Thesis (MScFor)--Stellenbosch University, 2014. / ENGLISH ABSTRACT: Several studies have shown that stands of invasive alien plants in the natural forest environment can facilitate the rehabilitation and recovery of such forests and challenged the general and global perception that such invasive species threaten the biodiversity and functioning of natural vegetation systems. The aim of this study was to develop an understanding of the dynamics of the spread and establishment of natural forest species in a large stand of the invasive alien plant species Acacia mearnsii (Black wattle) along the Buffeljagsrivier, Western Cape, South Africa. Several patches of Moist forest, Dry forest and Riparian forest occur along the Buffeljagsrivier, above the Buffeljagsrivier dam. The stand of Black wattle consists of 90 ha for a distance of 3.12 km. The main objective was pursued through four specific objectives: (i) to map and assess the patterns in the distribution, size and species composition of the natural forest clusters within the Black wattle stand; (ii) to determine the relationship between natural forest clusters establishing within the Black wattle stand and the neighboring natural forest patches as potential seed sources for the developing forest clusters; (iii) to determine the subsequent spread of natural forest species from the developing forest clusters into the rest of the Black wattle stand; and (iv) to synthesize the information on the dynamics of the spread and establishment of natural forest species into the Black wattle stand as a basis for developing general guidelines for the conversion of invasive alien plant stands in the forest environment towards regrowing natural forest. In total, 329 clusters of natural forest species were GPS recorded and mapped (Arc-GIS) in three zones (Proximal, Intermediate and Distant in relation to the forest patches) within the Black wattle stand: 266 small clusters (one to three reproductively mature trees), 36 medium sized clusters (four to nine trees) and 27 large clusters (more than 10 trees). Large clusters were abundant in the zone close to the natural forest patches and the number of small clusters increased with increasing distance from the forest patches. A total of 28 species of 20 families were recorded. Natural forest species are therefore able to establish within a Black wattle stand. The relationship between natural forest clusters establishing within the Black wattle stand and the natural forest patches as potential seed sources was studied by sampling the stand composition along transects through the stands. A total of 55 rectangular plots (20 m x 10 m, 200 m2) were sampled across forest patches and forest clusters. Hierarchical clustering analysis, using number of stems of a species per plot, identified three main groups and 10 sub-groups. All the sampled forest clusters were included in four of the five Riparian forest sub-groups. Most Moist and Dry forest species were absent from the forest clusters. The three main forest types differed in their general characteristics and site conditions, and this was supported by the ordination analyses: aspect, slope and canopy closure. The developing forest clusters within the Black wattle stand related more to the Riparian forest in terms of similar very gentle south-westerly slope and mean stem diameter. This suggested Riparian forest to be the primary seed source of the establishing forest clusters within the Black wattle stand. However, the large-sized stems of common species were not significantly different between Riparian forest patches and forest clusters, suggesting that large-sized stems in the forest clusters could be part of remnant forest patches, which could act as local seed sources. Detailed evaluation of species importance values and stem diameter distributions showed that some important Moist and Dry forest species are present in the forest clusters. The conclusion was that every type of forest patch contributes to a greater or lesser degree to the development of forest clusters within the Black wattle stand. Seven large clusters were selected to sample the regeneration of natural forest species within 18 m from the forest cluster boundary. Two species lists were generated; one of species from adjacent natural forest patches, and another from 59 forest clusters of all sizes sampled throughout the wattle stand. The results indicated that (1) Mature trees of well-established forest clusters were the main seed sources for the cluster expansion in all directions; (2) Three different patterns were observed in terms of the distance of expansion of regeneration from the clusters: a decrease in regeneration with increasing distance from the cluster margin; increasing regeneration with increasing distance from the cluster; and no distinct pattern with a lack of regeneration of the dominant species of a forest cluster. The 40 species recorded within the Buffeljagsrivier site include a wide range of fruit and seed characteristics. Four main groups of woody species were identified, based on their presence/absence in forest patches and forest clusters. The presence/absence of most species can be explained in terms of their fruit/seed characteristics and dispersal mechanisms. The majority of recorded woody species were most likely dispersed by birds and mammal, particularly Rameron pigeons and baboons. In conclusion, a conceptual framework was developed to guide the rehabilitation of stands of light demanding invasive stands in the forest environment. Several topics for further research were identified. / AFRIKAANSE OPSOMMING: Verskeie studies het gewys dat opstande van uitheemse indringerplante in die woudomgewing kan die rehabilitasie en herstel van sulke woude fasiliteer, en daag die algemene en globale persepsie uit dat sulke indringerplantspecies die biodiversiteit en funksionering van natuurlike plantegroeisisteme bedreig. Die doel van hierdie studie was om ‘n beter begrip te ontwikkel van die dinamika vir die verspreiding en vestiging van inheemse woudspesies binne ‘n omvangryke opstand van die uitheemse indringerplant, Acacia mearnsii (swartwattel), langs die Buffeljagsrivier, Wes-Kaap, Suid Afrika. Verskeie kolle (‘patches’) van Vogtige, Droë- en Oewerwoud kom langs die Buffeljagsrivier voor. Die swartwattel opstand van ongeveer 90 ha groei langs die Buffeljagsrivier oor ‘n afstand van 3.12 km, tussen die dam en die woudkolle aan die stroom-op kant van die swartwattelopstand. Die hoofdoel is nagevolg deur vier spesifieke doelwitte: (i) om