Taxonomic homogenization and differentiation across Southern Ocean Islands differ among insects and vascular plants

Aim To investigate taxonomic homogenization and/or differentiation of insect and vascular plant assemblages across the Southern Ocean Islands (SOI), and how they differ with changing spatial extent and taxonomic resolution. Location Twenty‐two islands located across the Southern Ocean, further subdivided into five island biogeographical provinces. These islands are used because comprehensive data on both indigenous and non‐indigenous insect and plant species are available. Methods An existing database was updated, using newly published species records, identifying the indigenous and non‐indigenous insect and vascular plant species recorded for each island. Homogenization and differentiation were measured using Jaccard’s index (JI) of similarity for assemblages across all islands on a pairwise basis, and for island pairs within each of the biogeographical provinces. The effects of taxonomic resolution (species, genus, family) and distance on levels of homogenization or differentiation were examined. To explore further the patterns of similarity among islands for each of the taxa and groupings (indigenous and non‐indigenous), islands were clustered based on JI similarity matrices and using group averaging. Results Across the SOI, insect assemblages have become homogenized (0.7% increase in similarity at species level) while plant assemblages have become differentiated at genus and species levels. Homogenization was recorded only when pairwise distances among islands exceeded 3000 km for insect assemblages, but distances had to exceed 10,000 km for plant assemblages. Widely distributed non‐indigenous plant species tend to have wider distributions across the SOI than do their insect counterparts, and this is also true of the indigenous species. Main conclusions Insect assemblages across the SOI have become homogenized as a consequence of the establishment of non‐indigenous species, while plant assemblages have become more differentiated. The likely reason is that indigenous plant assemblages are more similar across the SOI than are insect assemblages, which show greater regionalization. Thus, although a suite of widespread, typically European, weedy, non‐indigenous plant species has established on many islands, the outcome has largely been differentiation. Because further introductions of insects and vascular plants are probable as climates warm across the region, the patterns documented here are likely to change through time.     

Measuring beta‐diversity from taxonomic similarity

Question: The utility of beta (β‐) diversity measures that incorporate information about the degree of taxonomic (dis)similarity between species plots is becoming increasingly recognized. In this framework, the question for this study is: can we define an ecologically meaningful index of β‐diversity that, besides indicating simple species turnover, is able to account for taxonomic similarity amongst species in plots? Methods: First, the properties of existing measures of taxonomic similarity measures are briefly reviewed. Next, a new measure of plot‐to‐plot taxonomic similarity is presented that is based on the maximal common subgraph of two taxonomic trees. The proposed measure is computed from species presences and absences and include information about the degree of higher‐level taxonomic similarity between species plots. The performance of the proposed measure with respect to existing coefficients of taxonomic similarity and the coefficient of Jaccard is discussed using a small data set of heath plant communities. Finally, a method to quantify β‐diversity from taxonomic dissimilarities is discussed. Results: The proposed measure of taxonomic β‐diversity incorporates not only species richness, but also information about the degree of higher‐order taxonomic structure between species plots. In this view, it comes closer to a modern notion of biological diversity than more traditional measures of β‐di‐versity. From regression analysis between the new coefficient and existing measures of taxonomic similarity it is shown that there is an evident nonlinearity between the coefficients. This nonlinearity demonstrates that the new coefficient measures similarity in a conceptually different way from previous indices. Also, in good agreement with the findings of previous authors, the regression between the new index and the Jaccard coefficient of similarity shows that more than 80% of the variance of the former is explained by the community structure at the species level, while only the residual variance is explained by differences in the higher‐order taxonomic structure of the species plots. This means that a genuine taxonomic approach to the quantification of plot‐to‐plot similarity is only needed if we are interested in the residual system's variation that is related to the higher‐order taxonomic structure of a pair of species plots.     

