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Density dependence

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266:, density-dependent processes can act at the host/vector interface or within the host/vector life-cycle stages. At the host/vector interface, density-dependence may influence the input of L3 larvae into the host's skin and the ingestion of microfilariae by the vector. Within the life-cycle stages taking place in the vector, density-dependence may influence the development of L3 larvae in vectors and vector life expectancy. Within the life-cycle stages taking place in the host, density-dependence may influence the development of microfilariae and host life expectancy. 251: 194: 215: 64:(separate sex) obligatory parasites, mated female worms are required to complete a transmission cycle. At low parasite densities, the probability of a female worm encountering a male worm and forming a mating pair can become so low that reproduction is restricted due to single sex infections. At higher parasite densities, the probability of mating pairs forming and successful reproduction increases. This has been observed in the population dynamics of 337:
Contrariwise, positive density-dependent or facilitation processes make elimination of a parasite population more likely. Facilitation processes cause the reproductive success of the parasite to decrease with lower worm burden. Thus, control measures that reduce parasite burden will automatically
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Anderson and Gordon (1982) propose that the distribution of macroparasites in a host population is regulated by a combination of positive and negative density-dependent processes. In overdispersed distributions, a small proportion of hosts harbour most of the parasite population. Positive
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infection promotes immunosuppressive processes within the human host that suppress immunity against incoming infective L3 larvae. This suppression of anti-parasite immunity causes parasite establishment rates to increase with higher parasite burden.
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refers to minimum parasite density level for the parasite to persist in a population. Interventions that reduce parasite density to a level below this threshold will ultimately lead to the extinction of that parasite in that population.
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density-dependent processes occur in the life cycles of parasites. However, the extent to which one process predominates over the other vary widely according to the parasite, vector, and host involved. This is illustrated by the
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processes tend to restrict population growth rates and contribute to the stability of these populations. Interventions that lead to a reduction in parasite populations will cause a relaxation of density-dependent
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require at least seven days to mature into infective L3 larvae in the black fly, the worm load is restricted to levels that allow the black fly to survive for long enough to pass infective L3 larvae onto humans.
163:. Greater competition means an individual has a decreased contribution to the next generation i.e. offspring. Density-dependent mortality can be overcompensating, undercompensating or exactly compensating. 376:
of parasite populations. As mean parasite burden increases, negative density-dependent processes become more prominent and the distribution of the parasite population tends to become less overdispersed.
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life cycles, density-dependent processes can influence parasite fecundity, survival, and establishment. Density-dependent processes can act across multiple points of the macroparasite life cycle. For
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and its rates of egg production decrease as host infection intensity increases. Thus, the per-capita contribution of each worm to transmission decreases as a function of infection intensity.
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Duerr, H.P. (2003). "Density-dependent parasite establishment suggests infection-associated immunosuppression as an important mechanism for parasite density regulation in onchocerciasis".
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Snow, L.C. (2006). "Transmission dynamics of lymphatic filariasis: vector-specific density dependence in the development of Wuchereria bancrofti infective larvae in mosquitoes".
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Consequently, interventions that lead to a reduction in parasite burden will tend to cause the parasite distribution to become overdispersed. For instance, time-series data for
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Anderson, R.M. (1982). "Processes influencing the distribution of parasite numbers within host populations with special emphasis on parasite-induced host mortalities".
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Negative density-dependence, or density-dependent restriction, describes a situation in which population growth is curtailed by crowding, predators and competition.
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Basanez, M.G. (1996). "Density-dependent processes in the onchocerciasis: relationship between microfilarial intake and mortality of the simuliid vector".
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processes predominate. Thus, the number of L3 larvae per mosquito declines as the number of ingested microfilariae increases. Conversely, in
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vector. In this life-cycle, the life expectancy of the black fly vector decreases as the worm load ingested by the vector increases. Because
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exists, where the birth rate falls as competition increases. In the context of gastrointestinal nematodes, the weight of female
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infection demonstrates that 10 years of vector control lead to reduced parasite burden with a more overdispersed distribution.
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processes increase the extinction threshold, making it easier to achieve using parasite control interventions. Conversely,
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reduce per-capita reproductive success and increase the likelihood of elimination when facilitation processes predominate.
