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  1. von Seth J, Dussex N, Díez-Del-Molino D, van der Valk T, Kutschera VE, Kierczak M, et al.
    Nat Commun, 2021 Apr 26;12(1):2393.
    PMID: 33896938 DOI: 10.1038/s41467-021-22386-8
    Small populations are often exposed to high inbreeding and mutational load that can increase the risk of extinction. The Sumatran rhinoceros was widespread in Southeast Asia, but is now restricted to small and isolated populations on Sumatra and Borneo, and most likely extinct on the Malay Peninsula. Here, we analyse 5 historical and 16 modern genomes from these populations to investigate the genomic consequences of the recent decline, such as increased inbreeding and mutational load. We find that the Malay Peninsula population experienced increased inbreeding shortly before extirpation, which possibly was accompanied by purging. The populations on Sumatra and Borneo instead show low inbreeding, but high mutational load. The currently small population sizes may thus in the near future lead to inbreeding depression. Moreover, we find little evidence for differences in local adaptation among populations, suggesting that future inbreeding depression could potentially be mitigated by assisted gene flow among populations.
    Matched MeSH terms: Borneo
  2. van der Meij SE
    Zookeys, 2015.
    PMID: 25987871 DOI: 10.3897/zookeys.500.9244
    A new species of gall crab is described from the free-living stony coral Trachyphylliageoffroyi. Specimens were collected during field work in Lembeh Strait (Indonesia) and off Kudat (Malaysian Borneo). This new species, here named Lithoscaptussemperi sp. n., is the ninth species assigned to the genus. It can be separated from its congeners by not having the internal orbital angle extending beyond the external orbital angle, and by the stout female P2 merus with prominent distomesial projection. In addition, the carapace surface appears smooth, despite having small tubercles on the anterior half, and is without noticeable spines, other than those on the frontal margin. The distinctive carapace pattern in life is a diagnostic character in male specimens.
    Matched MeSH terms: Borneo
  3. van der Ent A, Nkrumah PN, Aarts MGM, Baker AJM, Degryse F, Wawryk C, et al.
    BMC Plant Biol, 2021 Sep 27;21(1):437.
    PMID: 34579652 DOI: 10.1186/s12870-021-03190-4
    BACKGROUND: Some subspecies of Dichapetalum gelonioides are the only tropical woody zinc (Zn)-hyperaccumulator plants described so far and the first Zn hyperaccumulators identified to occur exclusively on non-Zn enriched 'normal' soils. The aim of this study was to investigate Zn cycling in the parent rock-soil-plant interface in the native habitats of hyperaccumulating Dichapetalum gelonioides subspecies (subsp. pilosum and subsp. sumatranum). We measured the Zn isotope ratios (δ66Zn) of Dichapetalum plant material, and associated soil and parent rock materials collected from Sabah (Malaysian Borneo).

    RESULTS: We found enrichment in heavy Zn isotopes in the topsoil (δ66Zn 0.13 ‰) relative to deep soil (δ66Zn -0.15 ‰) and bedrock (δ66Zn -0.90 ‰). This finding suggests that both weathering and organic matter influenced the Zn isotope pattern in the soil-plant system, with leaf litter cycling contributing significantly to enriched heavier Zn in topsoil. Within the plant, the roots were enriched in heavy Zn isotopes (δ66Zn ~ 0.60 ‰) compared to mature leaves (δ66Zn ~ 0.30 ‰), which suggests highly expressed membrane transporters in these Dichapetalum subspecies preferentially transporting lighter Zn isotopes during root-to-shoot translocation. The shoots, mature leaves and phloem tissues were enriched in heavy Zn isotopes (δ66Zn 0.34-0.70 ‰) relative to young leaves (δ66Zn 0.25 ‰). Thisindicates that phloem sources are enriched in heavy Zn isotopes relative to phloem sinks, likely because of apoplastic retention and compartmentalization in the Dichapetalum subspecies.

