Uncoupling ecological innovation and speciation in sea snakes (Elapidae, Hydrophiinae, Hydrophiini)

The viviparous sea snakes (Hydrophiini) are by far the most successful living marine reptiles, with ～ 60 species that comprise a prominent component of shallow-water marine ecosystems throughout the Indo-West Pacific. Phylogenetically nested within the ～ 100 species of terrestrial Australo-Melanesian elapids (Hydrophiinae), molecular timescales suggest that the Hydrophiini are also very young, perhaps only ～ 8-13 Myr old. Here, we use likelihood-based analyses of combined phylogenetic and taxonomic data for Hydrophiinae to show that the initial invasion of marine habitats was not accompanied by elevated diversification rates. Rather, a dramatic three to six-fold increase in diversification rates occurred at least 3-5 Myr after this transition, in a single nested clade: the Hydrophis group accounts for ～ 80% of species richness in Hydrophiini and ～ 35% of species richness in (terrestrial and marine) Hydrophiinae. Furthermore, other co-distributed lineages of viviparous sea snakes (and marine Laticauda, Acrochordus and homalopsid snakes) are not especially species rich. Invasion of the oceans has not (by itself) accelerated diversification in Hydrophiini; novelties characterizing the Hydrophis group alone must have contributed to its evolutionary and ecological success.     

Multilocus phylogeny and recent rapid radiation of the viviparous sea snakes (Elapidae: Hydrophiinae)

The viviparous sea snakes (Hydrophiinae: Hydrophiini) comprise a young but morphologically and ecologically diverse clade distributed throughout the Indo-Pacific. Despite presenting a very promising model for marine diversification studies, many relationships among the 62 species and 16 genera in Hydrophiini remain unresolved. Here, we extend previous taxonomic and genomic sampling for Hydrophiini using three mitochondrial fragments and five nuclear loci for multiple individuals of 39 species in 15 genera. Our results highlight many of the impediments to inferring phylogenies in recent rapid radiations, including low variation at all five nuclear markers, and conflicting relationships supported by mitochondrial and nuclear trees. However, concatenated Bayesian and likelihood analyses, and a multilocus coalescent tree, recovered concordant support for primary clades and several previously unresolved inter-specific groupings. The Aipysurus group is monophyletic, with egg-eating specialists forming separate, early-diverging lineages. All three monotypic semi-aquatic genera (Ephalophis, Parahydrophis and Hydrelaps) are robustly placed as early diverging lineages along the branch leading to the Hydrophis group, with Ephalophis recovered as sister to Parahydrophis. The molecular phylogeny implies extensive evolutionary convergence in feeding adaptations within the Hydrophis group, especially the repeated evolution of small-headed (microcephalic) forms. Microcephalophis (Hydrophis) gracilis is robustly recovered as a relatively distant sister lineage to all other sampled Hydrophis group species, here termed the ‘core Hydrophis group’. Within the ‘core Hydrophis group’, Hydrophis is recovered as broadly paraphyletic, with several other genera nested within it (Pelamis, Enhydrina, Astrotia, Thalassophina, Acalyptophis, Kerilia, Lapemis, Disteira). Instead of erecting multiple new genera, we recommend dismantling the latter (mostly monotypic) genera and recognising a single genus, Hydrophis Latreille 1802, for the core Hydrophis group. Estimated divergence times suggest that all Hydrophiini last shared a common ancestor ～6 million years ago, but that the majority of extant lineages diversified over the last ～3.5 million years. The core Hydrophis group is a young and rapidly speciating clade, with 26 sampled species and 9 genera and dated at only ～1.5–3 million years old.     

Independent innovation in the evolution of paddle-shaped tails in viviparous sea snakes (Elapidae: Hydrophiinae)

The viviparous sea snakes (Hydrophiinae) comprise ~90% of living marine reptiles and display many physical and behavioral adaptations for breathing, diving, and achieving osmotic balance in marine habitats. Among the most important innovations found in marine snakes are their paddle-shaped (dorsoventrally expanded) tails, which provide propulsive thrust in the dense aquatic medium. Here, we reconstruct the evolution of caudal paddles in viviparous sea snakes using a dated molecular phylogeny for all major lineages and computed tomography of internal osteological structures. Bayesian ancestral state reconstructions show that extremely large caudal paddles supported by elongated vertebral processes are unlikely to have been present in the most recent common ancestor of extant sea snakes. Instead, these characters appear to have been acquired independently in two highly marine lineages of relatively recent origin. Both the Aipysurus and Hydrophis lineages have elongated neural spines that support the dorsal edge of their large paddles. However, whereas in the Aipysurus lineage the ventral edge of the paddle is supported by elongated haemapophyses, this support is provided by elongated and ventrally directed pleurapophyses in the Hydrophis lineage. Three semi-marine lineages (Hydrelaps, Ephalophis, and Parahydrophis) form the sister group to the Hydrophis clade and have small paddles with poorly developed dorsal and ventral supports, consistent with their amphibious lifestyle. Overall, our results suggest that not only are the viviparous hydrophiines the only lineage of marine snakes to have acquired extremely large, skeletally supported caudal paddles but also that this innovation has occurred twice in the group in the past ~2-6 million years.     