die groepies (‘clusters’) vestigende inheemse houtagtige plantsoorte binne die swartwattelbos te karteer en die patrone in hul verspreiding, grootte en spesies samestelling binne die swartwattelopstand te evalueer; (ii) om die verwantskap tussen natuurlike woudgroepies wat binne die swartwattelopstand vestig en die aangrensende woudkolle as potensiële saadbronne vir die ontwikkelende woudgroepies te bepaal; (iii) om die daaropvolgende verspreiding van inheemse woudspesies vanaf die ontwikkelende woudgroepies binne die res van die swartwattelopstand te bepaal; en (iv) om die inligting oor die dinamika van die verspreiding en vestiging van die woudspesies binne-in die swartwattelopstand saam te vat as ‘n basis vir die ontwikkeling van algemene riglyne vir die omskepping van uitheemse indringerplantopstande in die woudomgewing na hergroeiende inheemse woud. In totaal is 329 groepies van inheemse woudspesies aangeteken (via GPS) en gekarteer (Arc-GIS) in drie sones (nabygeleë, intermediêre en afgeleë) binne die swartwattel opstand: 266 klein groepies (een tot drie voortplantingsvolwasse bome), 36 medium-grootte groepies (vier tot nege bome) en 27 groot groepies (10 of meer bome). Groot groepies was volop in die sone naby aan die woudkolle en die aantal klein groepies het toegeneem met toenemende afstand vanaf die woudkolle. ‘n Totaal van 28 species van 20 families is aangeteken, en spesies soos Canthium inerme, Celtis africana, Gymnosporia buxifolia, Rapanea melanophloeos en Vepris lanceolata was algemeen binne die swartwattelopstand. Dit is daarom moontlik vir inheemse woudspesies om binne die swartwattelopstand te vestig. Die verhouding tussen inheemse woudgroepies wat binne die swartwattelopstand vestig en die woudkolle as potensiële saadbronne is bestudeer deur die opstandsamestelling langs transekte deur die opstande te bemonster. ‘n Totaal van 55 reghoekige persele (20 x 10 m, 200 m2) is opgemeet: nege in Droeëwoudkolle, 17 in Vogtige woudkolle, 20 in Oewerwoudkolle, en nege in die woudgroepies binne the swartwattelopstand. Hiërargiese Groeperingsanalise, gebaseer op aantal stamme van ‘n spesies per plot, het drie hoofgroepe en 10 sub-groepe ge-identifiseer: Vogtige woud met drie sub-groepe, Oewerwoud met vyf sub-groepe, en Droeëwoud met twee sub-groepe. Al die gemete woudgroepies is in vier van die Oewerwoud sub-groepe ingesluit. Die meeste Vogtige en Droeëwoud spesies was afwesig van die woudgroepies. Die hoof woudtipes (Droog, Vogtig, Oewer) verskil in hul algemene eienskappe en groeiplektoestande, en dis ondersteun deur die ordinasie-analises: aspek, helling en kroonsluiting. Die ontwikkelende woudgroepies binne die swartwattelopstand was nouer verbonde met Oewerwoud in terme van soortglyke baie geleidelike suid-westelike helling en gemiddelde stamdeursneë. Dit veronderstel dat Oewerwoud is die primêre saadbron van die vestigende woudgroepies binne die swartwattelopstand. Baie inheemse woudspesies kom egter oor die verskillende groepe en sub-groepe voor, met goeie verjonging oor die gemeenskappe. Die stamdeursneë van goter stamme van algemene spesies was niebeduidend verskillend tussen die Oewerwoudkolle en die woudgroepies, en dit veronderstel dat die groter stamme in die woudgroepies kan deel wees van oorblywende woudkolle, wat as plaaslike saadbronne kan dien. Gedetaileerde ontleding van spesies belangrikheidswaardes en stamdeursneëklasverdelings het getoon dat sommige belangrike Vogtige en Droeëwoud spesies is wel teenwoordig in die woudgroepies. Die gevolgtrekking was dat elke tipe woudkol in a mindere of meerdere mate bydra tot die ontwikkeling van die woudgroepies binne die swartwattelopstand. Sewe groot woudgroepies is geselekteer om die woudverjonging binne 18 m vanaf die woudgroepierand te bemonster. Twee spesieslyste is saamgestel; een van spesies van die aangrensende woudkolle, en een van spesies van 59 woudgroepies van alle groottes wat deur die hele swartwattelopstand aangeteken is. Die resultate het aangetoon dat (1) volwasse bome van goed-gevestigde woudgroepies was die hoof saadbronne vir die uitbreiding van die woudgroepies in all rigtings; (2) drie verskillende patrone is waargeneem in terme van die afstand van uitbreiding van verjonging weg vanaf die woudgroepies: ‘n afname in verjonging met toenemende afstand vanaf die groepierand; toenemende verjonging met toenemende afstand vanaf die woudgroepie; en geen beduidende patroon met ‘n gebrek aan verjonging van die dominante spesies van die woudgroepie. Die 40 spesies wat binne die Buffeljagsrivierstudiegebied aangeteken is het ‘n wye reeks vrug- en saadeienskappe ingesluit. Vier hoofgroepe van houtagtige spesies is geïdentifiseer, gebaseer op hul aan- of afwesigheid in die woudgroepies and woudkolle: spesies algemeen in die woudkolle en teenwoordig in die woudgroepies; spesies spesifiek to bepaalde woudkolle en teenwoordig in die woudgroepies; spesies teenwoordig in die woudkolle maar afwesig van die woudgroepies; en spesies afwesig van die woudkolle maar teenwoordig in die woudgroepies. Die aan- of afwesigheid van die meeste spesies kan verduidelik word in terme van hul vrug/saadeienskappe en verspreidingsmeganismes. Die meerderheid van aangetekende houtagtige spesies was meeswaarskynlik deur voëls en soogdiere versprei, veral Geelbekbosduiwe en Bobbejane. Ten slotte, ‘n konsepsuele raamwerk is ontwikkel om die rehabilitasie van opstande van ligafhanklike indringerspesies in die woudomgewing te rig. Verskeie onderwerpe vir verdere navorsing is geïdentifiseer. / RESUME: Plusieurs études ont montré que les peuplements de plantes exotiques envahissantes dans le milieu de forêt naturelle peuvent favoriser la réhabilitation de ces forêts contestant par-là, la perception générale et globale que ces espèces envahissantes menacent la biodiversité et le fonctionnement des systèmes de végétation naturelle. L’objective de cet étude était de mieux comprendre le dynamisme d’établissement et de propagation dans une forêt naturelle des espèces forestières au sein d’un large peuplement de l'espèce de plantes exotiques envahissantes Acacia mearnsii (acacia noir) le long de Buffeljagsrivier, Western Cape, en Afrique du Sud afin d’élaborer des axes directives de réhabilitation relative aux recouvrements d’espèces invasive. La forêt de Buffeljagsrivier s’étale sur un gradient de petites parcelles de forêt allant de forêt sèche, humide et marécageux. Au milieu de ces parcelles de Buffeljagsrivier, on circonscrit un peuplement d’acacia noir estimé à 90 ha sur une distance de 3,12 km. L’objective