1167:
1177:
20:
1172:
526:
36:
578:. Selectively timed culls, or removal of a percentage of the breeding individuals, of grey squirrel populations can help control and prevent tree damage. Although grey squirrel populations readily recover from culls, their coincidence with annual periods of the greatest damage to trees, typically between April and September when the squirrels are most active in
516:
groups filed a lawsuit. During the three-year suspension, the grey squirrel population had grown so much that it was no longer feasible to attempt eradication by the methods initially used. If the species spreads significantly (the three populations in Italy invade France or
Switzerland and establish
137:
can be classified as a
Disease Mediated Invasion (DMI). More specifically, the invasion of the grey squirrels and displacement of the red squirrels can be described by "spillover DMI", which is defined as a non-native species bringing diseases with them into the new area that will infect the native
586:
populations, can minimize tree damage. Alternatively, non-lethal methods of population control, such as fertility treatments, are being developed, although the effectiveness of these treatments is unknown. In areas most threatened by spread of grey squirrels, eradication may be more effective than
419:
between 2051 and 2066. In a scenario where poor-year seed crops were not included, the "worst-case" scenario, grey squirrels will reach the
Western Alps by 2015, France by 2026, and Switzerland between 2031 and 2041. Along the border between France and Italy, the forest cover is mainly composed of
222:
between grey and red squirrels. In areas of large core overlap between red and grey squirrel populations, a reduction in body mass of red squirrels during the spring has been observed. This is most likely due to the fact that reds and greys participate in direct competition for limited resources,
537:
is a natural predator of squirrels, including the eastern grey squirrel, but has been eradicated from large parts of the area affected by the grey squirrel such as
England and Wales. The European pine marten's population is currently expanding and there is evidence that grey squirrel populations
562:
countries, but both illegal and allowed importations and escapes of grey squirrels from captivity pose threats to native red squirrel and tree populations. In addition to preventing introduction of grey squirrels, containment and eradication could serve to control the species’ invasive actions.
213:
One of the biggest concerns about the grey squirrel as an invasive species is the effect introduction has on populations of red squirrels. Grey and red squirrels compete for resources, and greys will pilfer food caches of red squirrels. Reduction in red squirrel individual growth, juvenile
243:
from grey squirrels to reds is thought to be a major factor in the decline of red squirrel populations. Grey squirrels do not die from the squirrelpox virus and can infect red squirrels causing the appearance of scabs and lesions on the face, feet, and genitals and eventually death. In
844:
Lawton, C., Cowan, P., Bertolino, S., Lurz, P.P.W. & Peters, A.R. (2010) The consequences of introducing non-indigenous species: two case studies, the grey squirrel in Europe and the brushtail possum in New
Zealand. Rev. sci. tech. Off. int. Epiz., 29 (2),
925:
Craig M. Shuttleworth, C. M.; Lurz, P.; Geddes, N.; Browne, J. (2012) Integrating red squirrel (Sciurus vulgaris) habitat requirements with the management of pathogenic tree disease in commercial forests in the UK. Forest
Ecology and Management 279,
199:
typically are not present in the area that the species has been introduced or has invaded. Because the grey squirrel poses such a threat to the trees and red squirrel in Europe, there is great effort to stop the further spread of the grey squirrels.
892:
Gossa, A., La Morgia, V., Marchisio, P., et al. (2012) Does a larger carrying capacity for an exotic species allow environment invasion? - Some considerations on the competition of red and grey squirrels. Journal of
Biological Systems, 20,
126:). Both species have similar diets, use similar locations of the environment and have similar activity patterns; however, there is no evidence of noticeable aggression between the two species. These eastern grey squirrels are considered an
879:
Bertolino, S., Lurz, P.W.W., Sanderson, R. & Rushton, S.P. (2008) Predicting the spread of the
American grey squirrel (Sciurus carolinensis) in Europe: A call for a co-ordinated European approach. Biological Conservation, 141,
648:
Bertolino, S. & Piero, G. (2003) Spread and attempted eradication of the grey squirrel (Sciurus caolinensis) in Italy, and consequences for the red squirrel (Sciurus vulgaris) in
Eurasia. Biological Conservation,
402:
populations. The models examined potential corridors that would allow grey squirrels to invade France and
Switzerland and the time scale of such invasions. In a scenario based on a random pattern of good, poor and
340:. Bark stripping makes branches weaker and more susceptible to being broken off by wind or animals and can also cause secondary infection that may lead to death. Bark stripping makes trees more susceptible to
936:
Emma Sheehy, Colin Lawton (March 2014). "Population crash in an invasive species following the recovery of a native predator: the case of the American grey squirrel and the European pine marten in Ireland".
