Universidade Federal De Mato Grosso Instituto De Biociências Programa De Pós-Graduação Em Ecologia E Conservação Da Biodiversidade

UNIVERSIDADE FEDERAL DE MATO GROSSO INSTITUTO DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA E CONSERVAÇÃO DA BIODIVERSIDADE

REVIEW OF THE BRACHYPTEROUS SPECIES OF THE GENUS Dichotomius HOPE (COLEOPTERA: SCARABAEIDAE: SCARABAEINAE)

RAFAEL VIEIRA NUNES

CUIABÁ – MT 2012

UNIVERSIDADE FEDERAL DE MATO GROSSO INSTITUTO DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ECOLOGIA E CONSERVAÇÃO DA BIODIVERSIDADE

REVIEW OF THE BRACHYPTEROUS SPECIES OF THE GENUS Dichotomius HOPE (COLEOPTERA: SCARABAEIDAE: SCARABAEINAE)

RAFAEL VIEIRA NUNES

Dissertação apresentada ao Programa de Pós-Graduação, do Instituto de Biociências, para obtenção do título de Mestre em Ecologia e Conservação da Biodiversidade

CUIABÁ – MT

2012

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ORIENTADOR: Fernando Zagury Vaz de Mello

I dedicate this work to my family

Our days are precious but we gladly see them going If in their place we find a thing more precious growing A rare, exotic plant, our gardener's heart delightning A child whom we are teaching, a booklet we are writing

Hermann Hesse, Magister Ludi: The Glass Bead Game, p.250

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ACKNOWLEDGEMENTS

First and foremost, I am grateful to my parents, Maria de Lourdes Nunes and Manoel Nunes for psychological, moral and financial support since my arrival in Cuiabá – MT. Academically, the productive times that I spent in Cuiabá I owe to Fernando Vaz-de- Mello, my advisor; to all teachers, colleagues and researchers from the Ecology and Biodiversity Conservation Post-gradutaion Program of UFMT, specially Thiago Izzo, Alexandre Ribeiro Cunha, Katiane Mara Ferreira and Rogério Vieira Rossi; to CAPES – Comissão de Aperfeiçoamento de Pessoal de Nível Superior, that provided me a Scolarship throughout my Reasearch Program, essential to cover my expenses while living in Mato Grosso. Besides them, my colleagues and friends of “Dung-Beetle Lab” (known by us as ‘Laboratório de Rola-Bostologia’): Letícia ‘Bagunça’ Bavutti, Michelle Ferreira, Marcelo Pessoa, Luis ‘Bibi’ Gabriel, Liliane ‘Lilica’ Silva, Ricardo Silva, Ana Sílvia Tissiani, Jorge ‘da Capadócia’ Silva and Maria Eduarda ‘Duda’ Maldaner. I am eternally thankful to Marcela Gigliotti for all time of learning and companionship, in all areas, every time I needed it. I also thank to my friends Mateus ‘Ventania’ Souza, Wesley Dátillo ‘Carioca’, Fernando Tortato ‘Galdério’ and Max.

I also thank Cristiano Lopes Andrade (UFV) for the important suggestions on this manuscript. Helder Borges Faria (UFMT) for taking important pictures for the second chapter and Maria Eduarda Maldaner (UFMT) for editing pictures for the first chapter. Helena Carolina Onody (MZSP) for providing information about the locality of a new species. Darren Mann, from Oxford University Museum and Paolo Gandini, for sending me valuable specimens. David Edmonds, for the valuable help with the English translation for the first chapter. Ricardo Andreazze (UFRN) and Tito Vidaurre (UFLA) for showing and sending valuable specimens. Graciela de Oliveira (UFMT) for providing useful information on the second Chapter.

Over the past two years, the following people were present, somehow, in every good time that I have spent, they probably know why their names are here, they are: Barbara Becker; Mariana Nunes; Manoel Nunes; Weldon Nunes; William Nunes; Arturo Gonzalez; Jorge Arias; Fernando ‘Besourinho’ Silva; Jéssica Falcão; Pedro Diniz; Desirée Ramos Marques; Renato Gama, Marina Regina Frizzas; Patrícia Leite; Vitor Azarias Campos; Caroline Dal Mas; Robert Nesta Marley and Jorge Ben Jor.

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SUMMARY

DISCLAIM ………………………………………………………………………………. 11 LIST OF FIGURES ……………………………………………………………………… 12 ABSTRACT ……………………………………………………………………………... 14 RESUMO ……………………………………………………………………………….. 15 General Introduction …………………………………….………………………………. 16 1. Bibliographic References ……………………………………………………………... 18 CHAPTER ONE: New brachypterous species of Dichotomius Hope, with taxonomic notes on the subgenus Luederwaldtinia Martínez (Coleoptera, Scarabaeidae, Scarabaeinae) … 21 1. Abstract ……………………………………………………………………………….. 22 2. Resumo ………………………………………………………………………………... 22 3. Introduction ………………………………………………………………………...…. 22 4. Material and Methods …………………………………………………………………. 23 5. Dichotomius (Luederwaldtinia) sp.1 n. sp. ………………………………...…………. 24 6. Notes on other species of Luederwaldtinia ………………………………………….... 26 7. Key for complexes of the subgenus Luederwaldtinia ……………………………….... 30 8. Discussion …………………………………………………. ……………………………….. 33 9. Bibliographic References …………………………………………..…………………. 35 CHAPTER TWO: Brachypterous species of the subgenus Selenocopris Burmeister (Coleoptera, Scarabaeidae, Scarabaeinae) ……………………………………………….. 41 1. Abstract …………………………………………………………………………….…. 42 2. Resumo ………………………………………………………………………………... 42 3. Introduction …………………………………………………………………………… 42 4. Material and Methods …………………………………………………………………. 43 5. Key for brachypterous species of the subgenus Selenocopris and definition of two complexes in the subgenus ………………………………………………………………. 43 6. Dichotomius (Selenocopris) sp.2 n. sp. ……………………………………….……… 44 7. Dichotomius (Selenocopris) sp.3 n. sp. …………………………………………….… 46 8. Dichotomius (Selenocopris) sp.4 n. sp. ……………………………………………….. 48 9. Dichotomius (Selenocopris) sp.5 n. sp. ……………………………………………….. 49 10. Redescription of Dichotomius (Selenocopris) ingens (Luederwaldt, 1935) ………… 50 11. References ………………………………………………………………………….... 52

CHAPTER THREE: Conservation of the brachypterous species of the genus Dichotomius Hope (Coleoptera: Scarabaeidae: Scarabaeinae): Conservation status , Environmental legislation and Management Tactics. ……………………………………………………. 60 1. Abstract ……………………………………………………………………………….. 61 2. Resumo ……………………………………………………………………………...… 61 3. Introduction ………………………………………………………………………….... 62 4. Material and Methods ……………………………………………………………….… 63 5. Results and Discussion ………………………………………………………………... 63 6. Conclusions ………………………………………………………………………….... 67 7. Bibliographic References …………………………………………………………...… 68 Conclusion …………………………………………………………..…………………… 72 1. Key for brachypterous species of the genus Dichotomius and Holocephalus ………... 73 2. Bibliographic Reference ………………………………………………………………. 76 ATTACHMENT 1: Guidelines for authors – Zootaxa ………………………………….. 77 ATTACHMENT 2: Guidelines for authors – Zookeys ………………………………….. 80

DISCLAIM

This dissertation does not intend to be a valid publication under articles 7-9 of the International Code of Zoological Nomenclature, so new zoological names and synonyms proposed here are not valid or available until their publication in other sources.

LIST OF FIGURES

CHAPTER ONE Figure 1. Dorsal habit (A); dorsal view of genitalia (B); hind wing (C) and tarsi and tarsal claw (indicated by black arrow) (D) of Dichotomius mysticus . Dorsal habit (E); dorsal view of genitalia (F); and hind wing of Dichotomius sp.1 sp. nov. Scale bar = 1.5 mm (G) ……………………………………………………………………………………………. 40

Figure 2. Black arrow indicating crenulated clypeal margin of Dichotomius eucranioides (A); black arrow indicating tooth or protuberance on clypeo-genal junction of D. sexdentatus . It also can be seen on D. eucranioides (B); black arrow indicating the assimetry on the paramera of D. nisus (C); black arrows indicating the pilosity on the hypomeron of D. crinicollis (D); white arrow indicating gibosities on the pronotum of D. superbus (E); white arrow indicating a fovea on the posterior angle of the pronotum of D. schiffleri (sericeus complex) (F); black arrows indicating cephalic transversal carina of D. speciosus (G); black arrow indicating male’s cephalic carina of D. agenor (H); elytra micro-sculpture of D. glaucus (carbornarius complex) (I). ……………………….…..… 41

CHAPTER TWO Figure 1. 1 . Arrows indicating the lacking of angulationon the clypeo-genal junction on the species of bicuspis complex. 2. Arrows indicating the angles on the Clypeo-genal junction on the species of ascanius complex. 3. Arrow indicating the humeral callus on the macropterous species of Selenocopris . 4. Arrow indicating the absence of humeral callus on the brachypterous species of Selenocopris. 5. Arrows indicating the equally on the width of pronotum and elytra. Elytra feebly convex. 6. Arrows indicating the pronotum wider than elytra. Elytra strongly convex. 7. Dorsal habitus of Dichotomius sp.4 n. sp. 8. Arrows indicating the ocellate punctures on the pronotal disc of D. sp.4. 9. Arrows indicating enlarged striae on the elytra of D. sp.4. 10. Arrows indicating the prolongation of last abdominal sternite beyond the apex of pygidium on a female of D. sp.4. 11 . Dorsal view of D. sp.4 paramera ………………………………………………………..………. 55

Figure 2. 12 . Dorsal habitus of Dichotomius sp.2 n.sp. 13. Head transversal carina of D. sp.3. 14 . Dorsal view of D. sp.3 paramera. 15. Dorsal habitus of Dichotomius sp.3 n. sp. 16. Arrows indicating ocellate punctures on the anterior lobe of the pronotum of D. sp.3. 17 . Dorsal view of D. sp.3 paramera ………………………………………..…………… 57

Figure 3. 18 . Dorsal habitus of Dichotomius sp.5 n. sp. 19. Arrow indicating sparse and small ocellate punctures on the anterior lobe of the pronotum of D. sp.5. 20. Head transversal carina of D. sp.5. 21. Dorsal view of D. sp.5 paramera. 22. Dorsal habitus of Dichotomius ingens (Luederwaldt, 1935). 23. Conical horn on the head of D. ingens . 24. Dorsal view of D. ingens paramera. …………………………………………………….. 58

Figure 4 . Distribution map of Dichotomius ingens, D. sp.5 , D. sp.2 , D. sp.4 and D. sp.3. ……………………………………………………………………………………………. 60 12

CHAPTER THREE Figure 1. Distribution map for nine brachypterous species of the genus Dichotomius (Coleoptera, Scarabaeidae, Scarabaeinae . …………………………………………….…. 67

ABSTRACT

The genus Dichotomius Hope (Coleoptera, Scarabaeidae, Scarabaeinae) comprises about 170 species endemic to the Americas mainly distributed in the tropics, and divided in four subgenera: Dichotomius Hope, Selenocopris Burmeister, Homocanthonides Luederwaldt and Luederwaldtinia Martinez. Some species of this genus are brachypterous, which means they have atrophy on the hind (or membranous) wing, therefore, they are flightless. This character is found on three subgenera of Dichotomius what makes the brachypterous Dichotomius an artificial group. We had three main aims in this dissertation: 1) to do a taxonomic review of the brachypterous species of the genus Dichotomius ; 2) to discuss evolutionary aspects of the brachypterism in the genus and 3) to access conservation status of the brachypterous species of the genus. There are nine described or new species of brachypterous Dichotomius: Dichotomius (D.) comarapensis Génier , placed in a homonymous complex; Dichotomius (S.) ingens (Luederwaldt), Dichotomius (S.) sp.5 n. sp. , Dichotomius (S.) sp.3 n. sp. , Dichotomius (S.) sp.4 n. sp., Dichotomius (S.) sp.2 n. sp., these last five species positioned in ascanius complex; Dichotomius (L.) eucranioides Pereira and D’Andretta, positioned in a homonymous complex, Dichotomius (L.) mysticus (Luederwaldt) positioned into geminatus complex and Dichotomoius (L.) sp.1 n. sp. placed in the batesi complex. We believe that brachypterism evolved independently at least five times in the genus, but there is the necessity of a phylogenetic analysis to test this hypothesis. Among brachypterous species of the genus, according to the criteria of IUCN, D. periotoi and D. mysticus can be considered Critically Endangered (CR) while the occurrence of the species is classified as Endangered (EN), Vulnerable (VU), Near Threatened (NT) and Data Deficient (DD). Population monitoring and respect to the specific environmental legislation need to be prioritized in order to conserve conserve the populations of the former species. In the conclusion chapter, a key for the brachypterous species of the genus Dichotomius and Holocephalus is provided.

Key-words: Dung beetles, Scarabs, taxonomy, conservation.

RESUMO

O gênero Dichotomius Hope (Coleoptera, Scarabaeidae, Scarabaeinae) está posicionado na subfamília Scarabaeinae, família Scarabaeidae, ordem Coleoptera. Possui cerca de 170 espécies exclusivamente encontradas no continente Americano, principalmente na região tropical e está dividido em quatro subgêneros: Dichotomius Hope, Selenocopris Burmeister, Homocanthonides Luederwaldt e Luederwaldtinia Martinez. Algumas espécies desse gênero são braquípteras, o que significa que elas possuem atrofiamento das asas posteriores (ou membranosas) e, portanto, são incapazes de voar. Esse caractere é encontrado em três subgêneros de Dichotomius , o que faz dos Dichotomius braquípteros um grupo artificial. Nossos três objetivos principais foram: 1) fazer uma revisão taxonômica das espécies braquípteras do gênero Dichotomius ; 2) discutir aspectos da evolução do braquipterismo no gênero e 3) acessar o status de conservação das espécies braquípteras de Dichotomius . Existem nove espécies braquípteras de Dichotomius: Dichotomius (D.) comarapensis Génier, posicionada em um complexo homônimo; Dichotomius (S.) ingens (Luederwaldt), D. (S.) sp.5 sp. nov , D. (S.) sp.2 sp. nov. , D. (S.) sp.4 sp. nov. , D. (S.) sp.3 sp. nov. , as últimas cinco listadas posicionadas no complexo ascanius; Dichotomius (L.) eucranioides Pereira e D’Andretta, posicionada em um complexo homônimo, D. (L.) mysticus (Luederwaldt) posicionada no complexo geminatus e .D.(L.) sp.1 sp. nov. , posicionada no complexo batesi. Nós acreditamos que o braquipterismo evoluiu independentemente no gênero pelo menos em cinco ocasiões, mas são necessárias análises filogenéticas para testar essa hipótese. Entre as espécies braquípteras do gênero, de acordo com os critérios da IUCN, D. periotoi e D. mysticus podem ser consideradas como Criticamente Ameaçadas (CR) enquanto outras espécies estão classificadas como Ameaçadas (EN), Vulneráveis (VU), Quase Ameaçadas (NT) e Deficiente em dados (DD). Nós propomos que o monitoramento das populações e o cumprimento de legislações específicas devem ser priorizadas para a conservação dessas espécies. No capítulo de conclusão, fornecemos uma chave para as espécies braquípteras dos gêneros Dichotomius e Holocephalus .

Palavras chave: Rola-bosta, Escaravelho, taxonomia, conservação.

General Introduction

Precision is an obsession. For which reason someone refines, at maximum, the acuity of knowledge? Precision is a search in which, like Zeno predicted, the travelers go until the half of the way, then until the half of the way again, and then successively, never reaching the final destiny … Scientific knowledge is a prize that falls back as we advance.. Ken Alder, The Measure of All Things, 2003, p. 249.