principal s’est poursuit à travers quatre sous objectives à savoir: (i) cartographier et évaluer les formes d’ invasions dans la distribution, la taille et la composition des recouvrements des espèces de forêt naturelle à l’intérieur du recouvrement d’acacia noir servant comme preuve possible de l’existence et établissement des espèces de forêt naturelle sous un recouvrement d'une espèces invasive; (ii) de déterminer la relation entre les recouvrements de forêt naturelle s’établissant á l’intérieur du peuplement d’acacia noir et ceux de la forêt naturelle avoisinante humide, sèche et marécageuse pouvant être considéré comme sources potentiel de semences conduisant au développement des premiers recouvrement cités; (iii) de déterminer l’étalement subséquent des espèces de la forêt naturelle à partir des recouvrements d’ acacia noir se développant à l’intérieur du reste du peuplement de l’Acacia noir; (iv) Synthétiser les informations sur la dynamique de la propagation et l'établissement d'espèces forestières naturelles dans l’acacia noir comme une base pour l'élaboration des lignes directrices générales pour la conversion de peuplements de plantes exotiques envahissantes se trouvant dans l'environnement de la forêt vers des repousses des forêts naturelles. Au total, 329 recouvrements d'espèces forestières naturelles ont été GPS enregistrées et cartographiées (Arc -GIS) dans trois zones (Proximale, Intermédiaire et Eloignée par rapport aux parcelles de forêt avoisinantes) dans le peuplement de l'acacia noir: 266 petits recouvrements avaient un à trois arbres matures, 36 recouvrements de taille moyenne avaient quatre à neuf arbres matures et 27 larges recouvrements avaient plus de 10 arbres matures. La visualisation de la carte a révélé que les larges recouvrements de forêt mature étaient abondant dans la zone proche des parcelles de forêt naturelle avoisinantes et que de petits recouvrements de jeunes arbres de forêt naturelles croissaient en nombre à mesure que la distance depuis les parcelles de forêts naturelles adjacentes s’élargissait. Un total de 28 espèces appartenant à 20 familles a été répertorié. Deux familles, Rubiaceae suivit de Anacardiaceae ont dominé l’echantillon. Les espèces forestières naturelles sont donc en mesure de s’établir dans un peuplement d'acacia noir. L’échantillon qui a servi à l’évaluation des sources de semences pour le développement des recouvrements de forêt naturelle à travers l’investigation de la relation entre les recouvrements de forêt naturels qui s’établissent à l’intérieur du peuplement d’acacia noir et les parcelles de forêt avoisinantes humide, sèche et marécageuse était constitué de 55 placettes rectangulaires de 200m2 le long de layons à travers le site d’étude. Une analyse hiérarchique des recouvrements réalisée sur la composition des espèces de l’échantillon a révèle que la forêt marécageuse concentre les espèces les plus fréquentes des recouvrements de forêts, et que la majorité des espèces présentes sur les parcelles des forêts sèches et humides étaient absente dans l’ensemble des recouvrements des forêts de l’acaci noir. Les trois principaux types de forêts différaient dans leurs caractéristiques générales et les conditions du site, et cela a été appuyé par l’analyse d’ordinants à partir de trois facteurs environnementaux, la pente, l’angle d’inclinaison par rapport au soleil, et la fermeture de la canopée. Les recouvrements de foret en développement au sein de l'acacia noir se sont trouvés davantage liés à la forêt marécageuse en termes de pente plate orientée sud-ouest et en diamètre moyen des arbres. Ces résultats ont désigné la forêt marécageuse comme la première source potentielle de semences permettant l’établissement des recouvrements de forêt à l’intérieur des peuplements d’acacia noir. Cependant, les souches d’arbres de grande taille des espèces communes entre les parcelles de forêts marécageuses et les recouvrements de forêt dans l’acaci noir n'étaient pas significativement différents, suggérant ainsi que ces arbres de grande taille trouves dans les recouvrements de foret pourraient faire parties des parcelles de forêt subsistantes de la dernière turbulence. Elles pourraient constituer des sources locales de semences. Par ailleurs, une évaluation détaillée des valeurs d'importance des espèces et des distributions des diamètres des souches d’arbres a montré que certaines espèces importantes de forêts humides et sèches sont présentes dans les recouvrements de foret. Finalement l’on conclue que chaque type de parcelles de forêt contribue plus ou moins à l’établissement des recouvrements de forêt dans le peuplement de l'acacia noir. Sept grands recouvrements de foret ont été sélectionnés pour étudier la régénération des espèces forestières naturelles sur un rayon de 18 m depuis la limite du recouvrement de foret. Deux listes d'espèces ont été générées, l'une des espèces de parcelles de forêt naturelle adjacentes, et une autre venant des 59 recouvrements de foret, toutes tailles confondues le long de l'acacia noir. Les résultats ont indiqué que (1) Les recouvrements de forêts, lorsque bien matures et bien établies devenaient les principales sources de semences pour leur expansion sur toutes les directions; (2) Trois allures différentes ont été observées en fonction de la distance de l'expansion de la régénération autour des recouvrements: une diminution de la régénération avec l'augmentation de la distance; une croissance de la régénération avec une distance croissante depuis les revcouvrements de foret, et pas d’allure particulaire voire une absence de régénération de certaines especes pourtant dominant dans les recouvrements de foret. Les 40 espèces recensées sur le site deBuffeljagsrivier comprennent un large éventail de caractéristiques de fruits et de graines. Quatre principaux groupes d'espèces ligneuses ont été identifiées, en fonction de leur présence / absence dans des parcelles de forêt et les recouvrements de forêt. De façon générale, la présence / absence de la plupart des espèces a pu être expliquée en termes de caractéristiques de leur fruits/graines et les mécanismes de dispersion. La majorité des espèces ligneuses enregistrées sont susceptibles d’être dispersées par les oiseaux et les mammifères, en particulier les pigeons rameron et les babouins vues sur le terrain. En conclusion, un cadre conceptuel a été élaboré pour guider la réhabilitation de peuplements envahissants de lumière en milieu forestier. Plusieurs sujets pour des recherches plus poussées ont été identifiés.