    CONCLUSIONS: The findings of this study reveal Zn cycling in the rock-soil-plant continuum within the natural habitat of Zn hyperaccumulating subspecies of Dichapetalum gelonioides from Malaysian Borneo. This study broadens our understanding of the role of a tropical woody Zn hyperaccumulator plant in local Zn cycling, and highlights the important role of leaf litter recycling in the topsoil Zn budget. Within the plant, phloem plays key role in Zn accumulation and redistribution during growth and development. This study provides an improved understanding of the fate and behaviour of Zn in hyperaccumulator soil-plant systems, and these insights may be applied in the biofortification of crops with Zn.

    Matched MeSH terms: Borneo
  4. van der Ent A, Edraki M
    Environ Geochem Health, 2018 Feb;40(1):189-207.
    PMID: 27848090 DOI: 10.1007/s10653-016-9892-3
    The Mamut Copper Mine (MCM) located in Sabah (Malaysia) on Borneo Island was the only Cu-Au mine that operated in the country. During its operation (1975-1999), the mine produced 2.47 Mt of concentrate containing approximately 600,000 t of Cu, 45 t of Au and 294 t of Ag, and generated about 250 Mt of overburden and waste rocks and over 150 Mt of tailings, which were deposited at the 397 ha Lohan tailings storage facility, 15.8 km from the mine and 980 m lower in altitude. The MCM site presents challenges for environmental rehabilitation due to the presence of large volumes of sulphidic minerals wastes, the very high rainfall and the large volume of polluted mine pit water. This indicates that rehabilitation and treatment is costly, as for example, exceedingly large quantities of lime are needed for neutralisation of the acidic mine pit discharge. The MCM site has several unusual geochemical features on account of the concomitant occurrence of acid-forming sulphide porphyry rocks and alkaline serpentinite minerals, and unique biological features because of the very high plant diversity in its immediate surroundings. The site hence provides a valuable opportunity for researching natural acid neutralisation processes and mine rehabilitation in tropical areas. Today, the MCM site is surrounded by protected nature reserves (Kinabalu Park, a World Heritage Site, and Bukit Hampuan, a Class I Forest Reserve), and the environmental legacy prevents de-gazetting and inclusion in these protected area in the foreseeable future. This article presents a preliminary geochemical investigation of waste rocks, sediments, secondary precipitates, surface water chemistry and foliar elemental uptake in ferns, and discusses these results in light of their environmental significance for rehabilitation.
    Matched MeSH terms: Borneo
  5. van der Ent A, Mesjasz-Przybyłowicz J, Przybyłowicz WJ, Barnabas AD, de Jonge MD, Harris HH
    Metallomics, 2022 May 13;14(5).
    PMID: 35556136 DOI: 10.1093/mtomcs/mfac020
    Globally, the majority of Ni hyperaccumulator plants occur on ultramafic soils in tropical regions, and the genus Phyllanthus, from the Phyllanthaceae family, is globally the most represented taxonomical group. Two species from Sabah (Malaysia) are remarkable because Phyllanthus balgooyi can attain >16 wt% of Ni in its phloem exudate, while Phyllanthus rufuschaneyi reaches foliar concentrations of up to 3.5 wt% Ni, which are amongst the most extreme concentrations of Ni in any plant tissue. Synchrotron X-ray fluorescence microscopy, nuclear microbe (micro-PIXE+BS) and (cryo) scanning electron microscopy with energy dispersive spectroscopy were used to spatially resolve the elemental distribution in the plant organs of P. balgooyi and P. rufuschaneyi. The results show that P. balgooyi has extraordinary enrichment of Ni in the (secondary) veins of the leaves, whereas in contrast, in P. rufuschaneyi Ni occurs in interveinal areas. In the roots and stems, Ni is localized mainly in the cortex and phloem but is much lower in the xylem. The findings of this study show that, even within the same genus, the distribution of nickel and other elements, and inferred processes involved with metal hyperaccumulation, can differ substantially between species.
