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Román-Palacios, C, Damgaard J, Cheng L, Moreira FFF, Ikawa T, Weir TA, Zettel H.  2018.  Molecular phylogeny of sea-skaters (Halobates ), relationship between Halobatini and Metrocorini, and a catalogue of the subfamily Halobatinae (Hemiptera: Heteroptera: Gerridae). Insect Systematics & Evolution. :1-102., Leiden, The Netherlands: Brill   10.1163/1876312X-00002197   Abstract

We investigated the phylogenetic relationships among seven of the ten Halobatinae genera (Heteroptera: Gerridae) based on COI+II, 16S rRNA, and 28S rRNA genes. Our analyses recovered monophyly of Halobatinae, and suggested paraphyly of Metrocorini caused by the position of Ventidius and Esakia. Since our phylogenies did not infer monophyly of the subgenus Halobates (s.str.) within Halobatini, we synonymized Austrobates and Halobates (Hilliella) with Halobates. We confirmed that (1) the limnic lifestyle of Metrocorini was ancestral in Halobatinae, (2) the marine lifestyle evolved only once in the common ancestor of Asclepios + Halobates, (3) the limnic lifestyle of some members of Halobates was independently derived from marine ancestors, and that (4) the open ocean was colonized at least three times in Halobates. A catalogue of Halobatinae organized according to an updated classification is presented, including all known geographic distributions, bibliographical references, and additional notes to all species of the subfamily.

Tran, AD, Yang CM, Cheng L.  2015.  Water bugs of Singapore and Peninsular Malaysia. , Singapore: Lee Kong Chian Natural History Museum
Cheng, LN, Huang DW.  2014.  Discovery of the flightless marine midge Pontomyia (Diptera: Chironomidae) at Christmas Island, Australia. Raffles Bulletin of Zoology. :93-96. AbstractWebsite

We document the discovery of a unique flightless marine midge Pontomyia (Chironomidae) in the Indian Ocean, presenting descriptions and photographs of the insect and its environment at Christmas Island, Australia. The insect is identified as P. natans, with distinctive male genitalia and rudimentary claws on the hind legs. This species is widely distributed in the Pacific Ocean but the only previous record from the Indian Ocean was from the Maldives. It is highly likely that other populations are present along the continental coast of the latter ocean basin.

Engels, M, Correia L, Piwinski S, Cheng LN, Zettler E.  2014.  Seasonal and decadal changes in distribution patterns of Halobates (Hemiptera: Gerridae) populations in the eastern tropical Pacific. Marine Biology. 161:1241-1250.   10.1007/s00227-014-2414-x   AbstractWebsite

Five species of the marine insect Halobates share similar ecology but have distinct biogeographic ranges in the eastern tropical Pacific, a region from approximately 75A degrees W-160A degrees W and 10A degrees S-35A degrees N. Between 2001 and 2010, the Sea Education Association collected Halobates from 682 neuston tows (surface net 1 m x 0.5 m, 335-mu m mesh) during fifteen cruises between San Diego, USA, Mexico and Tahiti. Total Halobates spp. densities varied substantially from year to year, but our data do not show a sustained change from a data set collected 40 years earlier from 1967 to 1968 (Cheng and Shulenberger in Fish Bull 78(3):579-591, 1980). Halobates are sensitive to sea surface temperature and we observed significant differences in species distributions over time, but these were not due to differences in water temperature or climate change. Our analyses show that the patterns observed are attributable to substantial but previously undescribed seasonal shifts that occur each year in the ranges for both Halobates sobrinus and Halobates micans. There is substantial overlap in ranges during seasonal shifts, but very little co-occurrence of H. sobrinus and H. micans in individual net tows, suggesting biological mechanisms rather than physical factors are restricting distribution and co-occurrence of these two species.