A abertura de clareiras no dossel das florestas ocorre periodicamente pela queda ou morte de árvores, acarretando a formação de uma paisagem em mosaico de áreas com diferentes idades, condições ambientais e composição florística. A principal modificação que ocorre no ambiente de clareira é o aumento da intensidade de luz, interferindo no estabelecimento, crescimento e sobrevivência de muitos indivíduos. O objetivo desse estudo foi caracterizar a estrutura de habitat e descrever a composição florística de espécies arbustivo-arbóreas de 0,50 a 2m nas clareiras naturais em Floresta de Restinga na parcela permanente localizada no Parque Estadual da Ilha do Cardoso e as acompanhar as modificações na comunidade após um ano. Durante um ano os indivíduos foram acompanhados em relação à sobrevivência, o crescimento e o recrutamento. As espécies foram classificadas em categorias sucessionais com base na literatura. Foram analisadas onze clareiras com áreas entre 30,5 e 398,4m2, das quais 6 foram consideradas pequenas (<150m2) e 5 grandes (>150m2). Foram analisados 1.316 indivíduos pertencentes a 99 espécies arbustivo-arbóreas. O fator determinante do número de espécies e indivíduos encontrados nas clareiras foi o tamanho da área atingida pela abertura do dossel. As espécies mais abundantes foram: Euterpe edulis, Rudgea villiflora, Miconia sp1, Myrcia bicarinata, Dendropanax cuneatum e Astrocaryum aculeatissimum. Apenas Euterpe edulis foi encontrado em todas as clareiras. Do total de plantas amostradas nas clareiras, 42 (3,2%) foram consideradas pioneiras distribuídas em 7 espécies, 579 (46,1%) secundárias iniciais distribuídas em 26 espécies, 419 (33,4%) climáxicas distribuídas em 40 espécies e 215 (17,1%) não classificadas pertencentes a 26 famílias. Nas clareiras consideradas pequenas houve destaque no número de indivíduos pertencentes à categoria clímax (46,3%), enquanto nas clareiras grandes houve predominância das secundárias inicias (52%). Houve diferença na abundância de algumas espécies em relação ao tamanho das clareiras. Nas clareiras pequenas Myrcia bicarinata foi a espécie com maior densidade, já nas clareiras grandes Euterpe edulis foi a espécie de maior densidade. Na remedição dos indivíduos observou-se que apenas 671 indivíduos apresentaram crescimento, os outros indivíduos não apresentaram crescimento, principalmente devido à alta taxa de herbivoria. Euterpe edulis foi a única espécie que apresentou diferença nas taxas de crescimento entre as duas classes de tamanho de clareira. Após um ano foi observada a mortalidade de 84 indivíduos e o recrutamento de 105 indivíduos. Os indivíduos ingressantes pertencem, principalmente, as categorias secundárias iniciais (27%) e clímax (22%). Não há indícios de uma convergência das comunidade presentes nas diferentes clareiras em direção à uma comunidade que fosse típica ou exclusiva de clareira, ao contrário, mostraram-se inicialmente distintas e permaneceram distintas após um ano. / Canopy gap openings periodically occurs when a tree dies or falls, creating a mosaic landscape which result into patches with different ages, environmental conditions and floristic composition. Light enhancement is the main modification within a gap environment, causing modifications on the establishment, development and survival of many individuals. The purpose of this study was to describe shrub and tree species? composition from 0,5 to 2cm height, point out habitat structure and follow community modifications within a year under light gaps in a Restinga Forest, located in a permanent plot at Parque Estadual da Ilha do Cardoso. For an entire year these individuals were studied regarding their survival, growth and recruitment. Species were classified into sucessional categories as suggested by literature. Eleven gaps were analyzed and their total area ranged from 30,5 to 398,4m2, with 6 gaps considered as small (<150m2) and 5 as large (>150m2). The overall number of individuals analyzed in this study was of 1316, representing 99 tree and shrub species. The main factor acting on the species and individuals quantity within a gap was the area size affected by the canopy opening. The most abundant species were: Euterpe edulis, Rudgea villiflora, Miconia sp1, Myrcia bicarinata, Dendropanax cuneatum e Astrocaryum aculeatissimum. The only species sampled in every gap was Euterpe edulis. Regarding sucessional categories, 42 individuals (3,2%) of 7 species were classified as pioneers, 579 individuals (46,1%) of 26 species as early secondary species, 419 individuals (33,4%) of 40 species as climax and 215 individuals remained non-classified (17,1%), representing 26 botanical families. Small gaps presented a remarkable quantity of climax species (46,3%), while on the larger ones early secondary species prevailed (52%). Species abundance varied according to gap size. Myrcia bicarinata showed the highest densities within smaller gaps, while in the larger ones the dominant species was Euterpe edulis. Height measurements indicated growth for only 671 individuals, mainly because of herbivory attacks. Euterpe edulis was the only species with significant growth rate differences between small and large gaps. After a year, 84 individuals were dead, while 105 recruits were accounted. The latter ones were mainly represented by early secondary species (27%) and climax (22%). The communities present in the different sized gaps did not bring any evidence of convergence towards a typical or exclusive gap community. On the contrary, these communities appeared to be dissimilar either on initial phase or after a year.