    Matched MeSH terms: Borneo
  6. de Silva JR, Amir A, Lau YL, Ooi CH, Fong MY
    PLoS One, 2019;14(9):e0222681.
    PMID: 31536563 DOI: 10.1371/journal.pone.0222681
    The Duffy blood group plays a key role in Plasmodium knowlesi and Plasmodium vivax invasion into human erythrocytes. The geographical distribution of the Duffy alleles differs between regions with the FY*A allele having high frequencies in many Asian populations, the FY*B allele is found predominately in European populations and the FY*Bes allele found predominantly in African regions. A previous study in Peninsular Malaysia indicated high homogeneity of the dominant FY*A/FY*A genotype. However, the distribution of the Duffy genotypes in Malaysian Borneo is currently unknown. In the present study, the distribution of Duffy blood group genotypes and allelic frequencies among P. knowlesi infected patients as well as healthy individuals in Malaysian Borneo were determined. A total of 79 P. knowlesi patient blood samples and 76 healthy donor samples were genotyped using allele specific polymerase chain reaction (ASP-PCR). Subsequently a P. knowlesi invasion assay was carried out on FY*AB/ FY*A and FY*A/ FY*A Duffy genotype blood to investigate if either genotype conferred increased susceptibility to P. knowlesi invasion. Our results show almost equal distribution between the homozygous FY*A/FY*A and heterozygous FY*A/FY*B genotypes. This is in stark contrast to the Duffy distribution in Peninsular Malaysia and the surrounding Southeast Asian region which is dominantly FY*A/FY*A. The mean percent invasion of FY*A/FY*A and FY*A/FY*B blood was not significantly different indicating that neither blood group confers increased susceptibility to P. knowlesi invasion.
    Matched MeSH terms: Borneo
  7. Zhao TT, Han HL
    Zookeys, 2020;985:127-141.
    PMID: 33223877 DOI: 10.3897/zookeys.985.54047
    In this paper, four species of the genus Diduga Moore, [1887] from China (Chongqing and Guangdong) and Malaysia (Borneo, Sabah) are described as new to science, namely D. simianshanasp. nov., D. chebalingasp. nov., D. chewisp. nov., and D. hollowayisp. nov. Adults of these species are illustrated in color, and images of the male and female genitalia are provided. A distribution map of the new species is provided, together with an updated checklist of all species of Diduga.
    Matched MeSH terms: Borneo
  8. Zhang JX, Maddison WP
    Zootaxa, 2014;3852(2):252-72.
    PMID: 25284396 DOI: 10.11646/zootaxa.3852.2.5
    Six new species of marpissoid jumping spiders from Sarawak, Borneo, are described in the new genus Tisaniba Zhang & Maddison. They are the type species, T. mulu Zhang & Maddison sp. nov., as well as the species T. bijibijan Zhang & Maddison sp. nov., T. dik Zhang & Maddison sp. nov., T. kubah Zhang & Maddison sp. nov., T. selan Zhang & Maddison sp. nov., and T. selasi Zhang & Maddison sp. nov. The spiders are small and brown to black, living in leaf litter in the tropical forest. Phylogenetic analyses based on 28s and 16sND1 genes indicate that they are a distinctive group within the marpissoids. Diagnostic illustrations and photographs of living spiders are provided for all species.
    Matched MeSH terms: Borneo
  9. Zhang C, Martens J
    Zookeys, 2018.
    PMID: 30271242 DOI: 10.3897/zookeys.785.26389
    Lobonychiumpalpiplus Roewer, 1938, originally reported from Indonesian Borneo, is redescribed based on the specimens from Malaysia. The genitalia of this species are described for the first time and a new genital terminology is proposed. The rediscovery expands the known distribution of the species to Malaysian Borneo.
    Matched MeSH terms: Borneo
  10. Zahidin MA, Jalil NA, Naharuddin NM, Abd Rahman MR, Gani M, Abdullah MT
    Data Brief, 2019 Aug;25:104133.
    PMID: 31321260 DOI: 10.1016/j.dib.2019.104133
