Drosophilidae (Insecta, Diptera) in the state of Pará (Brazil)

Drosophilidae (Insecta, Diptera) no estado do Pará (Brasil)

Rosângela Santa-Brígida Hermes José Schmitz Marlúcia Bonifácio Martins About the authors

Abstract

This list contains information on the Drosophilidae that occur in the Brazilian state of Pará, Amazon biome, and an analysis of the current knowledge of Drosophilidae based on museum material and literature records. This list includes a detailed account of the material deposited in the entomological collections of the Museu Paraense Emílio Goeldi and Museu de Zoologia da Universidade de São Paulo, up to 2015. In total, 122 species of Drosophilidae were registered, including 27 new records for the state of Pará and 22 are new records for the Amazon; for instance, the genera Diathoneura and Rhinoleucophenga, and three new records for Brazil, (Drosophila fasciola, Diathoneura flavolineata and Drosophila neochracea). The state of Pará is the third state in Brazil in terms of numbers of species of Drosophilidae, with 17% locally native species. Despite the high species richness, there is still a lot to be known about the states's Drosophilidae fauna. This study highlights the importance of scientific collections, particularly as an aid to study regional biodiversity.

Keywords:
species composition; Amazon; Diathoneura; Rhinoleucophenga

Resumo

As informações reunidas nesta lista referem-se à ocorrência de Drosophilidae no estado do Pará, bioma Amazônia com uma análise do estado de conhecimento do grupo. Este documento contém uma revisão detalhada do material depositado na Coleção Entomológica do Museu Paraense Emílio Goeldi e Museu de Zoologia da Universidade de São Paulo até 2015, bem como a análise de toda a literatura relevante. Um total de 122 espécies de Drosophilidae foi registrado, com 27 novos registros para o estado do Pará e 22 são novas ocorrências para a Amazônia. Entre estes o primeiro registro para os gêneros Diathoneura e Rhinoleucophenga, além de três registros novos para o Brasil (Drosophila fasciola, Diathoneura flavolineata e Drosophila neochracea). O estado do Pará é o terceiro maior estado brasileiro em termos de número de espécies registradas, com 17% das espécies com ocorrência exclusiva no Estado. Ainda assim há grandes lacunas para o conhecimento da fauna dos Drosophilidae do estado. Este estudo destaca a importância das coleções científicas, acima de tudo, como um elemento crucial para compreensão da biodiversidade regional.