Predictability of reef fish diversity and abundance using remote sensing data in Diego Garcia (Chagos Archipelago)

The diversity, abundance and distribution of reef fish are related to heterogeneity and physical complexity of benthic habitat. However, the field effort required to evaluate these aspects of the benthos in situ, at the scale of entire reefscapes, is greatly constrained by logistical and resource limitations. With moderate ground truthing, both substratum type and seabed topography are amenable to monitoring using satellite data. Here, remote sensing imagery was used to resolve the bathymetry and benthic character of a reef system in Diego Garcia (British Indian Ocean Territory). Replicate fish counts were made at seven measurement stations across the study area using visual census. Monte Carlo simulation revealed that species richness and abundance of several guilds and size groupings of reef fish appraised in situ were correlated with the satellite-derived seabed parameters over areas of seafloor as large as 5,030 m². The study suggests that satellite remote sensing is capable of predicting habitat complexity at a scale relevant to fish. Furthermore, as larger size classes of fish were better predicted with the satellite habitat complexity data, this technique could be used to predict fish stocks and identify potential sites for marine protected areas where intensive field surveys are not practical.     

Distribution pattern, endemism, threat status and conservation measures of fishes in the Tunga and Bhadra rivers of Western Ghats, India

This study documents the distribution patterns, endemism and uniqueness (species richness) of fishes in the Tunga and Bhadra rivers of Western Ghats, India. We recorded 77 species represented by 7 orders, 16 families and 44 genera, of which 36 species are endemic to Western Ghats, 12 species endemic to India and 26 species endemic to the Indian Subcontinent. Based on our analysis on the distribution patterns, the Tunga River is richer in diversity and higher in endemism than the Bhadra. We calculated the similarity of the species composition among sites within these two rivers using the Jacquard index. The similarity index between the sampling sites of these rivers revealed that the similarity decreases with increasing distance between the sampling sites. Of the 77 fish species we collected, 8 species (11.1%) are in the Critical category, 10 species (13.8%) are in the High Risk category, 36 species (50%) are in the Moderate category and the remaining 18 species (25%) are at lower risk. The threat status of fishes found in the Tunga and Bhadra rivers strongly suggests the need for effective conservation measures to conserve the fish species richness of these rivers.     

The assembly of ecological communities inferred from taxonomic and functional composition

Among-site variation in metacommunities (beta diversity) is typically correlated with the distance separating the sites (spatial lag). This distance decay in similarity pattern has been linked to both niche-based and dispersal-based community assembly hypotheses. Here we show that beta diversity patterns in community composition, when supplemented with functional-trait information, can be used to diagnose assembly processes. First, using simulated data, we show how the relationship between distance decay patterns in taxonomic and functional measures of community composition can be used to predict the influence of a given trait on community assembly. We then use the patterns generated by the simulation as a template to show that the sorting of benthic macroinvertebrate metacommunities in headwater streams is likely influenced by different sets of functional traits at regional and local scales. We suggest that functional-trait databases and spatially referenced taxonomic surveys can be used to predict the spatial scales at which different aspects of interspecific functional variation are involved in niche-based community assembly while accounting for the influence of dispersal-based community assembly processes.     

Is spectral distance a proxy of beta diversity at different taxonomic ranks? A test using quantile regression

Beta diversity represents a powerful indicator of ecological conditions because of its intrinsic relation with environmental gradients. In this view, remote sensing may be profitably used to derive models characterizing or estimating species turnover over an area. While several examples exist using spectral variability to estimate species diversity at several spatial scales, most of these have relied on standard correlation or regression approaches like the common Ordinary Least Square (OLS) regression which are problematic with noisy data. Moreover, very few attempts were made to derive beta diversity characterization models at different taxonomic ranks. In this paper, we performed quantile regression to test if spectral distance represents a good proxy of beta diversity considering different data thresholds and taxonomic ranks. We used plant distribution data from the North and South Carolina including 146 counties and covering a variety of vegetation formations. The dissimilarity in species composition at different taxonomic ranks (using Sørensen distance) among pairs of counties was compared with their distance in NDVI values derived from 23 yearly MODIS images. Our results indicate that (i) spectral variability represents a good proxy of beta diversity when appropriate statistics are applied and (ii) a lower taxonomic rank is important when changes in species composition are examined spatially using remotely sensed data.     