Goldstein, MC, Rosenberg M, Cheng LN.  2012.  Increased oceanic microplastic debris enhances oviposition in an endemic pelagic insect. Biology Letters. 8:817-820.   10.1098/rsbl.2012.0298   AbstractWebsite

Plastic pollution in the form of small particles (diameter less than 5 mm)-termed `microplastic'-has been observed in many parts of the world ocean. They are known to interact with biota on the individual level, e.g. through ingestion, but their population-level impacts are largely unknown. One potential mechanism for microplastic-induced alteration of pelagic ecosystems is through the introduction of hard-substrate habitat to ecosystems where it is naturally rare. Here, we show that microplastic concentrations in the North Pacific Subtropical Gyre (NPSG) have increased by two orders of magnitude in the past four decades, and that this increase has released the pelagic insect Halobates sericeus from substrate limitation for oviposition. High concentrations of microplastic in the NPSG resulted in a positive correlation between H. sericeus and microplastic, and an overall increase in H. sericeus egg densities. Predation on H. sericeus eggs and recent hatchlings may facilitate the transfer of energy between pelagic-and substrate-associated assemblages. The dynamics of hard-substrate-associated organisms may be important to understanding the ecological impacts of oceanic microplastic pollution.

Appeltans, W, Ahyong ST, Anderson G, Angel MV, Artois T, Bailly N, Bamber R, Barber A, Bartsch I, Berta A et al..  2012.  The Magnitude of Global Marine Species Diversity. Current Biology. 22:2189-2202.   10.1016/j.cub.2012.09.036   AbstractWebsite

Background: The question of how many marine species exist is important because it provides a metric for how much we do and do not know about life in the oceans. We have compiled the first register of the marine species of the world and used this baseline to estimate how many more species, partitioned among all major eukaryotic groups, may be discovered. Results: There are similar to 226,000 eukaryotic marine species described. More species were described in the past decade (similar to 20,000) than in any previous one. The number of authors describing new species has been increasing at a faster rate than the number of new species described in the past six decades. We report that there are similar to 170,000 synonyms, that 58,000-72,000 species are collected but not yet described, and that 482,000-741,000 more species have yet to be sampled. Molecular methods may add tens of thousands of cryptic species. Thus, there may be 0.7-1.0 million marine species. Past rates of description of new species indicate there may be 0.5 +/- 0.2 million marine species. On average 37% (median 31%) of species in over 100 recent field studies around the world might be new to science. Conclusions: Currently, between one-third and two-thirds of marine species may be undescribed, and previous estimates of there being well over one million marine species appear highly unlikely. More species than ever before are being described annually by an increasing number of authors. If the current trend continues, most species will be discovered this century.

Leo, SST, Cheng L, Sperling FAH.  2012.  Genetically separate populations of the ocean-skater Halobates sericeus (Heteroptera: Gerridae) have been maintained since the late Pleistocene. Biological Journal of the Linnean Society. 105:797-805.   10.1111/j.1095-8312.2011.01836.x   AbstractWebsite

The oceanic water strider (or ocean-skater) Halobates sericeus Eschscholtz has a disjunct distribution in the Pacific Ocean, with northern and southern populations widely separated by an equatorial zone. It is sensitive to sea surface conditions and, consequently, its distribution and population structure may provide an insight into environmental changes on the ocean surface on both recent and historical time scales. We assessed the genetic diversity and population structure of H. sericeus in the Pacific Ocean using three gene markers cytochrome oxidase subunit I (COI), elongation factor 1a and internal transcribed spacer 1 (ITS-1). These markers indicate that both populations are evolutionarily distinct with limited gene flow, having separated 20 00050 000 years ago. This suggests that physical conditions and/or biotic interactions on the surface of the Pacific Ocean have provided significant barriers to gene flow since the late Pleistocene or earlier, creating biotic stability over large geographical and temporal scales in spite of a long history of global climate change. (c) 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 797805.