Através da amostragem de 28 sítios em diferentes estádios de regeneração em uma área de Mata Atlântica contínua, procuramos nesta dissertação contribuir para o entendimento dos mecanismos relacionados às mudanças faunísticas observadas durante o processo de regeneração em florestas tropicais, e, assim, melhor compreender o valor das florestas secundárias para a conservação da biodiversidade tropical. Para isso, na primeira parte da dissertação investigamos a influência da regeneração sobre características de estrutura da floresta e disponibilidade de alimento consideradas importantes para diversos grupos da fauna. Encontramos maior profundidade do folhiço, volume de galhadas e disponibilidade de frutos de uma abundante palmeira de sub-bosque nas florestas mais maduras, enquanto que nas áreas em estádio mais inicial de regeneração encontramos maior conexão da vegetação, biomassa de artrópodes no solo e disponibilidade de frutos no total e da espécie de planta mais abundante no sub-bosque. Essas modificações podem estar relacionadas às mudanças na fauna observadas durante a regeneração florestal. As espécies que dependem de características só encontradas nas matas mais maduras, como espaços abertos para movimentação, maior complexidade do chão da floresta, ou determinados recursos alimentares poderiam encontrar limitações em áreas em estádios mais iniciais de regeneração, ao passo que espécies que não dependem desses recursos poderiam se beneficiar da maior disponibilidade total de alimentos, ou da maior conexão da vegetação para movimentação, encontradas nessas áreas. Na segunda parte do trabalho, avaliamos a influência da regeneração e de características estruturais e de disponibilidade de alimento sobre espécies endêmicas (especialistas de floresta) e não-endêmicas (generalistas de habitat) de pequenos mamíferos terrestres. Como previsto, observamos que um grupo de espécies (generalistas de habitat) prolifera nas áreas mais jovens, enquanto que o outro grupo (especialistas de floresta) foi mais comum, embora de maneira mais sutil, nas áreas mais maduras. Esses padrões foram em parte explicados pelas variáveis mensuradas, principalmente a disponibilidade de recursos alimentares. Aparentemente, as espécies generalistas estão se beneficiando da maior disponibilidade de alimentos encontrada nas áreas em estádio mais inicial de regeneração, enquanto que as especialistas possuem maior capacidade de ocupação das áreas mais maduras, onde esses recursos são mais escassos. Portanto, esses resultados sugerem um compromisso (trade-off) entre capacidade competitiva e capacidade de utilização de recursos 101 abundantes, como prevê o mecanismo de nicho sucessional, proposto inicialmente para explicar a sucessão de espécies vegetais. As características encontradas nas florestas mais jovens favoreceram a proliferação de espécies de pequenos mamíferos terrestres generalistas de habitat. Entretanto, o efeito positivo da regeneração florestal sobre as espécies especialistas, de maior interesse para a conservação, foi menos acentuado, de forma que as florestas secundárias abrigaram uma assembléia de pequenos mamíferos terrestres rica, podendo, portanto, representar um importante instrumento de aumento de área e conectividade em paisagens altamente modificadas, como as encontradas na Mata Atlântica. Entretanto, esses resultados não reduzem o valor das florestas maduras, principalmente em paisagens fragmentadas e para grupos mais sensíveis da fauna. Essas florestas devem ser protegidas, assim como deve ser garantida (e se preciso auxiliada) a regeneração das florestas secundárias, para que possam adquirir em longo prazo as condições necessárias à manutenção das espécies e dos grupos de espécies da fauna mais severamente afetados pela secundarização das florestas tropicais. / By sampling 28 sites in different regeneration stages in a continuous Atlantic forest area, in this master thesis we aimed to contribute to the understanding of the mechanisms associated with the wildlife changes observed during tropical forest regeneration, and thereby better understand the value of secondary forests for the conservation of tropical biodiversity. In the first part of the thesis we investigated the influence of regeneration on aspects of forest structure and food availability considered to be important to several wildlife groups. We found deeper leaf litter, and higher woody debris volume and fruit availability of an abundant understorey palm in the older-growth areas, while in areas in earlier stages we found higher vegetation connection, higher ground-dwelling arthropod biomass, total fruit availability, and availability of fruits of the most abundant understorey plant species. These modifications may be related to the wildlife changes observed during forest regeneration. Species that rely on features only found in older-growth forests, such as open space for movement, higher complexity of the forest floor, or certain food resources, could find limitations in younger forests, whereas species which do not depend on these resources could benefit from the higher total food availability, or higher 102 vegetation connection for movements, found in these areas. In the second part of the study, we assessed the effect of regeneration and of structural attributes and food availability on endemic (forest specialist) and non-endemic (habitat generalist) terrestrial small mammal species. As expected, we found that a group of species (habitat generalists) proliferate in younger areas, while the other group (forest specialists) is more common, although more subtly, in older-growth areas. These patterns were partly explained by the measured variables, especially food availability. Apparently, the generalist species are benefiting from the increased food availability found in younger forests, while specialist species have greater ability to occupy older-growth areas, where these resources are scarcer. Therefore, our data suggest a trade-off between competitive ability and ability to use abundant resources, as predicted by the successional niche mechanism, initially proposed to explain the succession of plant species. The characteristics observed in younger forests favored the proliferation of habitat generalist terrestrial small mammals. However, the positive effect of forest regeneration on specialist species, of more conservation concern, was less pronounced, so that the secondary forests harbored a rich terrestrial small mammal assemblage, and may therefore be an important tool to increase the area and connectivity in highly modified landscapes, such as those found in the Atlantic forest. However, these results do not diminish the value of old-growth forests, especially in fragmented landscapes and for more sensitive wildlife groups. These forests should be protected, as well as the regeneration of the secondary forests should be guaranteed (and assisted, if necessary), so that these areas could acquire in the long term the necessary conditions to maintain the species and the groups of species most adversely affected by the secondarization of the tropical forests.

As communities and managers become aware of the long-term impacts of mangrove loss, estimated at 1-2% per year, interest in sediment erosion and mangrove rehabilitation has increased substantially. In this thesis project I 1) examine erosion rates within coastal fringing Rhizophora mangle ecosystems following mangrove clearing and compare these rates to accretion rates in intact mangroves; and 2) investigate the abiotic factors influencing mangrove seedling survival and regeneration of naturally colonizing R. mangle, in historic mangrove habitat after anthropogenic clearing. Differences in erosion were compared between patches of open-coast intact and anthropogenically cleared R. mangle to quantify the sediment trapping function provided by mangroves and its loss following clearing over a 24 month period. Growth rates of mangrove seedlings in intact forest were compared to seedlings in cleared areas. Seedling growth indicators were measured on 100 seedlings at five sites (50 in the intact and 50 in the cleared areas). To examine the limiting factors on seedling growth rates, nutrient addition and wave protection treatments were applied to seedlings in three disturbed areas. Sites within intact mangroves had sediment accretion (M= +3.83 mm) while areas cleared of mangroves had sediment erosion (M= -7.30 mm). Seedling growth (height) over the 2 year study period significantly differed between intact mangrove (M = 15.6 cm) and cleared (M = 10.24 cm) areas. Seedling mortality from the cleared areas (31%) differed from the intact areas (13%). Average seedling growth (height) was: greater with both nutrient/wave (M = 18.4 cm) and nutrient (M = 17.65 cm) treatments compared to controls (M = 10.8 cm), which suggests that providing nutrients and/or wave protection result in growth outputs comparable to seedlings found in intact mangroves. This study may prove to be useful in identifying areas that are most vulnerable to erosion following mangrove removal and ideal location of restoration following mangrove removal. Areas cleared of mangroves can lead to intensified erosion in areas where fringing reefs are not continuous. When managers are determining areas to focus resources for restoration, focusing on areas with nutrient rich habitat may result in higher survival rates and growth outputs.