Benthic cyanobacterial diversity of Iles Eparses (Scattered Islands) in the Mozambique Channel

The marine benthic cyanobacteria of the Iles Eparses, Mozambique Channel, were surveyed for the first time. A total of 39 species are reported: 29 from Europa, 17 from Glorioso and 23 from Juan de Nova Islands. The higher biodiversity in Europa is explained by greater habitat diversity on this Island with unique ecosystems (mangroves, fossil reefs, pools). Average species richness varied between the geomorphological habitat types with higher diversity in shallow environments (fossil reef pools, mangroves, reef flats), which are characterized by high temperatures and high irradiances. The most common species observed on the three islands were Hydrocoleum coccineum, Hydrocoleum glutinosum, Hydrocoleum lyngbyaceum, Phormidium laysanense, Lyngbya sordida, and Symploca hydnoides; which are also the dominant species observed in the Southwest Indian Ocean region. The most frequent species was Phormidium laysanense with extensive cover observed in the northwest of Juan de Nova Island. Our study provided a comparison between the cyanobacterial flora of Iles Eparses and the recorded surveys in the Southwest Indian Ocean region. The low similarity observed between these species lists could be explained by differences in sampling strategies and efforts, as well as by different taxonomic approaches employed in past regional studies.     

ITS1 versus ITS2 as DNA metabarcodes for fungi

The nuclear ribosomal Internal Transcribed Spacer ITS region is widely used as a DNA metabarcoding marker to characterize the diversity and composition of fungal communities. In amplicon pyrosequencing studies of fungal diversity, one of the spacers ITS1 or ITS2 of the ITS region is normally used. In this methodological study we evaluate the usability of ITS1 vs. ITS2 as a DNA metabarcoding marker for fungi. We analyse three data sets: two comprising ITS1 and ITS2 sequences of known taxonomic affiliations and a third comprising ITS1 and ITS2 environmental amplicon pyrosequencing data. Clustering analyses of sequences with known taxonomy using the bioinformatics pipeline ClustEx revealed that a 97% similarity cut‐off represent a reasonable threshold for estimating the number of known species in the data sets for both ITS1 and ITS2. However, no single threshold value worked well for all fungi at the same time within the curated UNITE database, and we found that the Operational Taxonomic Unit (OTU) concept is not easily translated into the level of species because many species are distributed over several clusters. Clustering analyses of the 134 692 ITS1 and ITS2 pyrosequences using a 97% similarity cut‐off revealed a high similarity between the two data sets when it comes to taxonomic coverage. Although some groups are under‐ or unrepresented in the two data sets due to, e.g. primer mismatches, our results indicate that ITS1 and ITS2 to a large extent yield similar results when used as DNA metabarcodes for fungi.     

Climate and geographic distance are more influential than rivers on the beta diversity of passerine birds in Amazonia

Variation in the spatial structure of communities in terms of species composition (beta diversity) is affected by different ecological processes, such as environmental filtering and dispersal limitation. Large rivers are known as barriers for species dispersal (riverine hypothesis) in tropical regions. However, when organisms are not dispersal limited by geographic barriers, other factors, such as climatic conditions and geographic distance per se, may affect species distribution. In order to investigate the relative contribution of major rivers, climate and geographic distance on Passeriformes beta diversity, we divided Amazonia into 549 grid cells (1° of latitude and longitude) and obtained data of species occurrence, climate and geographic position for each cell. Beta diversity was measured using taxonomic, phylogenetic and functional metrics of composition. The influence of climatic variables, geographic distance and rivers on these metrics was tested using regression analyses. Passerine beta diversity is characterized mainly by the change in species taxonomic identity and in phylogenetic lineages across climatic gradients and over geographic distance. However, species with similar traits are found throughout the entire Amazonia. The size of rivers was proportional to their effect on species composition. However, climate and geographic distance are relatively more important than rivers for Amazonian taxonomic and phylogenetic species composition.     

Spatial variation in species diversity and composition of flea assemblages in small mammalian hosts: geographical distance or faunal similarity?

Aim Spatial variation in the diversity of fleas parasitic on small mammals was examined to answer three questions. (1) Is the diversity of flea assemblages repeatable among populations of the same host species? (2) Does similarity in the composition of flea assemblages among populations of the same host species decay with geographical distance, with decreasing similarity in the composition of local host faunas, or with both? (3) Does the diversity of flea assemblages correlate with climatic variables? Location The study used previously published data on 69 species of small mammals and their fleas from 24 different regions of the Holarctic. Methods The diversity of flea assemblages was measured as both species richness and the average taxonomic distinctness of their component species. Similarity between flea assemblages was measured using both the Jaccard and Morisita–Horn indices, whereas similarity in the composition of host faunas between regions (host ‘faunal’ distance) was quantified using the Jaccard index. Where appropriate, a correction was made for the potentially confounding influence of phylogeny using the independent contrasts method. Results Flea species richness varied less within than among host species, and is thus a repeatable host species character; the same was not true of the taxonomic distinctness of flea assemblages. In almost all host species found in at least five regions, similarity in flea assemblages decreased with increases in either or both geographical and faunal distance. In most host species, the diversity of flea assemblages correlated with one or more climatic variable, in particular mean winter temperature. Main conclusions Spatial variation in flea diversity among populations of the same mammal species is constrained by the fact that it appears to be a species character, but is also driven by local climatic conditions. The results highlight how ecological processes interact with co‐evolutionary history to determine local parasite biodiversity.     