Cheng, LN, Damgaard J, Garrouste R.  2012.  The sea-skater Halobates (Heteroptera: Gerridae) - probable cause for extinction in the Mediterranean and potential for re-colonisation following climate change. Aquatic Insects. 34:45-55.   10.1080/01650424.2012.643024   AbstractWebsite

Sea-skaters in the genus Halobates Eschscholtz 1822 include some of the most specialised water striders and are found in tropical and subtropical seas around the world. Even though species of Halobates occur in both the Atlantic Ocean and the Red Sea, no extant sea-skater has been reported from the Mediterranean Sea. A fossil, Halobates ruffoi Andersen et al., 1994, described from Middle-Upper Eocene (45 Ma) Italy indicates that sea skaters were present in this part of the world in the past. Other geological evidence points to dramatic changes in the Mediterranean Sea during the Tertiary and Quaternary that may have led to their later extinction. In this paper we review briefly the distribution, systematics, evolution and ecology of Halobates, and discuss the potential for the Mediterranean to be recolonised following expected environmental changes due to global warming.

Ikawa, T, Okabe H, Cheng LN.  2012.  Skaters of the seas - comparative ecology of nearshore and pelagic Halobates species (Hemiptera: Gerridae), with special reference to Japanese species. Marine Biology Research. 8:915-936.   10.1080/17451000.2012.705848   AbstractWebsite

The insects overwhelm all other organisms of the world in species numbers and diversity. However, there are relatively few insects that inhabit the sea. Most of them are confined to the intertidal zone, with only five species of sea skaters, genus Halobates, having been successful in colonizing the open ocean. We discuss the ecology of both coastal and pelagic Halobates and the closely related genus Asclepios, focusing on their distribution ranges and adaptive strategies to marine environments accompanied by brief discussions of their biology. Updated information on the known localities for three species of Asclepios and some 40 coastal species of Halobates are presented. Many species live in vulnerable coastal habitats exposed to environmental pollution and coastal development. As a case study we trace the historical changes in populations of three Japanese sea skaters, Asclepios shiranui, Halobates matsumurai and H. japonicus, all designated as endangered species. For oceanic Halobates, we present an updated distribution map along with global current systems and sea-surface temperatures, examine interactions between distribution ranges and physical factors at the air-sea interface and discuss spatio-temporal variations in populations of each species. Finally, we infer the life history strategy of oceanic Halobates through theoretical considerations.

Huang, DW, Cheng LN.  2011.  The flightless marine midge Pontomyia (Diptera: Chironomidae): ecology, distribution, and molecular phylogeny. Zoological Journal of the Linnean Society. 162:443-456.   10.1111/j.1096-3642.2010.00680.x   AbstractWebsite

Pontomyia Edwards, 1926 (Diptera: Chironomidae) is a genus of exclusively marine flightless midges. There are four described species from the Indo- Pacific, and one undescribed species known only from females, pupal skins, and larvae from the Atlantic/Caribbean. They are poorly known owing to their small size (similar to 1.0 mm), extremely short adult life (< 3 h), and unusual habitat for an insect (coastal lagoons, bays, or rock pools). We reviewed scattered literature on their biology and systematics, presented photomicrographs of the male hypogium, and updated the geographic distribution of each species. We carried out the first molecular study to elucidate relationships among and within three of the species. Results from our four- gene phylogenetic reconstruction using combined gene tree and species tree approaches showed that Pontomyia natans, Pontomyia oceana, and Pontomyia pacifica are each well- supported clades, with P. natans as sister to P. oceana + P. pacifica. Genetic distances based on mitochondrial cytochrome oxidase I are extraordinarily large within P. natans and P. pacifica, which suggests that they may be cryptic species complexes. (C) 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 443- 456.

Cheng, L, Baars MA, Smith A.  2011.  Life on the High Seas: The Bug Darwin Never Saw. Antenna. 35:36-42. Abstract
Cheng, L, Spear LB, Ainley DG.  2010.  Importance of marine insects (Heteroptera: Gerridae, Halobates spp.) as prey of eastern tropical Pacific seabirds. Marine Ornithology. 38:91-95. AbstractWebsite

