Orange-spotted grouper, Epinephelus coioides) and snapper (Russell’s snapper, Lutjanus russellii). We found that both species were almost exclusively found in seagrass locations, with a preference for microhabitats of high complexity (dense and mixed microhabitats). L. russellii had a higher probability of catch and abundance during the night, most probably because of its ability to avoid the beam trawl during daytime sampling. In addition there was an effect of week and month on the presence and abundance of both species, but patterns were unclear, probably because of high fishing pressure on juvenile groupers and snappers by push net fishermen. Groupers and snappers mainly fed on abundant shrimps, and to a lesser extent on fish. Moreover, juveniles find protection against predators in seagrass, which confirmed the critical role of quantity and quality of seagrass areas for juvenile groupers and snappers in Banten Bay.     

Etelis boweni sp. nov., a new cryptic deepwater eteline snapper from the Indo-Pacific (Perciformes: Lutjanidae)

A new species of Etelis is described based on 16 specimens collected from the Red Sea and Western Australia, with confirmed genetic records throughout the Indo-West Pacific. It is similar to and was often misidentified as Etelis carbunculus Cuvier, with both species sharing the diagnostic character of low number of developed gill rakers. Nonetheless, the two species are genetically divergent and differ morphologically in adult body length; proportions of eye, snout, cheek and caudal fin; shape of head, opercular spine and sagittal otolith; and coloration of the tip of the upper caudal fin. Etelis boweni has a wide Indo-west Pacific distribution that largely overlaps with E. carbunculus, and the two species are often caught on the same fishing line.     

Global habitat suitability for framework-forming cold-water corals

Predictive habitat models are increasingly being used by conservationists, researchers and governmental bodies to identify vulnerable ecosystems and species' distributions in areas that have not been sampled. However, in the deep sea, several limitations have restricted the widespread utilisation of this approach. These range from issues with the accuracy of species presences, the lack of reliable absence data and the limited spatial resolution of environmental factors known or thought to control deep-sea species' distributions. To address these problems, global habitat suitability models have been generated for five species of framework-forming scleractinian corals by taking the best available data and using a novel approach to generate high resolution maps of seafloor conditions. High-resolution global bathymetry was used to resample gridded data from sources such as World Ocean Atlas to produce continuous 30-arc second (～1 km(2)) global grids for environmental, chemical and physical data of the world's oceans. The increased area and resolution of the environmental variables resulted in a greater number of coral presence records being incorporated into habitat models and higher accuracy of model predictions. The most important factors in determining cold-water coral habitat suitability were depth, temperature, aragonite saturation state and salinity. Model outputs indicated the majority of suitable coral habitat is likely to occur on the continental shelves and slopes of the Atlantic, South Pacific and Indian Oceans. The North Pacific has very little suitable scleractinian coral habitat. Numerous small scale features (i.e., seamounts), which have not been sampled or identified as having a high probability of supporting cold-water coral habitat were identified in all ocean basins. Field validation of newly identified areas is needed to determine the accuracy of model results, assess the utility of modelling efforts to identify vulnerable marine ecosystems for inclusion in future marine protected areas and reduce coral bycatch by commercial fisheries.     

Assessing Habitat Use by Snapper (Chrysophrys auratus) from Baited Underwater Video Data in a Coastal Marine Park

Baited Underwater Video (BUV) systems have become increasingly popular for assessing marine biodiversity. These systems provide video footage from which biologists can identify the individual fish species present. Here we explore the relevance of spatial dependence and marine park boundaries while estimating the distribution and habitat associations of the commercially and recreationally important snapper species Chrysophrys auratus in Moreton Bay Marine Park during a period when new Marine National Parks zoned as no-take or “green” areas (i.e., areas with no legal fishing) were introduced. BUV studies typically enforce a minimum distance among BUV sites, and then assume that observations from different sites are independent conditional on the measured covariates. In this study, we additionally incorporated the spatial dependence among BUV sites into the modelling framework. This modelling approach allowed us to test whether or not the incorporation of highly correlated environmental covariates or the geographic placement of BUV sites produced spatial dependence, which if unaccounted for could lead to model bias. We fitted Bayesian logistic models with and without spatial random effects to determine if the Marine National Park boundaries and available environmental covariates had an effect on snapper presence and habitat preference. Adding the spatial dependence component had little effect on the resulting model parameter estimates that emphasized positive association for particular coastal habitat types by snapper. Strong positive relationships between the presence of snapper and rock habitat, particularly rocky substrate composed of indurated freshwater sediments known as coffee rock, and kelp habitat reinforce the consideration of habitat availability in marine reserve design and the design of any associated monitoring programs.     

