A General Model of Population Dynamics Accounting for Multiple Kinds of Interaction

<div>Facultative-facultative mutualism with negative intraspecific interaction: (a) we have the case where <span class="nowrap"><svg height="12.4698pt" id="M222" style="vertical-align:-3.18147pt" version="1.1" viewbox="-0.0498162 -9.28833 94.4455 12.4698" width="94.4455pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M452 333C452 398 425 448 375 448C329 448 235 404 140 292H138L229 683C233 701 234 712 225 712C208 712 149 686 66 677L64 652H97C135 652 140 651 129 598L38 167C23 96 29 57 44 33C60 7 98 -12 132 -12C169 -12 232 3 290 41C383 102 452 223 452 333ZM365 316C365 199 308 87 239 49C221 39 200 35 183 35C137 35 99 70 112 163C116 191 123 215 129 233C190 316 290 392 330 392C348 392 365 372 365 316Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,4.992,3.132)"><path d="M389 0V32C297 38 291 46 291 118V635C234 613 175 595 109 583V556L161 554C203 552 207 547 207 497V118C207 46 201 38 110 32V0H389Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,9.424,3.132)"><path d="M389 0V32C297 38 291 46 291 118V635C234 613 175 595 109 583V556L161 554C203 552 207 547 207 497V118C207 46 201 38 110 32V0H389Z"></path></g><g transform="matrix(.013,0,0,-0.013,18.019,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,29.282,0)"><path d="M452 333C452 398 425 448 375 448C329 448 235 404 140 292H138L229 683C233 701 234 712 225 712C208 712 149 686 66 677L64 652H97C135 652 140 651 129 598L38 167C23 96 29 57 44 33C60 7 98 -12 132 -12C169 -12 232 3 290 41C383 102 452 223 452 333ZM365 316C365 199 308 87 239 49C221 39 200 35 183 35C137 35 99 70 112 163C116 191 123 215 129 233C190 316 290 392 330 392C348 392 365 372 365 316Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,34.274,3.132)"><path d="M414 144C384 79 371 75 317 75H135L276 221C367 316 408 376 408 465C408 570 327 635 237 635C179 635 131 609 100 575L42 494L67 471C94 510 138 565 205 565C277 565 321 517 321 435C321 348 258 270 195 195C146 137 88 81 33 26V0H411C423 44 433 88 446 135L414 144Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,38.705,3.132)"><path d="M414 144C384 79 371 75 317 75H135L276 221C367 316 408 376 408 465C408 570 327 635 237 635C179 635 131 609 100 575L42 494L67 471C94 510 138 565 205 565C277 565 321 517 321 435C321 348 258 270 195 195C146 137 88 81 33 26V0H411C423 44 433 88 446 135L414 144Z"></path></g><g transform="matrix(.013,0,0,-0.013,47.3,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,58.563,0)"><path d="M535 230V280H52V230H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,66.194,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,72.434,0)"><path d="M113 -12C146 -12 170 11 170 46C170 78 146 103 114 103S58 78 58 46C58 11 82 -12 113 -12Z"></path></g><g transform="matrix(.013,0,0,-0.013,75.398,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,81.638,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,87.878,0)"><path d="M412 140C382 77 369 73 315 73H129L270 222C362 320 402 379 402 466C402 571 322 635 234 635C177 635 130 609 99 576L42 495L64 475C90 514 133 568 201 568C274 568 318 519 318 435C318 349 255 267 193 193C144 135 87 78 32 23V0H405C417 45 427 89 440 131L412 140Z"></path></g></svg>,</span> which means that intraspecific competition is weaker than mutualism; (b) we have <span class="nowrap"><svg height="12.4698pt" id="M223" style="vertical-align:-3.18147pt" version="1.1" viewbox="-0.0498162 -9.28833 94.4455 12.4698" width="94.4455pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M452 333C452 398 425 448 375 448C329 448 235 404 140 292H138L229 683C233 701 234 712 225 712C208 712 149 686 66 677L64 652H97C135 652 140 651 129 598L38 167C23 96 29 57 44 33C60 7 98 -12 132 -12C169 -12 232 3 290 41C383 102 452 223 452 333ZM365 316C365 199 308 87 239 49C221 39 200 35 183 35C137 35 99 70 112 163C116 191 123 215 129 233C190 316 290 392 330 392C348 392 365 372 365 316Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,4.992,3.132)"><path