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processes predominate. Consequently, the number of L3 larvae per mosquito increases as the number of ingested
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Duerr, H.P. (2005). "Determinants of the eradicability of filarial infections: a conceptual approach".
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May, R.M. (1977). "Togetherness among Schistosomes: its effects on the dynamics of the infection".
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processes contribute to the resilience of macroparasite populations. At high parasite populations,
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Positive density-dependence processes may also occur in macroparasite infections that lead to
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This is also true for other organisms because an increased density means an increase in
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of parasite populations, whereas negative density-dependent processes contribute to
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Churcher, T.S. (2006). "Density dependence and the control of helminth parasites".
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Parasite-induced vector mortality is a form of negative density-dependence. The
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processes complicates control measures by decreasing the extinction threshold.
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An example of a density-dependent variable is crowding and competition.
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Positive density-dependence, density-dependent facilitation, or the
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Transactions of the Royal Society of Tropical Medicine and Hygiene
32:. This article will focus on density dependence in the context of 2283: 2090: 1960: 1955: 1582: 1530: 1190: 835: 171: 152:
available to each cell becomes insufficient to allow continued
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mosquitoes, which lack a well-developed cibarial armature,
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Density-dependence processes (red) in filariasis life cycle
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mosquitoes, which have well-developed cibarial armatures,
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Hixon, M (2009), "Density Dependence and Independence",
144:reaches a certain density, the amount of required 769:Modelling Onchocerciasis Transmission and Control 318:Implications for parasite persistence and control 2370: 356: 352: 331: 326: 322: 307: 291: 274: 270: 52:is facilitated by increased population density. 646: 644: 245: 73:Positive density-dependence processes occur in 124: 39: 1567: 820: 684: 682: 641: 103:and impede the development of infective L3 1788:Latitudinal gradients in species diversity 1574: 1560: 827: 813: 613:(4). Cambridge University Press: 331–355. 471: 469: 368:density-dependent processes contribute to 679: 581: 571: 418:, Chichester: John Wiley & Sons Ltd, 1686:Predator–prey (Lotka–Volterra) equations 1325:Tritrophic interactions in plant defense 766: 723: 650: 323:Negative density-dependent (restriction) 249: 213: 192: 1718:Random generalized Lotka–Volterra model 604: 466: 341: 228:life cycle involves transmission via a 2371: 1526:Herbivore adaptations to plant defense 553: 363:Implications for parasite distribution 1555: 808: 688: 518: 413: 1541:Predator avoidance in schooling fish 475: 1991:Intermediate disturbance hypothesis 445: 13: 1744:Ecological effects of biodiversity 800:Eradicability of filarial diseases 81:with a cibarial armature, such as 28:are regulated by the density of a 14: 2410: 1080:Generalist and specialist species 788: 771:. Rotterdam: Erasmus University. 