Palavras-chave:
Composição de espécies; Amazônia; Diathoneura; Rhinoleucophenga

Introduction

There are more than 4,200 Drosophilidae species in the world (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
). This number will most likely increase as new species are continuously being described, particularly in the Neotropical region (Silva & Martins 2004SILVA, A.A.R. & MARTINS, M.B. 2004. A new anthophilic species of Drosophila Fallén belonging to the bromeliae group of species (Diptera, Drosophilidae). Rev. Bras. Zool. 21(3): 435–437. http://www.scielo.br/pdf/rbzool/v21n3/21890.pdf
http://www.scielo.br/pdf/rbzool/v21n3/21...
, Vilela & Bächli 2005VILELA, C.R. & BÄCHLI, G. 2005. Three new species of Neotropical Drosophilidae (Diptera). Mitt. Schweiz. Entomol. Ges. 78:143–165., Ratcov & Vilela 2007RATCOV, V. & VILELA, C.R. 2007. A new Neotropical species of spot-thoraxed Drosophila (Diptera, Drosophilidae). Rev. Bras. Entomol. 51(3):305–311. http://www.scielo.br/pdf/rbent/v51n3/v51n3a09.pdf.
http://www.scielo.br/pdf/rbent/v51n3/v51...
, Culik & Ventura 2009CULIK, M.P. & VENTURA, J.A. 2009. New species of Rhinoleucophenga, a potencial predator of pineapple mealybugs. Pesq. Agropec. Bras. 44(4):417–420., Schmitz et al. 2009SCHMITZ, H.J., GOTTSCHALK, M.S. & VALENTE, V.L.S. 2009. Rhinoleucophenga joaquina sp. nov. (Diptera: Drosophilidae) from the Neotropical Region. Neotrop. Entomol. 38(6): 786–790. http://www.scielo.br/pdf/ne/v38n6/12.pdf
http://www.scielo.br/pdf/ne/v38n6/12.pdf...
, Figuero & Rafael 2011FIGUERO, M. & RAFAEL, V. 2011. Dos nuevas especies del grupo Drosophila onychophora (Diptera, Drosophilidae) em los bosques de Polylepis de Palallacta, Pichincha, Ecuador. Iheringia 101(4):342–349., Figuero et al. 2012FIGUERO, M.L., LEÓN, R., RAFAEL, V. & CÉSPEDES, D. (2012). Cuatro nuevas espécies del grupo Drosophila onychophora (Diptera, Drosophilidae) em el Parque Arqueológico Rumipamba, Pichincha, Ecuador. Iheringia. 102(2):212–220., Gottschalk et al. 2012GOTTSCHALK, M.S., MARTINS, M.B., PRAXEDES, C.L.B. & MEDEIROS, H.F. 2012. A new Amazonian species from the Drosophila annulimana species group (Diptera, Drosophilidae). Rev. Bras. Entomol. 56(4):431–435. http://www.scielo.br/pdf/rbent/v56n4/v56n4a06.pdf
http://www.scielo.br/pdf/rbent/v56n4/v56...
, Acurio et al. 2013ACURIO, A., RAFAEL, V., CESPEDES, D. & RUIZ, A. 2013. Description of a New Spotted-Thorax Drosophila: Drosophilidae) Species and its Evolutionary Relationships Inferred by a Cladistic Analysis of Morphological traits. Ann. Entomol. Soc. Am. 106(6):695–705., Poppe et al. 2014POPPE, J.L., SCHMITZ, H.J., GRIMALDI, D. & VALENTE, V.L.S. 2014. High diversity of Drosophilidae (Insecta, Diptera), in the Pampas Biome of South America, with descriptionsof new Rhinoleucophenga species. Zootaxa. 3779(2):215–2145., Junges & Gottschalk 2014JUNGES, J. & GOTTSCHALK, M.S. 2014. Two New Species of the New World Genus Rhinoleucophenga (Diptera: Drosophilidae). J. Insect Sci. 14(157):1–5., Guillín & Rafael 2015GUILLÍN, E.R. & RAFAEL, V. 2015. Three new species of Drosophila tripunctata group (Diptera: Drosophilidae) in the eastern Andes o Ecuador. Rev. peru. Boil. 22(3):289–296., Junges et al. 2016JUNGES, J., GOTTSCHALK, M.S., LORETO, E.L.S. & ROBE, L.J. 2016. Two new species of Mycodrosphila (diptera: Drosophilidae) proposed by molecular and morphological approaches, with a key to American species. Rev. Bras. Ent. 60:30–39., Vidal & Vilela 2015VIDAL, M.C. & VILELA, C.R. 2015. A New Species of Rhinoleucophenga (Diptera: Drosophilidae) From the Brazilian Cerrado Biome Associated with Extrafloral Nectaries of Qualea grandiflora (Vochysiaceae). Ann. Entomol. Soc. Am. 108(5):932–940.). In Brazil, the Drosophilidae family is represented by 16 genera and 305 species (Tidon et al. 2015TIDON, R., GOTTSCHALK, S.M., MARTINS, M.B. & SCHIMTZ, H.J. 2015. CTFB: Catálogo Taxonômico da Fauna do Brasil. Eletronic Database accessible at http://fauna.jbrj.gov.br/fauna/listaBrasil/ConsultaPublicaUC/ConsultaPublicaUC.do. (last access in 3 December 2015).
http://fauna.jbrj.gov.br/fauna/listaBras...
). Taxonomic inventories are needed to we fill the knowledge gaps on biodiversity (Hortal et al. 2015HORTAL, J., BELLO, F., DINIZ-FILHO, J.A.F, LEWINSOHN, T.M., LOBO, J.M. & LADLE, R.J. 2015. Seven shortfalls that beset large-scale knowledge of biodiversity. Ann. Ver. Ecol. Syst. 46:523–49.). Althought the first collections for the Amazon dating from 1920-1959 period (Duda 1927DUDA, O. 1927. Die sudamerikanischen Drosophiliden (Dipteren) unter Beruckssichtigung auch der anderen neotropischen sowie der nearktischen Arten. Archiv. Naturgesch. 91:1–228.; Hendel 1936HENDEL, F. 1936. Ergebnisse einer zoologischen Sammelreise nach Brazilien insbesondere in das Amazonasgebiet, ausgefuhrt von Dr. H. Zerny. X. Teil. Diptera. Muscidae acalyptratae (excl. Chloropidae). Ann. des K.K. Naturhist. Hofmus. 47:61–106.; Dobzhansky & Pavan 1943DOBZHANSKY, T. & PAVAN, C. 1943. Studies on Brazilian species of Drosophila. Bol. Facul. Fil. Ciên. E Letr. 36:7–72.; Pavan 1950PAVAN, C. 1950. Espécies Brasileiras de Drosophila. II. Bol. Facul. Fil. Ciên. e Letr. 111(8):1–38.; Pavan 1959PAVAN, C. 1959. Relações entre populações naturais de Drosophila e o meio ambiente. Facul. Fil. Ciên. e Letr. 11:1-81.), the richness of Drosophilidae fauna of the brazilian Amazon is still little known, states such as Acre, Amapá, Roraima, Rondônia e Tocantins have less than 12 recorded species (Gottschalk et al. 2008GOTTSCHALK, M.S., HOFMANN, P.R.P. & VALENTE, V.L.S. 2008. Diptera, Drosophilidae: historical occurrence in Brazil. Check List. 4(4):485–518.). Among the Amazonian states, only Amazonas and Pará have more consistent inventory Drosophilid (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
). As the result, the scenario of the distribution of species for this region is still very incomplete.