d="M389 0V32C297 38 291 46 291 118V635C234 613 175 595 109 583V556L161 554C203 552 207 547 207 497V118C207 46 201 38 110 32V0H389Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,9.424,3.132)"><path d="M389 0V32C297 38 291 46 291 118V635C234 613 175 595 109 583V556L161 554C203 552 207 547 207 497V118C207 46 201 38 110 32V0H389Z"></path></g><g transform="matrix(.013,0,0,-0.013,18.019,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,29.282,0)"><path d="M452 333C452 398 425 448 375 448C329 448 235 404 140 292H138L229 683C233 701 234 712 225 712C208 712 149 686 66 677L64 652H97C135 652 140 651 129 598L38 167C23 96 29 57 44 33C60 7 98 -12 132 -12C169 -12 232 3 290 41C383 102 452 223 452 333ZM365 316C365 199 308 87 239 49C221 39 200 35 183 35C137 35 99 70 112 163C116 191 123 215 129 233C190 316 290 392 330 392C348 392 365 372 365 316Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,34.274,3.132)"><path d="M414 144C384 79 371 75 317 75H135L276 221C367 316 408 376 408 465C408 570 327 635 237 635C179 635 131 609 100 575L42 494L67 471C94 510 138 565 205 565C277 565 321 517 321 435C321 348 258 270 195 195C146 137 88 81 33 26V0H411C423 44 433 88 446 135L414 144Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,38.705,3.132)"><path d="M414 144C384 79 371 75 317 75H135L276 221C367 316 408 376 408 465C408 570 327 635 237 635C179 635 131 609 100 575L42 494L67 471C94 510 138 565 205 565C277 565 321 517 321 435C321 348 258 270 195 195C146 137 88 81 33 26V0H411C423 44 433 88 446 135L414 144Z"></path></g><g transform="matrix(.013,0,0,-0.013,47.3,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,58.563,0)"><path d="M535 230V280H52V230H535Z"></path></g><g transform="matrix(.013,0,0,-0.013,66.194,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,72.434,0)"><path d="M113 -12C146 -12 170 11 170 46C170 78 146 103 114 103S58 78 58 46C58 11 82 -12 113 -12Z"></path></g><g transform="matrix(.013,0,0,-0.013,75.398,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,81.638,0)"><path d="M241 635C89 635 35 457 35 312C35 153 89 -12 240 -12C390 -12 443 166 443 312C443 466 390 635 241 635ZM238 602C329 602 354 454 354 312C354 172 330 22 240 22C152 22 124 173 124 313S148 602 238 602Z"></path></g><g transform="matrix(.013,0,0,-0.013,87.878,0)"><path d="M249 635C141 635 70 555 70 471C70 401 114 353 179 316C143 294 106 267 90 252C68 231 45 202 45 157C45 50 130 -12 237 -12C322 -12 435 52 435 169C435 256 372 304 303 343C349 374 375 398 383 407C401 429 411 458 411 487C411 569 344 635 249 635ZM238 603C285 603 337 567 337 482C337 422 310 385 276 358C205 393 145 426 145 500C145 552 179 603 238 603ZM248 20C183 20 125 70 125 163C125 218 158 268 206 300C284 261 355 217 355 143C355 66 308 20 248 20Z"></path></g></svg>,</span> which means that intraspecific competition is stronger than mutualism. Here, <span class="nowrap"><svg height="12.4894pt" id="M224" style="vertical-align:-3.181499pt" version="1.1" viewbox="-0.0498162 -9.3079 182.074 12.4894" width="182.074pt" xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink"><g transform="matrix(.013,0,0,-0.013,0,0)"><path d="M393 379C402 394 400 411 393 422C384 437 365 448 348 448C301 448 237 372 186 285H182L193 335C210 408 205 448 178 448C150 448 80 402 29 344L45 321C80 355 114 373 122 373C128 373 130 365 124 330C106 228 76 98 50 -5L57 -12C82 -5 112 3 132 6L172 203C196 256 234 304 254 329C275 355 293 367 306 367C318 367 330 360 342 348C347 343 355 343 365 350S386 367 393 379Z"></path></g><g transform="matrix(.0091,0,0,-0.0091,4.901,3.132)"><path d="M389 0V32C297 38 291 46 291 118V635C234 613 175 595 109 583V556L161 554C203 552 207 547 207 497V118C207 46 201 38 110 32V0H389Z"></path></g><g transform="matrix(.013,0,0,-0.013,13.479,0)"><path d="M535 323V373H52V323H535ZM535 138V188H52V138H535Z"></path></g><g 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Complexity

fig11

Figure 11

Figure 11: A General Model of Population Dynamics Accounting for Multiple Kinds of Interaction 