456:; for this reason, grey squirrel performance in this habitat will affect the spread of the species. Eventually, if populations are left unchecked, the grey squirrel will spread through a large portion of
913:
Bertolino, S. & Lurz, P. (2011) Callosciurus squirrels: worldwide introductions, ecological impacts and recommendations to prevent the establishment of new invasive populations. Mammal Review, 22-33.
117:
for grey squirrels. Genetic studies have shown that human intervention-- released pets and intentional dispersal--may play a much larger role in the spread of grey squirrels than previously believed.
452:
than red squirrels. It may be difficult for the grey squirrels to attain their higher energy requirement in this environment because they may have difficulty reaching the small seeds contained in the
563:
Methods for grey squirrel population control depend on the target of protection. In the presence of red squirrels, cage trapping or shooting can be effective. In the absence of red squirrels,
260:
is uncommon even though the disease may be highly prevalent in a red squirrel population. The full consequences of these interactions cannot be completely determined, though the difference in
288:. In those populations, the greys mainly feed on "seeds, fruits, buds, flowers and, occasionally, on insects, switching from one resource to another according to seasonal availability".
1018:
Shine, R. & Doody, J. S. (2011) Invasive species control: understanding conflicts between researchers and the general community. Frontiers in Ecology and the Environment 9: 400–406.
546:
Although the negative consequences of introduction of non-native species are generally known in the scientific community, the majority of the public is unaware of the effects on native
187:. From studying all of the known successful introductions of this species, 71.4% used fewer than 10 individuals. This efficiency in the spread of grey squirrels can be explained by the
176:
but were captured and removed in 1996. As stated before, not all grey squirrel introductions were intentional. For example, in 1966 two females and one male escaped from a game farm on
183:
Almost 80% of known grey squirrel introductions worldwide resulted in successful establishment in the new area. Typically, grey squirrels do not need large numbers to start a new
227:
occurring. This direct competition also explains a decrease in juvenile recruitment and reproductive success, as immature reds will die due to lack of resources before reaching
669:
Sandro, B. (2008) Introduction of the American grey squirrel (Sciurus carolinensis) in Europe: a case study in biological invasion. Current Science, vol. 95, No. 7, 903-906.
998:
Population crash in an invasive species following the recovery of a native predator: the case of the American grey squirrel and the European pine marten in Ireland
153:
They were first introduced into England, in a concerted way, in 1876, and through rapidly growing population and further introductions they spread to the rest of
101:
They were first introduced into England, in a concerted way, in 1876, and through rapidly growing population and further introductions they spread to the rest of
794:
Strauss, A., White, A., & Boots, M. (2012) Invading with biological weapons: the importance of disease-mediated invasions. Functional Ecology, 26, 1249-1261.
517:
successful populations outside of Italy) before preventative or eradicative efforts are taken, it will, in all probability, be too late for effective action.
1046:
1660:
980:
386:
Spatially explicit population dynamics models were used by Bertolino et al. 2008 to predict the spread of the grey squirrel populations present in
1645:
252:
of the red squirrel due to increased mortality of reds from the squirrelpox virus. In fact, the virus works so quickly in killing its host after
421:
165:
440:
in Switzerland, making the area along the Ticino a likely corridor for the grey squirrel to invade Switzerland. The upper portion of the
280:
bird species as they sometimes feed on bird eggs and chicks. Although the grey squirrel is often blamed in part for this decline in the
816:
Simberloff, D. (2008) Rats are not the only introduced rodents producing ecosystem impacts on islands. Biological Invasions, 1735-1742.