Bare-root, container, and root production method (RPM™) seedlings of two oak species (Nuttall (Quercus texana Buckley), cherrybark (Q. pagoda Ell.)) were planted on lands damaged by Hurricane Katrina in southern Mississippi to compare the height growth, groundline diameter growth and survival of the different planting stocks. Tree shelters were applied to half of the bare-root seedlings to determine their effect on the height and groundline diameter growth and survival of the seedlings. RPM seedlings exhibited significantly greater height and groundline diameter growth than bare-root or container seedlings after one growing season. Bare-root seedlings exhibited significantly greater height and groundline diameter growth than container seedlings. Tree shelters significantly increased height growth of bare-root seedlings; however, sheltered bare-root seedlings exhibited significantly less groundline diameter growth than non-sheltered seedlings. Cherrybark oak exhibited greater height growth than Nuttall oak, while Nuttall oak exhibited greater groundline diameter growth than cherrybark across all planting stocks.

No description available.

Os ecossistemas de ilhas, continentais ou oceânicas, são considerados os mais sensíveis à perturbação humana. A maioria das ilhas costeiras do Brasil sofreu fortes alterações na fauna e flora silvestre. A Ilha Anchieta (Ubatuba, SP) é uma Área Protegida com um longo histórico de perturbação, tendo sido amplamente desmatada no passado e sofrido a introdução de animais. Sabe-se que a fauna possui um papel chave na composição e estrutura da comunidade vegetal, favorecendo algumas espécies e prejudicando outras. Sendo assim, a perda ou alteração dos processos de interações entre animais e plantas afetam na estrutura e composição de espécies. Este estudo analisa as interações antagônicas entre planta e animais como possíveis fatores limitantes no processo de regeneração natural em três ambientes com distintos estágios sucessionais presentes na Ilha Anchieta: campo aberto (CA) e florestas ombrófilas rala (FOR) e densa (FOD). Para isso foram testados nos três ambientes os processos pós-dispersão de sementes: a predação de sementes pós-dispersas, a germinação do banco de sementes e a herbivoria. A predação de sementes apresentou forte efeito espacial (FOD>FOR>CA) e sazonal, com maiores proporções de predação de sementes no período de escassez de alimento (estação seca). Quantitativamente o banco de sementes não representa um fator limitante, porém foi composto por uma baixa diversidade de espécies lenhosas. A capacidade de germinação do banco de sementes do solo foi similar entre os ambientes florestais apresentando menor emersão de plântulas no campo aberto, provavelmente associado ao intenso escoamento superficial no local. Adicionalmente, a mortalidade de juvenis transplantados foi de 72,27%, sendo superior nas parcelas abertas aos vertebrados para todas as espécies em todos os ambientes, demonstrando um forte efeito negativo dos herbívoros vertebrados na comunidade vegetal. Sendo assim, foram detectados distintos gargalos atuando simultaneamente na regeneração natural nos três ambientes da Ilha Anchieta, sendo necessário o estabelecimento de práticas de manejo visando minimizar os efeitos abióticos no campo aberto e floresta ombrófila rala, favorecendo as interações mutualísticas e inibindo a atividade de animais antagônicos nas florestas ombrófilas rala e densa, acelerando assim o processo de regeneração natural na Ilha Anchieta. Considerar simultaneamente os obstáculos no processo de regeneração nos auxiliará a traçar práticas de restauração e recuperação de áreas degradadas mais efetivas e viáveis economicamente. / Island ecosystems, either continental or oceanic, are considered the most sensible to anthropogenic influences. Most Brazilian coastal islands have their original fauna and flora composition altered. The Anchieta Island (southeast Brazil) is a Protected Area that suffered deforestation and introduction of alien species. It is known that the fauna plays a key role in composing and structuring the vegetal community, favoring some species and harming others. Thus, the loss or change of animal-plant interactions affects the framework and composition of species. We simultaneously analyzed the antagonistic animal-plant interactions as possible limiting factors in the natural regeneration on tree successional stages of the Anchieta Island: the old fields (OF), the early secondary forest (ESF) and old growth forest (OGF). Therefore, in each habitat we are evaluating post-dispersal seed predation processes: post-dispersal seed predation, soil seed bank, and the herbivory. We found spatial (OGF>ESF>OF) and temporal effects on seed predation, with highest rates on dry season. Quantitatively, soil seed bank did not represent a limiting factor; but qualitatively, it was composed by few woody species. Our data pointed to the absence of a viable soil seed bank in the OF, probably due to an intense runoff. The forested habitats presented similar soil bank. Moreover, the total mortality of saplings was 72.27%, being higher on the unfenced treatment for all species and in all habitats, showing a very strong negative effect of vertebrate herbivores on the vegetal community. Therefore, we are detecting distinct bottlenecks acting simultaneously in the natural regeneration process in all of the successional stages of the Anchieta Island. Our results showed that management actions are required, aiming to both minimize the abiotic effects on the old fields and in the early secondary forest and to favor the mutual interactions and inhibit the activity of antagonic animals in the old growth and early secondary forests, accelerating, this way, the natural regeneration process of the Anchieta Island. Considering simultaneously the obstacles in the regeneration process will help to define restoration and recuperation procedures of degraded areas more effective and affordable.