Review of sea snakes of the genus <Emphasis Type="Italic">Hydrophis</Emphasis> sensu stricto (Serpentes: Hydrophiidae)

The results of the genus Hydrophis revision s. str. are presented. The genus is divided into three subgenera, including Mediohydrophis subgen. nov. The validity of Hydrophis atriceps and H. (Microcephalophis) gracilis microcephalus is confirmed. New data on the distribution of H. klossi is given.Original Russian Text Copyright © 2004 by Biologiya Morya, Kharin.     

Seeking the sun in deep,dark places: mesopelagic sightings of ocean sunfishes (Molidae)

Evidence is presented from publicly available remotely operated vehicle (ROV) footage that suggests deep‐water ranging in ocean sunfishes (family Molidae) is more common than typically thought, including a new maximum depth recorded for the southern sunfish Mola ramsayi.     

An annotated checklist of sea snakes of Vietnam, with notes on a new record of the yellow-lipped sea krait, Laticauda colubrina (Schneider, 1799) (Laticaudidae, Hydrophiidae)

The yellow-lipped sea krait, Laticauda colubrina (Schneider, 1799), is recorded in the waters of Vietnam for the second time. The morphological characteristics of this species are described. Identification keys, a checklist, and distribution data for sea snakes of Vietnam are provided, proceeding from museum collections and extensive analysis of the available literature. Thalassophis anomalus P. Schmidt, 1852 and Leioselasma melanocephala (Gray, 1849) are registered for the first time for Vietnamese waters.     

Terrestrial locomotion in sea snakes: the effects of sex and species on cliff-climbing ability in sea kraits (Serpentes, Elapidae, Laticauda)

Ecomorphological theory predicts a match between an organism's environment and its locomotor abilities, such that animals function most effectively under the conditions they experience in nature. However, amphibious species must simultaneously optimize performance in two different habitats posing incompatible demands on locomotor morphology and physiology. This situation may generate a mismatch between environment and locomotor function, with performance optimized only for the more important habitat type; alternatively, selection may fine-tune locomotor abilities for both types of challenges. Two species of sea kraits in New Caledonia offer an opportunity to examine this question: Laticauda laticaudata is more highly aquatic than L. colubrina , and males are more terrestrial than females within each taxon. We examined an aspect of locomotor performance that is critical to coming ashore on steep-walled rocky islets: the ability to climb steep cliffs. We also measured the muscular strength of these animals, a character that is likely critical to climbing performance. Laticauda colubrina was heavier-bodied and stronger (even relative to its body mass) than the more aquatic L. laticaudata ; and within each species, males were heavier-bodied and stronger than females. The same patterns were evident in cliff-climbing ability. Thus, the ability of different species and sexes of sea kraits to climb steep cliffs correlates with their body shape even though these primarily aquatic animals use terrestrial habitats only rarely.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 433–441.     

Testing the role of climate in speciation: New methods and applications to squamate reptiles (lizards and snakes)

Climate may play important roles in speciation, such as causing the range fragmentation that underlies allopatric speciation (through niche conservatism) or driving divergence of parapatric populations along climatic gradients (through niche divergence). Here, we developed new methods to test the frequency of climate niche conservatism and divergence in speciation, and applied it to species pairs of squamate reptiles (lizards and snakes). We used a large‐scale phylogeny to identify 242 sister species pairs for analysis. From these, we selected all terrestrial allopatric pairs with sufficient occurrence records (n = 49 pairs) and inferred whether each originated via climatic niche conservatism or climatic niche divergence. Among the 242 pairs, allopatric pairs were most common (41.3%), rather than parapatric (19.4%), partially sympatric (17.7%), or fully sympatric species pairs (21.5%). Among the 49 selected allopatric pairs, most appeared to have originated via climatic niche divergence (61–76%, depending on the details of the methods). Surprisingly, we found greater climatic niche divergence between allopatric sister species than between parapatric pairs, even after correcting for geographic distance. We also found that niche divergence did not increase with time, further implicating niche divergence in driving lineage splitting. Overall, our results suggest that climatic niche divergence may often play an important role in allopatric speciation, and the methodology developed here can be used to address the generality of these findings in other organisms.     