Species richness and range size of the terrestrial mammals of the world: biological signal within mathematical constraints

We explore global spatial diversity patterns for terrestrial mammals using as a tool range-diversity plots. These plots display simultaneously information about the number of species in localities and their spatial covariance in composition. These are highly informative, as we show by linking range-diversity plots with maps and by highlighting the correspondences between well defined regions of the plots with geographical regions or with taxonomic groups. Range-diversity plots are mathematically constrained by the lines of maximum and minimum mean covariance in species composition. We show how regions in the range-diversity plot corresponding to the line of maximum covariance correspond to large continental masses, and regions near the lower limit of the range-diversity plot correspond to archipelagos and mountain ranges. We show how curves of constant covariance correspond to nested faunas. Finally, we show that the observed distribution of the covariance range has significantly longer tails than random, with clear geographic correspondences. At the scale of our data we found that range-diversity plots reveal biodiversity patterns that cannot be replicated by null models, and correspond to conspicuous terrain features and taxonomic groupings.     

Phylogeography of the Crown-of-Thorns Starfish in the Indian Ocean

Background

Understanding the limits and population dynamics of closely related sibling species in the marine realm is particularly relevant in organisms that require management. The crown-of-thorns starfish Acanthaster planci, recently shown to be a species complex of at least four closely related species, is a coral predator infamous for its outbreaks that have devastated reefs throughout much of its Indo-Pacific distribution.

Methodology/Principal Findings

In this first Indian Ocean-wide genetic study of a marine organism we investigated the genetic structure and inferred the paleohistory of the two Indian Ocean sister-species of Acanthaster planci using mitochondrial DNA sequence analyses. We suggest that the first of two main diversification events led to the formation of a Southern and Northern Indian Ocean sister-species in the late Pliocene-early Pleistocene. The second led to the formation of two internal clades within each species around the onset of the last interglacial. The subsequent demographic history of the two lineages strongly differed, the Southern Indian Ocean sister-species showing a signature of recent population expansion and hardly any regional structure, whereas the Northern Indian Ocean sister-species apparently maintained a constant size with highly differentiated regional groupings that were asymmetrically connected by gene flow.

Conclusions/Significance

Past and present surface circulation patterns in conjunction with ocean primary productivity were identified as the processes most likely to have shaped the genetic structure between and within the two Indian Ocean lineages. This knowledge will help to understand the biological or ecological differences of the two sibling species and therefore aid in developing strategies to manage population outbreaks of this coral predator in the Indian Ocean.     

Sequence diversity and novelty of natural assemblages of picoeukaryotes from the Indian Ocean

Despite the ecological importance of marine pico-size eukaryotes, the study of their in situ diversity using molecular tools started just a few years ago. These studies have revealed that marine picoeukaryotes are very diverse and include many novel taxa. However, the amount and structure of their phylogenetic diversity and the extent of their sequence novelty still remains poorly known, as a systematic analysis has been seldom attempted. In this study, we use a coherent and carefully curated data set of 500 published 18S ribosomal DNA sequences to quantify the diversity and novelty patterns of picoeukaryotes in the Indian Ocean. Our phylogenetic tree showed many distant lineages. We grouped sequences in OTUs (operational taxonomic units) at discrete values delineated by pair-wise Jukes–Cantor (JC) distances and tree patristic distances. At a distance of 0.01, the number of OTUs observed (237/242; using JC or patristic distances, respectively) was half the number of sequences analyzed, indicating the existence of microdiverse clusters of highly related sequences. At this distance level, we estimated 600–800 OTUs using several statistical methods. The number of OTUs observed was still substantial at higher distances (39/82 at 0.20 distance) suggesting a large diversity at high-taxonomic ranks. Most sequences were related to marine clones from other sites and many were distant to cultured organisms, highlighting the huge culturing gap within protists. The novelty analysis indicated the putative presence of pseudogenes and of truly novel high-rank phylogenetic lineages. The identified diversity and novelty patterns among marine picoeukaryotes are of great importance for understanding and interpreting their ecology and evolution.     

