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INTRODUCTION
The general public wants to know more about natural environments and the organisms that live there (Quaresma 2003). Biodiversity professionals need to identify the groups that occur in the different biomes. An identification key that is updated and easy to use meets the demands of both the general and scientific audiences.
The Amazonian region holds a highly diverse fauna and flora (Hoorn and Wesselingh 2010). Large animals such as mammals (Sigrist 2012) and more conspicuous plants such as orchids (Luz and Oliveira 2012) are usually highlighted in Amazonian literature. Insects are mostly seen as a nuisance and/or risk (e.g. mosquitoes), with most people unaware of their great diversity (Rafael et al. 2009).
Syrphids, also known as flower/hover flies (Diptera: Syrphidae), are an example of poorly studied fauna in the Brazilian Amazon. The diversity of syrphid forms reaches extremes where they can be mistaken for wasps and bees (Hymenoptera) (Thompson et al. 2010). Most adults feed on nectar and pollen, with the larvae having habits and habitats much more diverse than the adults (Rotheray and Gilbert 1999; Pérez-Lachaud et al. 2014). The family is currently divided into four subfamilies: Eristalinae (mainly saprophagous larvae), Microdontinae (larvae that scavenge in ant nests), Pipizinae and Syrphinae (mainly predaceous larvae) (Thompson et al. 2010; Mengual et al. 2015). In Europe there is a long tradition in the study of syrphids, not only in academia (Bartsch 2009a; 2009b), but also with many amateurs/enthusiasts providing useful information on these flies (e.g. Owen 2010; Ball and Morris 2013). This encourages 'citizen-scientists' to develop an interest in the biodiversity around them, which can also aid on gathering data for ecological studies (Dickinson et al. 2010). The Amazon syrphid fauna could be explored in the same way as was done in Europe and in the Nearctic (Miranda et al. 2013). Thus, the objective of the current study is to develop a photographic identification key, as supplementary material, to the genera of Syrphidae from the Brazilian Amazon and present new records of distribution.
MATERIALS AND METHODS
Specimens and area studied
Material studied came from the following Brazilian institutions: Coleção de Invertebrados from the Instituto Nacional de Pesquisas da Amazônia (INPA)-Manaus/AM, Coleção Zoológica do Maranhão (CZMA)-Caxias/MA, Museu Nacional da Universidade Federal do Rio de Janeiro (MNRJ)-Rio de Janeiro/RJ and Museu Paraense Emílio Goeldi (MPEG)-Belém/PA. All voucher material remained at the institutions mentioned above. Terminology followed Thompson (1999) and Cumming and Wood (2009), with some terms simplified to ease understanding by the user (e.g. 'antennal base' instead of 'frontal prominence' or 'antennifer'). Some common terms for structures on syrphids (e.g. venation) and notes on orientation (e.g. anterior x posterior) can be found only in the supplementary material online.
The genera presented in this key were taken from the literature (Thompson et al. 1976; Thompson 1999; Reemer and Ståhls 2013a, 2013b) and later supplemented with new taxon records found in the material studied. Generic classification followed updated Neotropical Syrphidae literature (e.g. Miranda 2011; Reemer e Ståhls 2013a; 2013b; Miranda et al. 2014). Species lists were based in the literature (Thompson et al. 1976; Rotheray et al. 2000; Blatch et al. 2003; Borges and Pamplona 2003; Rotheray et al. 2007; Thompson 2007a, b; Mengual and Thompson 2008; Borges and Couri 2009; Mengual et al. 2009; Morales and Marinoni 2009; Reemer 2010; Thompson 2010; Miranda 2011; Montoya et al. 2012; Reemer 2012; Ricarte et al. 2012; Reemer and Ståhls 2013a; Miranda 2014; Miranda et al. 2014; Reemer 2014).
The study area was limited to the Brazilian Amazon, but took into consideration records for the Amazon of neighbouring countries (Bolivia, Colombia, French Guyana, Guyana, Peru, Surinam and Venezuela) since the taxa could occur in the Brazilian side as well. The 'Brazilian Amazon' considered in this study represents the 'Amazônia legal' (see figure 1 in Fearnside 2014), but limited to the rainforest areas (Acre, Amazonas, Amapá, western Maranhão, northern Mato Grosso, Pará, Rondônia, Roraima and northern Tocantins).