478:Medical and Veterinary Entomology 218:Parasite-induced vector mortality 1803:Occupancy–abundance relationship 665:10.1111/j.1365-2656.2006.01154.x 490:10.1111/j.1365-2915.2006.00629.x 174:or environmental conditions and 136:, it describes the reduction in 1823:Relative abundance distribution 1536:Plant defense against herbivory 1403:Competitive exclusion principle 1115:Mesopredator release hypothesis 760: 48:describes a situation in which 1408:Consumer–resource interactions 717: 598: 547: 512: 439: 424:10.1002/9780470015902.a0021219 407: 170:, where other factors such as 168:density-independent inhibition 1: 2394:Evolutionary biology concepts 2254:Biological data visualization 2081:Environmental niche modelling 1808:Population viability analysis 533:10.1016/S0035-9203(03)90132-5 416:Encyclopedia of Life Sciences 401: 188: 1739:Density-dependent inhibition 460:10.1016/0025-5564(77)90030-X 269:In reality, combinations of 246:In macroparasite life cycles 7: 2208:Liebig's law of the minimum 2043:Resource selection function 934:Metabolic theory of ecology 389: 197:Density-dependent fecundity 125:Negative density-dependence 55: 40:Positive density-dependence 10: 2415: 2108:Niche apportionment models 1828:Relative species abundance 1032:Primary nutritional groups 929:List of feeding behaviours 178:may affect a population's 2357: 2289:Ecosystem based fisheries 2231: 2131: 2056: 1929: 1901:Interspecific competition 1866: 1793:Minimum viable population 1726: 1651:Maximum sustainable yield 1636:Intraspecific competition 1631:Effective population size 1594: 1511:Anti-predator adaptations 1496: 1375: 1302: 1259: 1181: 1148: 1045: 1022:Photosynthetic efficiency 957: 851: 738:10.1017/S0031182000055347 653:Journal of Animal Ecology 619:10.1017/S003118200006649X 161:intraspecific competition 77:life cycles that rely on 2279:Ecological stoichiometry 2244:Alternative stable state 703:10.1016/j.pt.2004.11.011 448:Mathematical Biosciences 2123:Ontogenetic niche shift 1986:Ideal free distribution 1896:Ecological facilitation 1646:Malthusian growth model 1616:Consumer-resource model 1473:Paradox of the plankton 1438:Energy systems language 1158:Chemoorganoheterotrophy 1125:Optimal foraging theory 1100:Heterotrophic nutrition 767:Plaisier, A.P. (1996). 560:Parasites & Vectors 275:positive (facilitation) 26:population growth rates 2269:Ecological forecasting 2213:Marginal value theorem 2011:Landscape epidemiology 1946:Cross-boundary subsidy 1881:Biological interaction 1231:Microbial intelligence 919:Green world hypothesis 691:Trends in Parasitology 573:10.1186/1756-3305-2-11 271:negative (restriction) 255: 219: 198: 140:division. When a cell 2274:Ecological humanities 2173:Ecological energetics 2118:Niche differentiation 1981:Habitat fragmentation 1749:Ecological extinction 1696:Small population size 1448:Feed conversion ratio 1428:Ecological succession 1360:San Francisco Estuary 1274:Ecological efficiency 1216:Microbial cooperation 253: 217: 196: 24:processes occur when 2299:Evolutionary ecology 2264:Ecological footprint 2259:Ecological economics 2183:Ecological threshold 