The Drosophilidae fauna of the state of Pará has been sporadically studied since 1920, by pioneer systematists (Duda 1927DUDA, O. 1927. Die sudamerikanischen Drosophiliden (Dipteren) unter Beruckssichtigung auch der anderen neotropischen sowie der nearktischen Arten. Archiv. Naturgesch. 91:1–228.; Hendel 1936HENDEL, F. 1936. Ergebnisse einer zoologischen Sammelreise nach Brazilien insbesondere in das Amazonasgebiet, ausgefuhrt von Dr. H. Zerny. X. Teil. Diptera. Muscidae acalyptratae (excl. Chloropidae). Ann. des K.K. Naturhist. Hofmus. 47:61–106.; Burla et al. 1949BURLA, H., CUNHA, A.B. DA, CORDEIRO, A.R., DOBZHANSKY, T., MALOGOLOWKIN, C. & PAVAN, C. 1949. The willistoni group of sibling species of Drosophila. Evolution. 3:300–314.), evolutionary biologists (Pavan 1950PAVAN, C. 1950. Espécies Brasileiras de Drosophila. II. Bol. Facul. Fil. Ciên. e Letr. 111(8):1–38.) and more recently by ecologists (De Toni et al. 2005DE TONI, D.C., BRISSON, J.A. HOFMANN, P.R.P. MARTINS, M. & HOLLOCHER, H. 2005. First record of Drosophila parthenogenetica and D. neomorpha, cardini group, Heed, 1962 (Drosophila, Drosophilidae), in Brazil. DIS. 88: 33-38. http://www.ou.edu/journals/dis/DIS88/Research/R2.pdf#DeToni
http://www.ou.edu/journals/dis/DIS88/Res...
; Martins & Oliveira 2007MARTINS, M.B. & OLIVEIRA, L. 2007. Dinâmica espacial de Drosophila em remanescente de mata na Reserva do Mocambo e suas áreas adjacentes. In Mocambo: Diversidade e Dinâmica Biológica da Área de Pesquisa Ecológica do Guamá (APEG). (J.I. GOMES, M.B. MARTINS, R.C.V. MARTINS-DA-SILVA & S.S. ALMEIDA, Orgs.). Museu Paraense Emílio Goeldi: Centro de Pesquisa Agroflorestal da Amazônia Oriental, Belém, Pará, p. 389–404.; Martins & Santos 2007MARTINS, M.B. & SANTOS, R.C.O. 2007. Sítios de criação de Drosophila na Reserva Mocambo, Belém, Pará. In Mocambo: Diversidade e Dinâmica Biológica da Área de Pesquisa Ecológica do Guamá (APEG). (J.I. GOMES, M.B. MARTINS, R.C.V. MARTINS-DA-SILVA & S.S. ALMEIDA, Orgs.). Museu Paraense Emílio Goeldi: Centro de Pesquisa Agroflorestal da Amazônia Oriental, Belém, Pará, p. 315–329.; Silva & Martins 2009SILVA, A.A.R. & MARTINS, M.B. 2009. Polinização de cacauí Theobroma speciosum Willd. ex Spreng (Sterculiaceae) por drosofilídeos em área de ocorrência natural. In Caxiuanã: Desafios para Conservação de uma Floresta Nacional na Amazônia, P. Lisboa, Org.). 3° ed. Museu Paraense Emílio Goeldi, Belém, Pará, p. 431–440.; Praxedes & Martins 2014PRAXEDES, C.L.B. & MARTINS, M.B. 2014. Inventário biológico: riqueza, biodiversidade local e regional de Drosophilidae (Diptera) frugívoros. In Caxiuanã: Desafios para Conservação de uma Floresta Nacional na Amazônia, P. Lisboa, Org.). 4° ed. Museu Paraense Emílio Goeldi, Belém, Pará, p. 521–537. and Robe et al. 2014ROBE, L.J.R., MACHADO, S., BOLZAN, A.R., SANTOS, J.P.dos, VALER, F.B., SANTOS, A.P., BLAUTH, M.L. & GOTTSCHALK, M.S. 2014. Comparative ecological niche modeling and evolutionary ecology of Neotropical mycophagous Drosophilidae (Diptera) species. Stud. Neotrop. Fauna Environ. 2014.925370.). Currently, in Pará there are 75 Drosophilidae species from nine genera that are from two subfamilies Steganinae and Drosophilinae (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
).

