Single-island Endemic Species Research Articles

Myrsine cirrhosa (Primulaceae), a distinctive new shrub species from Kaua‘i, Hawaiian Islands

Myrsine cirrhosa Lorence & K.R.Wood (Primulaceae), a new single-island endemic shrub species from Kaua‘i, Hawaiian Islands, is described and illustrated. Notes on its distribution, ecology and conservation status are included. The new species is known from an area with ca. 45 individuals, where it is restricted to the remote central windward region of Kaua‘i in open bogs and along open windy ridges. Suggested IUCN Red List status is CR (Critically Endangered). It differs from its Kaua‘i congeners by its longer petals and narrowly elliptic leaves with strongly undulate margins and tendril-like apex. Phylogenetic analysis using RADseq data supports the recognition of this new species.

Timing of diversification, dispersal, and biogeography of parrots in thegenus Amazona throughout the Caribbean, visualized in GIS

Modern bird diversity for groups in the Greater Antilles, like parrots, is a fraction of what it once was prior to human arrival in the Caribbean. Among parrot groups thought to have become once more widespread are amazons (Amazona), a genus with almost 30 species, found across Central and South America. Phylogenetic work on Amazona has determined molecularly-dated divergence times for the evolution of different species. Today, the Greater Antilles have many single-island endemic amazon species and only one species, the Rose-Throated Parrot (Amazona leucocephala) being found on multiple islands. Divergence times of amazon parrots from molecular phylogenetic dating, can be correlated to known sea-level values from marine isotope stages (MIS) of the Pliocene and Pleistocene. Bathymetric data was altered using raster math (map algebra) to create seascapes – islands/landmasses reflecting sea-level conditions during the Late Pliocene and Pleistocene to determine optimal connectivity between islands which were then made into Routes using Network Analyst in ArcGIS Pro. Exposure vs inundation of the Nicaraguan Rise likely influenced island-hopping for these parrots between the mainland and Jamaica, leading to at least two colonization events, where after which amazons colonized the rest of the Greater Antilles.

A new species of scops-owl (Aves, Strigiformes, Strigidae, Otus) from Príncipe Island (Gulf of Guinea, Africa) and novel insights into the systematic affinities within Otus.

A new species of scops-owl (Aves, Strigiformes, Strigidae, Otus) is described from Príncipe Island, São Tomé and Príncipe (Gulf of Guinea, Africa). This species was discovered for science in 2016, although suspicions of its occurrence gained traction from 1998, and testimonies from local people suggesting its existence could be traced back to 1928. Morphometrics, plumage colour and pattern, vocalisations, and molecular evidence all support the species status of the scops-owl from Príncipe, which is described here as Otusbikegila sp. nov. Phylogenetic analyses suggest that this species descended from the first colonisation of the Gulf of Guinea islands, being sister to the clade including the mainland African Scops-Owl O.senegalensis, and the island endemics Sao Tome Scops-Owl O.hartlaubi and Pemba Scops-Owl O.pembaensis. The most diagnostic trait in the field is its unique call which, curiously, is most similar to a distantly related Otus species, the Sokoke Scops-Owl O.ireneae. The new species occurs at low elevations of the old-growth native forest of Príncipe, currently restricted to the south of the island but fully included within Príncipe Obô Natural Park. Otusbikegila sp. nov. takes the number of single-island endemic bird species of Príncipe to eight, further highlighting the unusually high level of bird endemism for an island of only 139 km2.

Collecting critically endangered cliff plants using a drone-based sampling manipulator

Kauaʻi, an island within the Hawaiʻi archipelago, is home of a unique flora that contains 250 single-island endemic plant species. Threats have led to a significant population decrease where 97% of these plant species are now listed as endangered, critically endangered, or extinct. Vertical cliff habitats on Kauaʻi work as refugia to protect plants from their stressors. However, this habitat makes conservation work particularly difficult, forcing scientists, and botanists to use risky and time-consuming methods such as abseiling to access remote plant populations. Here we present the Mamba, the first aerial system capable of sampling plants on cliffs. This system is operated by two pilots and consists of an actively controlled platform suspended by a long cable under a lifting drone. Eleven otherwise inaccessible samples from five critically endangered species were collected during the first field trials on Kauaʻi Island. The samples are currently surviving in nurseries, demonstrating that the Mamba can be used to complete the conservation life cycle for organisms located in difficult areas, from location to collection, then cultivation and outplanting.

