Cryptic species,taxonomic inflation,or a bit of both? New species phenomenon in Sri Lanka as suggested by a phylogeny of dwarf geckos (Reptilia,Squamata, Gekkonidae,Cnemaspis)

We evaluated the status of 16 of 22 recognized Sri Lankan Cnemaspis Strauch species, and flagged overlooked diversity with two mitochondrial (cyt b & ND2) and two nuclear markers (RAG1 & PDC) totalling 2829 base pairs. A fossil-calibrated timetree and sampling of other South Asian Cnemaspis provide insights into the diversification of the genus in peninsular India and Sri Lanka. Phylogenetic analyses consistently inferred two broad clades within South Asian Cnemaspis, with Sri Lankan species in two clades, which we call the podihuna and kandiana clades. Each Sri Lankan clade as a whole is sister to Indian taxa and nested within Indian lineages. Cnemaspis modigliani Das from Indonesia is a member of the kandiana clade. This suggests a minimum of two dispersal events between India and Sri Lanka and one between Sri Lanka/India and South-east Asia. South Asian Cnemaspis date back to at least the Eocene, in Sri Lanka to the early Miocene, with late Miocene diversification in the kandiana clade. All but one of the named species we sampled is likely to be valid, and 10 divergent unnamed lineages may warrant specific recognition. A resolution of Sri Lankan Cnemaspis taxonomy will require thorough sampling and the use of both morphological and molecular data.     

Genetic structure of the beaked whale genus Berardius in the North Pacific,with genetic evidence for a new species

There are two recognized species in the genus Berardius, Baird's and Arnoux's beaked whales. In Japan, whalers have traditionally recognized two forms of Baird's beaked whales, the common “slate‐gray” form and a smaller, rare “black” form. Previous comparison of mtDNA control region sequences from three black specimens to gray specimens around Japan indicated that the two forms comprise different stocks and potentially different species. We have expanded sampling to include control region haplotypes of 178 Baird's beaked whales from across their range in the North Pacific. We identified five additional specimens of the black form from the Aleutian Islands and Bering Sea, for a total of eight “black” specimens. The divergence between mtDNA haplotypes of the black and gray forms of Baird's beaked whale was greater than their divergence from the congeneric Arnoux's beaked whale found in the Southern Ocean, and similar to that observed among other congeneric beaked whale species. Taken together, genetic evidence from specimens in Japan and across the North Pacific, combined with evidence of smaller adult body size, indicate presence of an unnamed species of Berardius in the North Pacific.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

Review of the distribution of Mesoplodon species (order Cetacea, family Ziphiidae) in the North Atlantic

Four species of beaked whales of the genus Mesoplodon regularly occur in the North Atlantic. These are Sowerby's Beaked Whale Mesoplodon bidens, Blainville's Beaked Whale Mesoplodon densirostris, Gervais Beaked Whale Mesoplodon europaeus and True's Beaked Whale Mesoplodon mirus. These four species differ in their distribution within this area. M. bidens is the most northerly recorded species followed by M. mirus. M. densirostris and M. europaeus both generally occur further south and their distributions cross the equator. The distribution of Mesoplodon species may relate to variations in water temperature.     

Declining Abundance of Beaked Whales (Family Ziphiidae) in the California Current Large Marine Ecosystem