For records that indicated only Maranhão, Mato Grosso or Tocantins, without indicating where in those states, they were only considered part of the Brazilian Amazon if they had records in other Amazonian areas (e.g.: for a record indicating 'Mato Grosso, Venezuela' it was assumed that the species was part of the Amazonian part of Mato Grosso). Species records from the literature that indicated only 'Brazil' were not considered.
For cases where the species was assumed to occur in the Brazilian Amazon, the species name was followed by an asterisk (*) and either a neighbouring country name or 'Amazon'. A country name was used when the record was in an Amazonian region in that neighbouring country. 'Amazon' was used when the record in the literature only mentioned 'Amazon' or 'Amazon valley'.
New records found in the material studied were indicated after the species name by 'n. rec.' followed by 'BR', if they were new records for Brazil, and/or the acronym for the state (in bold) if it was a new record for that state, i.e.: AC-Acre, AM-Amazonas, AP-Amapá, MA-Maranhão, MT-Mato Grosso, PA-Pará, RO-Rondônia, RR-Roraima, and TO-Tocantins.
Images and key development
Most images were obtained by the author using a camera set on a stereomicroscope (M205, Leica, Wetzlar, Germany), and the software Leica Application Suite (v.3.6, Leica, Wetzlar, Germany), in the Laboratório de Entomologia Sistemática, Urbana e Forense (LESUF/CBio/INPA), or in the field with a DSLR camera (D3100, Nikon, Tokio, Japan) and macro lens (Macro 105mm/f2.8, Nikon, Tokio, Japan). Some images were obtained from other sources, which were indicated in the corner of the images. Images were edited in the Adobe Photoshop software (CS4, Adobe, San Jose, USA).
The steps of the key were made in the Adobe Illustrator software (CS4, Adobe, San Jose, USA) and exported as hyper-text pages (HTML). The format of the key followed Miranda et al. (2013), with steps leading to a diagnostic taxon page or to another step. There are two types of steps: (1) Multi-optional, where the user is shown several options of different character combinations that represent a group of taxa or a specific taxon; and (2) dichotomous, where there are only two options, of character combinations, to choose from. There may be links to species lists and additional images in the diagnostic taxon pages. Some taxa occurred more than once in the key due to the diversity of forms and intermediate cases that they possess.
Some images in the key had highlighted structures, that appeared in either of two ways: (1) a red frame highlighting the position of the structure on the specimen followed by another image on the side with the structure magnified and with a red border; or (2) an orange frame, highlighting the position of the structure on the specimen, which can be clicked to reveal a magnified view of the structure.
The PDF version was assembled from the individual HTML files through the 'Create PDF' (File/Create PDF/From Web Page...) tool from the Adobe Acrobat software (9 Pro, Adobe, San Jose, USA) by selecting the first step of the key and adjusting the 'Capture multiple levels' settings to 'Get the entire site'.
RESULTS
This study reports 64 taxa (61 genera and three subgenera) for the Brazilian Amazon region, and 36 were new records for Brazil and/or the Brazilian Amazon region (Table 1 and Appendix 1).
Table 1 New records of Syrphidae (Diptera) from the Brazilian Amazon. AC: Acre; AM: Amazonas; MA: Maranhão; PA: Pará; RR: Roraima.