    Tarsier is an endangered nocturnal primate in the family Tarsiidae and is an endemic to Sundaic islands of Philippine (Carlito syrichta), Sulawesi (Tarsius tarsier-complex) and Borneo (Cephalopachus bancanus). Recent records indicated that most molecular studies were done on the Eastern Tarsier and little information for the other group of tarsiers. Here, we present a partial cytochrome b data set of C. bancanus in Sarawak, Malaysian Borneo. Standard mist nets were deployed at strategic locations in various habitat types. A total of 18 individuals were caught, measured and weighed. Approximately, 2 × 2 mm of tissue samples were taken and preserved in molecular grade alcohol. Out of 18, only 11 samples were screened with partial mtDNA (cytochrome b) and the DNA sequences were registered in the GenBank (accession numbers: KY794797-KY794807). Phylogenetic trees were constructed with 20 additional mtDNA sequences downloaded from GenBank. The data are valuable for the management authorities to regulate the type of management units for the metapopulation to sustain population genetics integrity of tarsiers in the range countries across the Sunda Shelf.
    Matched MeSH terms: Borneo
  11. Zacaery Khalik M, P Hendriks K, J Vermeulen J, Menno Schilthuizen
    Zookeys, 2019;840:35-86.
    PMID: 31065227 DOI: 10.3897/zookeys.840.33326
    The Bornean representatives of the genus Georissa (Hydrocenidae) have small, dextral, conical, calcareous shells consisting of ca. three teleoconch whorls. Our recent study on the Georissa of Malaysian Borneo has revealed high intra- and inter-specific variation in the "scaly" group (a group of species with striking scale-like surface sculpture). The present study on the "non-scaly" Georissa is the continuation of the species revision for the genus. The "non-scaly" species are also diverse in shell sculptures. This informal group comprises Georissa with subtle spiral and/or radial sculpture. The combination of detailed conchological assessment and molecular analyses provides clear distinctions for each of the species. Conchological, molecular, and biogeographic details are presented for 16 species of "non-scaly" Georissa. Three of these are new to science, namely Georissacorrugatasp. n., Georissainsulaesp. n., and Georissatrusmadisp. n.
    Matched MeSH terms: Borneo
  12. Yusoff MM, Ibrahim H, Hamid NA
    Chem Biodivers, 2011 May;8(5):916-23.
    PMID: 21560240 DOI: 10.1002/cbdv.201000270
    Two poorly studied, morphologically allied Alpinia species endemic to Borneo, viz., A. ligulata and A. nieuwenhuizii, were investigated here for their rhizome essential oil. The oil compositions and antimicrobial activities were compared with those of A. galanga, a better known plant. A fair number of compounds were identified in the oils by GC-FID and GC/MS analyses, with large differences in the oil composition between the three species. The rhizome oil of A. galanga was rich in 1,8-cineole (29.8%), while those of A. ligulata and A. nieuwenhuizii were both found to be extremely rich in (E)-methyl cinnamate (36.4 and 67.8%, resp.). The three oils were screened for their antimicrobial activity against three Gram-positive and three Gram-negative bacteria and two fungal species. The efficiency of growth inhibition of Staphylococcus aureus var. aureus was found to decline in the order of A. nieuwenhuizii>A. ligulata ∼ A. galanga, while that of Escherichia coli decreased in the order of A. galanga>A. nieuwenhuzii ∼ A. ligulata. Only the A. galanga oil inhibited the other bacteria and the fungi tested.
    Matched MeSH terms: Borneo
  13. Yusah KM, Foster WA, Reynolds G, Fayle TM
    PeerJ, 2018;6:e4231.
    PMID: 29423344 DOI: 10.7717/peerj.4231
    Background: Competitive interactions in biological communities can be thought of as giving rise to "assembly rules" that dictate the species that are able to co-exist. Ant communities in tropical canopies often display a particular pattern, an "ant mosaic", in which competition between dominant ant species results in a patchwork of mutually exclusive territories. Although ant mosaics have been well-documented in plantation landscapes, their presence in pristine tropical forests remained contentious until recently. Here we assess presence of ant mosaics in a hitherto under-investigated forest stratum, the emergent trees of the high canopy in primary tropical rain forest, and explore how the strength of any ant mosaics is affected by spatial scale, time of day, and sampling method.