We analyzed the foraging ecology of seabirds in the eastern tropical Pacific Ocean during 1983–1991 on a series of oceanographic cruises during spring and fall of each year. We report details about the consumption of sea skaters Halobates spp., marine insects that are small, can hide well within sea foam, and can be very fast moving. One abundant sea skater of the four species present in the study area, H. sobrinus, is not taken by sea birds, and the reason is unknown. Among the predators, it appears that frigate storm-petrels, White-faced Storm-Petrel Pelagodroma marina and White-bellied Storm-Petrel Fregetta grallaria (likely also White-throated Storm-Petrel Nesofregetta fuliginosa), make directed efforts to consume sea skaters, a fact that may explain their unique flight and foraging behavior: slow, with extensive “kick splashing” against the sea surface, to incite movement in Halobates. The few other seabirds for which sea skaters constitute more than an incidental component of the diet (Herald Petrel Pterodroma heraldica, Bulwer’s Petrel Bulweria bulwerii) also move slowly across and close to the sea surface. In the case of the White Tern Gygis alba, it readily hovers, an ability that would be advantageous to taking these insects. Among the avian species that rarely take a sea skater, almost all are included in the guild of seabirds that associate with tuna, and as a result they must fly quickly to keep pace.

Garrouste, R, Cheng L.  2009.  Les punaises marines Halobates. Clipperton, Environement et Biodiversitè d'un Microcosme Océanique.. 68( Charpy L, Ed.).:119-126., Paris, IRD Marseilles: MHNH Abstract
Satoh, A, Cheng L.  2009.  A new record of Pontomyia pacifica (Diptera, Chironomidae) from Okinawa, Japan. Biological Magazine Okinawa. 47:47-48. Abstract
Cheng, LN.  2008.  Nils and Halobates (Heteroptera: Gerridae). Insect Systematics & Evolution. 39:475-479.   10.1163/187631208794760876   AbstractWebsite
Ikawa, T, Onodera S, Okabe H, Hoshizaki S, Cheng LN.  2007.  Occurrence and density of Halobates micans (Hemiptera : Gerridae) in the eastern South Indian Ocean. Entomological Science. 10:213-215.   10.1111/j.1479-8298.2007.00215.x   AbstractWebsite

Two species of ocean skaters, Halobates germanus and Halobates micans, live in the tropical and subtropical waters of the Indian Ocean. From December 1992 to December 1993, Halobates was intensively sampled in the easternmost region of the South Indian Ocean (13 - 18.5 degrees S, 114 - 121 degrees E), from which there have been a small number of records of Halobates. No H. germanus was caught, but a total of 1190 H. micans were collected, with densities estimated at 13 900 - 28 100 individuals/km(2). This suggests that H. micans lives in the study area at high densities comparable to those in the Atlantic and the Pacific Oceans. We also discuss the possible effects of ocean currents and winds on the geographic distributions of the two Halobates species in the eastern South Indian Ocean.

Cheng, LN, Yang CM, Li DQ, Liu HM.  2006.  Aquatic heteroptera (Insecta : Gerromorpha and Nepomorpha) from Xishuangbanna, Yunnan, China. Raffles Bulletin of Zoology. 54:203-214. AbstractWebsite

Eighty five species of aquatic and semi-aquatic Heteroptera belonging to 14 families are reported from Xishuangbanna and nearby counties in Yunnan. Collecting locations and habitats are given. Ten species (Hydrometra jaczewskii, H. longicapitis, H. orientalis, Amemboa (Amemboa) sexualis, A. (A.) lyra, Metrocoris ciliatus, Pleciobates pacholatkoi, Ptilomera burmana, Rhyacobates anderseni, Ventidus pulai) are new records for China. In addition, Nine new species (Timasius yunanensis, Ranatra lansburyi, R. sterea, Micronecta (Micronecta) erythra, M. (M.)janssoni, M. (M.) lobata, M (M.) ornitheia, M. (Unguinecta) melanochroa, Anisops pseudostali) based on material collected during this study have been described and published elsewhere.