Beaked whales are among the most diverse yet least understood groups of marine mammals. A diverse set of mostly anthropogenic threats necessitates improvement in our ability to assess population status for this cryptic group. The Southwest Fisheries Science Center (NOAA) conducted six ship line-transect cetacean abundance surveys in the California Current off the contiguous western United States between 1991 and 2008. We used a Bayesian hidden-process modeling approach to estimate abundance and population trends of beaked whales using sightings data from these surveys. We also compiled records of beaked whale stranding events (3 genera, at least 8 species) on adjacent beaches from 1900 to 2012, to help assess population status of beaked whales in the northern part of the California Current. Bayesian posterior summaries for trend parameters provide strong evidence of declining beaked whale abundance in the study area. The probability of negative trend for Cuvier''s beaked whale (Ziphius cavirostris) during 1991–2008 was 0.84, with 1991 and 2008 estimates of 10771 (CV = 0.51) and ≈7550 (CV = 0.55), respectively. The probability of decline for Mesoplodon spp. (pooled across species) was 0.96, with 1991 and 2008 estimates of 2206 (CV = 0.46) and 811 (CV = 0.65). The mean posterior estimates for average rate of decline were 2.9% and 7.0% per year. There was no evidence of abundance trend for Baird''s beaked whale (Berardius bairdii), for which annual abundance estimates in the survey area ranged from ≈900 to 1300 (CV≈1.3). Stranding data were consistent with the survey results. Causes of apparent declines are unknown. Direct impacts of fisheries (bycatch) can be ruled out, but impacts of anthropogenic sound (e.g., naval active sonar) and ecosystem change are plausible hypotheses that merit investigation.     

Encounters with the Cuvier’s beaked whale, <Emphasis Type="Italic">Ziphius cavirostris</Emphasis> Cuvier, 1823 (Cetacea: Ziphiidae), in waters off eastern Kamchatka and the Commander Islands

The biology of the Cuvier’s beaked whale, Ziphius cavirostris Cuvier, 1823, in Russian waters remains almost unstudied. The species is encountered rarely elsewhere in the seas of Russia, except for the waters off the Commander Islands. Currently, some researchers express doubt about the accuracy of the known Cuvier’s beaked whale records in the region. This report provides information on the confirmed encounters with Cuvier’s beaked whale off the Commander Islands and eastern Kamchatka.     

Does a lack of observed beaked whale strandings in military exercise areas mean no impacts have occurred? A comparison of stranding and detection probabilities in the Canary and main Hawaiian Islands

Anthropogenic activities must be monitored to determine effects on marine mammal species, but the difficulty lies in how to measure impact. Mass strandings of beaked whales have occurred in association with naval exercises, with two species most affected, Cuvier's (Ziphius cavirostris) and Blainville's (Mesoplodon densirostris) beaked whales. Six such events have occurred in the Canary Islands but there have been no reported mass strandings in Hawai‘i. We assess the hypothesis that factors that influence the likelihood of strandings occurring and/or being detected differ between the Canary and main Hawaiian Islands, such that beaked whale stranding/detection probabilities will be lower in Hawai‘i. On an archipelago-wide basis, nearshore bathymetric comparisons indicate that the Canaries have a greater proportion and a total greater amount of appropriate beaked whale habitat closer to shore, with a steeper slope. Hawaiian shorelines are more dominated by steep cliffs, human population density is much lower, and human population per kilometer of shoreline is 53% lower than in the Canaries. All of these factors suggest that there is a higher probability of a carcass washing onshore and being detected in the Canary Islands. It cannot be concluded that the lack of mass strandings in Hawai‘i is evidence of no impact.     

Two new species of Eutrombidium Verdun, 1909 (Acari: Prostigmata: Eutrombidiidae) from Asia and a new host record for this genus

Two new species of larval mites are described Eutrombidium laosanum n. sp. ectoparasitic on undetermined Gryllidae (Orthoptera) from Laos and Indonesia (Lingga Islands) and E. sigirijanum n. sp. ectoparasitic on undetermined Tenebrionidae (Coleoptera) from Sri Lanka.     

Diversity,phylogeny and biogeography of Systomus (Teleostei,Cyprinidae) in Sri Lanka