558:
as primary means to prevent new introductions of non-native species. Currently, the release of non-native species is illegal in most
407:
seed crops (vegetation produces significant abundance of food resources), in the "best-case" scenario, grey squirrels will reach the
769:"Don't blame grey squirrels: Their British invasion had much more to do with us | Imperial News | Imperial College London"
1665:
1166:
1129:
1039:
249:
489:
341:
574:
can be used in special feeding hoppers. These methods are currently effectively controlling the grey squirrel population in
168:
was said to have introduced the American squirrels into his estate in 1911. Then in 1966, five squirrels were introduced to
1024:
1002:
492:, decreasing the population of lodgepole pines exponentially, and that Scots pine populations will decrease slower due to
1563:
1554:
1540:
1532:
445:
1655:
1032:
480:
for the virus. Further, a separate model examining the effects of the spread of the grey squirrel on the health of
325:
768:
683:
1602:
1444:
1244:
1236:
1153:
538:
plummet where it becomes present, with the populations of the co-evolved red squirrel subsequently recovering.
188:
296:
Certain behaviour of the grey squirrel, such as bark stripping, also has negative effects on trees and timber
361:
is another fungus affecting tree health. Trees between 10 and 40 years of age and certain species, including
1670:
353:
160:
The first reported introduction of grey squirrels in continental Europe was in 1948, and they were taken to
1284:
1195:
1186:
1145:
150:
Grey squirrels have been introduced either on purpose or by accident in many places throughout the world.
1610:
1412:
1404:
1396:
1348:
1137:
265:
219:
71:
67:
24:
1328:
1292:
1468:
1452:
1420:
946:
726:
534:
345:
215:
139:
1476:
1428:
1388:
1368:
1252:
554:
will require a coordinated approach of many nations, and ecologists recommend public education and
366:
1625:
1512:
1300:
1228:
962:
750:
551:
493:
457:
449:
313:
261:
224:
110:
106:
1436:
1260:
997:
742:
583:
555:
473:
317:
240:
1650:
1594:
1492:
1176:
954:
734:
321:
309:
305:
177:
127:
122:
276:
Grey squirrels also bear some of the responsibility for the decline of select woodland and
130:
in Europe because of their presence contributing to the displacement of the red squirrels.
1276:
1268:
228:
172:, Italy in the park of Genoa. Later in 1994, three pairs were again released to a park in
950:
730:
1320:
620:
485:
477:
429:
301:
134:
113:
in 1948 and has quickly taken advantage of Europe's food sources, habitats and lack of
19:
1639:
1577:
966:
575:
568:
513:
437:
404:
357:. RBNB results in a necrotrophic infection that kills needle tissue in pine species.
245:
154:
102:
63:
754:
369:, are more vulnerable than others. Grey squirrels also damage trees by eating their
1484:
1460:
1112:
1101:
688:
592:
453:
408:
337:
329:
40:
612:
424:
trees; this represents a likely corridor to France for the grey squirrel. Mixed
416:
333:
1171:
958:
525:
509:
481:
297:
184:
35:
529:
European pine martens have kept grey squirrel populations in check in Ireland
448:. Grey squirrels, which are heavier than native red squirrels, have a higher
191:, which is when the invading species has better survival because its natural
1060:
616:
547:
425:
370:
253:
161:
114:
738:
587:
population containment. Eradication of the population of grey squirrels in
464:, potentially invading the entire global distribution of the red squirrel.
746:
138:
species. Spillover is especially an issue of when the invading species is
1083:
1064:
1056:
579:
571:
505:
433:
391:
277:
257:
196:
192:
164:, Italy, where there is still a growing population. In Ireland, however,
95:
248:, grey squirrels have been able to spread 17 to 25 times faster through
1209:
461:
173:
51:, which has largely been displaced by the grey squirrel in some areas.
1095:
1089:
1077:
714:
713:
Okubo, A.; Maini, P.K.; Williamson, M.H.; Murray, J.D. (1989-11-22).
608:
564:
559:
412:
399:
349:
91:
79:
55:
48:
300:. This is thought to be biggest effect introduced squirrels have on
604:
588:
550:. Effective control of the spreading of grey squirrels throughout
524:
496:
resulting from the bark-stripping behaviour of the grey squirrel.
395:
387:
362:
285:
169:
87:
83:
34:
18:
441:
105:
by the early to mid-20th century. The eastern grey squirrel was
75:
1028:
373:, suggesting this behaviour provides some nutritional benefit.
476:
is predicted to spread by the grey squirrels as they act as a
284:, this is probably not the case for populations introduced to
615:
for the next 100 years. While these efforts are supported by
328:
of seeds by grey squirrels causes notching in the acorns of
239:
In addition to competition for resources, the spread of the
512:; however, this effort was suspended for three years after
281:
611:
would eliminate the threat of grey squirrel invasion into
623:, governmental support, and awareness of the situation.
