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

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.

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 2010).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 km 2 , is completely immersed in the Amazon Biome (IBGE 2010).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. 2005).
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. 2004).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 2015).We also verified and analyzed each literature reference for the state of Pará.
1903, Diathoneura Duda, 1924, Drosophila Fallén, 1823, Hirtodrosophila Duda, 1923, Mycodrosophila Oldenberg, 1914, Neotanygastrella Duda, 1925, Scaptodrosophila Duda, 1923, Zaprionus Coquillett, 1901and Zygothrica Wiedemann, 1830) and two belong to Steganinae (Leucophenga Mik, 1886 andRhinoleucophenga 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.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 andDrosophila 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, 1927) only from Costa Rica; and Drosophila neochracea (Wheeler, 1959) from Bolivia and Ecuador (Bächli 2015).The vittimaculosa group of Zygothrica (Z.vittinubila Burla, 1956), 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 pallidipennis Dobzhansky & Pavan, 1943, Zygothrica aldrichi Sturtevant, 1920and Zygothrica dispar Wiedemann, 1830).Three species, Drosophila polymorpha Dobzhansky & Pavan, 1943, Drosophila mesostigma Frota-Pessoa, 1954and 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).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, 1956and 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).However, here, we considered this record as belonging to Paraliodrosophila antennata Wheeler, 1957, following Vilela & Bächli (2007), 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 2015).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 1929).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 1936).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 1963), 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 2004) 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. 2012).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. 2008).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.
http://www.scielo.br/bnhttp://dx.doi.org/10.1590/1676-0611-BN-2016-0179 1 New records for the state of Pará; 2 New records for Brazil; 3 Species that are recorded in Brazil only in the state of Pará; 4 New records for Amazon 1 New records for the state of Pará; 2 New records for Brazil; 3 Species that are recorded in Brazil only in the state of Pará; 4 New records for Amazon