Throughout the years a great volume of material has accumulated in scientific collections. Material from the state of Pará is, for the most part, deposited in the collections Museu Paraense Emílio Goeldi (MPEG) and Museu de Zoologia da Universidade de São Paulo (MZUSP). The Drosophilidae species in the MPEG deposit were mostly collected in the Amazon, from expeditions dating back to the 1970s. At MZUSP it is known to hold Brazil's largest Diptera collection, with specimens from large expeditions during the 1940s and 1950s, particularly in the Amazon, coordinated by Dr. Crodowaldo Pavan (Magalhães 2010MAGALHÃES L.E. 2010. Reminiscências do tempo de drosófilas. Pesq. Fapesp, Vol. 168, p.78–79.). We analyzed material from these two collections and using literature data to complement our survey. Our goal was to list the species of Drosophilidae that are found in the state of Pará, with the aim to fill a knowledge gap of Drosophilae found in the Amazon.

Material and Methods

The state of Pará, Northern Brazil, has an area of 1,248,000 km2, is completely immersed in the Amazon Biome (IBGE 2010IBGE 2010. IBGE Instituto Brasileiro de Geografia e Estatística. Eletronic Database accessible at http://www.ibge.gov.br/estadosat/perfil.php?sigla=pa. (last access in 08 December 2015).
http://www.ibge.gov.br/estadosat/perfil....
). The climate is Equatorial (Am) and Tropical (As), according to Köppen classification, with a mean temperature of 25 °C and minimum precipitation of 1,300 mm per year and maximum of 2,400 mm per year (Moraes et al. 2005MORAES, B.C., COSTA, J.MN., COSTA, A.C.L. & COSTA, M.H. 2005. Variação espacial e temporal da precipitação no estado Pará. Acta Amazon. 35(2):207–214.).