719:
Proceedings of the Royal Society B: Biological Sciences
715:"On the Spatial Spread of the Grey Squirrel in Britain"
504:
In 1991, eradication of a grey squirrel population in
142:
to the native one. This can have devastating effects.
981:
How to eradicate grey squirrels without firing a shot
607:, and eradication of the grey squirrel population in
468:
Predicted spread of the poxvirus and damage to trees
264:
and weight of the two species may have an effect on
16:
American eastern grey squirrels acclimated in Europe
1575:
1552:
1207:
1184:
1110:
684:"If you want red squirrels, you have to kill greys"
120:
The native squirrel in Europe is the red squirrel (
909:
907:
905:
903:
901:
899:
875:
873:
871:
812:
810:
808:
806:
804:
802:
800:
869:
867:
865:
863:
861:
859:
857:
855:
853:
851:
644:
642:
640:
638:
636:
1006:, March 2014, Volume 23, Issue 3, pp. 753-774.
1040:
790:
788:
786:
784:
782:
780:
778:
677:
675:
488:pines showed that grey squirrels will spread
8:
1014:
1012:
921:
919:
888:
886:
840:
838:
836:
834:
832:
830:
828:
826:
824:
822:
665:
663:
661:
659:
657:
655:
500:Continued spread after attempted eradication
708:
706:
1047:
1033:
1025:
542:Proposed plans for control or eradication
133:The invasion of the grey squirrel in the
632:
591:would protect two native subspecies of
146:Introduction of grey squirrel to Europe
432:; this area connects with contiguous
324:including the grey squirrel. Scatter-
7:
382:Predicted spread throughout Europe
157:by the early to mid-20th century.
14:
1661:Invasive animal species in Europe
1175:
1170:
1165:
1130:Central American dwarf squirrel
996:Emma Sheehy & Colin Lawton,
256:that seeing a red squirrel with
619:, their success will depend on
508:was attempted via trapping and
195:are not present. Also, natural
1646:1876 establishments in England
1:
1541:Southern Amazon red squirrel
1533:Northern Amazon red squirrel
1156:(Microsciurus santanderensis)
1003:Biodiversity and Conservation
939:Biodiversity and Conservation
601:Sciurus vulgaris meridionalis
342:red band needle blight (RBNB)
272:Woodland and songbird species
682:Hickman, Leo (2012-09-05).
415:between 2066 and 2071, and
336:, which can hinder natural
1687:
1603:Southwestern red squirrel
1564:Bangs's mountain squirrel
1237:South Yungas red squirrel
1198:(Rheithrosciurus macrotis)
1140:(Microsciurus flaviventer)
66:, where they are known as
1622:
1613:(Tamiasciurus hudsonicus)
1445:Calabrian black squirrel
1269:Atlantic Forest squirrel
1245:Yellow-throated squirrel
1163:
1154:Santander dwarf squirrel
1072:
1055:Extant species of family
959:10.1007/s10531-014-0632-7
597:Sciurus vulgaris italicus
428:woodland lines the river
344:, which is caused by the
266:energy flow in ecosystems
31:, native to North America
1597:(Tamiasciurus douglasii)
1566:(Syntheosciurus brochus)
189:enemy release hypothesis
1666:Invasive mammal species
1605:(Tamiasciurus fremonti)
1479:(Sciurus variegatoides)
1196:Tufted ground squirrel
1146:Western dwarf squirrel
411:between 2026 and 2036,
354:Dothistroma septosporum
223:suggesting there is no
204:Impact on other species
140:morphologically similar
70:(to differentiate from
1611:American red squirrel
1495:(Sciurus yucatanensis)
1455:(Sciurus nayaritensis)
1447:(Sciurus meridionalis)
1415:(Sciurus carolinensis)
1413:Eastern gray squirrel
1405:Mexican gray squirrel
1397:Arizona gray squirrel
1349:Western gray squirrel
1247:(Sciurus gilvigularis)
1148:(Microsciurus mimulus)
1138:Amazon dwarf squirrel
739:10.1098/rspb.1989.0070
530:
490:red band needle blight
444:is covered largely by
72:western gray squirrels
68:eastern gray squirrels
52:
32:
1535:(Sciurus igniventris)
1453:Mexican fox squirrel
1407:(Sciurus aureogaster)
1399:(Sciurus arizonensis)
1255:(Sciurus granatensis)
1239:(Sciurus argentinius)
1132:(Microsciurus alfari)
528:
250:competitive exclusion
38:
22:
1477:Variegated squirrel
1303:(Sciurus stramineus)
1285:Richmond's squirrel
1279:(Sciurus pucheranii)
1253:Red-tailed squirrel
535:European pine marten
359:Phytophthora ramorum
332:. Notching prevents
320:and the presence of
216:reproductive success
209:Competition for food
60:Sciurus carolinensis
29:Sciurus carolinensis
1543:(Sciurus spadiceus)
1513:Caucasian squirrel
1331:(Sciurus pyrrhinus)
1329:Junín red squirrel
1323:(Sciurus flammifer)
1301:Guayaquil squirrel
1293:Sanborn's squirrel
1287:(Sciurus richmondi)
1229:Brazilian squirrel
987:, January 30, 2015.
