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

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

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 2015). This number will most likely increase as new species are continuously being described, particularly in the Neotropical region (Silva & Martins 2004, Vilela & Bächli 2005, Ratcov & Vilela 2007, Culik & Ventura 2009, Schmitz et al. 2009, Figuero & Rafael 2011, Figuero et al. 2012, Gottschalk et al. 2012, Acurio et al. 2013, Poppe et al. 2014, Junges & Gottschalk 2014, Guillín & Rafael 2015, Junges et al. 2016, Vidal & Vilela 2015). In Brazil, the Drosophilidae family is represented by 16 genera and 305 species (Tidon et al. 2015). Taxonomic inventories are needed to we fill the knowledge gaps on biodiversity (Hortal et al. 2015). Althought the first collections for the Amazon dating from 1920-1959 period (Duda 1927; Hendel 1936; Dobzhansky & Pavan 1943; Pavan 1950; Pavan 1959), 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. 2008). Among the Amazonian states, only Amazonas and Pará have more consistent inventory Drosophilid (Bächli 2015). 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 1927; Hendel 1936; Burla et al. 1949), evolutionary biologists (Pavan 1950) and more recently by ecologists (De Toni et al. 2005; Martins & Oliveira 2007; Martins & Santos 2007; Silva & Martins 2009; Praxedes & Martins 2014 and Robe et al. 2014). Currently, in Pará there are 75 Drosophilidae species from nine genera that are from two subfamilies Steganinae and Drosophilinae (Bächli 2015).

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 km², 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.

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.