    Methods: To test whether these factors might impact the detection of ant mosaics in pristine habitats, we sampled ant communities from emergent trees, which rise above the highest canopy layers in lowland dipterocarp rain forests in North Borneo (38.8-60.2 m), using both baiting and insecticide fogging. Critically, we restricted sampling to only the canopy of each focal tree. For baiting, we carried out sampling during both the day and the night. We used null models of species co-occurrence to assess patterns of segregation at within-tree and between-tree scales.

    Results: The numerically dominant ant species on the emergent trees sampled formed a diverse community, with differences in the identity of dominant species between times of day and sampling methods. Between trees, we found patterns of ant species segregation consistent with the existence of ant mosaics using both methods. Within trees, fogged ants were segregated, while baited ants were segregated only at night.

    Discussion: We conclude that ant mosaics are present within the emergent trees of the high canopy of tropical rain forest in Malaysian Borneo, and that sampling technique, spatial scale, and time of day interact to determine observed patterns of segregation. Restricting sampling to only emergent trees reveals segregatory patterns not observed in ground-based studies, confirming previous observations of stronger segregation with increasing height in the canopy.

    Matched MeSH terms: Borneo
  14. Yunos NE, Sharkawi HM, Hii KC, Hu TH, Mohamad DSA, Rosli N, et al.
    Sci Rep, 2022 Oct 14;12(1):17284.
    PMID: 36241678 DOI: 10.1038/s41598-022-21439-2
    Plasmodium knowlesi infections in Malaysia are a new threat to public health and to the national efforts on malaria elimination. In the Kapit division of Sarawak, Malaysian Borneo, two divergent P. knowlesi subpopulations (termed Cluster 1 and Cluster 2) infect humans and are associated with long-tailed macaque and pig-tailed macaque hosts, respectively. It has been suggested that forest-associated activities and environmental modifications trigger the increasing number of knowlesi malaria cases. Since there is a steady increase of P. knowlesi infections over the past decades in Sarawak, particularly in the Kapit division, we aimed to identify hotspots of knowlesi malaria cases and their association with forest activities at a geographical scale using the Geographic Information System (GIS) tool. A total of 1064 P. knowlesi infections from 2014 to 2019 in the Kapit and Song districts of the Kapit division were studied. Overall demographic data showed that males and those aged between 18 and 64 years old were the most frequently infected (64%), and 35% of infections involved farming activities. Thirty-nine percent of Cluster 1 infections were mainly related to farming surrounding residential areas while 40% of Cluster 2 infections were associated with activities in the deep forest. Average Nearest Neighbour (ANN) analysis showed that humans infected with both P. knowlesi subpopulations exhibited a clustering distribution pattern of infection. The Kernel Density Analysis (KDA) indicated that the hotspot of infections surrounding Kapit and Song towns were classified as high-risk areas for zoonotic malaria transmission. This study provides useful information for staff of the Sarawak State Vector-Borne Disease Control Programme in their efforts to control and prevent zoonotic malaria.
    Matched MeSH terms: Borneo
  15. Yule CM, Lim YY, Lim TY
    Carbon Balance Manag, 2018 Feb 07;13(1):3.
    PMID: 29417248 DOI: 10.1186/s13021-018-0092-6
    BACKGROUND: Tropical peat swamp forests (TPSF) are globally significant carbon stores, sequestering carbon mainly as phenolic polymers and phenolic compounds (particularly as lignin and its derivatives) in peat layers, in plants, and in the acidic blackwaters. Previous studies show that TPSF plants have particularly high levels of phenolic compounds which inhibit the decomposition of organic matter and thus promote peat accumulation. The studies of phenolic compounds are thus crucial to further understand how TPSF function with respect to carbon sequestration. Here we present a study of cycling of phenolic compounds in five forests in Borneo differing in flooding and acidity, leaching of phenolic compounds from senescent Macaranga pruinosa leaves, and absorption of phenolics by M. pruinosa seedlings.

    RESULTS: The results of the study show that total phenolic content (TPC) in soil and leaves of three species of Macaranga were highest in TPSF followed by freshwater swamp forest and flooded limestone forest, then dry land sites. Highest TPC values were associated with acidity (in TPSF) and waterlogging (in flooded forests). Moreover, phenolic compounds are rapidly leached from fallen senescent leaves, and could be reabsorbed by tree roots and converted into more complex phenolics within the leaves.