What strategies really work against the intense loss of native habitat? How can we improve forested areas around the world? The creation of protected areas - natural reserves aiming at preserving biodiversity and ecological relations in pristine environments - has been the main strategy against land surface transformation and the loss of natural capital. Recently, financial incentives have promptly become the most promising and accepted new strategies for conservation, and several experiences were implemented, particularly in the tropics. However, we urge to systematically evaluate the effectiveness of these mechanisms, both old and new, and several researchers have argued that econometric methods, traditionally used in impact evaluation of development, health, and education policies, should be applied in assessing the impacts of interventions for environmental conservation. Our goal in the present study is to estimate the effectiveness of two popular financial incentives against native vegetation loss in Brazil: payments for ecosystem services (PES) and the intergovernmental fiscal transfer based on ecological criteria - the Ecological ICMS. We used a counterfactual approach, which seeks adequate control groups in order to estimate what would have happened in the absence of the intervention - i.e., we search for additional gain or additionality. Payment for ecosystem services programs were evaluated regarding the promotion of additional native forest areas while the Ecological ICMS was assessed for its ability to promote new legally protected areas for conservation. We evaluated two of the oldest and most well established payment for ecosystem services programs in the Atlantic Forest, located in the Cantareira Water System, near the metropolis of São Paulo: the \\textit{Produtor de Água} (Water Producer) in Joanópolis and Nazaré Paulista (SP), and the \\textit{Conservador das Águas} (Water Conservationist), in Extrema (MG). In total, 83 rural properties contracted by the programs, and 83 properties selected as control units, were mapped using satellite images available in Google since 2003 and evaluated for the presence of regeneration and deforestation in three moments: before the program, at the time of program implementation and after program implementation. We show that PES programs positively affect the area under regeneration inside contracted rural properties, but does not affect deforestation. The average effect on regeneration was 0.69\\% of property area per year, which means that in 0.69\\% of the evaluated property area there would be no regeneration of native vegetation were it not for PES programs intervention. This result shows the additional effect of PES programs on regeneration of native forest but also shed light that this is a slow process. Secondly, the effect of the Ecological ICMS mechanism was evaluated regarding the expansion of Conservation Units - protected areas for conservation. We compared 2,481 municipalities from eight states in the Atlantic Forest region, in a panel data (i.e., longitudinal database) covering the period from 1987 to 2016. We show that municipalities under the incentive of the ecological ICMS law, when compared to control municipalities not subject to the same mechanism, present a small increase in area of new conservation units in the first years of law existence. However, this effect decreases over time disappearing about 10 years after the approval of the law. In addition to the temporary effect, we show that protected areas created, particularly by local governments, are preferably Environmental Protection Areas (APA) - a category that imposes few restrictions on land use and promotes little contribution to the conservation of natural resources. In order to estimate the effect of the Ecological ICMS on regeneration and deforestation processes, we accessed a national database on land use change between 1985 and 2017. However, observing deforestation patterns throughout this historical data series, it called our attention the possible existence of electoral cycles affecting deforestation - which gave rise to the third chapter of the thesis. We analyze deforestation (percentage deforested over previous forested area) in a panel with 2,277 municipalities within the Atlantic Forest region, from seven states of the South and Southeast, considering the period from 1991 to 2014. We demonstrate the existence of electoral deforestation cycles, linked to state elections in five observed states and municipal elections in three states. We also show evidence that elections in which the incumbent is losing by a large margin of votes tend to be the events with greater observed deforestation, and that party alignment between mayor and governor parties can also contribute to an increase in deforestation during municipal elections. Financial mechanisms, at least in the way they have been designed to date, are limited as major instruments in achieving the goals of conservation and restoration of native forests. In addition, eventhough there is a major effort to create new mechanisms and increase conservation gains, it is of concern that these gains can be offset by losses due to political motivations, even in the region of the country with one of the most stringent legislations for tropical forests in the world / Quais estratégias realmente funcionam contra a intensa perda de habitat nativo? Como podemos aumentar nossas áreas de florestas? A criação de áreas protegidas - reservas naturais com o objetivo de preservar a biodiversidade e as relações ecológicas de ambientes pristinos - tem sido a principal estratégia contra as transformações na superfície da Terra e a perda de habitat nativo. Recentemente, incentivos financeiros se tornaram rapidamente promissores e populares instrumentos para a conservação ambiental, e diversas experiências foram implementadas, particularmente nas florestas tropicais. No entanto, carecemos de avaliações sistemáticas sobre a efetividade destes mecanismos, antigos e novos, e diversos pesquisadores argumentam que métodos da econometria, tradicionalmente usados na avaliação de políticas de desenvolvimento, saúde e educação, devem ser aplicados na avaliação dos impactos das intervenções para a conservação ambiental. O objetivo do presente trabalho foi estimar a efetividade de dois dos mais populares incentivos financeiros no combate à perda de vegetação nativa no Brasil: os pagamentos por serviços ambientais (PSA) e a transferência fiscal intergovernamental com base em critérios ecológicos - o ICMS Ecológico, concebido e implementado no Brasil de forma inédita. Para estas avaliações, nos baseamos na abordagem contrafactual, que busca adequados grupos de controle com o objetivo de estimar o que haveria ocorrido na ausência da intervenção estudada, isto é, se houve ganho adicional ou adicionalidade. Os programas de pagamento por serviços ambientais foram avaliados quanto a promoção de áreas adicionais de floresta nativa enquanto que o ICMS Ecológico foi avaliado quanto a sua capacidade de promover novas áreas legalmente destinadas à conservação de recursos naturais - as Unidades de Conservação. Avaliamos dois dos mais antigos e bem estabelecidos programas de pagamento por serviços ambientais na Mata Atlântica, localizados no Sistema Cantareira, ao redor da região metropolitana de São Paulo: o Produtor de Água, em Joanópolis e Nazaré Paulista (SP), e o Conservador das Águas, em Extrema (MG). No total, 83 propriedades rurais contratadas pelos programas, e 83 propriedades selecionadas como controle, foram acompanhadas por imagens de satélite disponíveis no Google desde 2003 e avaliadas quanto a presença de regeneração e desmatamento, em três momentos: antes dos programas, no momento da implementação e após um período de operação dos programas. Mostramos que os programas de PSA afetam positivamente a área em regeneração dentro das propriedades rurais contratadas, mas não afeta o desmatamento. O efeito médio sobre a regeneração foi de 0.69\\% da área da propriedade ao ano, o que significa que em 0.69\\% da área das propriedades avaliadas não haveria regeneração de vegetação nativa não fosse a intervenção dos programas avaliados. Esse resultado