Coleoptera of the family Scarabaeidae, subfamily Scarabaeinae, also known as dung beetles, comprises about 7.000 species divided in about 240 genera in the world (SCHOOLMEESTERS, 2011). A considerable part of this diversity is found at the New World, where several genera are endemic (DAVIS et al., 2002). One of these genera, Dichotomius , has 167 valid species up to date and which can be classified in four subgenera ( sensu Martínez (1951)): Dichotomius Hope, Homocanthonides Luederwaldt, Luederwaldtnia Martínez and Selenocopris Burmeister. Despite having a high number of described species and being relatively common in many inventories done in the Americas (KOHLMAN; SOLÍS, 1997; VAZ-DE-MELLO, 1999; VAZ-DE-MELLO, 2000; MEDINA et al., 2001; PERAZA; DELOYA, 2006; ALMEIDA; LOUZADA, 2009), the occurrence of new species and unreliable identifications is still very common for Dichotomius . Much of the taxonomic knowledge on the genus can be attributed to Herman Luederwaldt who, between 1929 and 1936, did a synopsis of the Brazilian species of Dichotomius (LUEDERWALDT, 1929, 1931, 1935, 1936). Posteriorly, some complexes (PEREIRA, 1947), isolated descriptions (PEREIRA; D’ANDRETTA, 1955) and regional reviews (KOHLMAN; SOLÍS, 1997) were published. A recent growing interest on the taxonomy of Neotropical Scarabaeinae unleashes the beginning of a taxonomic review of Dichotomius and, currently, the first complexes (or group of species or sections as defined by Luederwaldt (1929)) in subgenera Luederwaldtinia and Dichotomius are being revised. The urgent need of a taxonomic revision in the genus associated with the occurrence of new species in private and public collections motivate us to start a taxonomic review of the brachypterous species of Dichotomius . Brachypterism can be defined as the atrophy of hind (or membranous) wing, thus, brachypterous species are flightless. Until the beginning of our revision, there were described four brachypterous species of Dichotomius : D. (S. ) ingens (Luederwaldt 1935), D. (L.) eucranioides Pereira & D’Andretta 1955, D.(L.) mysticus (Luederwaldt 1935) and D. (?) comarapensis Génier 2000. Therefore, since these species clearly belong to distinct

17 subgenera and complexes, it seems that they do not constitute a monophyletic group. The apparent polyphyletism of brachypterous Dichotomius motivate us to not simply describe new species but also to propose their positioning in subgenera and complexes and, consequently, to comment important taxonomic issues involving Dichotomius . It also challenges us to discuss some hypothesis for the evolution of brachypterism in the genus Dichotomius . Finally, we deal with conservation and management of populations of these species, since all of them have a narrow geographic range, due to their limitations on displacement, fitting on the threatened statuses according to IUCN criteria. We had three main aims: 1) to conduct a taxonomic revision of the brachypterous Dichotomius ; 2) to discuss the evolution of brachypterism in the genus; 3) to access conservation status of these species proposing management actions to conserve their populations. This manuscript is organized so that the first and second chapters are related to the first and second objective while the third chapter is related to the third objective. In the first chapter, all attached to the subgenus Luederwaldtnia , we describe a new brachypterous species and redescribe D. mysticus, until now, known by a single female. We also comment on other species of this subgenus and propose a separation of Luederwaldtnia in complexes providing an illustrated identification key for them. The second chapter brings a review of the brachypterous species in the subgenus Selenocopris , with description of four new species and redescription of D. ingens . On the third chapter we propose to access conservation status of the brachypterous Dichotomius and suggest management tactics for conservation of their populations. Finally, general conclusions of this study and an illustrated key for identification of the brachypterous species in Dichotomius and Holocephalus are presented. The conclusion chapter also synthesizes the new taxonomic knowledge achieved concerning the brachypterous species of Dichotomius.

Bibliographic references

ALMEIDA, S.S.P.; LOUZADA, J.N.C. Estrutura da Comunidade de Scarabaeinae (Scarabaeidae: Coleoptera) em Fitofisionomias do Cerrado e sua Importância para Conservação. Neotropical Entomology, v.38, n.1, p.32-43, 2009.

DAVIS, A.L.V. et al. Historical Biogeography of Scarabaeinae dung beetles. Journal of Biogeography, v.29, p.1217-1256, 2002.

KOHLMAN, B.; SOLÍS, A. El género Dichotomius (Coleoptera: Scarabaeidae) em Costa Rica. Giornale Italiano di Entomologia, v.8, p.343-382, 1997.

LUEDERWALDT, H. As espécies Brasileiras do gênero Pinotus . Revista do Museu Paulista, tomo XVI, p.1-173, 1929.

LUEDERWALDT, H. Três espécies novas de Pinotus , supplementos a Memória "As espécies Brasileiras do gênero Pinotus " na "Revista do Museu Paulista" Tomo XVI, 1929, P.603 SS. Revista do Museu Paulista, v.17, p.353-361, 1931.

LUEDERWALDT, H. Zur kenntnis der gattung Pinotus . Revista de Entomologia, v.5,n.3, p.334-342, 1935.

LUEDERWALDT, H. Notas complementares "As espécies Brasileiras do gênero Pinotus ". Revista do Museu Paulista, v.20, p.207-216, 1936.

MARTÍNEZ, A. La invalidez del nombre genérico Pinotus Erichson y dos nuevas sinonímias (Col. Scarab.). Anales de La Sociedad Científica Argentina. Notas Coleopterológicas, V, p. 138–142, 1951.

MEDINA, C.A. et al. Escarabajos Coprófagos (Coleoptera: Scarabaeidae: Scarabaeinae) de Colombia. Biota Colombiana, v.2, n.2, pp. 131-144, 2001.

PERAZA, L.N.; DELOYA, C. Uma Nueva Especie Mexicana de Dichotomius Hope (Coleoptera: Scarabaeidae) y Clave para Identificación de las Especies del Grupo carolinus . Neotropical Entomology, v.35, n.5, pp. 629–631, 2006.

PEREIRA, F.S. Pinotus da seção batesi . Arquivos do museu Paranaense, v.6, pp.317–328, 1947.

PEREIRA, F.S.; D’ANDRETTA, M.A.V. Novos Escarabeídeos e Novas Sinonímias (Col. Scarabaeidae). Papéis Avulsos do Departamento de Zoologia, v.12, n.11, pp. 247–264, 1955.

SCHOOLMEESTERS, P. ScarabNet’s Global Taxon Database updated to v.1.5. Available at: http://216.73.243.70/scarabnet/results.htm. Access on: 1 oct. 2011.

VAZ-DE-MELLO, F.Z. Scarabaeidae s. str . (Coleoptera: Scarabaeoidea) de um Fragmento de Floresta Amazônica no Estado do Acre, Brasil. 1. Taxocenose. Anais da Sociedade Entomológica do Brasil, v.28,n.3,pp.447–453, 1999.

VAZ-DE-MELLO, F.Z. Estado atual de conhecimento dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: MARTÍN-PIERA, F. et al. (Eds.), Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. Monografias Tercer Milênio, vol 1., Zaragoza, pp. 183–195, 2000.

CHAPTER ONE: New brachypterous species of Dichotomius Hope, with taxonomic notes on the subgenus Luederwaldtinia Martínez (Coleoptera: Scarabaeidae: Scarabaeinae)

Submitted to Zootaxa on 24 August 2011

New brachypterous species of Dichotomius Hope, with taxonomic notes on the subgenus Luederwaldtinia Martínez (Coleoptera: Scarabaeidae: Scarabaeinae)

RAFAEL VIEIRA NUNES¹ & FERNANDO Z. VAZ-DE-MELLO²

1 - Universidade Federal de Mato Grosso, Instituto de Biociências. Programa de Pós- graduação em Ecologia e Conservação da Biodiversidade. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, BRASIL. [email protected]

2 - Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoologia. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, BRASIL. [email protected]

Abstract: D. sp.1 n. sp. ., from Bolivia, a new brachypterous species of the subgenus Luederwaldtinia is described. Another brachypterous species, from Brazil, D. mysticus (Luederwaldt) is re-described. A division of this subgenus into species-groups is proposed and a provisional key is provided for identification of these groups. Dichotomius paraguayanus Gandini & Aguilar is synonymized with Canthidium kelleri (Martínez, Halffter & Pereira). We also discuss some aspects of evolution of flightlessness in Dichotomius . Key words: Dichotomius mysticus, Dichotomius vidaurrei, brachypterism, dung beetles

Resumo: D. sp.1 sp. nov. , da Bolívia, uma nova espécie braquíptera do subgênero Luederwaldtinia é descrita. Outra espécie braquíptera, do Brasil, D. mysticus (Luederwaldt) é re-descrita. Propõe-se uma divisão desse subgênero em grupos de espécies e uma chave de identificação provisória para esses grupos é fornecida. Dichotomius paraguayanus Gandini & Aguilar é sinonimizado com Canthidium kelleri (Martínez Halffter & Pereira). Alguns aspectos da evolução do braquipterismo em Dichotomius são discutidos Palavras chave: mysticus, vidaurrei, braquipterismo, rola-bosta

Introduction Dichotomius is a widespread dung beetle genus in the Americas; it comprises 167 species assigned to four subgenera ( sensu Martínez 1951), Dichotomius Hope, Homocanthonides

Luederwaldt, Luederwaldtnia Martínez and Selenocopris Burmeister. Despite being a common, widespread (Kohlman & Solís 1997; Vaz-de-Mello 1999; Vaz-de-Mello, 2000; Medina et al . 2001; Peraza & Deloya 2006; Almeida & Louzada 2009), its last synopsis (with emphasis on Brazilian species) was done more than 80 years ago (Luederwaldt 1929). Current taxonomic work on the genus includes attempts to clarify relationships among its species groups ("sections" of Luderwaldt) and with other genera, and the call for a modern revision of the genus has been voiced by several taxonomists, including Vaz-de- Mello (1999), Medina et al . (2001); and Gandini & Aguilar (2009). Luederwaldtinia was proposed by Martínez (1951) as a substitute name for the group improperly called Selenocopris by Luederwaldt (1929). Luederwaldtinia presently has 62 valid species distributed among the twelve species-groups established by Luederwaldt (1929, 1931, 1936): agenor, assifer, batesi, calcaratus, carbonarius, crinicollis, inachus, interstitialis, micans, nisus, sericeus and speciosus groups. Members of Luederwaldtinia can be distinguished from other Dichotomius by 1) bearing at least two distinct clypeal teeth separated by emargination; 2) having the head margin rounded and lacking an angulation at the clypeo-genal junction (except for some species that have a teeth on the clypeo-genal angle); 3) having metasternal pilosity usually abundant laterally, and 4) never having last abdominal sternite of female prolonged beneath apex of pygidium nor with pair of central tubercles (Luederwaldt 1929; Vaz-de-Mello et al., 2011). Presently there are four described brachypterous species of Dichotomius : D. (Selenocopris) ingens (Luederwaldt, 1935); Dichotomius (Luederwaldtinia) mysticus (Luederwaldt, 1935); Dichotomius (Luederwaldtinia ) eucranioides Pereira & D’Andretta, 1955; and Dichotomius (Dichotomius ) comarapensis Génier 2000. These species have atrophied hind wings and are flightless. All known brachypterous species have strongly convex and rounded elytra which lack humeral callus while pronotum is wider than the elytra. In D. eucranioides and D. ingens the elytra are fused. Few of biology of brachypterous Dichotomius is known. Evolutionally, it seems that this condition appeared independently at least four times in the genus. The aim of this paper is four-fold: 1) to describe a new brachypterous species of Dichotomius belonging to the subgenus Luederwaldtinia ; 2) to re-describe D. mysticus , until now, known from a unique exemplar; 3) to provide a provisional key for identification of the species groups comprising the subgenus ; and 4) to discuss some aspects of evolution of brachypterism in the subgenus Luederwaldtnia .

Material and Methods Examined specimens belong to the following collections (curators in parenthesis): APRES – Museo Noel Kempff Mercado, Santa Cruz, Bolivia (Tito Vidaurre and Julieta Ledezma). CEMT – Setor de Entomologia da Coleção Zoológica da Universidade Federal de Mato Grosso, Departamento de Biologia e Zoologia, Cuiabá, Mato Grosso, Brazil (Fernando Vaz-de-Mello). NHML – Natural History Museum, London, England (Maxwell Barclay) MZSP – Museu de Zoologia da Universidade de São Paulo, São Paulo, Brasil (Sônia Casari and Carlos Campaner). UFRN – Coleção Entomológica da Universidade Federal do Rio Grande do Norte, Natal, Brazil (Ricardo Andreazze).

Taxonomy Description of Dichotomius (Luederwaldtinia) sp.1 n. sp. Material studied: Holotype: m# BOLIVIA: Santa Cruz : Prov.: Cordillera. Loc: Palmar de las Islas. 29/I/2007 / Leg. T. Vidaurre / Dichotomius vidaurrei sp. nov., Holotype. R.V. Nunes and F.Z. Vaz-de-Mello, 2011 [CEMT]. Paratypes: 7 m#m#, 2 f#f# same data as holotype [APRES]; 1 m# same data but 1-6/II/2007 [CEMT]. Diagnosis: since this species has the pronotal disc covered with ocellate punctures (Fig. 1E), it belongs to the batesi group of Luederwaldtinia. Within this group, D. sp.1 can be distinguished by the presence of three smooth knobs on the head, pronotum wider than elytra, lacking of humeral callus and atrophiated hind wing, that reaches, at most, one third the size of elytra (Fig. 1E). Holotype: male: length: 12.8 mm. Maximum width (pronotum): 8 mm. Black, barely shinning. Head: triangular, surface transversally ridged, except on three smooth knobs at clypeo-frontal region. Ventral clypeal process bifurcated. Clypeal teeth separated by a strong U-shaped emargination. Clypeo-genal suture indistinct. Dorsal clypeal and genal margins with a single row of short setae, the length of these setae increase up to twice on clypeal teeth margin. Head surface lacking horns, acute tubercles or carinas, with three smooth knobs on clypeo-frontal region. Antennal club with light red tumescence. Pronotum: wider than long and wider than elytra. Disc with ocellate punctures, separated by its diameter. Ocellate punctures present also along posterior margin, lateral margin and

24 on anterior angle. Ill defined posterior longitudinal sulcus. Pronotal disc with feeble anterior declivity. Anterior angles rounded. Hypomeron: with a lateral band of ocellate punctures, wider anteriorly and posteriorly and reduced to a single row medially. Prosternum: with elliptical ocellate setose punctures (setae have half the length of hypomeral setae). Mesosternum: narrowed and shining with foveolate microsculpture medially. Ocellate punctures present laterally. Mesepisternum: ocellate setose punctures separated by twice its diameter. Metasternum: portion between mesocoxae one half wider than the rest of the lobe. Anterior lobe completely glabrous with few ocellate punctures on lateral margins. Elytra: barely shining. Discal striae distinctly punctuated. Punctures with the same width of stria and separated by 3/4 its diameter. Lateral (ninth) striae reaching the basis. Interstriae convex, barely shining. Legs: bearing sparse, black pilosity, denser on meso and hind tibiae. Apical fourth of anterior tibial spur strongly curved inward. Anterior tibiae with four lateral teeth. Hind tibial spur truncated. Hind wing: non articulated and non foldable (length 4 mm). Basal sclerites glabrous. Membrane extending up to the point equivalent to the lacking folding articulation. Sub-costal vein joining costal axis at its basal quarter. Medial vein extending until the one third of coastal axis. Distinct pilosity present on the apical third of costal axis. Cubital and anal veins absent. Sub-venations absent (Fig. 1G). Abdomen: sixth sternite strongly narrowed medially. Posterior margin of sixth sternite with complete margination. Pygidium: as long as wide, shinning with sparse ocellate punctures on the basal half. Phallobase: dorsally, with apical border strongly curved, medial angulation present. Parameres: almost symmetrical, apically flattened (on lateral view). Dorso-ventrally apex with medially directed hooks (Fig. 1F). Morphological variation: length varies from 9 to 14 mm. In some individuals, clypeo- genal suture as well the posterior longitudinal sulcus on pronotum can be more or less distinct. Female differ from males as follows: Legs: Anterior tibial spur feelbe curved at fourth apical. Abdomen: Medially, sixth abdominal sternite with twice the width than other sternites. Distribution: known only from Palmar de las Islas, Cordillera province, Santa Cruz, Bolivia. Etymology: named after Tito Vidaurre Sánchez, scarabaeoidologist who collected the type series. Tito’s enthousiastic interest and knowledge of scarabs is responsible for a large amount of new data on Bolivian scarabs.

Systematics: the batesi complex, in which D. sp.1 is inserted, despite has been through a taxonomic synopsis (Pereira 1947), has many taxonomic problems to be solved. Thus, is hard to take conclusions on which species are morphological similar to D. vidaurrei . Ecological information: The type-series was collected in a project whose ecological results are published by Vidaurre et al . (2008), where it is under the name of Dichotomius sp.2. Dichotomius sp.1 n. sp. was collected with pitfall traps baited with human feces. Specimens were collected at “Bosque Chaqueño” and “Bosque de Palmas”, both ecosystems are considered disturbed. Within the study site, D. sp.1 is rare; only 22 of 3382 collected individuals belonging to 57 species were D. sp.1 . This species is sympatric with a non-identified species of the carbonarius group and with Dichotomius nisus (Olivier, 1789). Three paratypes were collected manually during the day.