    CONCLUSIONS: Extreme conditions-waterlogging and acidity-may facilitate uptake and synthesis of protective phenolic compounds which are essential for impeded decomposition of organic matter in TPSF. Conversely, the ongoing drainage and degradation of TPSF, particularly for conversion to oil palm plantations, reverses the conditions necessary for peat accretion and carbon sequestration.

    Matched MeSH terms: Borneo
  16. Yu S, Park KT, Wang S
    Zootaxa, 2019 Jun 18;4619(1):zootaxa.4619.1.7.
    PMID: 31716319 DOI: 10.11646/zootaxa.4619.1.7
    Seven new species of the genus Deltoplastis Meyrick are described: D. acutangulata Wang et Yu, sp. nov., D. anatoliana Wang et Park, sp. nov., D. multidentalis Wang et Yu, sp. nov. and D. similihoristis Wang et Yu, sp. nov. from China; D. aculeata Wang et Yu, sp. nov. and D. spatuliunca Wang et Yu, sp. nov. from Malaysian Borneo; and D. ovidiscalis Park et Wang, sp. nov. from Vietnam. Deltoplastis horistis (Meyrick, 1910) is newly recorded in China and its female is described for the first time. Images of adults and genitalia of the new species are provided.
    Matched MeSH terms: Borneo
  17. Young KI, Buenemann M, Vasilakis N, Perera D, Hanley KA
    Ecosphere, 2021 Apr;12(4).
    PMID: 33996190 DOI: 10.1002/ecs2.3463
    Deforestation precipitates spillover of enzootic, vector-borne viruses into humans, but specific mechanisms for this effect have rarely been investigated. Expansion of oil palm cultivation is a major driver of deforestation. Here, we demonstrate that mosquito abundance decreased over ten stepwise distances from interior forest into conterminous palm plantations in Borneo. Diversity in interior plantation narrowed to one species, Aedes albopictus, a potential bridge vector for spillover of multiple viruses. A. albopictus was equally abundant across all distances in forests, forest-plantation edge, and plantations, while A. niveus, a known vector of sylvatic dengue virus, was found only in forests. A. albopictus collections were significantly female-biased in plantation but not in edge or forest. Our data reveal that the likelihood of encountering any mosquito is greater in interior forest and edge than plantation, while the likelihood of encountering A. albopictus is equivalent across the gradient sampled from interior plantation to interior forest.
    Matched MeSH terms: Borneo
  18. Young KI, Mundis S, Widen SG, Wood TG, Tesh RB, Cardosa J, et al.
    Parasit Vectors, 2017 Aug 31;10(1):406.
    PMID: 28859676 DOI: 10.1186/s13071-017-2341-z
    BACKGROUND: Mosquito-borne dengue virus (DENV) is maintained in a sylvatic, enzootic cycle of transmission between canopy-dwelling non-human primates and Aedes mosquitoes in Borneo. Sylvatic DENV can spill over into humans living in proximity to forest foci of transmission, in some cases resulting in severe dengue disease. The most likely vectors of such spillover (bridge vectors) in Borneo are Ae. albopictus and Ae. niveus. Borneo is currently experiencing extensive forest clearance. To gauge the effect of this change in forest cover on the likelihood of sylvatic DENV spillover, it is first necessary to characterize the distribution of bridge vectors in different land cover types. In the current study, we hypothesized that Ae. niveus and Ae. albopictus would show significantly different distributions in different land cover types; specifically, we predicted that Ae. niveus would be most abundant in forests whereas Ae. albopictus would have a more even distribution in the landscape.

    RESULTS: Mosquitoes were collected from a total of 15 sites using gravid traps and a backpack aspirator around Kampong Puruh Karu, Sarawak, Malaysian Borneo, where sylvatic DENV spillover has been documented. A total of 2447 mosquitoes comprising 10 genera and 4 species of Aedes, were collected over the three years, 2013, 2014 and 2016, in the three major land cover types in the area, homestead, agriculture and forest. Mosquitoes were identified morphologically, pooled by species and gender, homogenized, and subject to DNA barcoding of each Aedes species and to arbovirus screening. As predicted, Ae. niveus was found almost exclusively in forests whereas Ae. albopictus was collected in all land cover types. Aedes albopictus was significantly (P = 0.04) more abundant in agricultural fields than forests. Sylvatic DENV was not detected in any Aedes mosquito pools, however genomes of 14 viruses were detected using next generation sequencing.