Coral Reef Habitat Response to Climate Change Scenarios

Coral reef ecosystems are threatened by both climate change and direct anthropogenic stress. Climate change will alter the physico-chemical environment that reefs currently occupy, leaving only limited regions that are conducive to reef habitation. Identifying these regions early may aid conservation efforts and inform decisions to transplant particular coral species or groups. Here a species distribution model (Maxent) is used to describe habitat suitable for coral reef growth. Two climate change scenarios (RCP4.5, RCP8.5) from the National Center for Atmospheric Research’s Community Earth System Model were used with Maxent to determine environmental suitability for corals (order Scleractinia). Environmental input variables best at representing the limits of suitable reef growth regions were isolated using a principal component analysis. Climate-driven changes in suitable habitat depend strongly on the unique region of reefs used to train Maxent. Increased global habitat loss was predicted in both climate projections through the 21^st century. A maximum habitat loss of 43% by 2100 was predicted in RCP4.5 and 82% in RCP8.5. When the model is trained solely with environmental data from the Caribbean/Atlantic, 83% of global habitat was lost by 2100 for RCP4.5 and 88% was lost for RCP8.5. Similarly, global runs trained only with Pacific Ocean reefs estimated that 60% of suitable habitat would be lost by 2100 in RCP4.5 and 90% in RCP8.5. When Maxent was trained solely with Indian Ocean reefs, suitable habitat worldwide increased by 38% in RCP4.5 by 2100 and 28% in RCP8.5 by 2050. Global habitat loss by 2100 was just 10% for RCP8.5. This projection suggests that shallow tropical sites in the Indian Ocean basin experience conditions today that are most similar to future projections of worldwide conditions. Indian Ocean reefs may thus be ideal candidate regions from which to select the best strands of coral for potential re-seeding efforts.     

A Potential Distribution Model and Conservation Plan for the Critically Endangered Ecuadorian Capuchin, Cebus albifrons aequatorialis

Conservation actions that effectively and efficiently target single, highly threatened species require current data on the species’ geographic distribution and environmental associations. The Ecuadorian capuchin (Cebus albifrons aequatorialis) is a critically endangered primate found only in the fragmented forests of western Ecuador and northern Peru, which are among the world’s most severely threatened ecosystems. We use the MAXENT species distribution modeling method to model the potential distribution and environmental associations of Cebus albifrons aequatorialis, using all known presence localities recorded within the last 2 decades as well as 13 climate, topography, vegetation, and land-use data sets covering the entire geographic range of the subspecies. The environmental conditions that our model predicted to be ideal for supporting Cebus albifrons aequatorialis included ≥20% tree cover, mild temperature seasonality, annual precipitation <2000 mm, and low human population density. Our model identified 5028 km² of suitable habitat remaining, although many of these forest fragments are unprotected and are unlikely to support extant populations. Using the median population density across all sites for which data are available, we estimate the total carrying capacity of the remaining habitat to be 12,500 total individuals. The true number of remaining individuals is likely to be considerably lower due to anthropogenic factors. We highlight four critical regions of high predicted suitability in western Ecuador and northern Peru on which immediate conservation actions should focus, and we lay out clear priorities to guide conservation actions for ensuring the long-term survival of this gravely threatened and little known primate.     

Gerres limbatus Cuvier and G. lucidus Cuvier from the Indo-Malay Archipelagos, the latter corresponding to young of the former (Perciformes: Gerreidae)

Taxonomic status of Gerres limbatus Cuvier in Cuvier and Valenciennes, 1830 and G. lucidus Cuvier in Cuvier and Valenciennes, 1830 was studied. The two species were reassessed as the same species, the latter corresponding to young of the former, on the basis of examination of their eight syntypes and other comparative specimens. Gerres limbatus clearly differs from other congeners in having 2½–3 (usually 2½) scales between the 5th dorsal fin spine base and the lateral line, and four or five diffuse, dark saddle patches mainly along the back in life (more apparent in small specimens less than ca. 65 mm in standard length, or preserved and stressed live specimens) (vs. usually more than 3½ scales between fifth dorsal fin spine base and lateral line, and and no such body color pattern of G. limbatus in other congeners). The species is currently known from the southern and western coasts of India including Sri Lanka, the Malay Peninsula, Gulf of Thailand, and Indonesia, becoming rare in occurrence eastward.     