951:2014BiCon..23..753S
731:1989RSPSB.238..113O
338:forest regeneration
218:has been linked to
58:'s grey squirrels (
1515:(Sciurus anomalus)
1487:(Sciurus vulgaris)
1471:(Sciurus oculatus)
1469:Peters's squirrel
1437:Japanese squirrel
1423:(Sciurus colliaei)
1421:Collie's squirrel
1295:(Sciurus sanborni)
1261:Bolivian squirrel
1231:(Sciurus aestuans)
771:. 26 January 2016.
556:trade restrictions
552:continental Europe
531:
458:continental Europe
450:energy requirement
262:population density
225:niche partitioning
111:Continental Europe
74:). They eat large
53:
33:
1656:Rodents of Europe
1633:
1632:
1595:Douglas squirrel
1493:Yucatan squirrel
1429:Deppe's squirrel
1389:Allen's squirrel
1369:Abert's squirrel
1351:(Sciurus griseus)
1271:(Sciurus ingrami)
1263:(Sciurus ignitus)
1120:(Dwarf squirrels)
725:(1291): 113–125.
474:squirrelpox virus
446:coniferous forest
318:urban development
302:island ecosystems
241:squirrelpox virus
214:recruitment, and
94:and occasionally
1678:
1588:
1587:
1586:
1585:(Pine squirrels)
1529:
1509:
1431:(Sciurus deppei)
1391:(Sciurus alleni)
1385:
1371:(Sciurus aberti)
1365:
1345:
1317:
1277:Andean squirrel
1225:
1179:
1174:
1169:
1123:
1122:
1121:
1049:
1042:
1035:
1026:
1019:
1016:
1007:
994:
988:
979:George Monbiot,
977:
971:
970:
933:
927:
923:
914:
911:
894:
890:
881:
877:
846:
842:
817:
814:
795:
792:
773:
772:
765:
759:
758:
710:
701:
700:
698:
697:
679:
670:
667:
650:
646:
603:, in south-east
521:Natural predator
377:Predicted spread
322:invasive species
310:Vancouver Island
178:Vancouver Island
128:invasive species
123:Sciurus vulgaris
62:) originated in
1686:
1685:
1681:
1680:
1679:
1677:
1676:
1675:
1636:
1635:
1634:
1629:
1618:
1584:
1583:
1582:
1581:
1571:
1548:
1522:
1502:
1463:(Sciurus niger)
1378:
1358:
1338:
1321:Fiery squirrel
1310:
1218:
1203:
1187:Rheithrosciurus
1180:
1161:
1119:
1118:
1117:
1116:
1106:
1068:
1053:
1023:
1022:
1017:
1010:
995:
991:
978:
974:
935:
934:
930:
924:
917:
912:
897:
891:
884:
878:
849:
843:
820:
815:
798:
793:
776:
767:
766:
762:
712:
711:
704:
695:
693:
681:
680:
673:
668:
653:
647:
634:
629:
544:
523:
502:
470:
454:conifers' cones
436:on the side of
422:mixed broadleaf
384:
379:
306:Garry oak trees
304:. For example,
294:
274:
237:
229:sexual maturity
211:
206:
148:
17:
12:
11:
5:
1684:
1682:
1674:
1673:
1671:Urban wildlife
1668:
1663:
1658:
1653:
1648:
1638:
1637:
1631:
1630:
1623:
1620:
1619:
1617:
1616:
1608:
1600:
1591:
1589:
1573:
1572:
1570:
1569:
1560:
1558:
1555:Syntheosciurus
1550:
1549:
1547:
1546:
1538:
1530:
1519:
1518:
1510:
1499:
1498:
1490:
1482:
1474:
1466:
1458:
1450:
1442:
1434:
1426:
1418:
1410:
1402:
1394:
1386:
1375:
1374:
1366:
1355:
1354:
1346:
1342:Hesperosciurus
1335:
1334:
1326:
1318:
1307:
1306:
1298:
1290:
1282:
1274:
1266:
1258:
1250:
1242:
1234:
1226:
1215:
1213:
1205:
1204:
1202:
1201:
1192:
1190:
1182:
1181:
1164:
1162:
1160:
1159:
1151:
1143:
1135:
1126:
1124:
1108:
1107:
1105:
1104:
1098:
1092:
1086:
1080:
1073:
1070:
1069:
1054:
1052:
1051:
1044:
1037:
1029:
1021:
1020:
1008:
989:
972:
945:(3): 753–774.