The embryonic development of the Egyptian cobra Naja h. haje (Squamata: Serpentes: Elapidae)

The Egyptian cobra, Naja h. haje, is the largest of the African cobras and is a member of a successful and medically important species complex found throughout Africa, north and south of the Sahara, as well as across the Arabian Peninsula to Oman. Although its phylogenetic position and venom characteristics have been well studied, its development has not. Here, we present a normal staging table for N. h. haje, based on external features. Comparison with firstly the Asian monocled cobra, Naja kaouthia, and then with the small number of other oviparous snake species, allowed us to examine whether differences between two species in the same genus were of the same type and magnitude as those between unrelated genera. In fact, at least with respect to external features, we found a similar level of disparity. N. h. haje embryos lagged behind those of N. kaouthia in body and head scale development, size in ovo and hatchling length, despite having a slightly shorter incubation period and a somewhat larger adult size. Some of these differences may have been the result of differing incubation temperatures. Nonetheless, there does appear to be a broadly conserved pattern of in ovo development in at least macrostomatan snakes.     

Demographics and spatio‐temporal signature of the biotoxin domoic acid in California sea lion (Zalophus californianus) stranding records

California sea lions (Zalophus californianus) in otherwise good nutritional condition have been consistently affected by the marine biotoxin domoic acid since the late 1990s. In this study we evaluated the temporal and spatial stranding patterns of suspected and confirmed cases of domoic acid intoxicated sea lions from 1998 to 2006, using records of strandings along the California coast obtained from members of the California Marine Mammal Stranding Network. The majority of domoic acid cases were adult females (47%–82% of the total annual domoic acid cases), a contrast to strandings that were not related to domoic acid, which were generally dominated by juveniles and pups. Exposure to this biotoxin led to a 6.67‐fold increase in adult female strandings in 2000, and a 5.44‐fold increase in adult female deaths in 2006, relative to strandings and deaths of adult female not affected by domoic acid. Domoic acid cases have occurred annually since 1998 (except for 1999) between April and August, with clusters centered primarily at Pismo Beach (San Luis Obispo County), as well as at other beaches in San Luis Obispo, Monterey, Santa Cruz, Santa Barbara, Orange, and San Diego counties. The larger ecological and population level implications of increased domoic acid strandings and deaths, particularly among adult female sea lions, warrant further attention and need to be investigated.     

First records of Hippocampus algiricus in the Canary Islands (north‐east Atlantic Ocean) with an observation of hybridization with Hippocampus hippocampus

Morphometric and genetic analyses confirmed the first records of the West African seahorse Hippocampus algiricus at Gran Canaria Island (north‐east Atlantic Ocean), and also the first evidence of interspecific hybridization in seahorses. These results provide additional data on the distribution of H. algiricus that may help to establish future conservation strategies, and uncover a new potential sympatric scenario between H. algiricus and Hippocampus hippocampus.     

The fossil history of palms (Arecaceae) in Africa and new records from the Late Oligocene (28–27 Mya) of north-western Ethiopia

The African palm fossil record is limited but the data provide an outline of palm evolution from the Late Cretaceous through the Neogene. Pollen attributed to palms is reported from the Aptian (125–112 Mya), but the earliest unequivocal record in Africa is Campanian (83.5–70.6 Mya). Palms diversified 83.5–65.5 Mya and became widespread, although most records are from the west and north African coasts. Many taxa were shared between Africa and northern South America at that time, but a few were pantropical. Extirpations occurred throughout the Palaeogene, including a notable species turnover and decline at the Eocene–Oligocene boundary (33.9 Mya), a change that resulted in the elimination of nypoid palms from Africa. The Neogene plant macrofossil record is better sampled than the Palaeogene, although few palms are documented. Thus, the low diversity of African palms today is more likely the result of Palaeogene, rather than Neogene extinctions. Newly discovered palm fossils of leaves, petioles and flowers from the Late Oligocene (27–28 Mya) of north-western Ethiopia document the abundance and dominance of palms in some communities at that time. The fossils represent the earliest records of the extant genera Hyphaene (Coryphoideae) and Eremospatha (Calamoideae).  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 69–81.     

Functional morphology of the palato‐maxillary apparatus in “Palatine dragging” snakes (Serpentes: Elapidae: Acanthophis,Oxyuranus)