Areas of endemism in the Nearctic: a case study of 1339 species of Miridae (Insecta: Hemiptera) and their plant hosts

Areas of endemism are essential first hypotheses in investigating historical biogeography, but there is a surprising paucity of such hypotheses for the Nearctic region. Miridae, the plant bugs, are an excellent taxon to study in this context, because this group combines high species diversity, often small distribution ranges, a history of modern taxonomic revisions, and comprehensive electronic data capture and data cleaning that have resulted in an exceptionally error‐free geospatial data set. Many Miridae are phytophagous and feed on only one or a small number of host plant species. The programs ndm/vndm are here used on plant bug and plant data sets to address two main objectives: (i) identify areas of endemism for plant bugs based on parameters used in a recent study that focused on Nearctic mammals; and (ii) discuss hypotheses on areas of endemism based on plant bug distributions in the context of areas identified by their host plant species. Given the narrow distribution ranges of many species of Miridae, the analytical results allow for tests of the prediction that areas of endemism for Miridae are smaller and more numerous, especially in the Western Nearctic, than are those of their host plants. Analyses of the default plant bug data set resulted in 45 areas of endemism, 35 of them north of Mexico and many located in the Western Nearctic; areas in the Nearctic are more numerous and smaller than those identified by mammals. The host plant data set resulted in ten areas of endemism, and even though the size range of areas is similar between the Miridae and plant data sets, the average area size is smaller in the Miridae data set. These results allow for the conclusion that the Miridae indeed present a valuable model system to investigate areas of endemism in the Nearctic.     

Micro‐evolutionary diversification among Indian Ocean parrots: temporal and spatial changes in phylogenetic diversity as a consequence of extinction and invasion

Almost 90% of global bird extinctions have occurred on islands. The loss of endemic species from island systems can dramatically alter evolutionary trajectories of insular species biodiversity, resulting in a loss of evolutionary diversity important for species adaptation to changing environments. The Western Indian Ocean islands have been the scene of evolution for a large number of endemic parrots. Since their discovery in the 16th century, many of these parrots have become extinct or have declined in numbers. Alongside the extinction of species, a number of the Indian Ocean islands have experienced colonization by highly invasive parrots, such as the Ring‐necked Parakeet Psittacula krameri. Such extinctions and invasions can, on an evolutionary timescale, drive changes in species composition, genetic diversity and turnover in phylogenetic diversity, all of which can have important impacts on species potential for adaptation to changing environmental and climatic conditions. Using mtDNA cytochrome b data, we resolve the taxonomic placement of three extinct Indian Ocean parrots: the Rodrigues Psittacula exsul, Seychelles Psittacula wardi and Reunion Parakeets Psittacula eques. This case study quantifies how the extinction of these species has resulted in lost historical endemic phylogenetic diversity and reduced levels of species richness, and illustrates how it is being replaced by non‐endemic invasive forms such as the Ring‐necked Parakeet. Finally, we use our phylogenetic framework to identify and recommend a number of phylogenetically appropriate ecological replacements for the extinct parrots. Such replacements may be introduced once invasive forms have been cleared, to rejuvenate ecosystem function and restore lost phylogenetic diversity.     

Long‐standing environmental conditions,geographic isolation and host–symbiont specificity influence the relative ecological dominance and genetic diversification of coral endosymbionts in the genus Symbiodinium