Occurrence data for Drosophilidae was compiled based on literature research, and the examination of material deposited in the entomological collection of the MPEG and MZUSP up to 2015. Data was plotted on a map of Brazil's municipalities. The identification of all Drosophilidae at MPEG was rechecked. For this, we dissected the genitalia of, and examined, up to five males of each species (following the procedure described by Bächli et al. 2004BÄCHLI, G., VILELA, C.R., ESCHER, S.A. & SAURA, A. 2004. The Drosophilidae (Diptera) of Fennoscandia and Denmark. Fauna Entomol. Scand. 39: 362 p.). The identity of species represented only by females was checked using external morphological characters whenever possible. Drosophilidae specimens that were in 70% ethanol and from the wet collection at MPEG, were mounted on entomological pins after being dissected. The specimen terminalia were stored in microtubes containing glycerin and pinned together with the specimen. Specimens deposited in the MZUSP had already been dissected and their terminalia were stored in microtubes containing glycerin, thus facilitating confirmation. The individual number of each species deposited in the MPEG collection was recorded. The total number of individual specimens of each species found in MZUSP was not accounted.

We searched Drosophilidae literature using the website compiled by Gerhard Bächli (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
). We also verified and analyzed each literature reference for the state of Pará.

Results and Discussion

In total, 122 species of Drosophilidae in 11 genera were found, distributed in 24 of the 144 municipalities of the state of Pará (Figure 1). Nine genera belong to the subfamily Drosophilidae (Chymomyza Czerny, 1903, Diathoneura Duda, 1924, Drosophila Fallén, 1823, Hirtodrosophila Duda, 1923, Mycodrosophila Oldenberg, 1914, Neotanygastrella Duda, 1925, Scaptodrosophila Duda, 1923, Zaprionus Coquillett, 1901 and Zygothrica Wiedemann, 1830) and two belong to Steganinae (Leucophenga Mik, 1886 and Rhinoleucophenga Hendel, 1917). The actualized list of all species can be found in Table 1. References in the column ‘record’ of Table 1 represent the oldest record for the species. The table also shows the municipalities of Pará where each species was recorded from, and the total number of specimens deposited in the collection of the Museu Paraense Emílio Goeldi. The map (Figure 1) shows the distribution of Drosophilidae in Pará, indicating the known species richness of each municipality. The municipality of Belém had the greatest number of species, 65, followed by Melgaço (MEL) and Portel (POR), with 55 species, and Santarém (SAN), with 27. More than half of the species (66) occurred in less than three municipalities. Drosophila malerkotliana, an invasive species present in the Amazon since the 1980s, was the most widespread species, occurring in 15 municipalities. The second most abundant species was the native species Drosophila nebulosa, Drosophila sturtevanti and Drosophila willistoni, which were found in 13 of the 24 municipalities.