Redescription of Dichotomius (Luederwaldtinia) mysticus (Luederwaldt, 1935) - Pinotus (Selenocopris) mysticus, Luederwaldt, 1935, Rev. de Entomol. 5(3): 340 - Pinotus mysticus, Blackwelder, 1944, U.S. Nat. Mus. Bull., 185: 207 - Dichotomius (Luederwaldtinia) mysticus, Vaz-de-Mello, 2000, p. 193

Material studied: Holotype: f# Natal (Brasilien). W.M. Mann [NHML]; 2f#F#, m# BRASIL: Rio Grande do Norte: Natal. Parque das Dunas. 5º48’35’’S; 35º11’28’’W. hum faec. 18.V.2011. MEMaldaner [CEMT]; 1 m# same data but 15.III.1998. O.Câmara. [UFRN]; 4 m#m#, 2 f#f# same but 15.III.1991. [UFRN]; 2 m# m#, 5 f#f# same data but 16.III.1991. [UFRN]; 1 m#, 1 f# same data but 16.III.1991 [UFRN]; 1 m#, 1 f# same data but 13.IV.1991[UFRN]; 2 m#m# same data but 17. V.1991[UFRN]; 2 f#f# same data but 15.VI.1991 [UFRN]. Diagnosis: Dichotomius mysticus belongs to the geminatus complex due the combination of the following characters: head with single central tubercle, pronotum with an anterior single central lobe and hypomeron, laterally, with dense long setae (Fig 1A). Among this complex, D. mysticus is the unique brachypterous species, thus, its pronotum is wider than the elytra, humeral callus is absent, the elytra are strongly convex and hind wing never reaches the size of the elytra. Besides, meso and meta tarsi and tarsal claws are covered by a dense red pilosity (Fig. 1D). Males: Length: 11 to 13 mm. Maximum width (pronotum): 6.0 to 7.5 mm. Black, shining, with feeble blue sheen on base of elytra of some specimens. Some males do not bear this blue reflection. Head: semicircular, surface almost smooth, with few ocellate punctures

26 near each eye. Clypeogenal junction evenly rounded, lacking angulation, with weak emargination. Ventral clypeal process bifurcated. Clypeo-frontal region with a single, central tubercle. Antennal club with dark red or light brown tumescence. Clypeal and genal lateral margins with a ventral row of small red setae that increase in length from the middle of head laterally. Pronotum: length equal to width; wider than elytra. Disc smooth, shining, with indistinct punctures (only seem under 20x magnification); ocellate punctures present along posterior border and on lateral and anterior angles. Anterior margin widened in the middle forming a triangular projection. Pronotal disc with central knob anteriorly. Anterior angles rounded. Hypomeron: with a lateral band of ocellate punctures bearing abundant red pilosity, narrowest medially and reduced to single medial row (hairs in the single row about one half the hairs of anterior and posterior portion). The lateral red pilosity may vary in abundance. Hypomeron glabrous, shining medially. Prosternum: with ocellate, setose punctures. Mesosternum: weakly narrowed, shining, glabrous between mesocoxae, with ocellate, punctures and pilosity above coxae. Mesepisternum: with ocellate, setose punctures evenly distributed on surface, Metasternum: portion between mesocoxae widened apically, width twice that between apices of coxae, with ocellate punctures anteriorly and laterally and posterior depression. Elytra: shining with blue sheen anteriorly on some males, discal striae distinctly punctated, punctures wider than striae and separated by twice of its. Lateral (ninth) striae not reaching the basis. Elytral interstriae shining, feebly convex. Legs: bearing abundant red pilosity on tibia, tarsi and tarsal claws (Fig. 1D). Anterior tibiae with three well defined teeth, and feebly developed fourth (basal) indicated by a sinuosity. Ventral surface of anterior femur with tree rows of setose ocellate punctures. Apical fifth of anterior tibial spur strongly curved inward. Ventral surface of middle femur with ocellate punctures near tibial articulation. Hind tibial spur flattened, subapically dentate. Hind wing: articulated and foldable, reduced (length 6.5 mm). Basal sclerites glabrous. Subcostal vein intersecting costal axis at its basal quarter. Medial vein extending until the folding articulation. Distinct pilosity present on the basal third of costal axis and on folding region. Cubital vein present, anal vein absent. Small developed membranous portion apical to folding articulation with incomplete or weakly-marked venations (Fig. 1C). Abdomen: with sixth sternite strongly narrowed medially and bearing small medial protuberance. Pygidium: wider than long, shining and glabrous. Ocellate punctures present only basally. Phallobase: dorsally, with apical border curved. Parameres: almost symmetrical, truncated and apically flattened (on lateral view).

Females: length varies from 11 to 12.5 mm. Pronotum width varies from 6.0 to 7.5 mm. Females differ from males as follows: Head: clypeus completely transversally ridged. Ocellate punctures more conspicuous near eyes. Fronto-clypeal junction with a pair of closely-positioned medial tubercles. Frons with median curved depression delimited anteriorly by a C-shaped transverse elevation. Pronotum: lacking knobs or declivities. Hypomeron: Hairs longer and denser than in males. Abdomen: Medially, sixth abdominal sternite with twice the width than other abdominal segments. Distribution and conservation: Natal, Rio Grande do Norte, northeastern Brazil. Except for the holotype, a female that was collected at an undefined location at Natal, other examined specimens were collected at Parque Estadual das Dunas Costeiras, a conservation unit in the city of Natal. Parque das Dunas is severely impacted by urban growth and tourism (Jesus 2003). Besides, D. mysticus , because it cannot fly, may have a restricted habitat. Because it is an endemic to this region, and occupies a narrow, compromised habitat, this species can be regarded as threatened in accordance with IUCN criteria (IUCN 2010) (unpublished data). Systematics: at the time of its description, this species was positioned at inachus group (Luederwaldt 1935). However, with a new definition of Luederwaldtinia groups herein this work, we suggest that this species belongs to the geminatus group due the characters of pronotum, head and pilosity (see diagnosis and key for the groups). Besides, D. mysticus, D. geminatus and D. opacipennis compose this group. Ecological information: Three known specimens are associated with environments with trees on sand dunes in the city of Natal. They were caught using pitfall traps baited with human faeces. Other specimens were collected between March and June but lack ecological data. Dichotomius mysticus is sympatric with unidentified species of the sericeus and carbonarius complexes .

Notes on other species of the subgenus Luederwaldtinia Dichotomius eucranioides Pereira and D’Andretta, 1955 Since the time of its description, D. eucranioides has maintained a doubtful position within Luederwaldtinia . Pereira & D’Andretta (1955) did not assign it to a group. Vulcano et al. (1967) later placed it in the assifer species-group due the presence of a transversal carina on males head. However, we believe that D. eucranioides should be considered an isolate hence monobasic group within the subgenus for the following reasons: it is the unique species in the genus that combine following characters: crenulated clypeal margin

(Fig. 2A); head transversal carina width subequal to the distance between the eyes while; head transversal carina is wider than high; pronotal disc of males is delimited, anteriorly, by four gibosities; on pronotal declivity of males, there are two lateral depressions filled with ocellate punctures. Besides, contrary to the species of assifer group, D. eucranioides hypomeron do not have an anterior excavation delimited medially by a dense brush of red setae. Despite D. eucranioides being a Luederwaldtinia brachypterous species, we decided not to redescribe herein since the description of the female (type, Pereira & D’Andretta 1955) and of the male (Vulcano et al. 1976) are detailed enough.

Dichotomius paraguayanus Gandini and Aguilar, 2009, new synonym of Canthidium kelleri (Martínez, Halffter and Pereira, 1964) Gandini and Aguilar (2009) assigned Dichotomius paraguayanus to the subgenus Luederwaldtinia in a species-group (section) of its own. Comparison of a paratype of D. paraguayanus (Paraguay: Dpto Cordillera: Naranjo. 15.X.2005. J.Rivas, CEMT) and a paratype of C. kelleri (Município: Rio Verde, Goyaz. Dr Nick, CEMT) leads us to conclude that the two are synonymous. The reassignment of this species to Canthidium is supported by the absence of a ventral clypeal process, the angulation of the apices of the protibiae (less than 90°), a short and convex mesosternum and ovoid body shape. Canthidium paraguayanus belongs to a group of Canthidium that is commonly found mixed with Dichotomius in collections around the world. This group also includes C. gigas Balthasar, 1939 and C. bokermanni (Martínez, Halffter and Pereira, 1964) and is currently under review by the second author and a collaborator. We have consulted Paolo Gandini about this synonym and he agrees with reassignment to Canthidium .

Dichotomius sexdentatus (Luederwaldt 1925) and D. simplicicornis (Luederwaldt 1935) Luederwaldt (1936) created the section calcaratus to accommodate D. calcaratus (Arrow 1913), D. sexdentatus (Luederwaldt, 1925) and D. simplicicornis (Luederwaldt 1935). Vulcano et al . (1976) transfered D. calcaratus to the buqueti species-group, then to the subgenus Dichotomius. For this reason, calcaratus group does not make any sense in subgenus Luederwaldtinia and two remaining species, D. sexdentatus and D. simplicicornis should compose other groups. After examining the type [MZSP], we suggest that D. sexdentatus composes crinicollis species group for the following reasons: it has a tooth on clypeal genal junction

(Fig. 2B) and its hypomeron bears dense, long and curved red pilosity (Fig. 2D). After examining the type of D. simplicicornis [NHML] and scrutinize its original description, we suggest that this species should compose agenor species group since the cephalic horn in males constitutes a transverse carina with a strong central tooth and two lateral teeth (Fig. 2H), characteristic to other species of this complex.

Dichotomius interstitialis (Luederwaldt 1931) Luederwaldt (1931) proposed D. interstitialis to comprise a species-group for its own. As a result of the examination of the holotype [NHML] and scrutiny of the original description, where the author compares it to the carbonarius species-group, we are inclined to consider this species as part of this later group for the following reasons: elytra microsculpture (Fig. 2I), two tubercles on cephalic carina, pronotum without horns, tubercles or knobs and body shape very close to other species of this group. Following description, D. interstitialis should be recognized within the species in the group due the hind tibial spur, which is not bifurcated.

Dichotomius guaranii Gandini and Aguilar, 2009 This species was considered by its authors to be closely related to D. nisus (Olivier 1789). No paratypes were examined. However, a close examination of the photograph in the original publication (Gandini and Aguilar, 2009), and of the original description, lead us to the conclusion that this species in fact belongs to the subgenus Selenocopris , and that it may be morphological similar to D. ascanius (Harold 1869). Our conclusion is based in the glabrous metasternal sides, oblong form of the center of the sixth abdominal ventrite, head and pronotal sculpture, and in overall similarity of the photographed specimen to females in the D. ascanius complex. Paolo Gandini (pers. comm.) agrees with our conclusion. The ascanius species complex is in need of revision. Several possibly undescribed species in this complex are known to us only by females, which do not always express the main character defining the subgenus Selenocopris (angled form of the clypeo- genal border).

Key for identification of species-groups in the subgenus Luederwaldtinia We define here thirteen species-groups, named after the older (or only) included species: agenor, assifer, batesi, carbonarius, crinicollis, eucranioides, geminatus, inachus, muticus, nisus, sericeus ,speciosus and superbus. The main changes in these groups from

30 previous treatments are 1. the incorporation of Luederwaldt’s (1936) micans group into crinicollis group; 2. the incorporation of interstitialis group into the carbonarius group (see above); 3. the splitting of the geminatus group from the inachus group; and 4. the addition of four monobasic groups, D. superbus (see above) D. muticus (from the inachus group), and D. eucranioides (from the assifer group). We stress that we consider those groups as a starting point for further taxonomic reviews in Luederwaldtinia . Currently, the crinicollis , speciosus and assifer species- groups, and the positioning of D. superbus are under review by other authors. We do not include a list of species of each group since there are many types to be examined. However, we are confident on the names used here, either by direct type examination, examination of photographs of types, or conclusive information from original descriptions.

Identification key for males and females of the complexes in the subgenus Luderwaldtinia

1 Tooth delimiting clypeo-genal junction (Fig. 2A, 2B) … 2 - Clypeo-genal junction lacking tooth … 3

2 Clypeal margin crenulated (Fig. 2A). Males with strong transversal carina with width subequal to interocular space. Male pronotal disc delimited anteriorly by four gibosities. Female pronotal disc with two lateral groups of ocellate punctures. Pronotum of both genders with longitudinal sulcus filled with ocellate punctures; pronotum wider than elytra. Metasternal borders, mesepisternum and hypomeron with dark brown or balck pilosity. Pilosity of hypomeron not curved upwards. Elytral suture fused. Hind wing reduced, brachypterous species … Dichotomius eucranioides Pereira & D’Andretta, 1955 - Clypeal margin smooth. Males head transversal carina higher than wide; Both genders pronotal disc smooth, at most transversally roughened; pronotum with simple longitudinal sulcus lacking ocellate punctures; pronotum width equal to elytra width. Metasternal borders, mesepisternum and hypomeron with abundant red pilosity (Fig. 2D). Pilosity of hypomeron generally curved upwards. Elytral suture not fused. Hind wing macropterous, not reduced. Macropterous species.... crinicolllis group

3 Posterior pronotal margin with well-defined fovea with sharp edges at each posterior angle. Pronotum evenly rounded in both sexes (Fig. 2F) … sericeus group

- Posterior pronotal margin lacking foveae … 4

4 Hypomeron with a well defined anterior excavation delimited by a dense brush of long, red setae. Male head with high transverse carina, pronotum with one or two median lobes delimiting anterior depression … assifer group - Hypomeron lacking antero-medial excavation, or excavation very shallow and without medial setal brush ... 5

5 Body shining metallic red, purple, blue, green, and/or gold. Metasternal disc concave. Male head with transverse carina bearing single horn at each end (Fig. 2G); pronotum with a central, well-defined, usually bifurcated lobe. Pronotal disc finely punctured. Length less than 16 mm … speciosus group - Variable color and shining. Matasternal disc convex. If transversal carina is present on head, then it does not own single horns at each end. Variable length (from 12 to 25 mm) … 6

6 Green to copper color. Pronotum with strong anterior depression with bilobed summit (Fig. 2E). Male head with conical horn, female with two closely set tubercles … Dichotomius superbus (Felsche, 1901) - Other combined characteristics. If body color is green to copper, than pronotum is simple, lacking lobe or anterior depression. If pronotal has anterior depressions or lobes, then body color is black or brown … 7

7 Male head with high transverse carina bearing strong medial tooth (Fig. 2H). Pronotum simple, at most with a small knob on anterior portion … agenor group - Male head without carina, or if carinate, then carina lacks medial tooth … 8

8 Metasternal sides, mesepisternum and hypomeron with dense, long red to tan pilosity. Pronotum of large males with well defined anterior depression bearing large punctures anteriorly delimited posteriorly by bilobed knob. Left paramere of aedeagus with strong apical hook (Fig. 2C) … nisus group: Dichotomius nisus (Olivier 1789) - Dense and long red to tan pilosity lacking in, at least one of mestasternal sides, mesepisternum or hypomeron. Pronotum of large males with at most ill-defined

32 anterior depression, lacking puncturing and delimited posteriorly by ill-defined knob. Left paramere lacking apico-lateral hook … 9

9 Pronotal disc with ocellate punctures (Fig. 1E, see pronotal disc) … batesi group - Pronotal disc without ocellate punctures. Ocellate punctures might occur on anterior angles or pronotal margins ... 10

10 Length from 15 to 25 mm. Dull black to green-copper metallic species. Pronotum (except sometimes on disc) and elytra evenly chagreened (20X) (Fig. 2I). Head with either conical horn, a pair of closely-set tubercles, or a short transverse carina, … carbonarius group - Length generally less than 15 mm. Dorsal surface smooth and shining. Head always with a conical horn or tubercle, or simply flat. ... 11

11 Hypomeron laterally with sparse long setae. Hypomeral disc not covered by setae. Head always with central horn, tubercle or carina. Pronotum with feeble declivity, never with central lobe … inachus group - Hypomeron laterally with dense long setae. Most of surface of the hypomeral disc covered by setae. Head with central horn, tubercle or lacking any structure. Pronotum with anterior declivity delimited by a central lobe (on bigger males) ... 12

12 Head with a central horn, tubercle or knob … geminatus group - Head flat, without tubercle, horn or knob … muticus group ... Dichotomius muticus (Luederwaldt 1922)

Discussion With the description of D. sp.1, the number of described brachypterous species in the genus Dichotomius increases to five, in three subgenera ( D. eucranioides, D. mysticus and the new species in Luederwaldtinia , D. ingens (Luederwaldt 1935) in Selenocopris , and D. comarapensis Génier 2000 in Dichotomius s.str. ). However, at least three other species are in process of description by us, all of them in Selenocopris . These species are included in at least five supposedly different lineages (species-groups) in three subgenera,

33 what supplies a strong evidence for independent evolution events of brachypterism within this genus, as already suggested by Génier (2000). Luederwaldtinia would then include three independent appearances of brachypterism, one in Dichotomius eucranioides, one in the geminatus species-group ( D. mysticus ) and other in the batesi species-group ( D. sp.1 n. sp. ). External morphology, male genitalia and, mainly, hind wings, are very different among those species. Hind wings venation and articulations support our belief that wing atrophy occurred independently in these species (Figs. 1C and 1D). According to Zunino and Halffter (1988), the evolution of flightless Scarabaeinae species is related to the arid environments (e.g.: deserts) and high altitude zones. At arid environments, there is a great necessity to conserve water, which brachypterous species can do in different way. For Scarabaeus Linnaeus, an Afroasian- genus with some species living at arid environments including deserts, the very convex elytra increases the volume of the subelytral chamber, where water may be conserved during activity (Chown et al ., 1998). This kind of elytra is also found on flightless species of other genera living at arid environments (Chown et al , 1998, Duncan, 2002). Furthermore, fused elytra may also prevent water losses from the subelytral chamber. This adaptation occurs on D. ingens and D. eucranioides . Indeed, simply avoiding flight reduces water loss. The water-conservation hypothesis may fit for Dichotomius mysticus, D. sp.1 and flightless species of Selenocopris since they occur at environments with high temperatures and evapotranspiration rates. However, in these environments, sympatric macropterous Dichotomius species tend to be very abundant what would be a great disadvantage for flightless species since the main resource (mammals feces) is unforeseeable at the environment and would be better and faster (and easier) found by flight. Besides, D. eucranioides occurs at riparian forests associated to Paraná River, thus to be considered a humid and non-persistent environment where brachypterism is very rare and may occur only on high specialized behavior species (eg:. Termitophilous or myrmecophilous beetles, Roff (1990)). For those species, the occurrence of brachypterism remains an open question that may be answered collecting behavioral and physiological data.