Ethnobiology of snappers (Lutjanidae): target species and suggestions for management

In this study, we sought to investigate the biology (diet and reproduction) and ethnobiology (fishers knowledge and fishing spots used to catch snappers) of five species of snappers (Lutjanidae), including Lutjanus analis, Lutjanus synagris, Lutjanus vivanus, Ocyurus chrysurus, and Romboplites saliens at five sites along the northeast (Riacho Doce, Maceió in Alagoas State, and Porto do Sauípe, Entre Rios at Bahia State) and the southeast (SE) Brazilian coast (Paraty and Rio de Janeiro cities at Rio de Janeiro State, and Bertioga, at São Paulo State.). We collected 288 snappers and interviewed 86 fishermen. The stomach contents of each fish were examined and macroscopic gonad analysis was performed. Snappers are very important for the fisheries of NE Brazil, and our results indicated that some populations, such as mutton snapper (L. analis) and lane snapper (L. synagris), are being caught when they are too young, at early juvenile stages. Local knowledge has been shown to be a powerful tool for determining appropriate policies regarding management of target species, and artisanal fishermen can be included in management processes. Other suggestions for managing the fisheries are discussed, including proposals that could provide motivation for artisanal fishermen to participate in programs to conserve resources, such as co-management approaches that utilize local knowledge, the establishment of fishing seasons, and compensation of fishermen, through 'payment for environmental services'. These suggestions may enhance the participation of local artisanal fishermen in moving to a more realistic and less top-down management approach of the fish population.     

Reef‐coral refugia in a rapidly changing ocean

This study sought to identify climate‐change thermal‐stress refugia for reef corals in the Indian and Pacific Oceans. A species distribution modeling approach was used to identify refugia for 12 coral species that differed considerably in their local response to thermal stress. We hypothesized that the local response of coral species to thermal stress might be similarly reflected as a regional response to climate change. We assessed the contemporary geographic range of each species and determined their temperature and irradiance preferences using a k‐fold algorithm to randomly select training and evaluation sites. That information was applied to downscaled outputs of global climate models to predict where each species is likely to exist by the year 2100. Our model was run with and without a 1 °C capacity to adapt to the rising ocean temperature. The results show a positive exponential relationship between the current area of habitat that coral species occupy and the predicted area of habitat that they will occupy by 2100. There was considerable decoupling between scales of response, however, and with further ocean warming some ‘winners’ at local scales will likely become ‘losers’ at regional scales. We predicted that nine of the 12 species examined will lose 24–50% of their current habitat. Most reductions are predicted to occur between the latitudes 5–15°, in both hemispheres. Yet when we modeled a 1 °C capacity to adapt, two ubiquitous species, Acropora hyacinthus and Acropora digitifera, were predicted to retain much of their current habitat. By contrast, the thermally tolerant Porites lobata is expected to increase its current distribution by 14%, particularly southward along the east and west coasts of Australia. Five areas were identified as Indian Ocean refugia, and seven areas were identified as Pacific Ocean refugia for reef corals under climate change. All 12 of these reef‐coral refugia deserve high‐conservation status.     

Predicting the distribution of the crested tinamous, Eudromia spp. (Aves, Tinamiformes)

A bioclimatic analysis of the crested tinamous was conducted to explore climatic factors underpinning the distribution of both Eudromia elegans and E. formosa and to evaluate its potential application in paleontological studies. The study utilized records throughout the entire known range of Eudromia spp. in southern South America. Relationships between 20 environmental parameters and the presence of Eudromia species were established, mapping and characterizing their spatial distribution in a geographic information system using BIOCLIM and MAXENT algorithms. The MAXENT prediction map shows a more homogeneous pattern while BIOCLIM showed a patchier pattern. The models applied here generated maps that adjust to the well-known previous distributions of both species. Nevertheless, for Eudromia elegans, the distribution predicted by MAXENT includes areas where it is actually considered absent, and the BIOCLIM prediction does not include some areas where it is presumed present. Eudromia formosa were found in warmer and wetter sites than E. elegans. Low precipitation areas were identified as suitable for Eudromia elegans. Strong differences between the climatic profiles for both Eudromia spp distributions occurred, with the precipitation the most important influence. E. formosa tolerates the highest maximum temperatures, whereas E. elegans supports the lowest temperatures.     

New records of two deep-sea eels collected from the Western Pacific Ocean based on COI and 16S rRNA genes

Background

Two deep-sea eels collected from the Western Pacific Ocean are described in this study. Based on their morphological characteristics, the two deep-sea eel specimens were assumed to belong to the cusk-eel family Ophidiidae and the cutthroat eel family Synaphobranchidae.