Figure 1
Distribution of Drosophilidae in Pará, indicating the known species richness of each municipality. Map of South America at the upper left with the state of Pará highlighted in gray. The numbers on the map correspond to the following municipalities. In parenthesis correspond to the number of species for each locality. 1. Almeirim (1°31’22” S, 52°34’55” W); 2. Altamira (3°2’10” S, 52°12’21” W); 3. Aveiro (3°36’21” S, 55°19’55” W); 4. Belém (1°26’27” S, 48°24’50” W); 5. Bragança (1°3’46” S, 46°46’22” W); 6. Breves (1°40’55” S, 50°28’48” W); 7. Castanhal (1°17’49”S, 47°55’19” W); 8. Curuça (0°44’2” S, 47°51’18” W); 9. Igarapé-Açu (1°7’37” S, 47°37’4” W); 10. Itaituba (4°16’33” S, 55°59’2” W); 11. Juruti (2°9’7” S, 56°5’31” W); 12. Marituba (1°21’18” S, 48°20’31” W); 13. Melgaço (1°48’14” S, 50°42’43” W); 14. Nova Ipixuna (4°55’15” S, 49°4’37” W); 15. Óbidos (1°55’4” S, 55°31’4” W); 16. Oriximiná (1°45’57” S, 55°51’57” W); 17. Pacajá (3°50’16” S, 50°38’16” W); 18. Paragominas (2°59’42” S, 47°21’10” W); 19. Parauapebas (6°4’4” S, 49°54’7” W); 20- Portel (1°56’9” S, 50°49’15” W); 21. Santarém (2°26’34” S, 54°42’28” W); 22. Salvaterra (0°45’10” S, 48°31’1” W); 23. Viseu (1°11’49” S, 46°8’24” W); 24. Tucuruí (3°46’4” S, 49°40’ 22” W).
Table 1
Drosophilidae recorded in the state of Pará, Brazil. Municipalities: ALM = Almeirim, ALT = Altamira, AVE = Aveiro, BEL = Belém, BRA = Bragança, BRE = Breves, CAS = Castanhal, CUR = Curuça, IGA = Igarapé-Açu, ITA = Itaituba, JUR = Juruti, MAR = Marituba, MEL= Melgaço, NIP = Nova Ipixuna, OBI = Óbidos, ORI = Oriximiná, PAC = Pacajá, PARG = Paragominas, PARP = Parauapebas, POR = Portel, SAN= Santarém, SAL= Salvaterra, TUC= Tucuruí and VIS= Viseu. Records: MPEG= Museu Paraense Emílio Goeldi and MZUSP= Museu de Zoologia da Universidade de São Paulo collections. # in col: Total Drosophilidae specimens deposited in the MPEG.

The scientific collections contributed 27 new records for Pará and 22 for the Amazon. Of these, 24 were in the MPEG collection, two in the MZUSP collection and one recorded in both collections. Among the new records, Drosophila fasciola (Williston, 1896), Diathoneura flavolineata Duda, 1927 and Drosophila neochracea Wheeler, 1959 are new records for Brazil. D. fasciola Williston, 1896 had been recorded from the United States, Mexico, El Salvador, Caribbean, Panama, Colombia and Guyana; Drosophila flavolineata (Duda, 1927DUDA, O. 1927. Die sudamerikanischen Drosophiliden (Dipteren) unter Beruckssichtigung auch der anderen neotropischen sowie der nearktischen Arten. Archiv. Naturgesch. 91:1–228.) only from Costa Rica; and Drosophila neochracea (Wheeler, 1959WHEELER, M.R. 1959. A Nomenclatural Study of the Genus Drosophila. Univ. Texas Publi. 5914:181–205.) from Bolivia and Ecuador (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
). The vittimaculosa group of Zygothrica (Z. vittinubilaBurla, 1956BURLA, H. 1956. Die Drosophilidengattung Zygothrica und ihre beziehung zur Drosophilauntergattung Hirtodrosophila. Mitt. Zool. Mus. Berl. 32:189–321.), the subgenus Siphlodora of Drosophila (D. flexa), the genera Rhinoleucophenga (R. punctulata) and Diathoneura (D. flavolineata) are new records for Pará. Seven cosmopolitan or semi-cosmopolitan species were found in the state; Drosophila ananassae, D. kikkawai, Drosophila malerkotliana Parshad & Paika, 1964, Drosophila melanogaster Meigen, 1830, D. simulans Sturtevant, 1919, Scaptodrosophila latifasciaeformis (Duda, 1940) and Zaprionus indianus Gupta, 1970.