Acknowledgments: We extend our thanks to Paolo Gandini, enthusiastic scarabaeidologist, for sending specimens of Dichotomius paraguayanus for comparison; to François Génier (Canadian Museum of Nature) for enlightenment and photos of type material; to Dr. Alexandre Cunha Ribeiro, Dra. Katiane Mara Ferreira and Dr. Rogério

Vieira Rossi, members of first author mastership qualification exam (by Universidade Federal de Mato Grosso) for valuable suggestions on this article; to Maria Eduarda Maldaner (Universidade Federal de Mato Grosso) for photo editing. We acknowledge with gratitude the collaboration of the curators of the collections cited above for the loan of specimens used in this study, especially Ricardo Andreazze, who showed to us the specimens of D. mysticus , and Tito Vidaurre, responsible for all known specimens of D. sp.1 . RVN receives a CAPES (Comissão de Aperfeiçoamento de Pessoal de Nível Superior) grant. FZVM is a CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) fellow and most of this work was supported by a grant from FAPEMAT (Fundação de Amparo à Pesquisa do Estado de Mato Grosso FAPEMAT, 570847/2008, 447441/2009, 283731/2010).

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Figure 1. Dorsal habit (A); dorsal view of genitalia (B); hind wing (C) and tarsi and tarsal claw (indicated by black arrow) (D) of Dichotomius mysticus . Dorsal habit (E); dorsal view of genitalia (F); and hind wing of Dichotomius sp.1 sp. nov. Scale bar = 1.5 mm (G).

CHAPTER TWO: Brachypterous species of the subgenus Selenocopris Burmeister (Coleoptera, Scarabaeidae, Scarabaeinae).

To be submitted to Zookeys

Brachypterous species of the subgenus Selenocopris Burmeister (Coleoptera, Scarabaeidae, Scarabaeinae)

Rafael Vieira Nunes¹, Fernando Zagury Vaz-de-Mello²

1 – Universidade Federal de Mato Grosso, Instituto de Biociências, Programa de Pós Graduação em Ecologia e Conservação da Biodiversidade. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, BRASIL.

2 – Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoologia. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, BRASIL.

Abstract Four new brachypterous species of the subgenus Selenocopris : Dichotomius sp.5 n.sp. , Dichotomius sp.4 n. sp. , Dichotomius sp.2 n. sp., and Dichotomius sp.3 n. sp. are described and Dichotomius ingens (Luederwaldt) is re-described. A key for brachypterous species in Selenocopris and pre-definition of two complexes in the subgenus are provided.

Keywords dung beetles, Scarabaeoidea, Coprini, flightless, wing, Cerrado

Resumo Quatro novas espécies braquípteras do subgênero Selenocopris : Dichotomius sp.5 sp. nov. , Dichotomius sp.4 sp. nov. , Dichotomius sp.2 sp. nov., e Dichotomius sp.3 sp. nov. são descritas e Dichotomius ingens (Luederwaldt) é redescrita. Uma chave para identificação das espécies braquípteras de Selenocopris e a pré-definição de dois complexos no subgênero são fornecidas.

Palavras-chave rola-bosta, Scarabaeoidea, Coprini, braquipterismo, asa membranosa, Cerrado

Introduction Dichotomius is a Neotropical genus with about 170 valid species divided into four subgenera: Dichotomius Hope, Homocanthonides Luederwaldt, Luederwaldtinia Martínez and Selenocopris Burmeister. Selenocopris has 21 valid species and two subspecies that can be easily recognized among other Dichotomius by bearing two distinct clypeal teeth separated by strong emargination; having short and scattered metasternum pilosity and having the last abdominal sternite of females prolonged beneath apex of pygidium (Luederwaldt 1929; Vaz-de-Mello et al 2011). Despite Selenocopris can be easily recognized forming a well defined subgenus in Dichotomius , taxonomic, phylogenetical, behavioral and ecological data are poor known. Among Dichotomius and related genera (e.g.: Holocephalus Hope, Chalcocopris Burmeister), the positioning of Selenocopris still remains unclear. Last species of Selenocopris was described more than fifty years ago ( Dichotomius enrietti , Vernalha 1952). Besides, considerable part of material received for identification and found at museums constitutes new species, mainly those proceeding from southeast and central Brazil. Brachypterism, the condition of atrophy of hind wing, evolved at least five times in Dichotomius (Nunes & Vaz-de-Mello, in press). One of these events comprises Selenocopris and, up to now, Dichotomius ingens (Luederwaldt 1935) was the unique 43 known flightless species in the subgenus (regardless this characteristic was noticed by Genier 2000 and not by Luederwaldt in the time of description). Mainly related to the necessity of describing new species in Selenocopris , our objective is three-fold: 1) to describe four new brachypterous species of Selenocopris; 2) to redescribe Dichotomius ingens L. and 3) to provide a key for identification of brachypterous species in this subgenus.

Collections consulted Examined specimens belong to the following collections (curators in paranthesis): CEMT – Setor de Entomologia da Coleção Zoológica da Universidade Federal de Mato Grosso, Departamento de Biologia e Zoologia, Cuiabá, Mato Grosso, Brazil (Fernando Vaz-de-Mello). IBSP – Coleção Entomológica do Instituto Biológico, São Paulo, Brazil (Nelson Perioto). CMCN – Canadian Museum of Nature (Fraçois Génier). OUMHN – Oxford University Museum of Natual History, Hope Entomological Collections, Oxford, United Kingdom (Darren Mann) NHML – Natural History Museum, London, England (Maxwell Barclay)

Taxonomy Definition of two complexes in Selenocopris and key for brachypterous species in the subgenus

In order to provide a start point for a further taxonomic review of Selenocopris we consider that the subgenus can be separated in two complexes: ascanius e bicuspis. This separation is artificial and it is based on the clypeo-genal angulation, character that Luederwaldt (1929) used to define this subgenus. The ascanius complex, relative to “section” fissus in Luederwaldt (1929) is composed by species that bear angulation on the clypeo-genal junction (Fig.1.2). On the other hand, species belonging to bicuspis complex do not have the angulation on the clypeo-genal junction (Fig.1.1). All brachypterous species described here are positioned in ascanius complex and only a complete revision and phylogenetic analysis of the subgenus would define if the brachypterous species of Selenocopris constitutes a monophyletic group.

1 Males with angulation on clypeo-genal junction (Fig.1.2) … ascanius complex (2) ’ Males usually lacking angulation on clypeo-genal junction (Fig.1.1) … bicuspis complex

2 Pronotum as large as elytra (Fig.1.5). Humeral callus generally present (Fig.1.3). Elytra feebly convex (Fig.1.5). Hind wings length superior to the length of elytra … macropterous species of ascanius complex ’ Pronotum usually wider than elytra (Fig.1.6). Humeral calus absent (Fig.1.4). Elytra strongly convex (Fig.1.6). Hind wings length never higher than to the elytra length … 3

3 Pronotal disc, posteriorly, with ocellate punctures (Figs.1.7 and 1.8). Elytral striae wide, lacking punctures (Fig.1.9). Brasil, Mato Grosso … Dichotomius paresi sp. nov. ’ Pronotal disc lacking any ocelatte puncture (Figs.2.12, 2.15, 3.18 and 3.22). Ocelate punctures restricted to the margins, anterior angles and anterior lobe. Elytral striae bearing distinct punctures … 4 44

4 Clypeo-frontal region of males developed into a single conical horn or tubercle (Fig.3.23). Females bearing transversally ridged surface on clypeo-frontal region … Dichotomius ingens (Luederwaldt 1935) ’ Clypeo-frontal region developed in a transversal carina of variable size (Figs.2.13 and 3.20). Both males and females bearing smooth surface on clypeo-frontal region… 5

5 Clypeo-frontal transversal carina strongly emarginated medially and bearing two tubercles laterally at the apex (Fig.2.13). Length of fifth metatarsomere three times the length of fourth metatarsomere. Brasil, São Paulo … Dichotomius periottoi sp. nov. ’ Clypeo-frontal transversal carina feebly emarginated medially, lacking tubercles at the apex or having small developed tubercles (like on Fig.3.20). Length of fifth metatarsomere never reach three times the length of fourth metatarsomere. … 6

6 Anterior lobe of pronotum filled with dense ocellate puncture (Fig.2.16). Species of smaller length (at most 20 mm) (Fig.2.15). Foveae of anterior portion of paramera widened, almost reaching the lateral margin of paramera (Fig.2.17) … Dichotomius darwini sp. nov. ’ Anterior lobe of pronotum filled with small spaced non-ocellate puncture. Species of bigger length (25 mm) (Fig.3.18). Foveae of anterior portion of paramera not widened, never almost reaching the lateral margin of paramera (Fig.3.21) … Dichotomius manni sp. nov.

Dichotomius (Selenocopris) sp.2 n. sp. (Fig.1.12) Material studied [collection deposited]: Holotype: (male) BRASIL: São Paulo: Descalvado. Fazenda Itaúnas. 21°54’09”S; 47°43’55”W. Mata-Citrus. Pitfall. 17-XI-2007. NWPerioto (leg.).[CEMT]. Paratype: 3 males, 2 females, same data of holotype [CEMT]; 6 males, 5 females, same data of holotype but 2.XI-2004 [IBSP]; 4 males, 3 females, same data of holotype but 3-I-2005 [IBSP]; 3 males, 2 females, same data of holotype but 16- XII-2004 [IBSP]; 4 males, 5 females, same data of holotype but 23-III-2008 [CEMT]; 4 males, same data but 23-II-2006 [CEMT]; 2 males, 1 female, same data but 27-II-2008 [CEMT]; 1 female, same data but 27-XI-2006 [CEMT]; 4 males, 8 females, same data but 22-I-2007 [CEMT]; 5 males, 2 females, same data but 6-IV-2006 [CEMT]; 1 male, 1 female, same data but 7-IV-2005 [CEMT]; 1 male, 2 females, same data but 13-I-2005 [CEMT]; 2 females, same data but 13-XI-2006 [CEMT]; 1 male, same data but 13-II-2008 [CEMT]; 4 males, 1 female, same data but 24-III-2005 [CEMT]; 3 males, 1 female, same data but 15-XII-2005 [CEMT]; 3 machos, same data but 9-II-2006 [CEMT]; 1 female, same data but 9-II-2007 [CEMT]; 1 male, 2 females, same data but 9-I-2007 [CEMT]; 3 females, same data but 11-XII-2006 [CEMT]; 2 males, same data but 16-I-2008 [CEMT]; 1 male, 2 females, same data but 1-XII-2005 [CEMT]; 1 male, 3 females, same data but 20-IV-2005 [CEMT]; 4 males, 5 females, same data but 4-II-2007 [CEMT]; 1 male, same data but 5-I-2006 [CEMT]; 1 male, same data but 19-X-2006 [CEMT]; 1 male, same data but 4-II-2007 [CEMT]; 1 male, same data but 3-XI-2005 [CEMT]; 1 female, same data but 10-II-2005 [CEMT]; 1 female, same data but 2-II-2009 [CEMT]; 1 female, same data but 18-V-2006 [CEMT]; 1 female, same data but 11-XII-2006 [CEMT]; 1 female, same data but 25-II-2005 [CEMT]. Diagnosis: this species belongs to the ascanius complex due the angulation of clypeo-genal junction (Fig.1.2). In ascanius complex, D. sp.2 can be separated of macropterous species by the characters related to brachypterism (pronotom wider than 45 elytra, lack of humeral calus and strongly convex elytra) (Figs.1.4 and 1.6). Between brachypterous species this complex, D. sp.2 can be differentiated by the clypeo-frontal carina – strongly emarginated medially and bearing two lateral tubercles at the apex (Fig.2.13). It can be also recognized by the length of fifth metatarsomere, three times the length of fourth metatarsomere. Description of holotype: Male. Lenght: 19.5 mm. Maximum width (pronotum): 13 mm. Black, shining. Head: surface smooth, with few ocellate punctures near each eye. Clypeogenal junction with obtuse angulation. Clypeal teeth separated by a strong V-shaped emargination. Dorsally, apex of clypeal teeth bearing bunch of setae. Ventral clypeal process strongly bifurcated, forming two distinct teeth. Clypeo-frontal carina higher than wide, with two lateral tubercles on apex separated by a central emargination. Clypeo-genal suture distinct. Small knob present on the posterior portion of clypeo genal suture. Antennal club with light brown tumescence. Pronotum: wider than long and wider than elytra. Anteriorly, with strong declivity and a central lobe delimited by two knobs. Anterior lobe with small ocellate punctures – diameter of punctures half the diameter of ocellate puncture on metasternum, anterior angles and margins of pronotum. Pronotal disc smooth, shining. Ocellate punctures present on pronotal posterior border, near anterior angles and at anterior central lobe. Anterior angle acute. Hypomeron: anterior portion covered laterally with dark brown setae and, medially with ocellate punctures. Posterior portion almost glabrous, with few ocellate setose punctures. Hypomeral disc shining, with a lateral row of ocellate setose puncture. Prosternum: with ocellate setose punctures (setae of this puncture do not reach one quarter of longer hypomeron setae). Mesosternum: strongly narrowed and glabrous in the middle. Ocellate small-setose punctures present in the lateral portion. Mesepisternum: ocellate punctures more abundant on anterior portion. Superior margin of posterior portion with a triangular projection that almost reach the pseudoepipleuron. Metasternum: with mesometasternal suture indistinct. Between mesocoxae, with almost twice the width of medial portion. Ocellate punctures present on borders of anterior portion. Long and sparse setae present along the lateral margins of mestasternal lobe. Mestasternal disc with longitudinal sulcus and a feeble posterior concavity. Elytra: shining. Lacking humeral callus. Discal striae with feeble punctures, punctures with the same width of striae and separated by twice its diameter. Lateral (ninth) striae not reaching the basis. Elytral feebly convex. Legs: anterior tibial spur strongly curved inward at apical fourth. Anterior tibiae with four lateral teeth. Ventral surface of anterior femur smooth, glabrous. Base of median and hind tibiae strongly sinuous, with a brush of red setae. Length of 5th hind tarsomere three times the length of 4th hind tarsomere. Abdomen: all sternites glabrous. Pygidium: shining, glabrous, wider than long. Phallobasis: dorsally with apical border strongly curved with angulation present. Parameres: dorsally symmetrical. Laterally, strongly flattened beyond the apical half. Anterior to this excavation, dorsally, parameres bears a distinct fovea, widened, almost reaching the lateral margin of paramera. Apex with right angle. Laterally, longitudinal sulcus parallel to lateral margins (Fig.2.14). Morphological variation: lenght varies from 15 to 21.5 mm in males and from 17 to 21.5 mm in females. Pronotum width varies from 7.5 to 13 mm in males and from 9.5 to 12.5 mm in females. Both males and females have variations in following characters: Head: height and central emargination of clypeo-frontal carina. In one male (from the same type locality), a simple conical horn is present. Pronotum: anterior declivity as well the pair of central knobs may vary in size. In most females, these structures are smaller and less conspicuous. Elytra: both males and females have variations on the color of elytra base, that can be black, brown or blue. Abdomen: Only on females, apex of sixth abdominal segment is prolonged beneath pygidium apex. This prolongation results in two medial tubercles separated by a feeble emargination. 46

Distribution: Descalvado, middle-west of São Paulo state. All known specimens were collected at municipality of Descalvado. Despite this locality is close to Estação Ecológica de Itirapina, a well studied Research Station 30 km near Descalvado, there are no records for D. sp.2 from this area (Fig.4). Etymology: named after Nelson Wanderley Perioto (Universidade Estadual Júlio Mesquita Filho, Jaboticabal, São Paulo), Hymenoptera taxonomist, who collected the whole type series. Discussion: between brachypterous species in ascanius complex, D. sp.2 is close to D. sp.3 (new species described below) due the shape of transversal carina on head and strong similarities on the length and excavations on paramera and its foveae. The whole D. sp.2 type-series was collected with pitfall traps, without attractive baits. It is sympatric to other non-identififed Selenocopris species that also belongs to ascanius complex. This species was collected in cerrado sensu stricto (sensu Ribeiro & Walter) and Citrus plantation next to a Cerrado environment. Females proportion in traps is equal to males. Between the years of 1962 and 1993, São Paulo state lost 87% of Cerrado vegetation (Cavassan 2002), which, associated with apparent restricted distribution, regards D. sp.2 as a threatened species in accordance to IUCN criteria (IUCN 2010) (unpublished data).