    CONCLUSIONS: Land cover type affects the abundance and distribution of the most likely bridge vectors of sylvatic DENV in Malaysia Borneo. Conversion of forests to agriculture will likely decrease the range and abundance of Ae. niveus but enhance the abundance of Ae. albopictus.

    Matched MeSH terms: Borneo/epidemiology
  19. Yoshida T, Hirowatari T
    Zookeys, 2018.
    PMID: 30584392 DOI: 10.3897/zookeys.805.28757
    A new silvanid genus Borneophanusgen. n. is described based on specimens collected from Malaysian Borneo. A new species, B.spinosussp. n., is described herein. Digitiform sensilla on the apical maxillary palpomere is reported in Silvanidae for the first time.
    Matched MeSH terms: Borneo
  20. Yonge CM
    Biol. Bull., 1968 Oct;135(2):378-405.
    PMID: 28368764 DOI: 10.2307/1539789
    1. Study of the structure and mode of life of Malleus regula provides the basis for consideration of the "hammer" species, M. malleus and M. albus.
    2. M. regula occurs byssally attached, vertically disposed on rocky substrates associated with mud and is widely distributed in the tropical Indo-Pacific.
    3. The distal two-thirds of the elongated shell is exclusively prismatic. By means of special pallial retractors the mantle lobes can be withdrawn within the nacreous region.
    4. The massive opisthodetic ligament has a short secondary extension of fusion layer.
    5. A promyal chamber on the right side proximal to the adductor increases water flow into the exhalant chamber.
    6. The long filibranch ctenidia provide a vertically extended food-collecting surface.
    7. The foot is concerned with planting of the massive byssus which emerges through a notch in the right valve but on the under ( i.e., dorsal) surface. There is also a unique and very long accessory foot, ventrally grooved, everywhere ciliated and in constant writhing activity due to blood pressure and intrinsic muscle. Moving freely throughout the lower mantle cavity it can only be concerned with cleansing.
    8. Pseudofaeces are ejected from the distal tip of the mantle cavity.
    9. M. malleus, the black hammer shell, occurs vertically embedded in coarse sand or sandy gravel. Byssus threads are attached to fragments within the substrate. The great anterior and posterior elongations of the hinge line (also exclusively prismatic) are separated by the byssal notch (now affecting both valves).
    10. Shells are usually excessively irregular due to the great exposure to damage and the almost unlimited powers of rapid repair by the three pallial extensions.
    11. M. albus, the white hammer shell, is stouter and inhabits muddy sand. During growth it loses the byssus with reduction of the foot and byssal retractors and closure of the byssal notch. The accessory foot is not affected. The animal becomes anchored in the substrate exclusively by the anterior and posterior extensions of the shell.
    12. Description of the typically rounded Isognomon ephippium leads to that of the elongated I. isognomon which occupies precisely the same habitat as M. regula.
    13. It lacks the pallial retractors, promyal chamber and accessory foot of Malleus, the nacreous region is more extended distally and the ligament is multivincular (the formation of which is discussed), producing some posterior, but never any anterior, extension of the hinge line.
    14. A remarkably dense mixed bed of M. regula and I. isognomon in Darvel Bay, Sabah (Borneo), is described. Different spawning periods, by preventing competition during settlement, may account for this complete intermixing of sympatric species, which, however, are generalized herbivores where numbers are not limited by food supply.
    15. From a basic epifaunal habit (Pteria, Isognomon, M. regula), members of the Pteriacea have become adapted for infaunal life within sponges (Vulsella, Crenatula) or within soft substrates ( M. malleus, M. alba and the Pinnidae).
    16. There is final discussion about elongation in monomyarians, i.e., in the genus Malleus and in I. isognomon.
    Matched MeSH terms: Borneo
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