Of the 29 species listed in Table 1 and which were found in neither collections, only 23 of the species are only known from their original descriptions. Three were recently collected in Pará, but the reference specimens were not preserved (Drosophila pallidipennisDobzhansky & Pavan, 1943DOBZHANSKY, T. & PAVAN, C. 1943. Studies on Brazilian species of Drosophila. Bol. Facul. Fil. Ciên. E Letr. 36:7–72., Zygothrica aldrichi Sturtevant, 1920 and Zygothrica dispar Wiedemann, 1830). Three species, Drosophila polymorpha Dobzhansky & Pavan, 1943, Drosophila mesostigma Frota-Pessoa, 1954 and Drosophila mediostriata Duda, 1925 were listed as being in the MPEG collection, but upon re-examination of the respective material, their identification was not confirmed. In the case of D. polymorpha, the males were identified as Drosophila neomorpha Heed & Wheeler, 1957. The similarity between their terminalia and the possibility of introgression between these two species was discussed by De Toni et al. (2005)DE TONI, D.C., BRISSON, J.A. HOFMANN, P.R.P. MARTINS, M. & HOLLOCHER, H. 2005. First record of Drosophila parthenogenetica and D. neomorpha, cardini group, Heed, 1962 (Drosophila, Drosophilidae), in Brazil. DIS. 88: 33-38. http://www.ou.edu/journals/dis/DIS88/Research/R2.pdf#DeToni
http://www.ou.edu/journals/dis/DIS88/Res...
. Upon re-examination of material identified as D. mediostriata, we found Drosophila paramediostriata Townsend & Wheeler, 1955 and Drosophila frotapessoai Vilela & Bächli, in addition to underscribed species.

The species Drosophila mediocris Frota-Pessoa, 1954, Drosophila medioimpressa Frota-Pessoa, 1954, Drosophila neoelliptica Pavan & Magalhães, 1950, Drosophila pseudosaltans Magalhães, 1956 and D. milleri Magalhães, 1962, mentioned in the literature, are represented in the MPEG collection. However, only by females preserved in ethanol, precluding detailed examination of the parts that are necessary to confirm identification. Among these, only D. milleri was not known from other parts of Brazil, being historically restricted to Puerto Rico.

Wheller (1957), suggests that the presence of Hirtodrosophila thoracis (Williston, 1896) in Pará, based on Burla (1956)BURLA, H. 1956. Die Drosophilidengattung Zygothrica und ihre beziehung zur Drosophilauntergattung Hirtodrosophila. Mitt. Zool. Mus. Berl. 32:189–321.. However, here, we considered this record as belonging to Paraliodrosophila antennata Wheeler, 1957, following Vilela & Bächli (2007)VILELA, C.R & BÄCHLI, G. 2007. Revision of the Neotropical genus Paraliodrosophila (Diptera, Drosophilidae). Mitt. Schweiz. Entomol. Ges. 80:291–317., according to whom the genitalia of the holotype of P. antennata was illustrated by Burla as if it was H. thoracis.

Among the species recorded from Pará, 21 species have been nowhere else in Brazil, but have been recorded or observed in other countries (Table 1). Most of those species had only been previously found in Northern localities of the Neotropical region, particularly Colombia, Central America and Caribbean (Bächli 2015BÄCHLI, G. 2015. TAXODROS: The database on taxonomy of Drosophilidae. Eletronic Database accessible at http://www.taxodros.uzh.ch/. (last access in 19 November 2015).
http://www.taxodros.uzh.ch/....
). This finding suggests an affinity between the Amazon fauna and the fauna of those other regions.