| Species | New Brazilian record | New state record |
|---|---|---|
| Eristalinae | ||
| Cepa margarita (Thompson, 1999) | X | AM |
| Mallota sp. | X | AM |
| Meromacrus milesia Hull, 1942 | X | AM |
| M. pachypus (Wiedemann, 1830) | AM | |
| Nausigaster bonariensis Lynch-Arribalzaga, 1892 | PA | |
| Orthonevra sp. | AM, MA | |
| Palpada aemula (Williston, 1891) | X | PA |
| P. agrorum (Fabricius, 1787) | AM | |
| P. fasciata (Wiedemann, 1819) | X | AM, PA |
| Polybiomyia bigotii (Williston, 1888) | RR | |
| Quichuana longicauda Ricarte & Hancock, 2012 | X | AM |
| Microdontinae | ||
| Aristosyrphus (Aristosyrphus) sp. | AM | |
| Carreramyia sp. | X | AM |
| Ceratophya carinifacies (Curran, 1934) | MA | |
| Domodon zodiacus Reemer, 2013 | X | AM |
| Hypselosyrphus trigonus Hull, 1937 | AM | |
| Masarygus planifrons Brèthes, 1908 | PA | |
| Menidon falcatus (Williston, 1887) | AM | |
| Microdon macquartii Lynch-Arribalzaga, 1891 | X | AM |
| Pseudomicrodon polistoides Reemer, 2013 | X | AM |
| P. smiti Reemer, 2013 | X | AM |
| Schizoceratomyia barretoi Carrera, Lopes & Lane, 1947 | PA | |
| Stipomorpha apicula (Curran, 1930) | X | AM |
| S. goettei (Shannon, 1927) | AM | |
| S. guianica (Curran, 1925) | AM | |
| S. mackiei (Shannon, 1927) | AM | |
| Surimyia minutula (Doesburg, 1966) | X | AM |
| Pipizinae | ||
| Trichopsomyia polita Williston, 1888 | RR | |
| Syrphinae | ||
| Calostigma elnora Shannon, 1927 | X | AM |
| Eosalpingogaster cochenillivora (Guérin-Méneville, 1848) | PA | |
| Hybobathus placivus (Williston, 1888) | PA | |
| Ocyptamus icarus Reemer, 2010 | X | AC |
| O. obliquus (Curran, 1941) | AM | |
| O. prenes (Curran, 1930) | AM | |
| Pelecinobaccha pandora (Hull, 1941) | AC | |
| Xanthandrus plaumanni Fluke, 1937 | AM, RR | |
| Total: 36 taxa |
The most recent keys for identification of species, updated from Thompson (1999), are listed on Tables 2, 3 and 4. The key (online version and a "supplementary material" PDF version), instructions about orientation, and basic terminology can be accessed through the website http://keys.inpa.gov.br/?idkey=syrphidae.
Table 2 Available references with species keys for Eristalinae (Syrphidae) with records for the Brazilian Amazon.
| Genus | Reference |
|---|---|
| Eristalinae | |
| Alipumilio | Vockeroth (1964) |
| Cepa | Thompson (2007a) |
| Chalcosyrphus (Neplas) | Curran (1941) (as Planes) and Thompson (1981) |
| Copestylum | Curran (1939, 1953) (as Volucella), Fluke (1951) and Thompson (1981) |
| Habromyia | Curran (1934) |
| Lepidomyia | Hull (1946) (as Lepidostola) |
| Lycastrirhyncha | Doesburg (1963) |
| Mallota | Single species is illustrated in this study |
| Meromacrus | Hull (1942), Thompson (1981) and Blatch et al. (2003) |
| Monoceromyia | Curran (1941) and Thompson (1981) (as Ceriana) |
| Myolepta | Thompson (1968) |
| Nausigaster | Curran (1941) and Carrera et al. (1947) (species published post Curran (1941)) |
| Ornidia | Carvalho-Filho and Esposito (2009) |
| Orthonevra | Single species is illustrated in this study |
| Palpada | Lagrange (1992), Mengual and Thompson (2008) and Morales and Marinoni (2009) |
| Polybiomyia | Curran (1941) |
| Quichuana | Ricarte et al. (2012) |
| Rhingia | Fluke (1943) |
| Senogaster | Single species is illustrated in this study |
| Sphiximorpha | Curran (1941) |
| Sterphus (Ceriogaster) | Hippa and Thompson (1994) and Zumbado and Thompson (1997) |
| Sterphus (Crepidomyia) | Hippa and Thompson (1994) and Zumbado and Thompson (1997) |
Table 3 Available references with species keys for Microdontinae (Syrphidae) with records for the Brazilian Amazon.
| Genus | Reference |
|---|---|
| Microdontinae | |
| Aristosyrphus (Aristosyrphus) | No available species key |
| Aristosyrphus (Eurypterosyrphus) | No available species key |
| Carreramyia | Reemer (2013) |
| Ceratophya | Reemer (2013) |
| Ceriomicrodon | Miranda (2014) |
| Chrysidimyia | Reemer and Ståhls (2013a) |
| Domodon | Reemer and Ståhls (2013a) and Reemer (2014) (no key, only original descriptions) |
| Hypselosyrphus | Reemer (2013) |
| Laetodon | Doesburg (1966) (as Microdon geijskesi) |
| Masarygus | Reemer and Ståhls (2013a) (see discussion about the genus and the description of M. palmipalpus) |
| Menidon | Thompson (2007b) and Reemer and Ståhls (2013a) |
| Microdon (Chymophila) | Curran (1941) |
| Microdon (Microdon) | Curran (1941) |
| Mixogaster | Carrera and Lenko (1958) |
| Peradon | Curran (1941) (as Microdon) |
| Piruwa | Reemer and Ståhls (2013a) |
| Pseudomicrodon | Curran (1941) (as Microdon) |
| Rhoga | Reemer (2012) |
| Rhopalosyrphus | Thompson (2012) |
| Schizoceratomyia | Papavero (1962) (as Masarygus) |
| Stipomorpha | Reemer (2013) |
| Surimyia | Reemer (2008) |
| Ubristes | Reemer (2013) |
Table 4 Available references with species keys for Pipizinae and Syrphinae (Syrphidae) with records for the Brazilian Amazon.