928:
915:
895:
882:
847:
818:
796:
774:
760:
702:
671:
651:
631:
630:
628:
625:
621:public opinion
584:broadleaf tree
543:
540:
522:
519:
501:
498:
469:
466:
383:
380:
378:
375:
293:
292:Bark stripping
290:
273:
270:
236:
233:
210:
207:
205:
202:
147:
144:
135:United Kingdom
15:
13:
10:
9:
6:
4:
3:
2:
1683:
1672:
1669:
1667:
1664:
1662:
1659:
1657:
1654:
1652:
1649:
1647:
1644:
1643:
1641:
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21:
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1578:Tamiasciurus
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1314:Hadrosciurus
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1111:
1102:Sciuromorpha
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694:. Retrieved
689:The Guardian
687:
600:
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409:Western Alps
385:
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312:have become
295:
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100:
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54:
47:, native to
44:
41:red squirrel
28:
1059:(subfamily
613:Switzerland
417:Switzerland
334:germination
298:plantations
220:competition
45:S. vulgaris
1640:Categories
1526:Urosciurus
1362:Otosciurus
1100:Suborder:
880:2564-2575.
696:2015-01-31
627:References
617:ecologists
548:ecosystems
510:euthanasia
330:white oaks
314:endangered
185:population
107:introduced
1524:Subgenus
1504:Subgenus
1380:Subgenus
1360:Subgenus
1340:Subgenus
1312:Subgenus
1220:Subgenus
1076:Kingdom:
1061:Sciurinae
1057:Sciuridae
967:254286641
565:poisoning
486:lodgepole
426:deciduous
405:mast-year
254:infection
197:parasites
193:predators
162:Stupinigi
115:predators
96:bird eggs
1626:Category
1096:Rodentia
1090:Mammalia
1084:Chordata
1082:Phylum:
1078:Animalia
1065:Sciurini
985:Guardian
926:167–175.
893:221-234.
845:287-297.
755:27529608
692:. London
580:foraging
572:Warfarin
560:European
506:Piedmont
434:hardwood
392:Piedmont
367:sycamore
326:hoarding
278:songbird
258:poxvirus
235:Poxvirus
82:, buds,
1651:Sciurus
1382:Sciurus
1210:Sciurus
1094:Order:
1088:Class:
947:Bibcode
747:2575748
727:Bibcode
649:351-358
576:Britain
567:by the
462:Eurasia
348:of the
316:due to
174:Trecate
92:insects
90:, some
80:flowers
1067:tribe)
965:
753:
745:
609:Ticino
582:among
478:vector
430:Ticino
413:France
400:Ticino
350:fungus
84:fruits
56:Europe
49:Europe
1506:Tenes
963:S2CID
751:S2CID
605:Italy
589:Genoa
482:Scots
396:Genoa
388:Italy
363:beech
286:Italy
170:Nervi
88:fungi
76:seeds
743:PMID
599:and
533:The
484:and
472:The
460:and
442:Alps
398:and
365:and
955:doi
735:doi
723:238
371:sap
308:on
109:to
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