2178:Ecological indicator 2048:Source–sink dynamics 2001:Land change modeling 1996:Insular biogeography 1848:Species distribution 1587:Modelling ecosystems 1246:Microbial metabolism 1085:Intraguild predation 874:Biogeochemical cycle 840:Modelling ecosystems 348:extinction threshold 342:Extinction threshold 208:Ascaris lumbricoides 96:Wuchereria bancrofti 2399:Population dynamics 2349:Theoretical ecology 2324:Natural environment 2188:Ecosystem diversity 2158:Ecological collapse 2148:Bateman's principle 2103:Limiting similarity 2016:Landscape limnology 1838:Species homogeneity 1676:Population modeling 1671:Population dynamics 1488:Trophic state index 117:Onchocerca volvulus 2384:Cellular processes 2360:Outline of ecology 2309:Industrial ecology 2304:Functional ecology 2168:Ecological deficit 2113:Niche construction 2076:Ecosystem engineer 1853:Species–area curve 1774:Introduced species 1589:: Other components 1521:Deimatic behaviour 1423:Ecological network 1355:North Pacific Gyre 1340:hydrothermal vents 1279:Ecological pyramid 1226:Microbial food web 1037:Primary production 982:Foundation species 795:Density dependence 554:Walker, M (2009). 256: 220: 201:Density-dependent 199: 166:There also exists 18:population ecology 2366: 2365: 2249:Balance of nature 2006:Landscape ecology 1891:Community ecology 1833:Species diversity 1769:Indicator species 1764:Gradient analysis 1641:Logistic function 1549: 1548: 1506:Animal coloration 1483:Trophic mutualism 1221:Microbial ecology 1012:Photoheterotrophs 997:Myco-heterotrophy 909:Ecosystem ecology 894:Carrying capacity 859:Abiotic component 180:carrying capacity 112:immunosuppression 50:population growth 22:density-dependent 2406: 2066:Ecological niche 2038:selection theory 1858:Umbrella species 1843:Species richness 1779:Invasive species 1759:Flagship species 1666:Population cycle 1661:Overexploitation 1626:Ecological yield 1576: 1569: 1562: 1553: 1552: 1458:Mesotrophic soil 1398:Climax community 1330:Marine food webs 1269:Biomagnification 1070:Chemoorganotroph 924:Keystone species 884:Biotic component 829: 822: 815: 806: 805: 783: 782: 764: 758: 757: 721: 715: 714: 686: 677: 676: 659:(6): 1313–1320. 648: 639: 638: 602: 596: 595: 585: 575: 551: 545: 544: 516: 510: 509: 473: 464: 463: 454:(3–4): 301–343. 443: 437: 436: 411: 93:mosquitoes. For 2414: 2413: 2409: 2408: 2407: 2405: 2404: 2403: 2369: 2368: 2367: 2362: 2353: 2339:Systems ecology 2227: 2198:Extinction debt 2163:Ecological debt 2153:Bioluminescence 2134: 2127: 2096:marine habitats 2071:Ecological trap 2052: 1932: 1925: 1868: 1862: 1818:Rapoport's rule 1813:Priority effect 1754:Endemic species 1722: 1681:Population size 1597: 1590: 1580: 1550: 1545: 1498: 1492: 1478:Trophic cascade 1388:Bioaccumulation 1371: 1298: 1255: 1177: 1144: 1041: 953: 914:Ecosystem model 847: 833: 791: 786: 779: 765: 761: 722: 718: 687: 680: 649: 642: 603: 599: 552: 548: 517: 513: 474: 467: 444: 440: 434: 412: 408: 404: 392: 374:underdispersion 365: 344: 320: 284:life cycle. 