Methods and results

To accurately identify the species of the deep-sea eel specimens, we sequenced the mitochondrial genes (cytochrome c oxidase subunit I [COI] and 16S ribosomal RNA [16S rRNA]). Through molecular phylogenetic analysis based on mtDNA COI and 16S rRNA gene sequences, these species clustered with the genera Bassozetus and Synaphobranchus, suggesting that the deep-sea eel specimens collected are two species from the genera Bassozetus and Synaphobranchus in the Western Pacific Ocean, respectively.

Conclusions

This is the first study to report new records of the genera Bassozetus and Synaphobranchus from the Western Pacific Ocean based on COI and 16S rRNA genes

     

Isolation and characterization of microsatellites in lane snapper (Lutjanus synagris), mutton snapper (Lutjanus analis), and yellowtail snapper (Ocyurus chrysurus)

Polymerase chain reaction primer pairs for a total of 25 nuclear‐encoded microsatellites (loci) were developed from genomic DNA libraries of lane snapper (Lutjanus synagris), mutton snapper (Lutjanus analis), and yellowtail snapper (Ocyurus chrysurus). The microsatellites include 24 perfect (21 dinucleotide and three trinucleotide) and one imperfect (combination tetranucleotide/tetranucleotide) repeat motifs. A total of 32 individuals of each species were assayed for allelic variation at all 25 microsatellites; reliable amplification products were generated for lane snapper (25 loci), mutton snapper (21 loci), and yellowtail snapper (24 loci). Significant deviations from Hardy–Weinberg expectations, following Bonferroni corrections, were found for one microsatellite in lane and yellowtail snappers, and for three microsatellites in mutton snapper. All pairwise comparisons of microsatellites (all three species) did not deviate significantly from genotypic equilibrium.     

The commonness of rarity in a deep-sea taxon

The generality and drivers of rarity, defined along the axes of geographic range, population size and habitat specificity, have received considerable scientific attention for well over a century. Yet, studies that examine rarity holistically among these three attributes are limited, especially among invertebrate and marine taxa. The perceived paradox of deep-sea species, with often low population size but large geographic ranges, remains poorly resolved and understood. Here I assess seven forms of rarity and their drivers in deep-sea bivalves across the Atlantic Ocean. Rarity appears to be a common trait among deep-sea bivalves, with nearly 85% of the species exhibiting some form of rarity. Bivalves also showed a strong bimodal pattern of very common and very rare species. Geographic range, population size and habitat specificity were all heavily right skewed. Taxonomic superfamilies, body size, energy availability, temperature, depth and latitude, all significantly predicted geographic range, population size and habitat specificity. In a few cases, these patterns were counter to theoretical expectations. The drivers of rarity appear to be predictable from knowledge of the intrinsic biological and extrinsic environmental context of the species. These findings have major implications for deep-sea conversation, especially as anthropogenic threats are increasing.     

基于MAXENT模型的贺兰山岩羊生境适宜性评价

生境评价对物种保护具有非常重要的意义。本研究在GIS空间技术的支持下,于2010年到2011年间,在贺兰山岩羊分布区采集其地理分布点数据以及环境变量数据,并运用MAXENT模型对贺兰山区域岩羊的生境适宜性进行评价分析。模型的评价结果达到优秀水平,研究结果表明：岩羊生境选择的主要影响因子为矿区、坡度、海拔和道路,生境适宜区主要分布于贺兰山东坡（宁夏贺兰山国家级自然保护区）的西南部,而西坡（内蒙古贺兰山国家级自然保护区）有少量分布,适宜生境面积总和为1 006.9 km2,占研究区域面积的28%。岩羊偏爱于海拔1 500－2 300 m的山势陡峭地带,建议对矿区和道路这两种人为干扰因素采取相应的措施,以提高岩羊的生境质量,促进其种群发展。     

Beta-diversity on deep-sea wood falls reflects gradients in energy availability

Wood falls on the deep-sea floor represent a significant source of energy into the food-limited deep sea. Unique communities of primarily wood- and sulfide-obligate species form on these wood falls. However, little is known regarding patterns and drivers of variation in the composition of wood fall communities through space and time, and thus, how wood falls contribute to deep-sea biodiversity. Eighteen Acacia logs varying in size were placed and retrieved after five years at a 3200 m site in the Pacific Ocean. We found that the taxonomic composition and structure of deep-sea wood fall communities varied considerably and equated with considerable differences in energy usage and availability. Our findings suggest that natural variability in wood falls may contribute significantly to deep-sea diversity.