Notes on Menisciopsis wailele (Thelypteridaceae), a Critically Endangered Rheophytic Fern Species from Kaua‘i, Hawaiian Islands

Menisciopsis wailele (Thelypteridaceae) is a single-island endemic fern species restricted to the Hawaiian island of Kaua‘i. All observations indicate it is an obligate rheophyte, preferring sites of fast-moving water along concave walls of remote streams and waterfall edges. This paper presents data on its morphology, evolution, taxonomy, distribution, abundance, and ecological characteristics. Based on phylogenetic and geographic evidence, the ancestor of the Hawaiian species of Menisciopsis may have dispersed from the mountains of continental East Asia. A formal IUCN assessment of the species has been completed and is reported here as Critically Endangered (CR B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)).

A new endemic species of Hemidactylus (Squamata: Gekkonidae) from São Nicolau Island, Cabo Verde.

A new species of gecko of the genus Hemidactylus (Squamata: Gekkonidae) is described from São Nicolau Island, Cabo Verde Archipelago, and the Sal and Boavista island populations of Hemidactylus boavistensis (i.e., Hemidactylus boavistensis boavistensis comb. nov. and Hemidactylus boavistensis chevalieri comb. nov.) are recognized as subspecies. Hemidactylus nicolauensis sp. nov. is genetically distinct from H. bouvieri, to which it has previously been referred, and from all other closely related endemic Hemidactylus from Cabo Verde Islands in mitochondrial (12S cyt b) and nuclear (RAG2, MC1R) markers. It is characterized morphologically by its distinct colouration and a diagnostically different arrangement of digital lamellae. With the description of this new species, São Nicolau is now known to harbour three single-island endemic gecko species, and the documented reptile diversity in Cabo Verde is raised to 23 endemic species. As a result of our taxonomic changes, existing conservation regulations should be updated and the conservation status of these taxa should be re-evaluated.

Cryptic diversity in a vagile Hawaiian moth group suggests complex factors drive diversification

Allopatric speciation should be the dominant model of diversification across archipelagos because islands naturally promote isolation. It also follows that ecologically similar, vagile species should be more resistant to this kind of isolation due to dispersal and unifying selection. In a closely-related group of endemic Hawaiian hawkmoths, we found confounding patterns of inter-island connectivity and speciation that did not correlate with vagility, ecological specialization, or island age. Speciation occurred both in allopatric and sympatric taxa, with only the oldest and youngest islands fostering single-island endemic species. The intermediately-sized, central islands supported a combination of endemic and more widely-occurring lineages, suggesting no clear pattern leading to the current diversity in Hawaii. While some species are relatively common, others are apparently extinct or very rare, even on the same island. Further research into the specific mechanisms for these patterns in Hyles may prove broadly informative for understanding both cladogenesis and improving conservation planning. Our study identifies one new species endemic to Kauai and unique mitochondrial lineages in H. perkinsi, which may prove to be new species.

Phylogenomics clarifies biogeographic and evolutionary history, and conservation status of West Indian tremblers and thrashers (Aves: Mimidae)

