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Universidade Estadual Do Maranhão Centro De Ciências Agrárias Programa De Pós-Graduação Em Agroecologia Curso De Doutorado Em Agroecologia

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Universidade Estadual Do Maranhão Centro De Ciências Agrárias Programa De Pós-Graduação Em Agroecologia Curso De Doutorado Em Agroecologia UNIVERSIDADE ESTADUAL DO MARANHÃO CENTRO DE CIÊNCIAS AGRÁRIAS PROGRAMA DE PÓS-GRADUAÇÃO EM AGROECOLOGIA CURSO DE DOUTORADO EM AGROECOLOGIA LUIS MANUEL HERNÁNDEZ GARCÍA -DIVERSIDADE DAS COMUNIDADES DE OLIGOQUETAS DA ÁREA DE ENDEMISMO DE BELÉM. São Luís, MA. 2019 LUIS MANUEL HERNÁNDEZ GARCÍA Biólogo -DIVERSIDADE DAS COMUNIDADES DE OLIGOQUETAS DA ÁREA DE ENDEMISMO DE BELÉM. Tese apresentada ao Curso de Doutorado do Programa de Pós-Graduação em Agroecologia da Universidade Estadual do Maranhão, para a obtenção do título de Doutor em Agroecologia. Orientador: Prof. Dr. Guillaume Xavier Rousseau São Luís - MA 2019 García, Luis Manuel Hernández. B-diversidade das comunidades de oligoquetas da Área de Endemismo de Belém / Luís Manuel Hernández García. – São Luís, 2019. 168 f Tese (Doutorado) – Curso de Agroecologia, Universidade Estadual do Maranhão, 2019. Orientador: Prof. Dr. Guillaume Xavier Rousseau. 1.Minhocas. 2.Taxonomia. 3.Macrofauna. 4.Conservação. I.Título CDU: 574.1(811.5) v Dedico A minha amada filha SOFÍA HERNÁNDEZ RODRÍGUEZ e a minha esposa LESBIA NEYLESS RODRIGUEZ GODOY. vi AGRADEÇO Á Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pela concessão da bolsa de estudos. Á Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) pelo financiamento da pesquisa. Á minha esposa Lesbia Rodríguez e minha filha Sofía por serem minhas fontes de inspiração, por terem me ajudado a ser uma melhor pessoa. Á meus padres e irmaõs cuja luta diária na Venezuela e Argentina serve de inspiração para eu trabalhar mais forte. Ao meu orientador Guillaume Xavier Rousseau por ter-me dado esta oportunidade. Á Danielle Celentano e o Christoph Gehring por serem mais do que pesquisadores comuns, eles abriram portas para desestressar as turmas. Aos meus amigos que são muitos, me perdoam se esqueço alguns: Jesus, Henry, Ernesto, Jhonatan, Robinson, Sandriel, André, Raymond, Silver, Nadja, Francisneydi, Vivian, Valeria, Lalia, Samara, Alexandra, Stefania, Marcelo, Raysa, Larissa, Kellen e etc. Gostaria de deixar um espaço em branco para aqueles que não lembro colocarem seus nomes. Aos meus colegas de trabalho de pesquisa por me ensinar, me guiar, me incentivar (George Brown, Samuel James, Bianca Santos, Sandriel Sousa). Aos habitantes das comunidades de Alcântara, Gurupi, Ribamar, e aos povos indígenas que são a razão de nosso trabalho. Lutamos para que essas terras sejam respeitadas. A aquelas pessoãs que de uma ou outra forma contribuíram para nossa pesquisa. vii LISTA DE ILUSTRACÕES REVISÃO BIBLIOGRÁFICA: Figura 1:Representação esquemática da anatomia de Pontoscolex corethrurus.. ........................ 3 Figura 2: Esquema básico da copulação de uma minhoca típica europeia (Eisenia foetida) e formação de casulos. A. Inseminação mútua; o espermatozóide do poro genital (somito 15) passa pelos sulcos seminais para os receptores seminais (somitos 9 e 10) de cada parceiro. B, C. Depois que as minhocas se separam, um tubo pastoso formado sobre o clitelo passa adiante para receber óvulos de ovidutos e espermatozoides de receptores seminais. D. O casulo desliza sobre a extremidade anterior e suas extremidades se fecham e selam. E. O casulo é depositado perto da entrada da galeria. F. As minhocas jovens emergem entre 2 a 3 semanas. ..... 5 Figura 3: Categorias ecológicas das minhocas. ....................................................................... 7 Figura 4: Rotas importantes através das quais grupos funcionais de minhocas influenciam os processos no solo e serviços ecossistêmicos. A espessura das setas representa a importância relativa.. ............................................................................................................................. 9 Figura 5: Índices de similaridades comumente usados para o cálculo de β diversidade. O Índice de Jaccard é expresso em uma matriz de presença e ausência e Bray-Curtis para abundância.. ..................................................................................................................... 16 Figura 6: Exemplo hipotético envolvendo 4 ilhas (A-D) com três localidades amostradas em cada uma. As biotas A1-A3 estão completamente aninhadas, as biotas mais pobres são subconjuntos da mais rica. Localidades B1-B3 tem a mesma riqueza (seis espécies cada uma) com três espécies em comum às três localidades e três espécies exclusivas de cada área. Localidades C1-C3 apresentam ambos padrões, porque C2 e C3 são subconjuntos de C1 (aninhamento), mas algumas espécies são substituidas entre C2 e C3, as quais são exclusivas dessa áreas. As localidades D1-D3 apresentam substituição espacial e não estão aninhadas, mas apresentam diferenças de riqueza.. ............................................................................... 