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1954: 1952: 1949: 1947: 1944: 1942: 1939: 1938: 1936: 1934: 1928: 1922: 1919: 1917: 1914: 1912: 1909: 1907: 1904: 1902: 1899: 1897: 1894: 1892: 1889: 1887: 1884: 1882: 1879: 1877: 1874: 1873: 1871: 1865: 1859: 1856: 1854: 1851: 1849: 1846: 1844: 1841: 1839: 1836: 1834: 1831: 1829: 1826: 1824: 1821: 1819: 1816: 1814: 1811: 1809: 1806: 1804: 1801: 1799: 1796: 1794: 1791: 1789: 1786: 1784: 1780: 1777: 1775: 1772: 1770: 1767: 1765: 1762: 1760: 1757: 1755: 1752: 1750: 1747: 1745: 1742: 1740: 1737: 1735: 1732: 1731: 1729: 1725: 1719: 1716: 1712: 1709: 1707: 1704: 1703: 1702: 1699: 1697: 1694: 1692: 1689: 1687: 1684: 1682: 1679: 1677: 1674: 1672: 1669: 1667: 1664: 1662: 1659: 1657: 1654: 1652: 1649: 1647: 1644: 1642: 1639: 1637: 1634: 1632: 1629: 1627: 1624: 1622: 1619: 1617: 1614: 1612: 1609: 1607: 1604: 1603: 1601: 1599: 1593: 1588: 1584: 1577: 1572: 1570: 1565: 1563: 1558: 1557: 1554: 1542: 1539: 1537: 1534: 1532: 1529: 1527: 1524: 1522: 1519: 1517: 1514: 1512: 1509: 1507: 1504: 1503: 1501: 1495: 1489: 1486: 1484: 1481: 1479: 1476: 1474: 1471: 1469: 1466: 1464: 1461: 1459: 1456: 1454: 1451: 1449: 1446: 1444: 1441: 1439: 1436: 1434: 1431: 1429: 1426: 1424: 1421: 1419: 1416: 1414: 1411: 1409: 1406: 1404: 1401: 1399: 1396: 1394: 1391: 1389: 1386: 1384: 1381: 1380: 1378: 1374: 1366: 1363: 1361: 1358: 1356: 1353: 1351: 1348: 1346: 1343: 1341: 1338: 1336: 1333: 1332: 1331: 1328: 1326: 1323: 1321: 1318: 1316: 1313: 1311: 1308: 1307: 1305: 1301: 1295: 1294:Trophic level 1292: 1290: 1287: 1285: 1282: 1280: 1277: 1275: 1272: 1270: 1267: 1266: 1264: 1262: 1258: 1252: 1251:Phage ecology 1249: 1247: 1244: 1242: 1241:Microbial mat 1239: 1237: 1234: 1232: 1229: 1227: 1224: 1222: 1219: 1217: 1214: 1212: 1209: 1207: 1204: 1202: 1199: 1197: 1196:Bacteriophage 1194: 1192: 1189: 1188: 1186: 1184: 1180: 1174: 1171: 1169: 1166: 1164: 1163:Decomposition 1161: 1159: 1156: 1155: 1153: 1151: 1147: 1141: 1138: 1136: 1133: 1131: 1128: 1126: 1123: 1121: 1118: 1116: 1113: 1111: 1110:Mesopredators 1108: 1106: 1103: 1101: 1098: 1096: 1093: 1091: 1088: 1086: 1083: 1081: 1078: 1076: 1073: 1071: 1068: 1066: 1063: 1061: 1058: 1056: 1055:Apex predator 1053: 1052: 1050: 1048: 1044: 1038: 1035: 1033: 1030: 1028: 1025: 1023: 1020: 1018: 1015: 1013: 1010: 1008: 1005: 1003: 1000: 998: 995: 993: 990: 988: 985: 983: 980: 978: 975: 973: 970: 968: 965: 964: 962: 960: 956: 950: 947: 945: 942: 940: 937: 935: 932: 930: 927: 925: 922: 920: 917: 915: 912: 910: 907: 905: 902: 900: 897: 895: 892: 890: 889:Biotic stress 887: 885: 882: 880: 877: 875: 872: 870: 867: 865: 862: 860: 857: 856: 854: 850: 845: 841: 837: 830: 825: 823: 818: 816: 811: 810: 807: 801: 798: 796: 793: 792: 780: 778:90-72245-68-7 774: 770: 763: 755: 751: 747: 743: 739: 735: 731: 727: 720: 712: 708: 704: 700: 696: 692: 685: 683: 674: 670: 666: 662: 658: 654: 647: 645: 636: 632: 628: 624: 620: 616: 612: 608: 601: 593: 589: 584: 579: 574: 569: 565: 561: 557: 550: 542: 538: 534: 530: 526: 522: 515: 507: 503: 499: 495: 491: 487: 483: 479: 472: 470: 461: 457: 453: 449: 442: 435: 429: 425: 421: 417: 410: 406: 397: 396:Plant density 394: 393: 387: 385: 384: 378: 375: 371: 360: 358: 354: 349: 339: 335: 333: 328: 324: 315: 313: 312:microfilariae 309: 305: 304: 299: 298: 293: 289: 288: 283: 282: 276: 272: 267: 265: 261: 260:macroparasite 252: 243: 240: 239:microfilariae 237: 236: 231: 227: 226: 216: 212: 210: 209: 204: 195: 186: 183: 181: 177: 173: 169: 164: 162: 157: 155: 151: 147: 143: 139: 135: 130: 122: 119: 118: 113: 108: 106: 102: 101:microfilariae 98: 97: 92: 91: 86: 85: 80: 76: 75:macroparasite 71: 69: 68: 63: 53: 51: 47: 37: 36:life cycles. 35: 34:macroparasite 31: 27: 23: 19: 2389:Epidemiology 2329:Regime shift 2314:Macroecology 2035: 2031: 1971:Edge effects 1941:Biogeography 1886:Commensalism 1738: 1734:Biodiversity 1611:Allee effect 1350:kelp forests 1303:Example webs 1168:Detritivores 1007:Organotrophs 987:Kinetotrophs 939:Productivity 768: 762: 729: 726:Parasitology 725: 719: 697:(2): 88–96. 