The South and South-East Asian freshwater fish genus Systomus (Cyprinidae) comprises 17 valid species. Six nominal species, including three endemics, have been reported from Sri Lanka, a continental island separated from India by a shallow-shelf sea. The species diversity of Systomus on the island has until now not been assessed; neither has an evaluation been made of their phylogenetic history. Here, based on an analysis of the nuclear recombination activating protein 1 (rag1), and mitochondrial cytochrome c oxidase subunit 1 (cox1) and cytochrome b (cytb) gene markers, and a morphological examination of 143 specimens from 49 locations in Sri Lanka, we reassess the diversity of Systomus on the island and analyse patterns of their evolution and biogeography. Divergence-time estimates, based on a substitution rate calibration, date the basal split between Systomus and its sister group, the Afrotropical small barbs, to 30.0 Ma (95% highest posterior density: 25.4–35.2 Ma). The species of Systomus belong to two distinct clades. The first includes the Sri Lankan endemics S. asoka, S. martenstyni and S. pleurotaenia, which comprise an insular diversification following the immigration of a common ancestor during the Oligocene. The second, which includes the remaining species of Indian, Sri Lankan and South-East Asian Systomus, has a crown age dating to the Late Miocene. Morphological and molecular species delimitation analyses failed to validate the two nominal species, S. spilurus and S. timbiri, previously reported from Sri Lanka: both are considered synonyms of S. sarana, as are the nomina S. chryseus, S. chrysopoma, S. laticeps, S. rufus, S. pinnauratus and S. subnasutus. Four genetically and geographically discrete lineages of S. sarana occur in the island, and three in India. Molecular species delimitation analysis suggests these all belong to a single species, S. sarana. The genetically distinct Sri Lankan populations of S. sarana result from Plio-Pleistocene dispersal or vicariance events between India and Sri Lanka—as a result of emergence and inundation of the now submerged isthmus connecting the two landmasses—as well as autochthonous insular diversification.     

Genus-level phylogeny of snakes reveals the origins of species richness in Sri Lanka

Snake diversity in the island of Sri Lanka is extremely high, hosting at least 89 inland (i.e., non-marine) snake species, of which at least 49 are endemic. This includes the endemic genera Aspidura, Balanophis, Cercaspis, Haplocercus, and Pseudotyphlops, which are of uncertain phylogenetic affinity. We present phylogenetic evidence from nuclear and mitochondrial loci showing the relationships of 40 snake species from Sri Lanka (22 endemics) to the remaining global snake fauna. To determine the phylogenetic placement of these species, we create a molecular dataset containing 10 genes for all global snake genera, while also sampling all available species for genera with endemic species occurring in Sri Lanka. Our sampling comprises five mitochondrial genes (12S, 16S, cyt-b, ND2, and ND4) and five nuclear genes (BDNF, c-mos, NT3 RAG-1, and RAG-2), for a total of up to 9582 bp per taxon. We find that the five endemic genera represent portions of four independent colonizations of Sri Lanka, with Cercaspis nested within Colubrinae, Balanophis in Natricinae, Pseudotyphlops in Uropeltidae, and that Aspidura + Haplocercus represents a distinct, ancient lineage within Natricinae. We synonymize two endemic genera that render other genera paraphyletic (Haplocercus with Aspidura, and Cercaspis with Lycodon), and discover that further endemic radiations may be present on the island, including a new taxon from the blindsnake family Typhlopidae, suggesting a large endemic radiation. Despite its small size relative to other islands such as New Guinea, Borneo, and Madagascar, Sri Lanka has one of the most phylogenetically diverse island snake faunas in the world, and more research is needed to characterize the island’s biodiversity, with numerous undescribed species in multiple lineages.     

Molecular genetic identification of southern hemisphere beaked whales (Cetacea: Ziphiidae)

To assist in the species-level identification of stranded and hunted beaked whales, we compiled a database of 'reference' sequences from the mitochondrial DNA control region for 15 of the 20 described ziphiid species. Reference samples for eight species were obtained from stranded animals in New Zealand and South Australia. Sequences for a further seven species were obtained from a previously published report. This database was used to identify 20 'test' samples obtained from incompletely documented strandings around New Zealand. Analyses showed that four of these 'test' specimens (20%) had initially been misidentified. These included two animals of particular interest: (i) a Blainville's beaked whale ( Mesoplodon densirostris) , the first record of this species in New Zealand waters; and, (ii) a juvenile Andrews' beaked whale ( Mesoplodon bowdoini ), a species known from just over 20 strandings worldwide. A published sequence from a beaked whale product purchased in the Republic of Korea was identified as a Cuvier's beaked whale ( Ziphius cavirostris ). Levels of intra- and interspecific variation were compared to determine the potential for misidentification when the database or taxonomy is incomplete. Intraspecific variation was generally <2%, and interspecific divergence was generally >4.7%. Exceptions were within-species variation in Hyperoodon planifrons , southern bottlenosed whale (4.12%), which exceeded the variation between the two species of Berardius (3.78%), and variation between the two specimens assigned to M. hectori , Hector's beaked whale (7.14%). The latter case appears to be an error in species identification, and could represent the discovery of a new species of beaked whale.     