Dichotomius (Selenocopris) sp.3 n. sp. (Fig.2.15) Material studied [collection deposited]: Holotype: (male) BRASIL: Goiás: Mineiros. Parque Nacional das Emas. 18°00’51”S; 52°58’17”W. 870mosl. Prc 54. III- 2011. Humfaec. MFSouza [CEMT]. Paratype: 3 females, same data of holotype [CEMT]; 1 male, 1 female, BRASIL: Goiás: Mineiros. Parque Nacional das Emas. I-XII-1999. G.Machado [CEMT]; 1 male, 1 female, same data of anterior [CEMT]; 1 male, 1 female, same data of anterior [CEMT]; 1 female BRASIL, Goiás: Mineiros. Parque Nacional das Emas. 17°55’47”S; 52°57’49”W. 838mosl. Prc7. III-2011. Hum faec. MFSouza. [CEMT]; 1 male, BRASIL, Goiás: Mineiros. Parque Nacional das Emas. 17°59’46”S; 52°56’11”W. 852mosl. Prc37. III-2011. Hum faec. MFSouza [CEMT]; 1 male, BRASIL, Goiás: Mineiros. Parque Nacional das Emas. 18°03’43”S; 52°56’35”W. 860mosl. Prc32. III-2011. Hum faec. MFSouza [CEMT]. Diagnosis: this species shall be placed in ascanius complex due the angulation of clypeo-genal junction (Fig.1.2). In ascanius complex, D. sp.3 can be separated of macropterous species by the characters related to brachypterism (as D. sp.2) (Figs. 1.4 and 1.6) . Between brachypterous of this complex, D. sp.3 can be differentiated by bearing dense and strong ocellate puncture at anterior lobe of pronotum (Fig.2.16) and clypeo- frontal carina, medially, with feeble emargination (Fig.2.15). Description of holotype: Male. Lenght: 19.5 mm. Maximum width (pronotum): 12.5 mm. Black, shining. Head: surface smooth, ocellate punctures present along intra- ocular space. Clypeogenal junction with obtuse angle. Clypeal teeth separated by a strong V-shaped emargination. Dorsally, apex of clypeal teeth bearing bunch of setae. Ventral clypeal process feebly bifurcated (? possibly broken). Clypeo-frontal carina higher than wide, with feeble central emargination at apex. Clypeo-genal suture distinct. Small knob present on posterior portion of clypeal suture. Antennal club with light red tumescence. Pronotum: wider than long and wider than elytra. Anteriorly, with strong declivity and a central lobe delimited by two knobs. Anterior central lobe with strong and dense ocellate punctures. Pronotal disc smooth, shnining. Ocellate punctures present on pronotal posterior border, near anterior angles and on anterior central lobe. Anterior angle acute. Hypomeron: anterior portion covered with ocellate setose punctures. Posterior portion with sparse setose ocellate punctures. Hypomeral disc shining and glabrous, except on lateral margin, with some sparse setae. Prosternum: glabrous, with ocellate punctures near posterior margin. Mesosternum: very narrowed and glabrous in the middle. Ocellate short-setose punctures 47 present on the lateral portion. Mesepisternum: whole structure with ocellate short-setose punctures – length of this setae reach one quarter the length of hypomeral setae. Superior margin of posterior portion with a triangular projection that almost reach the pseudoepipleuron. Metasternum: with mesometasternal suture indistinct. Between mesocoxae, with almost twice the width of medial portion. Borders of anterior part with ocellate punctures. Long and sparse setae present along lateral margins of metasternal lobe. Mestasternal disc with longitudinal sulcus and a feeble posterior concavity. Fine punctures near concavity. Elytra: shining. Lacking humeral callus. Discal striae with ocellate puncutes, separated by twice its diameter and sub-equal to striae widht. Lateral (ninth) striae not reaching elytral basis. Elytral interstriae feebly convex. Legs: anterior tibial spur strongly curved inward at apical fourth. Anterior tibiae with three well defined teeth, and a fourth basal one feebly indicated by a sinuosity. Ventral surface of anterior femur smooth, glabrous. Length of 5th hind tarsomere two times the length of 4th hind tarsomere. Apical median and hind tarsomeres with a small spur with the same length of tarsal claws. Abdomen: all sternites glabrous and without punctures. Pygidium: shining, smooth, wider than long. Phallobasis: dorsally with apical border strongly curved, angulation present. Parameres: dorsally symmetrical. Laterally, brusquely flattened beyond the apical half. Anterior to this excavation, dorsally, parameres bears a distinct fovea. Dorsally, these foveae almost reach the lateral margin of parameres. Apex with right angle. Laterally, longitudinal sulcus parallel to lateral margins (Fig.2.17). Morphological variation: size varies from 18.5 to 19.5 mm in males and from 18.0 to 21.0 mm in females. Pronotum width varies from 12.0 to 12.5 mm in males and from 11.5 to 12.0 mm in females. Both males and females have variations in following characters: Head: clypeo frontal carina higher or lower in some individuals. Central emargination of clypeo-frontal carina can be more or less distinct. Pronotum: anterior declivity as well pair of central gibosities may vary in size. In most females, the declivity is less conspicuous and there is one central knob instead of a pair. Elytra: both males and females have variations on the shining of elytra base, that can be black or brown. Abdomen: Only on females, apex of sixth abdominal segment is prolonged beneath pygidium apex. This prolongation results in two medial tubercles separated by a feeble emargination. Distribution: Cerrado of south-west of Goiás, at Chapadão do Céu, Mineiros municipality. The whole type series was collected at Parque Nacional das Emas, a major conservation unit of Brazilian Cerrado (Fig.4). Etymology: Named after Charles Darwin (1809 – 1882), one of the greatest naturalists of all time. This name is a sincere homage to Darwin who had a fondness for scarab beetles and, on his Origin of Species (Darwin 1859), makes brief comments about tarsal reduction on some Scarabaeinae (genus Scarabaeus and Onitis ). Coincidence or not, in the same chapter of Origin of Species (Chapter V, Effects of use and disuse), Darwin lists several cases of brachypterism on animals. Discussion: this species belongs to ascanius complex and seems to be close to D. periotoi (described above). D. sp.3 type-series were collected with pitfall traps, without baits or with human feces bait, always in open environments (Campo sujo and Cerrado sensu stricto , sensu Ribeiro & Walter (1998)). Like D. sp.2 and other brachypterous species of ascanius complex, proportion of females is equal to males. Despite its populations can be found in a major conservation unit (Emas National Park), the narrow distribution associated with growing expansion of agriculture on natural habitats regards D. darwini , according to IUCN criteria (IUCN 2010), as a threatened species (unpublished data).

Dichotomius (Selenocopris) sp.4 n. sp. (Fig.1.7) Material studied [collection deposited]: Holotype: (male) BRASIL: Mato Grosso: Tangará da Serra. Fazenda Aparados da Serra. 14°21’14”S; 57°43’54”W. Human feces. 17-I-2010. RJSilva (leg.) [CEMT]. Paratype: 1 male, 1 female, same data of holotype [CEMT]; 1 male, 1 female, same data of holotype [CEMT]; 6 males Tangará da Serra/MT_2008. Cerrado sensu stricto. Pitfall s/ isca. Tombo 13 [CEMT]; 1 male, 1 female, same data of anterior [CEMT]; 1 female, same data of anterior [CEMT]; 1 male, BRASIL: Mato Grosso. Tangará da Serra. Fazenda Aparados da Serra. 14°22’01”S; 57°43’30”O. 575mosl. Cerrado s. str. Pitfall s. isca. 8-16-II-2008 [CEMT]; 1 male, 1 female, same data of anterior [CEMT]. Diagnostic remarks: this species belongs to the ascanius complex due the angulation of clypeo genal junction(Fig.1.2). In this complex, D. sp.4 can be promptly separated from other species by the puncture on posterior portion of pronotal disc (Figs.1.7 and 1.8) and lacking of punctures on elytral striae (Fig.1.9). Between brachypterous Selenocopris , females of D. sp.4 bears the most developed tubercles on the prolongation of sixth abdominal segment(Fig.1.10). Description of holotype: Male. Length: 17.0 mm. Maximum width (pronotum): 10.5 mm. Black, shining. Head: surface smooth, ocellate punctures present along intra- ocular space. Clypeogenal junction with obtuse angulation. Clypeal teeth separated by a strong V-shaped emargination. Dorsally, apex of clypeal teeth bearing bunch of setae. Ventral clypeal process strongly bifurcated, forming two distinct teeth. Clypeo-frontal region with a single tubercle. Clypeo-genal suture distinct. Small knob present on the posterior portion of clypeo genal suture. Antennal club with dark brown tumescence. Pronotum: wider than long and sub equal to elytra in width. Anteriorly, with strong declivity delimited, superiorly, by a feeble emargination forming a central lobe. Central lobe filled with ocellate punctures. Pronotal disc shining and smooth anteriorly. Posterior region of the disc, mainly on lateral sides, with ocellate punctures with the same size of those on margins, anterior angle and central lobe. Anterior angle truncated. Hypomeron: with a lateral band of ocellate punctures bearing dark brown pilosity, wider anteriorly and posteriorly, and reduced to a single row medially. Hypomeral disk glabrous, shining. Prosternum: with ocellate-short setose punctures – length of these setae reach one quarter the length of hypomeral setae. Mesosternum: very narrowed, glabrous and shining medially. Ocellate punctures present laterally. Mesepisternum: whole structure with ocellate punctures, evenly distributed. Superior margin of posterior portion with projection that almost reach the pseudoepipleuron. Metasternum: mesometasternal suture distinct. Ocellate punctures present on the margins of anterior portion of the lobe. Anterior lobe and metasternal disc glabrous. Metasternal disc with feeble longitudinal sulcus and a feeble posterior concavity. Elytra: black, shining. Striae wide, without punctures, at most with some indication of points near elytra apex. Striae becomes wider near the elytra basis. Lateral (ninth) striae reaching elytral basis. Elytral interstriae feebly convex. Legs: anterior tibiae spur curved inward at apical third. Anterior tibiae with four teeth, Ventral surface of anterior femur smooth, glabrous. Length of 5th metatarsomere two times the length of 4th metatarsomere. Apical meso and metatarsomeres with a small spur with the same length of tarsal claws. Abdomen: all sternites glabrous. Pygidium: shining, smooth, wider than long. Phallobasis: dorsally with apical border strongly curved with angulation present. Parameres: dorsally symmetrical. Laterally, strongly flattened at apical half. Anterior to this excavation, dorsally, parameres bears a distinct fovea. Apex with right angle. Laterally, longitudinal sulcus parallel to lateral margins (Fig.1.11). Morphological variation: length varies from 17.0 to 17.5 mm in males and from 17.5 to 18.0 mm in females. Pronotum width varies from 10.5 to 11.5 mm in males and from 10.5 to 12.0 mm in females. Both males and females show variations in following 49 characters: Head: height of central conical horn on clypeo-frontal carina. Pronotum: anterior declivity as well pair of central gibosities may vary in size. In most females, the declivity is lower and central emargination delimiting exacation might be more or less distinct. Also in females, ocellate punctures at anterior lobe are smaller (3/4 the diameter of male punctures) and less dense. Elytra: both males and females have variations on the color of elytra base, that can be black or brown. Abdomen: only on females, apex of sixth abdominal segment is prolonged beneath pygidium apex. This prolongation results in two medial tubercles separated by a strong emargination. Distribution: Cerrado of Chapada dos Parecis, Mato Grosso. All type-series known from the same locality (Nova Marilândia municipality) (Fig.4). Etymology: Appositive substantive. Homage to Paresi tribe, originally found at Chapada dos Paresis and that was brutally exterminated by Bandeirantes colonizer, originating from São Paulo, Brazil. Currently, some ethnics groups of this tribe are confined in some Indigenous Reserves, which also ensure the conservation of some D. paresi populations. Discussion: this species belongs to the ascanius complex and seems to be isolated in this group, due the pronotal disc punctures and enlarged elytral striae. All known specimens were collected in pitfall traps, without baits or baited with human feces, always in Cerrado sensu stricto environment (sensu Ribeiro & Walter 1998). Due to the restriction on its distribution, quality of habitat (constant pressures of agriculture expansion), D. sp.4 can be considered a threatened species in accordance to IUCN criteria (IUCN 2010) (unpublished data).

Dichotomius (Selenocopris) sp.5 n. sp. (Fig.3.18) Material studied [collection deposited]: Holotype: (male) Serra da Canastra, M.G BRASIL / OUMNH-2003-051 W.D.Hamilton Colln. [OUMNH]. Diagnosis: this species belongs to the ascanius complex due the angulation of clypeo-genal junction (Fig.1.2). Within this complex, D. sp.5 can be differentiated of macropterous species due the characters associated to the brachypterism (Figs. 1.4 and 1.6). Within the brachypterous species in this complex, D. sp.5 is the largest species and is the unique that combine the following characters: transverse cephalic carina feebly emarginated (Fig.3.20); dorsally, foveae of paramera small and not widened (Fig.3.21) and anterior lobe of pronotum filled with small and sparse punctures (Figs.3.18 and 3.19). Description of holotype: Male. Length: 25.0 mm. Maximum width (pronotum): 14 mm. Black, shining. Head: surface smooth, ocellate punctures present near both eyes. Clypeogenal junction with obtuse angulation. Clypeal teeth separated by a strong V-shaped emargination. Dorsally, apex of clypeal teeth bearing bunch of setae. Ventral clypeal process strongly bifurcated, forming two distinct teeth. Clypeo-frontal region with a transversal carina feebly emarginated. Clypeo-genal suture distinct. Antennal club with light brown tumescence. Pronotum: wider than long and wider than elytra. Anteriorly, with strong declivity delimited, superiorly, by two weak developed knobs. Central anterior lobe filled with small and sparse non-ocellate punctures. Pronotal disc shining and smooth on all its extension. Anterior angle truncated. Hypomeron: with a lateral band of ocellate punctures bearing dark brown pilosity, wider anteriorly and posteriorly, and reduced to a single row medially.Posteriorly, most of the punctures do not bear pilosity. Hypomeral disk glabrous, shining. Prosternum: with ocellate punctures. Mesosternum: very narrowed, glabrous and opaque medially. Ocellate punctures present laterally. Mesepisternum: whole structure with ocellate punctures, evenly distributed. Superior margin of posterior portion with projection that almost reach the pseudoepipleuron. Metasternum: mesometasternal suture distinct. Ocellate punctures bearing dark brown pilosity present on the margins of the lobe. Metasternal disc glabrous, with a feeble longitudinal sulcus and a feeble posterior 50 concavity. Elytra: black, shining. Striae with punctures spaced by five times it diameter. Width of points is subequal to striae width. Elytral interstriae feebly convex. Legs: anterior tibiae spur curved inward at apical fifth. Anterior tibiae with four teeth, Ventral surface of anterior femur smooth, glabrous. Length of 5th metatarsomere two times the length of 4th metatarsomere. Apical meso and metatarsomeres with a small spur with the same length of tarsal claws. Abdomen: all sternites glabrous. Pygidium: shining, smooth, wider than long. Phallobasis: dorsally with apical border strongly curved with angulation present. Parameres: dorsally symmetrical. Laterally, brusquely flattened at apical half. Anterior to this excavation, dorsally, parameres bears a distinct fovea that never reach laterals of paramera. Apex with right angle (Fig.3.21). Distribution: One specimen known from Serra da Canastra, Minas Gerais, Brazil (Fig.4). Etymology: Named after Darren Mann, entomologist and Collection Manager of the Hope Entomological Collections at OUMHN, where the unique exemplar of D. manni is deposited and was kindly borrowed. Discussion: this species belongs to the ascanius complex and it seems to be morphological similar to D. sp.2, described above. We described this species from a single specimen and thus, it is not possible to take conclusions about its distribution, morphological variation and conservation status.