The West Indian avifauna has provided fundamental insights into island biogeography, taxon cycles, and the evolution of avian behavior. Our interpretations, however, should rely on robust hypotheses of evolutionary relationships and consistent conclusions about taxonomic status in groups with many endemic island populations. Here we present a phylogenetic study of the West Indian thrashers, tremblers, and allies, an assemblage of at least 5 species found on 29 islands, including what is considered the Lesser Antilles’ only avian radiation. We improve on previous phylogenetic studies of this group by using double-digest restriction site-associated DNA sequencing (ddRAD-seq) to broadly sample loci scattered across the nuclear genome. A variety of analyses, based on either nucleotide variation in 2223 loci recovered in all samples or at 13,282 loci confidently scored as present or absent in all samples, converged on a single well-supported phylogenetic hypothesis. Results indicate that the resident West Indian taxa form a monophyletic group, exclusive of the Neotropical–Nearctic migratory Gray Catbird Dumetella carolinensis, which breeds in North America; this outcome differs from earlier studies suggesting that Gray Catbird was nested within a clade of island resident species. Thus, our findings imply a single colonization of the West Indies without the need to invoke a subsequent ‘reverse colonization’ of the mainland by West Indian taxa. Additionally, our study is the first to sample both endemic subspecies of the endangered White-breasted Thrasher Ramphocinclus brachyurus. We find that these subspecies have a long history of evolutionary independence with no evidence of gene flow, and are as genetically divergent from each other as other genera in the group. These findings support recognition of R. brachyurus (restricted to Martinique) and the Saint Lucia Thrasher R. sanctaeluciae as two distinct, single-island endemic species, and indicate the need to re-evaluate conservation plans for these taxa. Our results demonstrate the utility of phylogenomic datasets for generating robust systematic hypotheses.

Subsistence lifestyles and insular forest loss in the Louisiade Archipelago of Papua New Guinea: an endemic hotspot

Insular areas of the south-west Pacific support high levels of global biodiversity and are undergoing rapid change. The Louisiade Archipelago of Papua New Guinea is a poorly known location with high levels of endemism. The largest island, Sudest Island, supports single-island endemic species and has the largest tract of forest remaining in this island group. The islands still support traditional subsistence lifestyles. This study investigated the patterns of forest loss since 1974 and predicted future forest loss to identify areas of conservation concern. We collected village population census data to assess population growth from 1979–2011. Historical vegetation mapping from 1974 was compared with Global Forest Change data from 2000–14. The geospatial drivers of forest loss were investigated using a generalised linear mixed model. Projected forest cover loss patterns in the islands were modelled in GEOMOD to the year 2030. Resident populations grew rapidly (6.0% per year, 1979–2011) but only a low rate of forest loss (e.g. −0.035% per year, Sudest Island) was observed between 1974 and 2014, restricted to low elevations near villages. Future modelling showed varied impacts on the remaining forest extents of the larger islands. The study offers a rare contemporary example of a biodiverse hotspot that has remained relatively secure. We concluded that local cultural and environmental settings of islands in the south-west Pacific can strongly determine the patterns and processes of forest cover change, and need to be considered in programs to conserve endemic diversity.

Climatic Niche Dynamics and Its Role in the Insular Endemism of Anolis Lizards

Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.

Using a new RAD-sequencing approach to study the evolution of Micromeria in the Canary islands

As found in other oceanic islands, the Canary Islands include a large number of single island endemic species, some of which form clades that are broadly distributed within the archipelago. The genus Micromeria (Lamiaceae), for instance, includes groups of morphologically similar but ecologically diverse species on each island, representing a great model to investigate niche shifts and adaptation within the Canary Archipelago. Previous attempts to reconstruct phylogenetic relationships within the genus did not lead to robust phylogenies, presumably due to introgression and/or incomplete lineage sorting. In this study, we use a newly developed RAD-sequencing method to improve phylogenetic resolution and to better understand relationships among the Canary Island endemic Micromeria. Overall, we obtained 3571 loci that were genotyped for a total of 46 individuals of Micromeria. Our data reconstructed a highly resolved phylogeny, and corroborated the latest species reclassification of the M. varia s.l. species complex, the taxonomically most complicated group within the genus. Furthermore, taxa occupying similar ecological conditions in different islands, were shown to be closely related. This is the case of taxa from the laurel forest from La Gomera and Gran Canaria, suggesting that the laurel forest likely worked as a filter, only allowing the establishment of colonizers already pre-adapted to these conditions. We also found introgression between these species so it is also possible that the genes that facilitated the adaptation to laurel forest were swapped between Gran Canaria and La Gomera. The observations obtained in this study also allowed us to explain the role of introgression in the origin of M. varia s.l. species complex.