Elapid snakes have previously been divided into two groups (palatine erectors and palatine draggers) based on the morphology and inferred movements of their palatine bone during prey transport (swallowing). We investigated the morphology and the functioning of the feeding apparatus of several palatine draggers (Acanthophis antarcticus, Oxyuranus scutellatus, Pseudechis australis) and compared them to published records of palatine erectors. We found that the palatine in draggers does not move as a straight extension of the pterygoid as originally proposed. The dragger palato‐pterygoid joint flexes laterally with maxillary rotation when the mouth opens and the jaw apparatus is protracted and slightly ventrally during mouth closing. In contrast, in palatine erectors, the palato‐pterygoid joint flexes ventrally during upper jaw protraction. In draggers, the anterior end of the palatine also projects rostrally during protraction, unlike the stability of the anterior end seen in erectors. Palatine draggers differ from palatine erectors in four structural features of the palatine and its relationships to surrounding elements. The function of the palato‐pterygoid bar in both draggers and erectors can be explained by a typical colubroid muscle contraction pattern, which acts on a set of core characters shared among all derived snakes. Although palatine dragging elapids share a fundamental design of the palato‐maxillary apparatus with all higher snakes, they provide yet another demonstration of minor structural modifications producing functional variants. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Deep‐sea bigscales,pricklefishes, gibberfishes and whalefishes (Teleostei: Stephanoberycoidei) off Brazil: new records,range extensions for the south‐western Atlantic Ocean and remarks on the taxonomy of Poromitra

The Stephanoberycoidei includes 23 genera and c. 94 species of deep‐sea teleosts commonly known as bigscales, pricklefishes, gibberfishes and whalefishes. Stephanoberycoidei is one of the least known groups of deep‐sea fishes, in spite of their apparent relative abundance in meso and bathypelagic depths. Nine species of the Stephanoberycoidei are reported here for the first time in Brazilian waters, and most of them represent new range extensions for the south‐western Atlantic Ocean. Those species are Melamphaes polylepis, Melamphaes typhlops, Poromitra sp. and Scopeloberyx robustus (Melamphaidae), Acanthochaenus luetkenii and Stephanoberyx monae (Stephanoberycidae), Rondeletia bicolor and Rondeletia loricata (Rondeletiidae) and Gyrinomimus sp. (Cetomimidae). Occurrences of the pricklefish Scopelogadus mizolepis (Melamphaidae), the gibberfish Gibberichthys pumilus (Gibberichthyidae) and the velvet whalefish Barbourisia rufa (Barbourisiidae) are confirmed in the Brazilian exclusive economic zone, but previously published records of Poromitra capito and Melamphaes simus (Melamphaidae) in the region most likely represent misidentifications. Validities of the recently described Poromitra kukuevi and Poromitra indooceanica are discussed in light of new specimens of the genus collected in the south‐western Atlantic Ocean. An identification key for the 13 species of Stephanoberycoidei reported off Brazil is also provided.     

Resistance of the heart sea urchin Echinocardium cordatum (Echinoidea: Spatangoida) to extreme environmental changes

The ability to survive under extreme environmental conditions was studied in the adults of the heart sea urchin Echinocardium cordatum (Pennant). At seawater temperatures of 13.3 to 14.8°C and salinity of 33.2–33.4‰, being devoid of the possibility to burrow into the sand or eat, some sea urchins died on day 5 and all individuals had perished by the end of day 8. At a temperature of 19°C, the salinity tolerance range of adults was limited to 33–28‰. Only 30 to 20% of sea urchins transferred to a solid substrate survived for 7 days at a salinity of 33 to 24‰, but all of them perished toward the end of day 8.     

Trace elements in the teeth of Steller sea lions (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>) from the North Pacific

The occurrence of trace elements in the teeth of Steller sea lions (Eumetopias jubatus) collected from the North Pacific from 1968 to 1999 is described. Using particle-induced X-ray emission analysis, we detected eight trace elements: Ca, Mn, Fe, Cu, Zn, Br, Sr and Pb. We used a general linear model to evaluate long-term trends of trace element accumulation. Zn (Zn/Ca) and Fe (Fe/Ca) levels both varied during the last three decades, and showed significant differences between animals of different age (Fe/Ca), season (Zn/Ca and Fe/Ca), and sex (Zn/Ca); both Zn/Ca and Fe/Ca ratios were higher in young males than older males. Pb was detected in 39% of the samples from Hokkaido (Japan) and 22% of those from Alaska. The high levels of Pb recorded in 1975 were thought to be related to the use of leaded gasoline during the 1970s. Our results indicate that the levels of trace elements in the teeth of Steller sea lions can be used as indicators of temporal and spatial variations of trace element pollution.     

First records of lice (Insecta: Phthiraptera) from wild southern pudu, <Emphasis Type="Italic">Pudu puda</Emphasis> (Mammalia: Artiodactyla: Cervidae)

The presence of two species of Phthiraptera, Bovicola caprae (Gurlt, 1843) (Ischnocera: Bovicoliidae) and Solenopotes binipilosus (Fahrenholz, 1916) (Anoplura: Linognathidae), is reported for the first time from Pudu puda (Molina, 1782).     

First records of two species of the genus Brachionycha Hübner (Lepidoptera: Noctuidae) in China

Two species of the genus Brachionycha Hübner –B. nubeculosa (Esper) and B. albicilia Sugi – are recorded for the first time from China. The adults and genital characteristics of the species are briefly redescribed with illustrations, and their distributions are given.