Colonization and speciation on volcanic islands: phylogeography of the flightless grasshopper genus Arminda (Orthoptera,Acrididae) on the Canary Islands

Abstract Volcanic archipelagos represent excellent areas to study colonization and speciation processes. The grasshopper genus Arminda is one of many endemic taxa of the Canary Islands. It consists of seven wingless species, most of which are single‐island endemics. We sequenced two mitochondrial (12s rRNA, ND5) and two nuclear gene fragments (28s rRNA, ITS2) to reconstruct the colonization pattern of the genus. Our results are in accordance with a stepping‐stone colonization model from east to west, corresponding to the prevailing ocean currents, but alternative hypotheses cannot be fully rejected. The populations of A. brunneri from Tenerife belong to two different lineages (east and west) consistent with the geological history of the island. It remains to be tested whether these lineages represent different species and whether further lineages exist on this island. The five clades of the four western islands (A. brunneri group) have approximately similar branch lengths. The short internodes between these lineages resulted in a poorer phylogenetic resolution. Specimens from La Palma were genetically distinct and are subsequently described as a new species, Arminda palmae sp.n. Our results suggest in situ speciation on Gran Canaria, which was accompanied by a stronger degree of morphological diversification than the inter‐island speciation processes. The aberrant species A. canariensis has formerly been assigned to a monotypic subgenus Chopardminda, which is now synonymized with Arminda syn.n. based on its phylogenetic position. Gran Canaria seems to be the only island where Arminda species occur sympatrically, although allopatric speciation seems likely due to the long history of volcanism and erosion on the island.     

Redescription of Amblyomma integrum adults and immature stages

Amblyomma integrum Karsch, 1879 (Acari: Ixodidae) is one of four Amblyomma Koch, 1844 species with eyes found in southern India and Sri Lanka. The immature stages of this species were poorly described. Therefore, accurate identification is difficult. Here we re‐describe the male, female, nymph and larva of A. integrum and illustrate all the stages in greater detail for the first time. A set of diagnostic morphological characters is defined to distinguish this species from other sympatric species of eyed Amblyomma in any parasitic stage of development. Adults of A. integrum parasitize mostly various larger mammals whereas nymphs and larvae use mostly larger and medium mammals. Amblyomma integrum is recorded from India (Andhra Pradesh, Goa, Karnataka, Orissa and Tamil Nadu States) and throughout Sri Lanka.     

Molecular evidence that the deadliest sea snake Enhydrina schistosa (Elapidae: Hydrophiinae) consists of two convergent species

We present a striking case of phenotypic convergence within the speciose and taxonomically unstable Hydrophis group of viviparous sea snakes. Enhydrina schistosa, the ‘beaked sea snake’, is abundant in coastal and inshore habitats throughout the Asian and Australian regions, where it is responsible for the large majority of recorded deaths and injuries from sea snake bites. Analyses of five independent mitochondrial and nuclear loci for populations spanning Australia, Indonesia and Sri Lanka indicate that this ‘species’ actually consists of two distinct lineages in Asia and Australia that are not closest relatives. As a result, Australian “E. schistosa” are elevated to species status and provisionally referred to Enhydrina zweifeli. Convergence in the characteristic ‘beaked’ morphology of these species is probably associated with the wide gape required to accommodate their spiny prey. Our findings have important implications for snake bite management in light of the medical importance of beaked sea snakes and the fact that the only sea snake anti-venom available is raised against Malaysian E. schistosa.     