Cheng, L.  2006.  A bug on the ocean waves (Heteroptera, Gerridae, Halobates Eschscholtz). Hug the bug – For love of true bugs. Festschrift zum 70. Geburtstag von Ernst Heiss.. 19( Rabitsch W, Ed.).:1033-1040.: Denisia Abstract
Ikawa, T, Okabe H, Hoshizaki S, Kamikado T, Cheng L.  2004.  Distribution of the oceanic insects Halobates (Hemiptera : Gerridae) off the south coast of Japan. Entomological Science. 7:351-357.   10.1111/j.1479-8298.2004.00083.x   AbstractWebsite

Specimens of ocean skaters Halobates were collected off the south coast of Japan in the East China Sea in 1995, and from the Kumano-nada Sea to the East China Sea in 1998 and 1999. Three species were identified: H. micans, H. germanus and H. sericeus. We found two species co-occurring in comparable densities in different years, a phenomenon not :hitherto reported in other regions of the ocean. We discuss distributions of the three Halobates species with special reference to the influence of the Kuroshio Current, temporal variations of sea-surface temperature, and monsoonal winds.

Cheng, L.  2004.  The Fightless Marine Midge, Pontomyia (Diptera : Chironomidae) with a First Record from the Republic of the Maldives, Indian Ocean. Contemporary Trends in Insect Science. ( Gujar GT, Ed.).:52-59., New Delhi, India: Campus Books International Abstract
Yang, C, Kovac D, Cheng L.  2004.  Insecta: Hemiptera, Heteroptera. Freshwater invertebrates of the Malaysian region. ( Yule CM, Yong H, Eds.).:457-490., Kuala Lumpur, Malaysia: Akademi Sains Malaysia Abstract
Andersen, NM, Cheng L.  2004.  The marine insect Halobates (Heteroptera : Gerridae): Biology, adaptations, distribution, and phylogeny. Oceanography and Marine Biology: An Annual Review, Vol 42. 42( Gibson RN, Atkinson RJA, Gordon JDM, Eds.).:119-179., Boca Raton: Crc Press-Taylor & Francis Group Abstract

Among the million or so insect species known, only a few thousand are found in marine habitats. The genus Halobates is almost exclusively marine and is unique in having the only known species to live in the open ocean. Of the 46 Halobates species described, only five are completely oceanic in habitat, with the majority of species living in coastal areas associated with mangroves or other marine plants. This review presents a brief historical account of the genus and provides information on various aspects of its life history, ecology, special adaptations, distribution, and biogeography. Distribution maps of the five oceanic species as well as several of the more widely distributed coastal species have been updated. The phylogeny and evolution of Halobates based on morphology and recent molecular data are also discussed. A key to all known species of Halobates and related genera and a checklist of all species and their distributions are included as appendices.

Cheng, L.  2004.  Marine insects and the sea-skater Halobates (Hemiptera: Gerridae). Encyclopedia of entomology. 2( Capinera JL, Ed.).:1346-1349., Dordrecht ; Boston: Kluwer Academic Abstract
Petrakis, PV, Tsoukatou M, Vagias C, Roussis V, Cheng LN.  2003.  Evolution probing for semiochemicals based on secondary metabolites in the cuticles of three species of Halobates (Heteroptera : Gerridae). Biological Journal of the Linnean Society. 80:671-688.   10.1111/j.1095-8312.2003.00267.x   AbstractWebsite

We developed methods for characterizing semiochemicals in the cuticles of the sea-skater Halobates (Gerridae). Such substances may be involved in chemical communication at the sea surface and have been shown to act as sexual pheromones for a related marine insect Trochopus plumbeus (Veliidae). We identified 86 volatile compounds in extracts from three Halobates species (H. micans Eschscholtz, H. hawaiiensis Usinger and H. sobrinus White) and used them to reconstruct the phylogeny of the clade. Sixty-seven of the compounds were found to be parsimony-informative. Suspected semiochemicals appear to be specific autapomorphies in the phylogram, and our results indicated that chemical profiles may be used to predict the phylogenetic history of Halobates. Moreover, the phylogenetic tree derived from our data is congruent with a previously published one based on morphology and mitochondrial DNA sequence (780 bp in the COI gene). The potential role of each compound to be used for chemical communication was also reviewed. At appropriate concentrations, some of the cuticular compounds, being partly hydrophobic and partly hydrophilic, could disperse on the sea surface and thereby play an energetically efficient role in chemical communication, e.g. mate location. Less dispersible components might act as 'sun-screens' and provide protection from UV-irradiation during daylight hours. (C) 2003 The Linnean Society of London.