mostra o efeito adicional dos programas de PSA sobre o processo de regeneração de floresta nativa mas nos revela também que se trata de um processo lento. Na segunda etapa da pesquisa, foi estimado o efeito do mecanismo do ICMS Ecológico sobre a expansão de Unidades de Conservação. Comparamos 2.481 municípios de oito estados na região da Mata Atlântica, em um painel (i.e. base de dados longitudinal) compreendendo o período de 1987 a 2016. Demonstramos que municípios sob o incentivo da lei do ICMS Ecológico, quando comparados com municípios controle não sujeitos ao mesmo mecanismo, apresentam um pequeno aumento em área de novas unidades de conservação nos primeiros anos de existência da lei. No entanto, este efeito decresce ao longo do tempo desaparecendo cerca de 10 anos após a aprovação da lei. Além do efeito temporário, mostramos que as áreas protegidas criadas, particularmente pelos governos locais, são preferencialmente Áreas de Proteção Ambiental (APA) - categoria que impõe poucas restrições ao uso da terra e que traz pequena contribuição à conservação dos recursos naturais. Com o intuito de estimar o efeito do ICMS ecológico sobre processos de regeneração e desmatamento, acessamos uma base de dados nacional sobre mudança no uso da terra no período entre 1985 e 2017. Observando os padrões de desmatamento ao longo desta série histórica, nos chamou a atenção a possível presença de ciclos eleitorais afetando o desmatamento - o que deu origem ao terceiro capítulo da tese. Analisamos o desmatamento (percentual desmatado sobre a área prévia de floresta) em um painel com 2.277 municípios na região da Mata Atlântica, dos sete estados do Sul e Sudeste, considerando o período de 1991 a 2014. Nós demonstramos a existência de ciclos eleitorais de desmatamento, ligados às eleições estaduais, em cinco dos estados observados, e municipais, em três estados. Também mostramos evidências de que as eleições nas quais o incumbente está perdendo por uma ampla margem de votos tendem a ser os eventos com maior desmatamento observado, e que o alinhamento entre o partido do prefeito e do governador também pode contribuir para o aumento do desmatamento durante as eleições municipais. Os mecanismos financeiros, ao menos na forma que foram concebidos até o momento, se mostram bastante limitados como instrumentos majoritários no alcance das metas de conservação e restauração de florestas nativas. Além disso, se existe um grande esforço para que novos mecanismos sejam criados e que se possa ampliar os ganhos em conservação, é preocupante que estes ganhos possam ser compensados por perdas decorrentes de motivações políticas, mesmo na região do país na qual vigora uma das legislações mais rígidas sobre florestas tropicais do mundo

Old-growth stands are important for management, conservation, wildlife, recreation, and maintaining biological diversity in forested landscapes. However, we are lacking the information needed to adequately identify and characterize old-growth stands. This is especially true for high elevation, interior forests. The characterization of stand structure and regeneration pattern will help in the development of site-specific guidelines for identifying old growth stands and restoring some of the old-growth characteristics in managed stands. This pamphlet presents a synopsis of a study investigating stand structure and regeneration of old-growth stands in the Moist Cold Engelmann Spruce - Subalpine Fir (ESSFmc) Subzone near Smithers, B.C. The three stands selected for the study were located on zonal sites, each in different watersheds, and the stands were established after fire. The criteria used for selection were: i) absence of lodgepole pine, ii) presence of advanced regeneration, and iii) abundant snags and coarse woody debris. These stands were presumed to represent the old-growth stage of stand development or the final (climax) stage of secondary succession.

Made available in DSpace on 2016-06-02T19:29:51Z (GMT). No. of bitstreams: 1 TeseMS.pdf: 839878 bytes, checksum: ce4d6d134c72bc480b1a92a8bc4ca303 (MD5) Previous issue date: 2003-05-27 / The present study evolved floristic composition, structure and regeneration dynamic aspects of a restinga forest (pluvial sand coastal plain forest), at Ilha do Cardoso, in Cananéia region, São Paulo State, Brazil. Three portions of restinga forest with same physiognomic aspects were selected: restinga of Trilha para o Morro das Almas (MA), Trilha Interpretativa (TI) and Estrada para a Captação (EC). Five strata of forest were sampled in each portion: Tree, Tree/shrub, Shrub/herbaceous, Herbaceous and Lianas. Each portion was analyzed individually and subsequently were analyzed the entirely of the three portions and compared them. At Trilha para o Morro das Almas were sampled 73 species, at Trilha Interpretativa 72, at Estrada para a Captação 56 and totally sample 112, inside them 31 species were common to three portions and 46 restrict only one of them. The three portions of a forest were floristically similar, MA and TI had the highest Sørensen similarity (70,3%), and TI and EC had the less Sørensen similarity (55,5%). When compared the strata in general Tree/shrub and Shrub/herbaceous strata had the highest value of Sørensen similarity, and the Tree and Herbaceous strata had the less similarity. The highest diversity and equability were found in herbaceous strata, and the less in tree strata. The plants of the three portions had majority of species with geographic distribution in south and southeast Brazil regions, and the majority individuals and species were secondary-climax category. The portions of studied forest were in advanced stage of regeneration, and shade-tolerant species were favored. The differences between three portions of a forest were due to natural process common in tropical forests, where the substitution of one tree produce a mosaic of distinct regenerative phases at community that differ in structure and floristic composition. Differences in environment physical conditions, especially edaphic factors, and distinct anthropic interventions are responsible too. / O presente estudo envolveu a composição florística, a estrutura e aspectos da dinâmica de regeneração de floresta de restinga (floresta pluvial da planície arenosa costeira), na Ilha do Cardoso, Cananéia, Estado de São Paulo, Brasil. Foram selecionadas três áreas com fisionomias semelhantes: restinga da Trilha para o Morro das Almas (MA), da Trilha Interpretativa (TI) e da Estrada para a Captação (EC). Foram amostrados cinco estratos em cada uma delas: Arbóreo, Arbóreo/arbustivo, Arbustivo/herbáceo, Herbáceo e Lianas. Cada uma das áreas foi analisada individualmente e posteriormente fez-se uma análise conjunta comparando as três áreas entre si. Na Trilha para o Morro das Almas foram amostradas 73 espécies, na Trilha Interpretativa 72, na Estrada para a Captação 56 e na amostragem total 112, sendo que 31 espécies foram comuns às três áreas e 46 foram restritas à apenas uma delas. Pelo índice de Sørensen as três áreas são floristicamente similares entre si, sendo que MA e TI apresentaram a maior similaridade (70,3%) e TI e EC a menor similaridade (55,5%). Na comparação entre os estratos de modo geral entre o arbóreo/arbustivo e o Arbustivo/herbáceo houve maior similaridade de Sørensen e, entre o Arbóreo e o Herbáceo menor. A maior diversidade florística e equabilidade couberam ao estrato herbáceo e as menores ao arbóreo. Nas três áreas amostradas a maioria das espécies apresentaram padrão de distribuição geográfica sul e sudeste do Brasil e, a maioria dos indivíduos e das espécies pertencia à categoria sucessional secundária tardia. Os trechos de floresta estudados estão em estágio avançado da sucessão e as espécies tolerantes à sombra estão sendo favorecidas. As diferenças entre os três trechos da floresta amostrados devem-se a processos naturais comuns nas florestas tropicais, onde a substituição de árvores produz um mosaico de distintas fases regenerativas na comunidade que diferem na estrutura e na composição floristica. As diferenças nas condições físicas ambientais, especialmente fatores edáficos e, as intervenções antrópicas diferenciadas também são responsáveis por essas diferenças.