A redescription ofDiaphanosoma modigliani Richard, 1894 (Crustacea : Daphniiformes : Sididae)and an evaluation of its occurence outside Lake Toba (Sumatra)

A pelagic species of Diaphanosoma, D. modigliani is redescribed on material from the type locality (Lake Toba, Sumatra, Indonesia). Other records of this species in tropical Asia (India, Philippine Islands, Malaysia, Sri Lanka, Thailand) are either unsubstantiated or erroneous.     

Phylogenetic position of the Ohiya rat (Srilankamys ohiensis) based on mitochondrial and nuclear gene sequence analysis

Buzan, E. V., Pagès, M., Michaux, J. & Krystufek, B. (2011). Phylogenetic position of the Ohiya rat (Srilankamys ohiensis) based on mitochondrial and nuclear gene sequence analysis. —Zoologica Scripta, 40, 545–553. We investigated the phylogenetic position the Ohiya rat, endemic to Sri Lanka, Srilankamys (Rodentia, Murinae), within the tribe Rattini based on the combined analysis of three independent genes (a mitochondrial one and two nuclear exons). Three major lineages (the Maxomys, the Dacnomys and the Rattus divisions) were retrieved as monophyletic groups within the tribe Rattini. Srilankamys was not affiliated to any of the representatives of the Dacnomys division as it was supposed based on morphological characters, but clearly appeared as the first genus to diverge among the Rattus division. The Mindanao Shrew Mouse, Crunomys melanius, emerged as a part of the Maxomys division raising questions about the validity of the Crunomys and the Maxomys divisions as currently defined. Molecular date of divergence between Srilankamys and the other representatives of the Rattus division falls within the interval 6.7 ± 0.74 Mya, coinciding with the time of the isolation of Sri Lanka from the Deccan peninsula and the aridification period owing to the climate change at the end of the Miocene epoch. We suggest that the isolation of Sri Lanka from the continent, reinforced by the action of a seasonal monsoon‐dominated climate, would have led to the isolation of some ancestral rodents of the Rattus division, which would have differentiated later into the Ohiya rat by a vicariant process. In a more general point of view, our study supports the previous results obtained on other organisms and evidence that Sri Lanka appears to be characterized by a particular fauna as compared to the Indian mainland. This island would therefore be considered as a specific distinct hotspot of biodiversity.     

The phylogenetic history of the old world butterfly subtribe Mycalesina extended: the Mycalesis (Lepidoptera: Nymphalidae) of Sri Lanka

Sri Lanka, together with the Western Ghats, is one of the world's biodiversity hotspots, yet little is known about the genetic diversity of the butterflies here. Within the framework of the recently reconstructed molecular phylogeny for the subtribe Mycalesina, we determined the phylogenetic relationships and biogeographic history of the five species of Mycalesis butterflies that are found in Sri Lanka. Sequences of Elongation Factor 1-α (EF1-α), Wingless (Wgl) and the barcode region of the Cytochrome c Oxidase sub unit 1 (COI) were used to confirm the identity of species, to resolve taxonomic queries and to infer the phylogenetic history of the group.Time-calibrated analysis of genetic data suggests that the Mycalesis species that occur in Sri Lanka diverged in the Miocene (5–23 million years ago). The results of phylogenetic analyses confirmed the following; the endemicity of Mycalesis rama to Sri Lanka; the subspecies classification of M. perseus typhlus and, the close sister relationship of the Sri Lankan M. patnia patnia to M. p. junonia of India. However, whether or not M. subdita of Sri Lanka and India are genetically similar remains unknown. Lastly, genetic evidence emerged suggesting that M. mineus forms a cryptic species complex in the Oriental region, and that in Sri Lanka, there may be occasional hybridization between M. mineus and M. perseus. As a case study of island colonization and diversification by the Mycalesis species, this study further extends our understanding of the Old World butterfly subtribe of Mycalesina.     