Aim This study examines the importance of geographic proximity, host life history and regional and local differences in environment (temperature and water clarity) in driving the ecological and evolutionary processes underpinning the global patterns of diversity and distribution of symbiotic dinoflagellates. By comparing and contrasting coral–algal symbioses from isolated regions with differing environmental conditions, we may assess the potential of coral communities to respond to significant changes in climate. Location Indian Ocean. Methods Community assemblages of obligate symbiotic invertebrates were sampled at numerous sites from two regions, the north‐eastern Indian Ocean (Andaman Sea, western Thailand) and the western Indian Ocean (Zanzibar, Tanzania). Molecular genetic methods, including denaturing gradient gel electrophoresis analysis of the ribosomal internal transcribed spacers, DNA sequencing and microsatellite genotyping, were used to characterize the ‘species’ diversity and evolutionary relationships of symbiotic dinoflagellates (genus Symbiodinium). Host–symbiont specificity, geographic isolation and local and regional environmental factors were evaluated in terms of their importance in governing the distribution and prevalence of certain symbiont taxa. Results Host‐generalist symbionts (C3u and D1‐4, formerly D1a now designated Symbiodinium trenchi) frequently occurred alone and sometimes together in hosts with horizontal modes of symbiont acquisition. However, the majority of Symbiodinium diversity consisted of apparently host‐specific ‘species’. Clade C Symbiodinium were diverse and dominated host assemblages from sites sampled in the western Indian Ocean, a pattern analogous to symbiont communities on the Great Barrier Reef with similar environmental conditions. Clade D Symbiodinium were diverse and occurred frequently in hosts from the north‐eastern Indian Ocean, especially at inshore locations, where temperatures are warmer, water turbidity is high and large tidal exchanges commonly expose coral populations to aerial desiccation. Main conclusions Regional and local differences in cnidarian–algal combinations indicate that these symbioses are ecologically and evolutionarily responsive and can thrive under various environmental conditions. The high temperatures and turbid conditions of the north‐eastern Indian Ocean partly explain the ecological success of Clade D Symbiodinium relative to Clade C. Phylogenetic, ecological and population genetic data further indicate that Clade D has undergone an adaptive radiation, especially in regions around Southeast Asia, during the Pleistocene.     

Do commonly used indices of β‐diversity measure species turnover?

Abstract. Indices of β‐diversity are of two major types, (1) those that measure among‐plot variability in species composition independently of the position of individual plots on spatial or environmental gradients, and (2) those that measure the extent of change in species composition along predefined gradients, i.e. species turnover. Failure to recognize this distinction can lead to the inappropriate use of some β‐diversity indices to measure species turnover. Several commonly‐used indices of β‐diversity are based on Whittaker's β_W (β_W = γ/α, where γ is the number of species in an entire study area and α is the number of species per plot within the study area). It is demonstrated that these indices do not take into account the distribution of species on spatial or environmental gradients, and should therefore not be used to measure species turnover. The terms ‘β‐diversity’ and ‘species turnover’ should not be used interchangeably. Species turnover can be measured using matrices of compositional similarity and physical or environmental distances among pairs of study plots. The use of indices of β‐diversity and similarity‐distance curves is demonstrated using simulated data sets.     

Multi‐locus sequence data reveal a new species of coral reef goby (Teleostei: Gobiidae: Eviota), and evidence of Pliocene vicariance across the Coral Triangle

Here, multi‐locus sequence data are coupled with observations of live colouration to recognize a new species, Eviota punyit from the Coral Triangle, Indian Ocean and Red Sea. Relaxed molecular clock divergence time estimation indicates a Pliocene origin for the new species, and the current distribution of the new species and its sister species Eviota sebreei supports a scenario of vicariance across the Indo‐Pacific Barrier, followed by subsequent range expansion and overlap in the Coral Triangle. These results are consistent with the ‘centre of overlap’ hypothesis, which states that the increased diversity in the Coral Triangle is due in part to the overlapping ranges of Indian Ocean and Pacific Ocean faunas. These findings are discussed in the context of other geminate pairs of coral reef fishes separated by the Indo‐Pacific Barrier.     

An updated checklist of the ants of India with their specific distributions in Indian states (Hymenoptera,Formicidae)

As one of the 17 megadiverse countries of the world and with four biodiversity hotspots represented in its borders, India is home to an impressive diversity of life forms. However, much work remains to document and catalogue the species of India and their geographic distributions, especially for diverse invertebrate groups. In the present study, a comprehensive and critical list of Indian ant species is provided with up-to-date state-wise distribution. A total of 828 valid species and subspecies names belonging to 100 genera are listed from India. Potential erroneous data, misidentifications and dubious distributional records that may exist in the literature are also identified. The present exhaustive listing of Indian ants will provide a holistic view about diversity and distribution and will also help to identify major undersampled areas where future sampling and taxonomic efforts should be directed.