Melicope stonei, section Pelea (Rutaceae), a new species from Kaua'i, Hawaiian Islands: with notes on its distribution, ecology, conservation status, and phylogenetic placement.

Melicope stonei K.R. Wood, Appelhans & W.L. Wagner (section Pelea, Rutaceae), a new endemic tree species from Kaua‘i, Hawaiian Islands, is described and illustrated with notes on its distribution, ecology, conservation status, and phylogenetic placement. The new species differs from its Hawaiian congeners by its unique combination of distinct carpels and ramiflorous inflorescences arising on stems below the leaves; plants monoecious; leaf blades (5–)8–30 × (4–)6–11 cm, with abaxial surface densely tomentose, especially along midribs; and very long petioles of up to 9 cm. Since its discovery in 1988, 94 individuals have been documented and are confined to a 1.5 km2 region of unique high canopy mesic forest. Melicope stonei represents a new Critically Endangered (CR) single island endemic species on Kaua‘i.

Allium symiacum(Amaryllidaceae), a new species from Symi Island (SE Aegean, Greece)

Abstract: Allium symiacum Galanos & Tzanoud., from the island of Symi (SE Aegean, Greece), is described as a species new to science. It is an autumn-flowering, single-island endemic species of A. sect. Codonoprasum (Amaryllidaceae) and is classified as Critically Endangered according to IUCN Red List categories and criteria. Considering the morphological and karyological characters of the new species, its possible relationships to other autumnal species of A. sect. Codonoprasum distributed in the E Mediterranean area are discussed. Citation: Galanos Ch. J. & Tzanoudakis D. 2017: Allium symiacum (Amaryllidaceae), a new species from Symi Island (SE Aegean, Greece). — Willdenowia 47: 107–113. doi: https://doi.org/10.3372/wi.47.47202 Version of record first published online on 17 May 2017 ahead of inclusion in August 2017 issue.

Island Biogeography, the Effects of Taxonomic Effort and the Importance of Island Niche Diversity to Single-Island Endemic Species

Island biogeography theory is fundamentally reliant on measuring the number of species per island and hence has taxonomy at its foundation. Yet as a metric used in tests of the theory, island species richness (S) has varied with time according to the level of taxonomic effort (a function of the rate of finding and describing species). Studies using a derivative of S, single-island endemic species richness (SIE S), may be prone to change in taxonomic effort. Decreases or increases in species numbers resulting from taxonomic revision or increased sampling are likely to have a large effect on values of SIE S, as they tend to be smaller than total S for the same island. Using simple biogeography models, we analysed estimates of SIE S in plants, land snails, beetles, and fungi from comprehensive data sets for eight island groups, produced species accumulation curves and applied Bayesian regression over five time periods. Explanatory power differed across taxa, but area and island age were not always the best explanatory variables, and niche diversity appeared to be important. Changing levels of SIE S over time had different effects on models with different taxa and between island archipelagos. The results indicated that the taxonomic effort that determines SIE S is important. However, as this cannot often be quantified, we suggest Bayesian approaches should be more useful than frequentist methods in evaluating SIE S in island biogeography theory. Fundamentally, the article highlights the importance of taxonomy to theoretical biogeography.

Elevation-driven ecological isolation promotes diversification on Mediterranean islands