Color patterns in white‐beaked dolphins (Lagenorhynchus albirostris) from Iceland

Few studies have examined the natural complex pigmentation patterns of white‐beaked dolphins. From 2002 to 2014, whale‐watching trips in Iceland provided a platform of opportunity to collect a large body of photographs of free‐ranging individuals from a single area of distribution for this species. Based on 823 images, 571 individuals showing one or more color components were identified, and assigned to the following four age classes: adults (n = 437), juveniles (n = 109), calves (n = 14), and neonates (n = 11). A total of 26 color components were observed and described: seven terms previously applied to white‐beaked dolphins, 12 previously applied to other dolphin species, and seven newly defined terms. Results showed that each age class could be positively identified by differences in specific color components, some of which were exclusive. Therefore, color patterns may prove useful in estimating maturity in free‐ranging white‐beaked dolphins. This tool could be further refined through assessment of a wide sample of freshly stranded specimens of known sex and age, which could reveal new age class‐specific components, as well as sexually dimorphic characteristics not seen here. Geographic variation should be investigated by comparing image data sets and stranded animals from different parts of the North Atlantic.     

Island hopping across the central Pacific: mitochondrial DNA detects sequential colonization of the Austral Islands by crab spiders (Araneae: Thomisidae)

Aim Phylogenetic studies concerning island biogeography have been concentrated in a fraction of the numerous hot‐spot archipelagos contained within the Pacific Ocean. In this study we investigate relationships among island populations of the thomisid spider Misumenops rapaensis Berland, 1934 across the Austral Islands, a remote and rarely examined southern Pacific hot‐spot archipelago. We also assess the phylogenetic position of M. rapaensis in relation to thomisids distributed across multiple Polynesian archipelagos in order to evaluate the proposed hypothesis that thomisid spiders colonized Polynesia from multiple and opposing directions. The data allow an examination of genetic divergence and species accumulation in closely related lineages distributed across four Polynesian archipelagos. Location The study focused on four Polynesian hot‐spot archipelagos: the Austral, Hawaiian, Marquesan and Society islands. Methods Mitochondrial DNA sequences comprising c. 1400 bp (portions of cytochrome oxidase subunit I, ribosomal 16S and NADH dehydrogenase subunit I) were obtained from thomisid spiders (64 specimens, representing 33 species) collected in the Australs, the Hawaiian Islands, the Society Islands, the Marquesas, Tonga, Fiji, New Zealand, New Caledonia and North and South America. Phylogenetic analyses using parsimony, maximum‐likelihood and Bayesian approaches were employed to resolve relationships of M. rapaensis to other Polynesian Misumenops and across the Austral Islands. Results Rather than grouping with other Misumenops spp. from the archipelagos of the Society Islands, Marquesas and Hawaiian Islands, M. rapaensis appears more closely related to Diaea spp. from Tonga, Fiji, New Zealand and New Caledonia. Phylogenetic analyses strongly support M. rapaensis as monophyletic across the Austral Islands. Misumenops rapaensis sampled from the two older islands (Rurutu and Tubuai) form reciprocally monophyletic groups, while individuals from the younger islands (Raivavae and Rapa) are paraphyletic. Across the Austral Islands, M. rapaensis exhibits a surprising level of genetic divergence (maximally 11.3%), an amount nearly equivalent to that found across the 16 examined Hawaiian species (14.0%). Main conclusions Although described as a single morphologically recognized species, our results suggest that M. rapaensis comprises multiple genetically distinct lineages restricted to different Austral Islands. Phylogenetic relationships among the island populations are consistent with sequential colonization of this lineage down the Austral archipelago toward younger islands. Analyses support the hypothesis that thomisid spiders colonized the central Pacific multiple times and suggest that M. rapaensis arrived in the Austral Islands from a westward direction, while Misumenops found in neighbouring archipelagos appear to be more closely related to New World congeners to the east. Finally, our data detect asymmetrical rates of morphological evolution and species diversification following colonization of four different Polynesian archipelagos.     

A new genus and species of Baetidae from Sri Lanka (Ceylon): Indocloeon primum gen.n., sp.n. (Insecta,Ephemeroptera)

The nymph of a new genus and species, Indocloeon primum gen.n., sp.n. from Sri Lanka is described, distinguishing characteristics are illustrated. This new genus is most closely related to Cloeon Leach and Centroptilum Eaton. Indocloeon gen.n. possesses a unique combination of morphological characters and two derived characters.