Redescription of Dichotomius ingens (Luederwaldt 1935) (Fig.3.22) Pinotus ingens , Luederwadlt 1935: 341. Pinotus ingens , Blackwelder 1944: 207. Dichotomius (Selenocopris) ingens, Vaz-de-Mello 2000: 195.

Material studied [collection deposited]: Holotype: 1 male, Chapada Brasilien, 2.600 Fuss, XI.1902, A. Robert leg. [BMNH]; 1 male, BRASIL: Mato Grosso. Rosário Oeste. Maller leg. Coll. Yarh’nez. Sept 1961 / Dichotomius (S.) ingens (Lued.). A.Martínez-Det. 1963 / Ingens. Det. F. Génier 2000 / Espéce Microptérr. Dét. F. Genier, 1999 / H & A Howden Collection. Ex A. Martínez coll. / Canadian Museum of Nature. LP2010-0080. [CMNC]; 1 female, no data [CEMT]; 1 male, BRASIL: Mato Grosso. Cuiabá. Fazenda Mutuca. Mata 4. 2-II-2009 [CEMT]; 1 female, BRASIL: Mato Grosso . Chapada dos Guimarães. Local não definido entre o portão do inferno e a cidade. Coleta Manual. WOSouza. 2002 [CEMT]; 1 male, BRASIL: Mato Grosso. Cuiabá. Fazenda Mutuca. 15°18’08”S; 55°57’54”W. Pitfall pulmão bovino. III-2011. LGOANunes [MNRJ]; 1 female, BRASIL: Mato Grosso. Cuiabá. Fazenda Mutuca. 15°18’25”S; 55°58’07”W. Cerrado chaquenho. Pitfall baço bovino. II-2011. LGOANunes [CEMT]; 2 males, 1 female, BRASIL: Mato Grosso . Chapada dos Guimarães. Local não definido entre o portão do inferno e a cidade. Coleta Manual. WOSouza. 2002 [CEMT]; 1 male, BRASIL: Mato Grosso. Cuiabá. Fazenda Mutuca. 15°18’25”S; 55°58’07”W. Cerrado chaquenho. Pitfall baço bovino. II-2011. LGOANunes [CEMT]; 1 male, 4 females; BRASIL: Mato Grosso. Cuiabá. Fazenda Mutuca. 15°18’08”S; 55°57’54”W. Pitfall pulmão bovino. III-2011. LGOANunes [CEMT]; 1 female, BRASIL: Mato Grosso do Sul. Dourados. Jardim Europa. 25-VI-2007. VHoras leg. [CEMT]. Diagnosis: this species belongs to the ascanius complex due the angulation of clypeo-genal junction (Fig.1.2). Among brachypterous species of this complex, D. ingens can be separated by combination of conical cephalic horn (Fig.3.23) and pronotum with ocellate punctures on posterior margin, near anterior angle and central lobe (Fig.3.22). Redescription: Male. Length: 21.0 mm. Maximum width (pronotum): 12.5 mm. Black, shining. Head: surface smooth, ocellate punctures present along intra ocular space. Clypeal junction with obtuse angle, almost right. Clypeal teeth separated by a strong V- 51 shaped emargination. Dorsally, apex of clypeal teeth bearing bunch of setae. Ventral clypeal process strongly bifurcated, forming two distinct teeth. Single cephalic horn. Clypeo-genal suture distinct. Small knob present on posterior portion of clypeo-genal suture. Antennal club with light red tumescence. Pronotum: wider than long and wider than elyra. Anteriorly, with strong declivity and a central lobe delimited by two median gibosities. Anterior lobe filled with ocellate punctures. Pronotal disc smooth, shining. Ocellate punctures present on pronotal posterior border, near anterior angles and on anterior lobe. Anterior angles truncated. Hypomeron: with a lateral band of ocellate punctures bearing dark brown pilosity, wider anteriorly and posteriorly, and reduced to a single row medially. Hypomeral disk glabrous, shining, smooth. Prosternum: posteriorly, with ocellates short-setose punctures – setae have one quarter the length of the hypomeral setae). Mesosternum: very narrowed, glabrous and shining medially. Ocellate punctures present laterally. Mesepisternum: ocellate short-setose punctures evenly distributed. Superior margin of posterior portion with projection that almost reach the pseudoepipleuron. Metasternum: with indistinct mesometasternal suture. Between mesocoxae, with almost the twice width of medial portion. Ocellate punctures present on borders of anterior portion of the lobe. Long sparse setae present along the margin of mestasternal lobe. Mestasternal disc lacking longitudinal sulcus and with a feeble posterior concavity. Elytra: shining. Striae with ocellate punctures separated by one half its diameter and quite wider than striae. Lateral (ninth) striae not reaching elytral basis. This striae is distinct only since the basal fifth. Elytral interstriae feebly convex. Legs: anterior tibial spur strongly curved inward at apical fourth. Anterior tibiae with four lateral teeth. Basis of median and hind tibiae sinuous, with a brush of red setae. Length of 5th metatarsomere two times and a half the length of 4th metatarsomere. Apical Meso and metatarsomeres with a small spur with the same length of tarsal claws. Abdomen: all sternites glabrous. Pygidium: shining, glabrous, smooth, wider than long. Phallobasis: dorsally with apical border strongly curved, angulation absent. Parameres: dorsally symmetrical. Laterally, brusquely flattened since the basal third. Anterior to this excavation, dorsally, paramera bears a distinct fovea. Apex with right angle. Laterraly, longitudinal sulcus parallel to lateral margins (Fig.3.24). Morphological variation: length varies from 18.0 to 21.0 mm in males and from 18.5 to 19.0 mm in females. Pronotum width varies from 12.0 to 12.5 mm in males and from 11.0 to 11.5 mm in females. Males cephalic horn on the head can vary in size as well the central pair of gibosities and anterior declivity on pronotum. Both in males and females, elytra basis can be black, brown or blue. Females differ in the following characters: Head: front transversely ridged. Cephalic horn lower or transversal carina with a feeble central emargination. Pronotum: anterior declivity as well pair of gibosities less developed. Ocellate punctures at anterior lobe denser and more conspicuous. Abdomen: Apex of sixth abdominal segment is prolonged beneath pygidium apex. This prolongation results in two medial tubercles separated by a feeble emargination.. Distribution: Chapada dos Guimarães (Rosário Oeste, Chapada dos Guimarães and Cuiabá municipalities) and surrounding areas at Baixada Cuiabana (Cuiabá municipality), Mato Grosso (Fig.4). Baixada Cuiabana localities where D. ingens is found comprise lower areas (at most 250 meters over sea level) while Chapada dos Guimarães and Rosário Oeste localities comprises higher areas (600-800 meters), including Chapada dos Guimarães National Park. We have a dubious records, one female from Mato Grosso do Sul state, Dourados municipality (about 900 km from where other specimens were found). After examining the external morphology of this female, we believe that this record is wrong. First, because scarab fauna of this region is well known and there are no records for D. ingens. Second, Dourados is far enough from Chapada dos Guimarães to avoid the

52 dispersal of a flightless species. As what happens with other flightless Scarabaeinae, distribution of D. ingens is very restricted. Discussion: this species belongs to the ascanius complex. D. ingens has been collected with pitfall traps baited with human dung and carrion. We also collected manually some exemplars during the day near bovine carcass (during a field experiment). This species is found in open environments like Cerrado sensu stricto (sensu Ribeiro & Walter 1998). Among flightless Selenocopris , this species has the largest extension of preserved environment (both on local farms as well in National Park). However, due to its narrow geographic range, it can be considered as threatened in accordance to IUCN criteria (IUCN 2010) (unpublished data).

Acknowledgments Darren Mann, collection manager of OUMNH for sending the specimen of D. manni. Helder Borges Faria (Universidade Federal de Mato Grosso) for photographing the dorsal habitus of all species in this work. Dra. Helena Onody (MZSP) for providing information about localities of D. periotoi . Mateus Souza (CEMT) for collecting valuable exemplars of D. darwini . Graciela de Oliveira (Universidade Federal de Mato Grosso) for providing information about Paresi tribe at Mato Grosso. RVN receives a CAPES (Comissão de Aperfeiçoamento de Pessoal de Nível Superior) grant. FZVM is a CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) fellow and most of this work was granted by FAPEMAT (Fundação de Amparo à Pesquisa do Estado de Mato Grosso FAPEMAT, 570847/2008, 447441/2009, 283731/2010).

References Blackwelder, R.E. 1944. Checklist of the Coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 2. United States Natl. Mus. Bull., 185: 189-341.

Cavassan, O. 2002. O cerrado do Estado de São Paulo. In: Klein, A.L. (Org.). Eugen Warming e o cerrado brasileiro - um século depois. 1 ed. São Paulo: Editora Unesp - Imprensa Oficial SP, v. 1, p. 93-106.

Darwin, C. 1859. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. London: John Murray. 502 p.

IUCN - International Union for Conservation of Nature. 2010. Guidelines for Using the IUCN Red List Categories and Criteria, version 8.1 . IUCN Standards and Petitions Subcommittee, 85 pp. Avaliable from: http://intranet.iucn.org/webfiles/doc/SSC/RedList/RedListGuidelines.pdf/.(18/10/2011).

Luederwaldt, H. 1929. As espécies Brasileiras do gênero Pinotus . Revista do Museu Paulista , Tomo XVI, 1-173.

Luederwaldt, H. 1935. Zur kenntnis der gattung Pinotus . Revista de Entomologia , 5(3), 334–342.

Ribeiro, J.F.; Walter, B.M.T. 1998. Fitofisionomias do bioma cerrado. In: Sano, S.M.; Almeida, S.P (Eds). Cerrado: ambiente e flora . Planaltina: EMBRAPA-CPAC. pp.89-166.

Vaz-de-Mello, F.Z.; Edmonds, W.D.; Ocampo, F.C.; Schoolmesters, P. 2011. A multilingual key to the genera and subgenera of the subfamiliy Scarabaeinae of the new world (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa , 2854: 1-73. 53

Vernalha, M.M. 1952. Descrição de uma espécie nova do gênero Dichotomius, Hope 1838 (Col. Scarabaeidae). Arquivos de Biologia e Tecnologia , v. 7, n. 8, p. 39-44.

Figure 1. 1 . Arrows indicating the lacking of angulationon the clypeo-genal junction on the species of bicuspis complex. 2. Arrows indicating the angles on the Clypeo-genal junction on the species of ascanius complex. 3. Arrow indicating the humeral callus on the macropterous species of Selenocopris . 4. Arrow indicating the absence of humeral callus on the brachypterous species of Selenocopris. 5. Arrows indicating the equally on the width of pronotum and elytra. Elytra feebly convex. 6. Arrows indicating the pronotum 55 wider than elytra. Elytra strongly convex. 7. Dorsal habitus of Dichotomius sp.4 n. sp. 8. Arrows indicating the ocellate punctures on the pronotal disc of D. sp.4. 9. Arrows indicating enlarged striae on the elytra of D. sp.4. 10. Arrows indicating the prolongation of last abdominal sternite beyond the apex of pygidium on a female of D. sp.4. 11 . Dorsal view of D. sp.4 paramera.

Figure 2. 12 . Figure 2. 12 . Dorsal habitus of Dichotomius sp.2 n.sp. 13. Head transversal carina of D. sp.3. 14 . Dorsal view of D. sp.3 paramera. 15. Dorsal habitus of Dichotomius sp.3 n. sp. 16. Arrows indicating ocellate punctures on the anterior lobe of the pronotum of D. sp.3. 17 . Dorsal view of D. sp.3 paramera.

Dorsal habitus of Dichotomius sp.5 n. sp. 19. Arrow indicating sparse and small ocellate punctures on the anterior lobe of the pronotum of D. sp.5. 20. Head transversal carina of D. sp.5. 21. Dorsal view of D. sp.5 paramera. 22. Dorsal habitus of Dichotomius ingens (Luederwaldt, 1935). 23. Conical horn on the head of D. ingens . 24. Dorsal view of D. ingens paramera.

Figure 4 . Distribution map of Dichotomius ingens, D. sp.5 , D. sp.2 , D. sp.4 and D. sp. 3.

CHAPTER FOUR: Conservation of the brachypterous species of the genus Dichotomius Hope (Coleoptera: Scarabaeidae: Scarabaeinae): Conservation status, Environmental legislation and Management Tactics.

ABSTRACT: Assessment of conservation status of biological species is one of the most important tools used in the science of Biological Conservation. This tool can help the management of species of insects, especially those with restricted distribution and thus, with populations more susceptible to extinction, case of brachypterous species of Dichotomius (Coleoptera: Scarabaeidae: Scarabaeinae). Our objectives in this chapter were: 1) Assess and discuss the conservation status of brachypterous species of the genus Dichotomius ; 2) discuss how environmental laws may affect the conservation and management of brachypterous species of Dichotomius and 3) propose management tactics for the studied species. Among species accessed in this work, Dichotomius comarapensis and D. sp.5 n.sp. could not be evaluated due the lack of data (DD). D. ingens and D. sp.4 n.sp. were evaluated as Near Threatened (NT) while D. sp.3 n. sp. was classified as Vunerable (VU). D. eucranioides and D. sp.1 n.sp. were considered Endagered (EN) and D. sp.2 n.sp. and D. mysticus were classified as Critically Endangered (CR). Part of environmental legislation has a seminal role in protecting the remnant populations of these species however some laws, like Brazilian Forest Code are not being observed. We also propose some possible management tactics for conserving brachypterous species of the genus Dichotomius.

RESUMO: O uso e avaliação do status de conservação de espécies biológicas é uma das ferramentas mais importantes utilizadas na ciência da Biologia da Conservação. Essa ferramenta pode auxiliar o manejo de espécies de insetos, especialmente aquelas que possuem distribuição restrita e que, por sua vez, estão mais suscetíveis a extinção, caso das espécies braquípteras de Dichotomius (Coleoptera; Scarabaeidae: Scarabaeinae). Nossos objetivos nesse capítulo foram: 1) Acessar o status de conservação das espécies braquípteras do gênero Dichotomius; 2) discutir como as leis ambientais afetam a conservação e manejo dessas espécies e 3) propor táticas de manejo para as espécies estudadas. Entre as espécies acessadas nesse trabalho, Dichotomius comarapensis e D. sp. 5 sp. nov. não puderam ser avaliadas devido a falta de informações (DD). D. ingens e D. sp.4 sp. nov. foram avaliadas como Quase Ameaçadas (NT) enquanto D. sp.3 sp. nov. foi classificada como Vunerável (VU). D. eucranioides e D. sp.1 sp. nov. foram consideradas como Ameaçadas (EN) e D. sp.2 sp. nov. e D. mysticus foram classificadas como criticamente ameaçadas (CR). Parte da legislação ambiental tem um papel fundamental na proteção das populações remanescentes, entretanto, algumas leis como o Código Florestal

Brasileiros não vem sendo observadas e cumpridas. Nesse trabalho, nós também propomos táticas de manejo para a conservação das espécies braquípteras do gênero Dichotomius .

1. INTRODUCTION

On the science of Biological Conservation, extinction is a key concept to understand the procces of biodiverity loss (EHRLICH; MOONEY, 1983; WILSON, 1989) and that has been much debated since 80's by many authors. Part of this debate begets new methodologies that help conservation scientists to understand and deal with extinction process. The study of minimum viable populations (GILPIN; SOULÉ, 1986), distribution predictive models (LOISELLE et al., 2003), monitoring of bioindicators (HALFFTER; FAVILLA, 1993; SPECTOR, 2006) and conservation status (IUCN, 2012) are some examples. In 1994, the International Union to Conservancy of Nature - IUCN published the first version of IUCN Red List Categories and Criterias (IUCN, 2012). The main objective of this document is to define clear and transparent ways to access species conservation status. Since them, Red List contained assessments for almost 60.000 species, yet a pitiful part of known biodiversity. On Coleoptera, the major order of nature (VANIN; IDE, 2002), until today, 212 assessments were done, many of them need updates or are deficiency on data (IUCN, 2012). The red list is an important initiative since it can be a start point for planning and acting the conservation of species, their environment and, indirectly the conservation of population of other species. Dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) are present on almost all terrestrial environments and having a great diversity in the tropics (HALFFTER; MATTHEWS, 1966). Dung beetles are essential to terrestrial ecosystems since they use, for feeding and nesting, mammals dung, promoting nutrient cycling on ecosystems (NICHOLS et al., 2008). Beyond a myriad of ecological functions provided by Scarabaeinae (NICHOLS et al., 2008), these group has being recently used to indicate the presence of mammals on environments and the health of ecosystems (ANDRESSEN; LAURANCE, 2007; NICHOLS et al., 2007; NICHOLS et al., 2008). One of many endemic genera at Neotropical Region, Dichotomius comprises about 170 species divided in four subgenera, being a very common genus in the inventories done in the American continent (KOHLMAN;SOLÍS, 1997; VAZ-DE- MELLO, 1999; VAZ-DE-MELLO, 2000; MEDINA et al., 2001; PERAZA; DELOYA,

2006; ALMEIDA; LOUZADA, 2009). In Dichotomius , some species are brachypterous, which means that they have the atrophy of hind (or membranous) wings making them unable to fly and, consequently, restricting their distribution. Currently, brachypterous Dichotomius are being revised by us and, as a complement for this revision, we decided to deal with conservation of brachypterous Dichotomius for the following reasons: there is no assessments for Scarabaeinae dung beetles in IUCN Red list, while a single species on the subfamily, Dichotomius shiffleri, is assessed on the Brazilian red list (VIEIRA et al., 2011); brachypterous Dichotomius have very restricted geographic distribution, seeming to be endemic of respective type localities and being more susceptible to losing its populations; these endemic species may indicate areas of endemism and, at last, some insect species are very sensible to ecosystem changes, thus, their conservation represents the conservation of whole environments. In this work we had three aims: 1) Assess and discuss the conservation status of brachypterous species of the genus Dichotomius ; 2) discuss how environmental laws may affect the conservation and management of brachypterous species of Dichotomius and 3) propose management tactics for the studied species.