17 Figura 7: Modelo de β-diversidade determinada por fatores físicos do solo. O gradiente geográfico não apresenta respostas dentro de florestas secundárias. ........................................ 19 CAPITULO 1: New earthworm species of Righiodrilus (Clitellata, Glossoscolecidae) from Eastern Amazonia FIGURE 1. Righiodrilus gurupi n. sp. A. Ventral view of clitellar region. B. Somatic setae A of xxi. C. Genital setae A of xvii. D. Ventral view of calciferous gland of xii; m: membranous region; g: glandular region. E. Ventro-lateral view of spermathecae. F. Nephridium as seen in xiv–xvii. ......................................................................................................................................... 35 FIGURE 2. Righiodrilus viseuensis n. sp. A. Lateral view of anterior body with clitellum. B. Ventral view of anterior body with clitellum. C. Photography of lateral view of anterior body with clitellum. D. Spermathecae of vi–viii. E. Dorsal view of calciferous gland of xii. F. Photography of dorsal view showing gizzard (segment vi), spermathecae (segments vi, vii and viii viii) and calciferous gland (segment xii); g=gizzard, s=spermathecae, c=calciferous gland. G. Somatic seta A of xv. ..................................................................................................................... 36 FIGURE 3. Righiodrilus moju n. sp. A. Ventral view of anterior body with clitellum. B. Lateral view of anterior body with clitellum. C. Spermatheca. D. Photography of the spermathecae (partly in segment ix and completely in segment x). E. Dorsal view of calciferous gland of xii. F. Photography of dorsal view showing internal structures: gizzard (segment vi), spermathecae (segments ix and x) and calciferous gland (segment xii); g=gizzard, s=spermathecae, c=calciferous gland. ......................................................................... 39 FIGURE 4. Records of species of the genus Righiodrilus, with data from Brown and James (2007), Feijoo and Celis (2010), and Celis and Rangel (2015). .................................................... 40 CAPITULO 2: Three new species of Holoscolex (Clitellata, Glossoscolecidae) from the Gurupi Biological Reserve, last forest remnant of the Belém Endemism Area, Eastern Amazon FIGURE 1. Localization of sampling site within Belém Endemism Area and Gurupi Biological Reserve in Maranhão state, Brazil. .............................................................................. 47 FIGURE 2. Holoscolex dossantosi sp. nov. (holotype). A. Ventral view of clitellar region. B. Genital setae of right papilla on A line of XVIII. C. Common seta of right A line of XXX. D. Dorsal view of right calciferous gland. BV, blood vessel; G, glandular region; M, membranous region. E. Frontal view of right post-clitellar nephridium. np, nephropore; npt, nephrostome; V, nepridium vesicle; Bd, bladder. F. Dorsal view of right spermatheca of VIII. 49 FIGURE 3. Holoscolex alatus sp. nov. (holotype). A. Ventral view of anterior region. B. Common seta on right A line of XXX. .......................................................................................... 52 FIGURE 4. Holoscolex fernandoi sp. nov. (holotype). A. Latero-ventral view of anterior region. a, A setae; b, B setae; c, C setae . B. Genital seta of papillae on right A line of XX. C. Common seta on right A line of XXX. D. Dorsal view of right calciferous gland. BV, blood vessel; G, glandular region; M, membranous region. E. Dorsal view of right spermathecae of VII and VIII. F. Lateral view of left post-clitellar nephidium. np, nephropore; npt, nephrostome; V, nepridium vesicle; Bd, bladder. G. Dorsal view of a left common atrial gland of XIX. ........................................................................................................................................... 53 FIGURE 5. Different arrangements of atrial glands of Holoscolex fernandoi, schematic. AG, atrial glands. .................................................................................................................................. 54 CAPITULO 3: Brasilisia n. gen. and Arraia n. gen., two new genera of Ocnerodrilidae (Annelida, Clitellata, Oligochaeta) from Eastern Amazonia, Brazil ix FIGURE 1. Sampling location at Alcântara and Rosário counties in Maranhão state, Brazil. ..... 61 FIGURE 2. Brasilisia punki n. gen. A. Ventral view of anterior body, with clitellar region. B. Post clitellar seta of a line of XXXV. C. Genital seta of b line of segment XVI. D. Dorsal view of left calciferous gland of IX. E. Dorsal view of left post clitellar nephridium. F. Dorsal view of left spermatheca of IX. sa = follicle of setae a, sb = follicles of setae b. ......................... 64 FIGURE 3. Arraia nelmae n. gen. A. Ventral
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