The percentage of single island neo-endemic species (an indicator for evolutionary diversification) was found to be independent of geographic distance to the continent in the case of the Aegean archipelago. It was concluded that speciation is independent of geographic isolation, while evolutionary processes are rather enhanced by habitat heterogeneity. An island's maximum elevation was used as an indicator for habitat heterogeneity. In contrast, we argue that habitat heterogeneity (=habitat diversity, i.e. the richness in different habitats) may be positively related to biotic richness, but a positive effect on speciation is yet to be proven. For any other type of heterogeneity, we propose a precise wording, especially when assessing its effect on speciation processes.Alternatively, we propose that elevation-driven ecological isolation causes the pattern of endemic species on high-elevation islands. Environmental filtering along an elevational gradient differentiates ecosystems, leading to an increase of isolation with elevation. The reason is that comparable ecosystems are much farther apart than is the case for lowland ecosystems. In addition, ecosystems on neighboring islands or on the continent that may be source regions for colonizing species are small in area in high elevations in comparison with low elevation ecosystems. Consequently, an increased speciation rate resulting in a larger percentage of single island endemic species can be expected for higher elevations on islands and high mountains. Support for this elevation-driven ecological isolation hypothesis comes from other islands in the Mediterranean region (e.g. Crete and Corsica), where an increase of the percentage of endemic species with elevation has been observed. Thus, the assessment of (genetic-) isolation should incorporate the distance to similar habitats instead of simple land-to-land connections.

Increase of island endemism with altitude – speciation processes on oceanic islands

Understanding speciation on oceanic islands is a major topic in current research on island biogeography. Within this context, it is not an easy task to differentiate between the influence of elevation as an indicator for habitat diversity and island age as an indicator for the time available for diversification. One reason for this is that erosion processes reduce the elevation of islands over time. In addition, the geographic distance to source ecosystems might differ among habitats, which could lead to habitat‐specific reduction of species immigration, niche occupation and diversification. We used the percentage of single island endemic species (pSIE) in five different zonal ecosystems (distributed in altitude) on the Canary Islands as an indicator for diversification. We tested whether diversification increases with altitude due to a greater ecological isolation of high elevation ecosystems on oceanic islands under the assumption of a low elevation source region on the mainland. In addition we tested whether the ‘hump‐shaped’ (unimodal) relationship between pSIE and island age as well as the linear relationship between species richness and pSIE is consistent across spatial scales. We also analyse a potential influence of island area and habitat area. We found that pSIE increases with elevation. The relations between species richness as well as age with pSIE are consistent across scales. We conclude that high elevation ecosystems are ecologically isolated. Surprisingly, the altitudinal belt with the strongest human influences has the highest values of pSIE. We successfully transfer the ‘general dynamic theory of island biogeography’ to the ecosystem scale, which provides multiple opportunities for future studies. With this approach we find that the effects of elevation on diversification can be separated from those of island age.

Biogeographical determinants of pteridophytes and spermatophytes on oceanic archipelagos

Using the data from six oceanic archipelagos, we investigated the species richness patterns on islands for all natives, archipelagic endemics and single-island endemics of pteridophytes and spermatophytes. We tested the hypothesis that the descriptive ability of biogeographical factors for species is reduced as we move from native species, to archipelagic endemics and to single island endemics, because of the increasing importance of island ‘idiosyncrasies’ (i.e. unique features of each island shaping its biota, such as catastrophic volcanic eruptions, random colonization events, mega-landslides) in controlling the species richness of endemic species. This hypothesis was addressed using two approaches: (1) the island species–area relationships (ISARs), and (2) a multiple regression approach with variable selection based on permutation, to test the combined effects of island area with other biogeographical factors. Area was an effective predictor of species richness for all native species (R 2 = 0.568 and R 2 = 0.624 for pteridophytes and spermatophytes respectively), but its predictive capacity decreased for archipelagic endemics (R 2 = 0.261 and R 2 = 0.531) and single-island endemics (R 2 = 0.084 and R 2 = 0.438). The reduction of R 2 from all native species, to archipelagic endemics and to single-island endemics was attributed to the increasing effects of the ‘idiosyncrasies’ of each island. The predictive capacity of multiple regression models increased with respect to ISARs, ranging from 27.3% (for single-island endemic pteridophytes) to 83.3% (all native pteridophytes), and included three to five predictors. Island area remained the most important variable for spermatophytes but was less important for pteridophytes. For pteridophytes, elevation was the most important predictor for native species, while isolation-related variables were the most important predictors for archipelagic endemics and single-island endemics. Our results support the hypothesis that as we move from native, to archipelagic endemic and to single-island endemic species the predictive ability of models is reduced, indicating an increased effect of the ‘idiosyncratic’ character of islands.