2. MATERIAL AND METHODS Species dealt here were theme of a taxonomic review named: review of brachypterous species of the genus Dichotomius Hope. Conservation status of the species were assessed according IUCN guidelines and IUCN guide for criteria and categories, both found on IUCN website (IUCN, 2012). Discussion on environmental law was mostly based on Brazilian laws, since the major part of brachypterous Dichotomius occurs exclusively in Brazil. For this, we consulted legislation found on Portal da Legislação (PLANALTO, 2012).

3. RESULTS AND DISCUSSION 3.1. Dichotomius comarapensis Génier, 2000 – Data Deficient (DD) (Figure 1). Last record: 2000. The known specimens were collected in a single locality (GÉNIER, 2000) while there are no dung beetle surveys or samplings at surrounding areas, making impossible to determine if this species is rare or if the region is poor sampled. This species occurs at high altitudes (1.800 meters) at the region of Comarapa, Bolivia, what may difficult the human access to these areas and consequently the occupation and degradation of them. Comarapa region also bears a critically endangered rodent, known only from a

64 near locality – Abrocoma boliviensis (IUCN, 2012). The same authors emphasizes that this region has no protected areas, what demands attention in relation to the populations of D. comarapensis . We propose that more samplings should be done in the region of Comarapa in order to increase the knowledge on the distribution of this species.

3.2. Dichotomius eucranioides Pereira and D’Andretta, 1955 – Endangered (EN), B1,a,b(i,ii,iii); B2,a,b(i,ii,iii) (Figure 1). Last Record: 2004. Restricted to gallery forests at the left margin of Parana River, state of Mato Grosso do Sul, between the municipalities of Três Lagoas and Selvíria, spaced approximately 80 kilometers. Besides the restricted area and extent of occurrence for this species, gallery forests of Parana River are severely fragmented due the predatory agriculture, urban occupation and the construction of hydroelectric power sources (POLETO et al., 2004). Brazil Forest Code predicts the maintenance of 500 meters of native vegetation on the margins of the rivers with more than 600 meters width (BRASIL, 1965). This area is called "Área de Preservação Permanente - APP" (Permanent Preservation Areas). Despite this legal determination, a brief look on satellite images reveals that few proprieties respected this regulation leaving D. eucranioides at few forest remnants. Currently, Brazilian Congress is in imminence to approve new regulations on the measures of APP length, what may reduce even more the protected native area for D. eucranioides . DEVELEY and PONGILUPPI 2010 comment on the possible negative impacts on Bird fauna if the changes on the Forest Code been accepted. Besides the necessary respect to Brazilian legislation by local farmers, we propose that the remnant populations of D. eucranioides should be monitored with capture- marking procedures in order to estimate the real size of population.

3.3. Dichotomius mysticus (Luederwaldt, 1935) – Critically Endangered (CR), B1, a,b(i,ii,iii), B2,a,b(i,ii,iii) (Figure 1). Last Record: 2011. Despite similar environments has been sampled at this region, this species has been recorded only in Parque Estadual das Dunas, inside the city of Natal, state of Rio Grande do Norte. It occurs associated to forest coastal vegetation, severely fragmented due tourism activities and urban expansion (JESUS, 2003). The creation of this park on 70's (RIO GRANDE DO NORTE, 1977) is seminal to the conservation of D. mysticus . The consolidation of “Sistema Nacional de Unidades de Conservação da Natureza – SNUC (National System of Natural Conservation Units) in Brazil (BRASIL, 2000) is also very important to the effectiveness of the conservation of local fauna in this and other stadual parks. However, the area of Parque

Estadual das Dunas is very small (1.172 hectares) to support viable populations on long terms. We propose that designation of new conservation areas in this region and estimate the size of population on mark-capture procedures.

3.4. Dichotomius sp.1 n.sp. – Endangered (EN), B1,a,b (ii,iii), B1,a,b(ii,iii) (Figure 1). Last Record: 2008. D. sp.1 extant and area of occurrence is located inside a Ramsar Site, humid areas known for bearing high biodiversity (BRASIL, 1996). Distribution of this species is very restricted and, despite it is found in a international preserved area, the quality of habitat may be low in some sites according to Vidaurre et al (2008). The Ramsar convention, created to protected humid areas of high diversity is being decisive to the conservation of this species. We propose the maintenance of protected are in order to ensure the conservation of this species.

3.5. Dichotomius ingens (Luederwaldt, 1935) – Near Threatened (NT) (Figure 1). Last Record: 2011. Despite having a restricted distribution, fitting on criteria B1 and B2 of vulnerable (VU) status , almost all known specimens can be found inside National Park of Chapada dos Guimarães, what makes this species having its whole distribution protected by a Conservation Unit. On the other hand, constant fire events on this rartea (FÁVERO et al., 2010) may reduce adult populations of this species. Chapada dos Guimarães National park was created in the 80's and has 32.630 hectares (BRASIL, 1989) while there are still many surrounding particular areas with native vegetation. The consolidation of the National Conservation Units System - SNUC is seminal to the effective protecting of this species since this area has a strong tourism and agricultural activity. Since the preservation of D. ingens area of occurrence is ensured, we propose that the effects of fire on the populations of D. ingens should be studied, these data would be useful for other species of Scarabaeinae and animals.

3.6. Dichotomius sp.4 n. sp. – Near Threatened (NT) (Figure 1). Last record: 2010. Our choice for this status follows the same reasons of D. ingens . Almost all extant of occurrence and predicted occurrence of D. sp.4 is located at a complex of Indigenous Reserves at the region of Chapada dos Parecis, in the state of Mato Grosso, Brazil. Despite being in a relative well protected area, agriculture expansion (SOUZA; MARTINI, 2000) and the disrespect of Legal Reserves (natural remnants that farmers should keep in their properties according Brazilian Forest Code) are the major threats for this species. We

66 propose that indigenous reserves should maintain their native remnants of Cerrado and a long-term monitoring with capture-marking procedures in order to estimate if there is declining on populations.

3.7. Dichotomius sp.2 n.sp. – Critically Endangered (CR), B1, a,b(i,ii,iii), B2,a,b(i,ii,iii) (Figure 1). Last record: 2004. This is species is only found in the municipality of Descalvado, state of São Paulo, where 87 % of Cerrado vegetation was lost (CAVASSAN, 2002). Despite being near a well studied Ecological State on the same region (Itirapina Ecological Station, about 30 km from Descalvado), there are no records for D. sp.2 there. The total disrespect of Legal Reserve legislation, found on Brazilian Forest Code (BRASIL, 1965) associated to agriculture and urban expansion are the causes for D. sp.2 situation. However, since this species can be found in disturbed environments (Cerrado with citrus plantation), we believe that the recuperation of disturbed Legal Reserves on the region may help increasing populations of D. sp.2 . We also propose mark-capture procedures in order to estimate the size of population.

3.8. Dichotomius sp.3 n.sp. - Vulnerable (VU), B1,b(iii),c(ii); B2,b(iii),c(ii) (Figure 1). Last record: 2011. Like D. ingens , D. darwini populations are protected by a National Park. However Emas National Park is surrounded by soybean plantations and there are constant events of fire while there is the presence of exotic vegetation (BATALHA; MARTINS, 2004). The consolidation of SNUC (BRASIL, 2000) is decisive to preserve the populations. However, it is important to remember that the neighborhood of Emas National Park is fully cultivated with soil bean, what lead us to conclude that this species is isolated and the Park. Despite Emas has more than one hundred thousand hectares, it is the last remnant for populations of D. sp.3. We propose that the effect of fire and exotic plant invasions should be studied in order to help the management of this species.

3.9. Dichotomius sp.5 n.sp. – Data Deficient (DD) (Figure 1). Last record: 1975. Probably found in the National Park of Serra da Canastra, state of Minas Gerais and apparently restricted from there. Since this species is known only for the holotype, any information could be inferred. We do not even know if this species is recorded from the National Park effective area or neighborhoods. In order to achieve more basic information, we propose extensive Scarabaeinae sampling in this region.

Figure 1. Distribution map for nine evaluated brachypterous species of the genus Dichotomius .

4. CONCLUSIONS Among species accessed in this work, Dichotomius comarapensis and D. sp.5 could not be evaluated due the lack of data (DD). Dichotomius ingens and D. sp.4 were evaluated as Near Threatened (NT) while D. sp.3 was classified as Vunerable (VU). Dichotomius eucranioides and D. sp.1 were considered Endagered (EN) and D. sp.2 and D. mysticus were classified as Critically Endangered (CR). Part of the environmental legislation, mainly SNUC (BRASIL, 2000) is seminal to the conservation of the species which distribution bears areas of Conservation Units and Indigenous Reserves. However, the legislation fails in preserving native areas located at APP’s and Legal Reserves, both expected on the Brazilian Forest Code (BRASIL, 1965). The monitoring of local populations, ecological studies on the effect of fire and exotic species, the maintenance of good quality native areas at Conservation Units and the effective implantation of legislation expected at Brazilian Forest Code are the main actions in order to conserve populations of brachypterous species of the genus Dichotomius . 68

5. BIBLIOGRAPHIC REFERENCES

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ANDRESSEN, E.; LAURANCE, S.G.W. Possible Indirect Effects of Mammal Hunting on Dung Beetle Assemblages in Panamá. Biotropica, v.39,n.1,p.141-146, 2007.

BATALHA, M.A.; MARTINS, F.R. Reproductive phenology of the cerrado plant community in Emas National Park (central Brazil). Australian Journal of Botany, v.52, p.149-151, 2004.

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EHRLICH, P.R.; MOONEY, H.A. Extinction, substitution and ecosystem services. BioScience, v.33,p.248-254, 1983.

FÁVERO, K.; BORDIGNON, L.; VECCHI-JUNIOR, K.; DINIZ, S. Efeito do tempo Pós- queimada Sobre Comunidades de Tephritidae (Diptera) em Áreas de Cerrado na Chapada dos Guimarães – MT. Entomobrasilis, v.3,n.2,p.29-33, 2010.

GILPIN, M.E.; SOULÉ, M.E. Minimum Viable Populations: Processes of species extinction. In: SOULÉ, M.E. (Ed.). Conservation Biology: The Science of Scarcity and Diversity. Sinauer Associations, Sunderland, 1986. P.19-34.

HALFFTER, G.; MATTHEWS, E.G. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomológica Mexicana, v.14,n.12,p.1- 312, 1966.

HALFFTER, G.; FAVILLA, M.E. The Scarabaeinae (Insecta: Coleoptera) an Animal Group for Analysing, Inventorying and Monitoring Biodiversity in Tropical Rainforest and Modified Landscapes. Biology International, v.27,p.15-21, 1993.

IUCN. IUCN Red List. Avaliable at: http://www.iucnredlist.org. Access at 02.Feb.2012.

JESUS, A.P. A importância do Parque das Dunas para a cidade do Natal. Diário de Natal - Da Vinci , 3, 2003.

KOHLMAN, B.; SOLÍS, A. El género Dichotomius (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, v.8,p.343–382, 1997.

LOISELLE, B.A.; HOWELL, C.A.; GRAHAM, C.H.; GOERCK, J.M.; BROOKS, T.; SMITH, K.G.; WILLIAMS, P.H. Avoiding Pitfalls of Using Species Distribution Models in Conservation Planning. Conservation Biology, v.17,n.6,p.1591-1600.

MEDINA, C.A.; LOPERA-TORO, A.; VÍTOLO, A.; GILL, B. Escarabajos Coprófagos (Coleoptera: Scarabaeidea: Scarabaeinae) de Colombia. Biota Colombiana, v.2.,n.2,p.131– 144, 2001.

NICHOLS, E.; LARSEN, T.; SPECTOR, S.; DAVIS, A.L.; ESCOBAR, F.;FAVILA, M.E.; VULINEC, K.; THE SCARABAEINAE RESEARCH NETWORK. Global dung beetle response to tropical forest modification and fragmentation: A quantitative literature review and meta-analysis. Biological Conservation, v.137,p.1-19, 2007.

NICHOLS, E.; SPECTOR, S.; LOUZADA, J.N.C.; LARSEN, T.; AMEZQUITA, S.; FAVILA, M.E.; THE SCARABAEINAE RESEARCH NETWORK. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biological Conservation v.141,p.1461-1474, 2008. PERAZA, L.N.; DELOYA, C. Uma Nueva Especies Mexicana de Dichotomius Hope (Coleoptera: Scarabaeidae) y Clave para Identificación de las Especies del Grupo carolinus. Neotropical Entomology, v.35,n.5,p.629–631, 2006.

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POLETO, C.; CARVALHO, S.L.; FREITAS LIMA, E.A.C. Problemas de degradação ambiental em uma microbacia hidrográfica situada no município de Ilha Solteira – S.P., Brazil., e sua percepção pelos propietários rurais. Holos Environment, v.4.n.1, p.68-80, 2004.

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Conclusion

Dichotomius is a widespread genus at American continent bearing about 170 species divided into four subgenera: Dichotomius Hope, Luederwaldtinia Martínez, Selenocopris Burmeister and Homocanthonides Luederwaldt. There are nine brachypterous species of Dichotomius ; into subgenus Luederwaldtinia , D. eucranioides Pereira e D'Andretta 1955 into a monobasic complex, D. mysticus (Luederwaldt, 1935) into geminatus complex and D. sp.1 n. sp. into batesi complex. Into subgenus Selenocopris , D. ingens (Luederwaldt, 1935), D. sp.5 n. sp ., D. sp.3 n. sp. , D. sp.2 n. sp . and D. sp.4 n .sp ., all into ascanius complex. As Génier (2000) we couldn't define a complex for D. comarapensis Génier, 2000 while the definition of this species into subgenus Dichotomius stills need a complete revision of this subgenus. By now, we consider that D. comarapensis is integrating subgenus Dichotomius within a homonymous complex, also composed by a new species from Chaco region. This decision is supported by the combination of following characters, as first defined and commented by Génier (2000): the configuration of anterior tibiae of males, elytra micro sculpture, width of posterior pronotum margin and presence of setae at the apex of paramera. The multiple occurrence of brachypterism in three subgenera and five complexes distributed among them lead us to conclude that brachypterism evolved several times in the genus. Thus, this hypothesis needs to be confirmed by phylogenetic analysis. On the conservation of this species populations, according to IUCN criteria (IUCN, 2012) D. sp.2 and D. mysticus status are considered as critically endangered (CR) while D. sp.1 and D. eucranioides are considered as endangered (EN), D. sp.3 vulnerable (VU), D. ingens and D. sp.4 considered Near Threatened (NT) while D. comarapensis, and D. mysticus are considered as data deficient (DD). We also suggest that population monitoring and the effective compliment of environmental laws are adequate management tactics to preserve these endangered species. Finally, as follows, we give a illustrated key for all brachypterous species of Dichotomius . This key also includes a species of the genus Holocephalus , a genus closely related to Dichotomius and that could cause misidentifications.

1. Antennal club spherical; second labial palpomere very large, flattened, oval, concealing smaller third palpomere in ventral view. Eyes reduced, ventral portion narrow. Posterior portion of cephalic horn strongly rugose, with well defined V-shaped carina.