| Genus | Reference |
|---|---|
| Pipizinae | |
| Trichopsomyia | Fluke (1937) |
| Syrphinae | |
| Allograpta | Fluke (1942) and Thompson (1981) |
| Argentinomyia | Fluke (1945) (as Rhysops) |
| Atylobaccha | Miranda et al. (2014) |
| Calostigma | Hull (1949) |
| Eosalpingogaster | Mengual and Thompson (2011) |
| Fazia | Fluke (1942) (as Epistrophe) |
| Hermesomyia | Vockeroth (1969) and Rotheray et al. (2000) |
| Hybobathus | Hull (1949) (as Baccha) and Reemer (2010) (as Ocyptamus) |
| Leucopodella | Thompson (1981) and Carvalho (2011) |
| Mimocalla | Thompson and Zumbado (2000) |
| Ocyptamus | Hull (1949) (as Baccha), Thompson (1981) and Reemer (2010) |
| Orphnabaccha | There are no appropriate current keys that cover the region |
| Pelecinobaccha | Miranda et al. (2014) |
| Pseudodoros | Kassebeer (2000) |
| Relictanum | Miranda et al. (2014) |
| Salpingogaster | Curran (1941) and Thompson (1981) |
| Toxomerus | Thompson (1981), Borges and Couri (2009) and Mengual (2011) |
| Xanthandrus | Borges and Pamplona (2003) |
The study of Reemer (2010) found a specimen of Paragus (Pandasyophthalmus) cf. haemorrhous in Surinam. The genus is present in North America, extending south at most into Costa Rica in Central America. Due to the single record in Reemer (2010), and the absence of specimens in the collections studied, this genus is not considered to be established in the Amazonian region and was not included in the key. Microdon (Syrphipogon) was not included in the key due to lack of conclusive evidences that its distribution could range into the Amazonian region (records only for Costa Rica, Panama and one record in the state of Paraná, Brazil).
A specimen of Polybiomyia, with a complete post-metacoxal bridge, was identified as Cerioides bigotii (Williston, 1888) in Curran (1941) key. Thompson (2010) recognizes it as Sphiximorpha bigotii (Williston, 1888), but the condition of the post-metacoxal bridge does not agree with Sphiximorpha (where it should be incomplete), so it is here placed in Polybiomyia.
DISCUSSION
This study discovered new taxon records for the Brazilian fauna of Syrphidae (Table 1), which reflects how much the fauna of this country, and specially the Amazonian region, is still underestimated. As an example, Mallota was only known from high altitude rain forests in South America, but this study found specimens in one of the few high altitude spots in the Brazilian Amazon. Another example was Carreramyia, unknown for Brazil but found in two different localities of the Amazonian lowlands (North and South of the Amazonas river), being captured only in canopy traps. These previously unknown taxa found in the Brazilian Amazon demonstrate the need to further study and definitely preserve this region and its still unknown biodiversity.
The key presented here will ease identification of syrphid genera in the Brazilian Amazon, allowing for a quicker assessment of new generic records, and also serving as an initial framework for regional taxonomic revisions and ecological studies. The compilation of references (Tables 2, 3 and 4) will direct the user to the most current species keys, and allied to the updated species lists in each taxon page, will also aid in identifying undescribed species or new records for the Brazilian Amazon.
CONCLUSIONS
Thirty-six new records of species and genera of Syrphidae are found, demonstrating the still unknown biodiversity present in the Brazilian Amazon. Furthermore, this study provides an updated identification key, and list of references to species keys, providing an initial study framework that will aid in future ecological studies and taxonomical revisions.