Additionally, six species occur only in Pará: D. caxiuana, D. speciosa, Z. somatia, D. hendeli, D. decemseriata and M. brunnescens. The last three are known only from their original descriptions; D. hendeli and D. decemseriata were collected more than 85 years ago, by the Austrian entomologist Hans Zerny in 1927, during an expedition to Fazenda Taperinha, in Santarém (Zerny 1929ZERNY, H. 1929. Ergebnisse einer zoologischen Sammelreise nach Brasilien, insbesondere in das Amazonasgebiet, ausgefürt von Dr. H. Zerny.Vorbemerkung. Ann. Naturhist. Mus. Wien. 43:243–255.). At that time, the specimens were deposited at the Naturhistorisches Museum Wien (NMW). There, in 1936, the entomologist Friedrich Hendel determined 11 specimens as D. annularis Sturtevant, 1916 and nine as D. decemseriata (Hendel 1936HENDEL, F. 1936. Ergebnisse einer zoologischen Sammelreise nach Brazilien insbesondere in das Amazonasgebiet, ausgefuhrt von Dr. H. Zerny. X. Teil. Diptera. Muscidae acalyptratae (excl. Chloropidae). Ann. des K.K. Naturhist. Hofmus. 47:61–106.). In 1990 Vilela and Bächli described D. hendeli, based on one of the specimens identified as D. annularis by Hendel (Vilela & Bächili 1990). D. decemseriata which was described in 1936 by Hendel has not been found anywhere else. The type of M. brunnescens was collected by the geneticist Theodosius Dobzhansky in Belém-PA, in 1952 (Wheeler & Takada 1963WHEELER, M.R. & TAKADA, H. 1963. A revision of the American species of Mycodrosophila (Diptera, Drosophilidae). Ann. Entomol. Soc. Am. 56:392–399.), and was apparently never collected again. D. caxiuana and D. speciosa, in contrast, have been recorded more recently. In 2001, D. speciosa was recorded from cacauí flowers (Thebroma speciosum Willd. Ex Spreng) (Silva & Martins 2004SILVA, A.A.R. & MARTINS, M.B. 2004. A new anthophilic species of Drosophila Fallén belonging to the bromeliae group of species (Diptera, Drosophilidae). Rev. Bras. Zool. 21(3): 435–437. http://www.scielo.br/pdf/rbzool/v21n3/21890.pdf
http://www.scielo.br/pdf/rbzool/v21n3/21...
) and was thus collected from flowers in various municipalities across the state, as observed in the MPEG (Table 1). D. caxiuana was collected in 2008 in banana traps (Gottschalk et al. 2012GOTTSCHALK, M.S., MARTINS, M.B., PRAXEDES, C.L.B. & MEDEIROS, H.F. 2012. A new Amazonian species from the Drosophila annulimana species group (Diptera, Drosophilidae). Rev. Bras. Entomol. 56(4):431–435. http://www.scielo.br/pdf/rbent/v56n4/v56n4a06.pdf
http://www.scielo.br/pdf/rbent/v56n4/v56...
). It is possible that these taxa are locally endemic to the Amazon Biome, however, it is possible that their rarity is due to collecting limitations.

There is still a lot to be learnt about the richness and composition of Drosophilidae species in the Amazon biome. Despite being the second largest state of Brazil, and being within the Amazon biome, Pará comes third in Drosophilidae species richness in the country, with 17% of locally native species, behind the states of São Paulo and Santa Catarina (Gottschalk et al. 2008GOTTSCHALK, M.S., HOFMANN, P.R.P. & VALENTE, V.L.S. 2008. Diptera, Drosophilidae: historical occurrence in Brazil. Check List. 4(4):485–518.). Considering the geographic coverage of the records in the collections and literature records, it is obvious that there is still much to be learned about the region's biodiversity. The known distribution of poorly sampled, diverse groups such as the Drosophilidae often reflects where past collecting efforts were more concentrated, rather than the real local diversity. Only 24% of the species known to occur in the state were not represented in the collections studied and those, in turn, have contributed to widen the known geographic distribution of 21% of the species in the list. This result highlights the importance of scientific collections as a source of information on local biodiversity. Most records, either in the literature or in the collections, are from material collected with banana baits. It is possible that the list presented here can be enriched not only by collecting in places that have not been sampled, but also by using other sources, such as flowers, fungi, decomposing leaves and other attractants.

Acknowledgements

We thank the MPEG technicians José A. Pena and Luís Augusto Quaresma, for information on the localities described on the labels of specimens, coordination and making available equipment necessary to conduct this work; Leonardo C. Trevelin plotted the maps; Ana Luiza C. Brasil and Angélico A. Flores provided references; and Marcos S. Fialho made corrections to the manuscript. We thank Bridget Johnson for providing a review of the English in this manuscript. The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) provided a scholarship (process number 380618/2010-9). We also thank Instituto Nacional de Ciência e Tecnologia (INCT) and Programa de Pesquisa em Biodiversidade Amazônia Oriental (PPBio).

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Publication Dates

  • Publication in this collection
    2017

History

  • Received
    05 Mar 2016
  • Reviewed
    07 Nov 2016
  • Accepted
    28 Feb 2017
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