694: 690: 656: 652: 610: 607:Parasitology 606: 600: 563: 559: 549: 524: 520: 514: 481: 477: 451: 447: 441: 415: 409: 381: 379: 366: 353:Facilitation 345: 336: 332:restrictions 321: 308:facilitation 301: 295: 285: 281:W. bancrofti 279: 268: 257: 233: 223: 221: 206: 200: 184: 176:disturbances 167: 165: 158: 134:cell biology 131: 128: 115: 109: 94: 88: 82: 72: 67:Schistosomes 66: 59: 46:Allee effect 43: 21: 15: 1966:Disturbance 1869:interaction 1691:Recruitment 1621:Depensation 1413:Copiotrophs 1284:Energy flow 1206:Lithotrophy 1150:Decomposers 1130:Planktivore 1105:Insectivore 1095:Heterotroph 1060:Bacterivore 1027:Phototrophs 977:Chemotrophs 949:Restoration 899:Competition 357:restriction 327:restriction 314:increases. 292:restriction 235:O. volvulus 154:cell growth 2379:Cell cycle 2373:Categories 2334:Sexecology 1911:Parasitism 1876:Antibiosis 1711:Resistance 1706:Resilience 1596:Population 1516:Camouflage 1468:Oligotroph 1383:Ascendency 1345:intertidal 1335:cold seeps 1289:Food chain 1090:Herbivores 1065:Carnivores 992:Mixotrophs 967:Autotrophs 846:components 402:References 142:population 30:population 2239:Allometry 2193:Emergence 1921:Symbiosis 1906:Mutualism 1701:Stability 1606:Abundance 1418:Dominance 1376:Processes 1365:tide pool 1261:Food webs 1135:Predation 1120:Omnivores 1047:Consumers 1002:Mycotroph 959:Producers 904:Ecosystem 869:Behaviour 566:(1): 11. 303:Anopheles 230:black fly 203:fecundity 150:nutrients 84:Anopheles 62:dioecious 2294:Endolith 2223:Xerosere 2135:networks 1951:Ecocline 1497:Defense, 1173:Detritus 1075:Foraging 944:Resource 754:19196511 711:15664532 673:17032363 635:42555004 592:19208229 541:14584385 506:23152390 498:17044876 390:See also 189:Examples 56:Examples 2284:Ecopath 2091:Habitat 1961:Ecotype 1956:Ecotone 1933:ecology 1931:Spatial 1867:Species 1727:Species 1598:ecology 1583:Ecology 1531:Mimicry 1499:counter 1443:f-ratio 1191:Archaea 879:Biomass 852:General 844:Trophic 836:Ecology 746:7145478 627:8873475 583:2672930 172:weather 79:vectors 1315:Rivers 1211:Marine 775:  752:  744:  709:  671:  633:  625:  590:  580:  539:  504:  496:  430:  105:larvae 2232:Other 2133:Other 2086:Guild 2058:Niche 1310:Lakes 750:S2CID 631:S2CID 502:S2CID 297:Aedes 287:Culex 90:Culex 1320:Soil 773:ISBN 742:PMID 707:PMID 669:PMID 623:PMID 588:PMID 537:PMID 494:PMID 428:ISBN 346:The 300:and 273:and 148:and 138:cell 734:doi 699:doi 661:doi 615:doi 611:113 578:PMC 568:doi 529:doi 486:doi 456:doi 420:doi 258:In 132:In 87:or 60:In 16:In 2375:: 1781:/ 1585:: 842:: 838:: 748:. 740:. 730:85 728:. 705:. 695:21 693:. 681:^ 667:. 657:75 655:. 643:^ 629:. 621:. 609:. 586:. 576:. 562:. 558:. 535:. 525:97 523:. 500:. 492:. 482:20 480:. 468:^ 452:35 450:. 426:, 182:. 156:. 114:. 70:. 20:, 2036:K 2034:/ 2032:r 1575:e 1568:t 1561:v 828:e 821:t 814:v 781:. 756:. 736:: 713:. 701:: 675:. 663:: 637:. 617:: 594:. 570:: 564:2 543:. 531:: 508:. 488:: 462:. 458:: 422::

Index

population ecology
population growth rates
population
macroparasite
Allee effect
population growth
dioecious
Schistosomes
macroparasite
vectors
Anopheles
Culex
Wuchereria bancrofti
microfilariae
larvae
immunosuppression
Onchocerca volvulus
cell biology
cell
population
growth factors
nutrients
cell growth
intraspecific competition
weather
disturbances
carrying capacity

fecundity
Ascaris lumbricoides

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