Elytra short, semicircular when viewed laterally Hind wing reduced, not reaching apex of elytron; brachypterous. Brazil (Mato Groaso), Paraguay … Holocephalus cristatus (Gillet, 1907) 1’. Antennal club elongate; second labial palpomere triangular, not covering the third palpomere in ventral view. Ventral portion of eyes wider than dorsal portion. United States to Argentina … 2

2. Clypeal margin rounded or weakly emarginated ... 3 2’. Clypeal margin bidentated, emarginated ... 4

3. Pronotum wider than elytra. Elytra strongly convex. Anterior tibiae, on males, emarginated, on females not emarginated. Elytra with shagrinated microsculpture. Hind wing atrophiated. Bolivia ... Dichotomius comarapensis Génier 2000 3' Pronotum as wide as elytra. Elytra rounded, not strongly convex. Others characters may be present, but not as the combination above ... other species of Dichotomius

4. Margin of head usually angled at the junction of clypeus and gena. Clypeal teeth strongly emarginated. Metasternal pilosity usually short and scattered or rarely absent. Last abdominal sternite of females produced beneath apex of pygidium ... 5 4'. Margin of head lacking angulation at the junction of clypeus and gena. Clypeal teeth emarginated, not as strongly as mentioned above. Metasternal pilosity usually abundant. Last abdominal sternite of females never produced beneath apexx of pygidium ... 9

5. Pronotal disc, posteriorly, with ocellate punctures. Elytral striae wide, lacking punctures. Brasil, Mato Grosso ... Dichotomius sp.4 n sp. 5'. Pronotal disc lacking any ocelatte puncture. Ocelate punctures restricted to the margins, anterior angles and anterior lobe. Elytral striae bearing distinct punctures .. 6

6. Clypeo-frontal region of males developed into a single conical horn or tubercle. Females bearing transversally ridged surface on clypeo-frontal region ... Dichotomius ingens (Luederwaldt 1935)

6'. Clypeo-frontal region developed in a transversal carina of variable size. Both males and females bearing smooth surface on clypeo-frontal region ... 7

7. Clypeo-frontal transversal carina strongly emarginated medially and bearing two tubercles laterally at the apex. Length of fifth metatarsomere three times the length of fourth metatarsomere. Dorsally, foveae of parameres (anterior to apical excavation) never reaching lateral margin of parameres. Brasil, São Paulo ... Dichotomius sp.2 n sp. 7’. Clypeo-frontal transversal carina feebly emarginated medially, lacking tubercles at the apex or having small developed tubercles. Length of fifth metatarsomere never reach three times the length of fourth metatarsomere. Dorsally, foveae of parameres may reach, or not, the lateral margin of parameres ... 8

8. Anterior lobe of pronotum filled with dense ocellate puncture. Length at most 20 mm. Dorsally, excavation at apex of paramera occupying half of the dorsal surface of paramera. Foveae of anterior portion of paramera strongly excavated, reaching the lateral margin of parâmeros ... Dichotomius sp.3 n. sp. 8'. Anterior lobe of pronotum filled with small spaced non-ocellate puncture. Length 25 mm. Dorsally, excavation at apex of paramera occupying third of the dorsal surface of paramera. Foveae of anterior portion of paramera never reaching the lateral margin of parâmeros ... Dichotomius sp.5 n. sp .

9. Clypeal margin crenulated. Males with strong transversal carina with width subequal to interocular space. Male pronotal disc delimited anteriorly by four gibosities. Female pronotal disc with two lateral groups of ocellate punctures. Pronotum of both sexes with longitudinal sulcus filled with ocellate puncutres. Pronotum wider than elytra. Elytral suture fused. ... Dichotomius eucranioides Pereira & D’Andretta, 1955 9'. Clypeal margin smooth. Males with single tubercle or rounded knobs. Pronotal disc smooth, at most with ocelate puncutres and feeble anterior declivity ... 10

10. Pronotal disc with ocellate puntures ... Dichotomius sp.1 n.sp. 10'. Pronotal disc lacking ocellate puncture ... Dichotomius mysticus (Luederwaldt 1935).

BIBLIOGRAPHIC REFERENCE

GÉNIER, F. Dichotomius comarapensis sp. nov., une nouvelle espèce bolivienne de scarabée brachyptère (Coleoptera: Scarabaeidae, Scarabaeinae). Fabreires, v.25, n.2-3, p.25-31, 2000.

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PEREIRA, F.S.; D’ANDRETTA, M.A.V. Novos Escarabeídeos e Novas Sinonímias (Col. Scarabaeidae). Papéis Avulsos do Departamento de Zoologia, v.12,n.11,p. 247–264, 1955.

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3) Figures. Please note that the journal has a matter size of 25 cm x 17 cm (printable area of the main text and plates) and is printed on A4 paper. If the final size of your plates is greater than this, please crop extra white areas around plates or reduce it to this size; this will reduce file size. Plates containing only black&white lines/dots are line art works and must be scanned as such (i.e. 1 bit, monochrome line art) at 600 dpi (maximum 1200 dpi). They should be saved in tiff file and LZW compression is recommended to be used to reduce file size for easy sub-mission by e-mail. Plates containing greyscale drawings and photographs should be scanned at 300 dpi and saved in TIFF (use LZW compression) or

78 jpeg at the highest quality. Please do not modify photographs in the jpeg files; the print quality would be severely altered if you did that. You can modify files in the TIFF file and when completed, convert to high quality jpeg for submission. If you have colour figures, it is best to group them together in plates, which will save cost.

4) Reference list. This must be formatted using Zootaxa style (see below for examples of main types of papers): Please note that (1) journal titles must be written in full (not abbreviated); (2) journal titles and volume numbers are followed by a “,”; (3) page ranges are connected by a “n dash”, not a hyphen “-”, which is used to connect two words. For websites, it is important to include the last date when you see that site, as it can be moved or deleted from that address in the future.

5) Submission of files. Please send final files by e-mail (or ftp) to your subject editor who will then forward files of accepted versions to Zootaxa office. Plates (if many) may be sent directly to Zootaxa office if the subject editor agrees. year in brackets, not followed by a period

Journal paper journal title in italics, spelt in full (never abbreviate), followed by comma volume/issue number followed by comma:

Gusarov, V.I. (2002) A revision of Nearctic species of the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa, 81, 1–88.

Connect author names by “&”, not “and”

Book chapter page ranges connected by n-dash

Newton, A.F., Thayer, M.K., Ashe, J.S. & Chandler, D.S. (2000) Staphylinidae Latreille, 1802. In: Arnett, R.H. & Thomas, M.C.

(Eds.), American Beetles. Vol.1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, pp. 272– 418. author family name first, followed by initials of given names (no space between initials) book title in italics publisher name first, followed by location

Book

Baker, E.W. & Tuttle, D.M. (1994) A Guide to Spider Mites (Tetranychidae) of the United States. Indira Publishing House, West Bloomfield, Michigan, 347 pp.

Internet resources

O’Connor, R.J., Dunn, E, Johnson, D.H., .Jones, S.L., Petit, D., Pollock, K., Smith, C.R., Trapp, J.L. & Welling, E. (2000) A programmatic review of the North American Breeding Bird Survey: report of a peer review panel. U.S. Geological Survey Patuxent Wildlife Research Center, Laurel, Maryland. Available from http://www.mp2- pwrc.usgs.gov/bbs//bbsreview/ (accessed 3 April 2003) include the last date when you see that site, as it can be moved or deleted from that address in the future.

ATTACHMENT 2: Guidelines for authors – Zookeys

Title: The title should be in a sentence case (only scientific, geographic or person names should be with a first capital letter, i.e. Elater ferrugineus L., Germany, etc.), and should include an accurate, clear and concise description of the reported work, avoiding abbreviations. The higher taxa within the title should be separated with commas and not with a semicolon, e.g.: (Coleoptera, Elateridae, Elaterini). Authors and Affiliations: Provide the complete names of all authors, and their addresses for correspondence, including e.g., institutional affiliation (e.g. university, institute), location (street, boulevard), city, state/province (if applicable), and country. One of the authors should be designated as the corresponding author. It is the corresponding author's responsibility to ensure that the author list, and the individual contributions to the study are accurate and complete. If the article has been submitted on behalf of a consortium, all consortium members and their affiliations should be listed after the Acknowledgements section. Abstract and Keywords: Please have your abstract and keywords ready for input into the submission module. Body Text: All papers should be in grammatically correct English. Non-native English speaking authors are required to have their manuscripts checked by a native English speaker prior to submission. Use either British/Commonwealth or American English provided that the language is consistent within the paper. A manuscript must be written with precision, clarity, and economy, whenever appropriate in active voice and first person. Avoid the use of parenthetical comments and italics or bold for emphasis. This journal discourages the use of quotation marks except for direct quotations, words defined by the author, and words used in unusual contexts. Short quotations should be embedded in the text and enclosed in double quotation marks (''). Long quotations should be on a separate line, italicized, but without quotation marks. Single quotation marks are to be used only for a quotation that occurs within another quotation. Spacing, Fonts, and Page Numbering: Single-space all material (text, quotations, figure legends, tables, references, etc.). Separate paragraphs with a blank line. Use a 12-point font (preferably Times New Roman or Arial). Capitals: First capital letters should be used only in the beginning of a sentence, in proper names and in headings and subheadings, as well as to indicate tables, graphs and figure/s within the text. Software programmes should be written with capital letters (e.g., ANOVA, MANOVA, PAUP). Italicization/Underlining: Scientific names of species and genera, long direct quotations and symbols for variables and constants (except for Greek

81 letters), such as p, F, U, T, N, r, but not for SD (standard deviation), SE (standard error), DF (degrees of freedom) and NS (non significant) should be italicized. These symbols in illustrations and equations should be in italics to match the text. Italics should not be used for emphasis, and not in abbreviations such as e.g., i.e., et al., etc., cf. Underlining of any text is not acceptable. Abbreviations: Abbreviations should be followed by ‘.' (full stop or period; for instance: i.e., e.g., cf., etc.). Note that you shouldn't add a full stop at the end of abbreviated words if the last letter of the abbreviation is the same as the last letter of the full word. For example, you should abbreviate „Eds”, „Dr”, „Mr” without full stop at the end. All measures, for instance mm, cm, m, s, L, should be written without full stop. On the use of dashes: (1) Hyphens are used to link words such as personal names, some prefixes and compound adjectives (the last of which vary depending on the style manual in use) (2) En dash or en-rule (the length of an 'n') is used to link spans. In the context of our journal en-dash should be used to link numerals, sizes, dates and page numbers (e.g., 1977–1981; figs 5–7; pp.237–258); geographic or name associations (Murray–Darling River; a Federal–State agreement); and character states combinations such as long– pubescent or red–purple. (3) Em-dash or em-rule (the length of an 'm') should be used rarely, only for introducing a subordinate clause in the text that is often used much as we use parentheses. In contrast to parentheses an em-dash can be used alone. En-dashes and em-dashes should not be spaced. Footnotes: Avoid footnotes in the body text of the manuscript. It is always possible to incorporate the footnote into the main text by rewording the sentences, which greatly facilitates reading. Additionally, footnotes are not always handled well by the journal software, and their usage may cause a failure of submission. Footnotes are acceptable only below tables; instead of numbers, please use (in order): †, ‡, §, |, ¶, #, ††, ‡‡, §§, ||, ¶, ##. Geographical coordinates : It is strongly recommended to list geographical coordinates as taken from GPS or online gazetteer, or georeferencer (http://wwold.gbif.org/prog/digit/Georeferencing). Geographical coordinates must be listed in one of the following formats: Definition: The locality consists of a point represented by coordinate information in the form of latitude and longitude. Information may be in the form of: Degrees, Minutes and Seconds (DMS); Degrees and Decimal Minutes (DDM) or Decimal Degrees (DD). Records should also contain a hemisphere (E or W and N or S) or, with Decimal Degrees, minus (–) signs to indicate western and/or southern hemispheres. Examples: 36° 31' 21" N; 114° 09' 50" W (DMS); Example 2: 36° 31.46’N; 114° 09.84’W (DDM); 36.5243° S; 114.1641° W (DD); -36.5243; -114.1641 (DD using minus signs to

82 indicate southern and western hemispheres). Note on accuracy: Because GPS units are very commonly used today to record latitude/longitude, many authors simply give the GPS readings for their localities. However, these readings are much too accurate. For example, a GPS unit might give the latitude in decimal seconds as 28°16'55.87"N. Since one second of latitude is about 30 m on the ground, the second figure after the decimal in 55.87 represents 30 cm, yet a typical handheld GPS unit is only accurate at best to a few metres. We therefore recommend two ways to report GPS-based locations. If you give the GPS reading without rounding off, make sure you include an uncertainty figure as a context for the over-accurate GPS reading. We recommend the Darwin Core definition of uncertainty (http://rs.tdwg.org/dwc/terms/index.htm#coordinateUncertaintyInMeters): "The horizontal distance (in meters) from the given decimalLatitude and decimalLongitude describing the smallest circle containing the whole of the Location."If you only give the GPS reading, please round it off to an implied precision appropriate to the error in the measurement, or to the extent of the area sampled. We suggest rounding off: to the nearest second in degree- minute-second format (28°16'56"N), which implies roughly ± 25-30 m at middle latitudes; to four decimal places in decimal degree format (28.2822°N), which implies roughly ± 10- 15 m at middle latitudes; to two decimal places in decimal minute format (28°16.93'N), which implies roughly 15-20 m at middle latitudes. Altitude: Many GPS users simply record the elevation given by their GPS unit. However, GPS elevation is NOT the same as elevation above sea level. GPS units record the elevation above a mathematical model of the earth's surface. The difference between this elevation and elevation above sea level can be tens of meters. In any case, the accuracy of a GPS elevation is often the same as the usual accuracy in horizontal position, so a GPS elevation such as '753 m' is much too accurate and should be rounded off to 'ca 750 m'. We strongly recommend the use of Example 2 (the DDM format). The other three are also possible but will be recalculated to DDM during the process of online mapping from the HTML version of the paper. The only restriction on format is in creating a KML (Keyhole Markup Language) file. KML latitudes and longitudes must be in the DD format shown above in Example 4. Please also consider submitting a table of localitie s with your manuscript, either as a spreadsheet or in CSV text format. By doing so you will make your specimen localities much more easily available for use in biodiversity databases and geospatial investigations. The geospatial table will be put online as supplementary material for your paper. A minimum table will have three fields: species (or subspecies) name, latitude and longitude. A full table will have the same data for each specimen lot as appears in the text of your paper. Please check

83 latitude/longitude carefully for each entry. Units: Use the International System of Units (SI) for measurements. Consult Standard Practice for Use of the International System of Units (ASTM Standard E-380-93) for guidance on unit conversions, style, and usage. Statistics: Use leading zeroes with all numbers, including probability values (e.g., P < 0.001). For every significant F-statistic reported, provide two df values (numerator and denominator). Whenever possible, indicate the year and version of the statistical software used. Web (HTML) links: Authors are encouraged to include links to other Internet resources in their article. This is especially encouraged in the reference section. When inserting a reference to a web-page, please include the http:// portion of the web address. Supplementary files: Larger datasets can be uploaded separately as Supplementary Files. Tabular data provided as supplementary files can be uploaded as an Excel spreadsheet (.xls), as an OpenOffice spreadsheets (.ods) or comma separated values file (.csv). As with all uploaded files, please use the standard file extensions. Headings and subheadings: Main headings: The body text should be subdivided into different sections with appropriate headings. Where possible, the following standard headings should be used: Introduction, Methods, Results, Discussion, Conclusions, Acknowledgements, References. These headings need to be in bold font on a separate line and start with a first capital letter. Please do not number headings or subheadings. Introduction - The motivation or purpose of your research should appear in the Introduction, where you state the questions you sought to answer, and then provide some of the historical basis for those questions. Methods - Provide sufficient information to allow someone to repeat your work. A clear description of your experimental design, sampling procedures, and statistical procedures is especially important in papers describing field studies, simulations, or experiments. If you list a product (e.g., animal food, analytical device), supply the name and location of the manufacturer. Give the model number for equipment used. Supply complete citations, including author (or editor), title, year, publisher, and version number, for computer software mentioned in your article. Results - Results should be stated concisely and without interpretation. Discussion - Focus on the rigorously supported aspects of your study. Carefully differentiate the results of your study from data obtained from other sources. Interpret your results, relate them to the results of previous research, and discuss the implications of your results or interpretations. Point out results that do not support speculations or the findings of previous research, or that are counter-intuitive. You may choose to include a Speculation subsection in which you pursue new ideas suggested by your research, compare and contrast your research with findings from other systems or

84 other disciplines, pose new questions that are suggested by the results of your study, and suggest ways of answering these new questions. Conclusion -This should state clearly the main conclusions of the research and give a clear explanation of their importance and relevance. Summary illustrations may be included. References - The list of References should be included after the final section of the main article body. A blank line should be inserted between single-spaced entries in the list. Authors are requested to include links to online sources of articles, whenever possible! Where possible, the standard headings should be used in the order given above. Additional headings and modifications are permissible. Subordinate headings : Subordinate headings (e.g. Field study and Simulation model or Counts, Measurements and Molecular analysis ), should be left-justified, italicized, and in a regular sentence case. All subordinate headings should be on a separate line.