Biogeographical determinants for total and endemic species richness in a continental archipelago

We examined the relationship between plant species richness and biogeo- graphical variables (island area, island maximum elevation, distance from nearest inhab- ited island, distance from nearest mainland) using a data set comprising 201 islands of the Aegean archipelago. We found that endemic species richness was strongly correlated to total species richness. Single-island endemic species richness was most strongly correlated to island maximum elevation, and then to island area, with an apparent small island effect for islands smaller than 47 km 2 . Total species richness was most strongly correlated to island area (with no apparent small island effect), and less strongly correlated to island maximum elevation. Distance from the mainland or other inhabited islands displayed limited predictive value in our data set. The slope of the relationship between species richness and geographical factors (island area, elevation, distance from island/mainland) was steeper for endemic species richness than for total richness. Finally, the different scales of endemicity (single-island endemics, island group endemics and Aegean regional endemics) displayed similar qualitative trends and only differed quantitatively. Thus, we conclude that different biogeographical factors act as drivers for total species richness than for endemic species richness.

Genetic tests of rapid parallel speciation of flightless birds from an extant volant ancestor

Prior to the extinction wave that followed the human colonization of Oceania, flightless rails (Aves: Rallidae) were among the largest radiations of island birds, and perhaps the most species-rich example of convergent evolution in vertebrates. Insular flightless species are thought to have evolved from extant, volant species that colonized from continental sources and rapidly followed parallel adaptive pathways to flightlessness. The present study provides the first test of this model of speciation using genetic data sampled throughout the range of a putative ancestral species. Mitochondrial control region sequences from 71 individuals of the Gallirallus philippensis species complex reveal essentially no geographic structure within archipelagos and only weak structure among archipelagos, with no major genetic breaks except for birds sampled in the Philippines. Demographic tests of coalescent models support a recent rapid expansion into Oceania (including Australia) out of the Philippines approximately 20 000 years ago. The estimated coalescence of G. philippensis mitochondrial alleles approximately 33 000 years ago closely corresponds to the expansion of humans into the archipelagoes of Near Oceania, suggesting that humans may have facilitated its colonization by exterminating flightless competitors and clearing lowland forests. Phylogenetic analyses that included all G. philippensis haplotypes and samples from 11 single-island endemic flightless species of Gallirallus indicate that G. philippensis is polyphyletic, but is not the ancestor of most of its flightless congeners, as previously thought. Nuclear gene sequences (β-actin inron 3) suggest that G. philippensis polyphyly is at least partly due to hybridization. The flightless condition evolves in rails before reproductive isolation is complete. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 601–616.

Phylogeography and ecology of an endemic radiation of Hawaiian aquatic case-bearing moths (Hyposmocoma: Cosmopterigidae)

The endemic moth genus Hyposmocoma (Lepidoptera: Cosmopterigidae) may be one of the most speciose and ecologically diverse genera in Hawaii. Among this diversity is the Hyposmocoma saccophora clade with previously unrecorded aquatic larvae. I present a molecular phylogeny based on 773 base pairs (bp) of the mitochondrial gene cytochrome c oxidase subunit I and 762 bp of the nuclear gene elongation factor 1-alpha. Topologies were constructed from data using maximum-parsimony, maximum-likelihood and Bayesian search criteria. Results strongly support the monophyly of the H. saccophora clade and the monophyly of the genus Hyposmocoma. The H. saccophora clade has single-island endemic species on Oahu, Molokai and West Maui. By contrast, there are three species endemic to Kauai, two being sympatric. The H. saccophora clade appears to follow the progression rule, with more basal species on older islands, including the most basal species on 11 Myr-old Necker Island, one of the Northwestern Hawaiian Islands. Aquatic behaviour either evolved recently in the species on the main Hawaiian Islands or was secondarily lost on the arid northwestern Necker Island. The phylogeny suggests that Hyposmocoma is older than any of the current main islands, which may, in part, explain Hyposmocoma's remarkable diversity.