Species	Municipalities	Records	# in col
Subfamily Drosophilinae
Genus Chymomyza
aldrichii group
3Chymomyza bicoloripes (Malloch, 1926)	ALM and SAN	Hendel (1936)
Chymomyza diatropa Grimaldi, 1986	MEL and POR	MPEG and Schmitz et al. (2013)	23
Genus Diathoneura
1,2,4Diathoneura flavolineata Duda, 1927	BEL	MZUSP
Genus Drosophila
Subgenus Drosophila
annulimana group
1Drosophila annulimana Duda, 1927	BEL, MEL and POR	MPEG	5
Drosophila araicas Pavan & Nacrur, 1950	ALM, BEL,MEL and POR	MPEG and Val (1982)	7
1,4Drosophila arapuan Da Cunha & Pavan, in Pavan & Da Cunha, 1947	PARP	MPEG	1
Drosophila ararama Pavan & Da Cunha, 1947	BEL and NIP	MPEG, Pavan & Cunha (1947)	14
3Drosophila caxiuana Gottshalk et al., 2012	MEL and POR	MPEG and Gottschalk et al. (2012)	5
bromeliae group
3Drosophila speciosa Silva & Martins, 2004	BEL,CUR, MEL and POR	MPEG and Silva & Martins (2004)	429
calloptera group
Drosophila atrata Burla & Pavan, 1953	AVE and PARP	MPEG and Burla & Pavan (1953)	2
Drosophila calloptera Schiner, 1868	BEL, MEL, POR and SAN	MPEG and Burla & Pavan (1953)	5
1,4Drosophila quadrum (Wiedemann, 1830)	OBI	MZUSP
canalinea group
3Drosophila annularis Sturtevant, 1916	SAN	Hendel (1936)
1,4Drosophila annulosa Duda, 1929	MEL and POR	MPEG	2
Drosophila canalinea Patterson & Mainland, 1944	BEL, MEL, PARP and POR	MPEG and Patterson & Stone (1952)	1
Drosophila davidgrimaldii Vilela & Bächli, 1990	MEL, SAN and POR	MPEG and Vilela & Bächli (1990)	1
3Drosophila hendeli Vilela & Bächli, 1990	SAN	Vilela & Bächli (1990)
caponei group
1,4Drosophila caponei Pavan & Cunha, 1947	BEL	MPEG	9
1,2,4Drosophila neochracea Wheeler, 1959	BEL and TUC	MPEG	5
cardini group
cardini subgroup
Drosophila cardini Sturtevant, 1916	BEL, NIP, PAC and PARP	MPEG and Heed & Russell (1971)	17
Drosophila cardinoides Dobzhansky & Pavan, 1943	AVE, BEL, IGA, MAR, NIP and PARP	MPEG and Streisinger (1946)	9
Drosophila neocardini Streisinger, 1946	MEL, NIP, PAC and POR	MPEG and Silva & Martins (2009)	2
Drosophila neomorpha Heed & Wheeler, 1957	ALM, BEL, JUR, MEL, NIP, PAC, PARP and POR	MPEG and De Toni et al. (2005)	11
Drosophila parthenogenetica Stalker, 1953	MEL, NIP, PAC and POR	MPEG and De Toni et al. (2005)	6
Drosophila polymorpha Dobzhansky & Pavan, 1943	BEL and BRE	Dobzhansky & Da Cunha (1955)
coffeata group
Drosophila coffeata Williston, 1896	ALM, BEL, MEL, PARP, POR and SAN	MPEG and Hendel (1936)	91
Drosophila fuscolineata Duda, 1925	BEL	MPEG and Dobzhansky & Pavan (1950)	2
dreyfusi group
Drosophila camargoi Dobzhansky & Pavan in Pavan, 1950	BEL BRE, MEL, NIP and POR	MPEG and Pavan (1959)	71
3Drosophila decemseriata Hendel, 1936	SAN	Hendel (1936)
flavopilosa group
3Drosophila melina Wheeler, 1962	BEL	MPEG and Santos & Martins 2000	256
guarani group
guaramunu subgroup
1,4Drosophila griseolineata Duda, 1927	PARP	MPEG	1
guarani subgroup
Drosophila ornatifrons Duda, 1927	BEL	MPEG and Pavan (1959)	1
pallidipennis subgroup
Drosophila pallidipennis Dobzhansky & Pavan, 1943	BEL	Dobzhansky & Pavan (1950)
peruensis subgroup
Drosophila peruviana Duda, 1927	AVE and BEL	MZSP and Pavan (1959)
repleta group
fasciola subgroup
Drosophila ellisoni Vilela, 1983	ALM, BEL, IGA, JUR, NIP and PAC	MPEG and Vilela (1983)	11
1,2,3,4Drosophila fasciola Williston, 1896	NIP, PAC and PARP	MPEG	7
Drosophila fascioloides Dobzansky & Pavan, 1943	BEL	Wasserman (1962)
1,4Drosophila ivai Vilela, 1983	ALM, NIP, PAC, PARP and POR	MPEG	19
1,4Drosophila mapiriensis Vilela & Bächli, 1990	BEL, MEL, NIP and POR	MPEG	6
Drosophila moju Pavan, 1950	BEL, JUR, NIP and PARP	MEPG and Vilela (1983)	11
3Drosophila mojuoides Wasserman, 1962	BEL	MPEG and Vilela (1983)	1
1,4Drosophila papei Bächli & Vilela, 2002	NIP and PARP	MPEG	12
1,4Drosophila querubimae Vilela, 1983	MEL, NIP, PAC and POR	MPEG	3
hydei subgroup
Drosophila hydei Sturtevant, 1921	BEL	MPEG and Martins & Oliveira (2007)	6
mercatorum subgroup
1Drosophila mercatorum Patterson & Wheeler, 1942	BEL	MPEG	1
1Drosophila paranaensis Barros, 1950	NIP and PARP	MPEG	7
mulleri group
1Drosophila aldrichi Patterson, in Patterson & Crow, 1940	ALT, MEL and POR	MPEG	10
repleta subgroup
Drosophila eleonorae Tosi et al., 1990	ALT	MPEG and Tosi et al. (1990)	41
Drosophila fulvimacula Patterson & Mainland, 1944	ALM, BEL, JUR, MEL, NIP, PAC, PARP, POR	MPEG and Dobzhansky & Da Cunha (1955)	52
1,4Drosophila limensis Pavan & Patterson in Pavan & Da Cunha, 1947	MEL and POR	MPEG	2
1Drosophila repleta Wollaston, 1858	MEL and POR	MPEG	11
tripunctata group
I subgroup
Drosophila mediocris Frota-Pessoa, 1954	MEL and POR	MPEG and Praxedes & Martins (2014)	1
Drosophila neoguaramunu Frydenberg, 1956	BEL	MPEG and Martins & Santos (2007)	1
1,4Drosophila setula Heed & Wheeler, 1957	PAC	MPEG	3
II subgroup
1,4Drosophila cuaso Bächli, Vilela & Ratcov, 2000	ALM, BEL, MEL, NIP, PARP and POR	MPEG	9
III subgroup
Drosophila addisoni Pavan, 1950	BEL	Pavan (1959)
Drosophila frotapessoai Vilela & Bächli, 1990	BEL, MEL and POR	MPEG and Martins & Oliveira (2007)	1
Drosophila medioimpressa Frota-Pessoa, 1954	MEL and POR	MPEG and Praxedes & Martins (2014)	1
Drosophila mediopicta Frota-Pessoa, 1954	MEL and POR	MPEG and Praxedes & Martins (2014)	1
Drosophila mediostriata Duda, 1925	BEL, MEL and POR	Martins & Oliveira (2007)
Drosophila mesostigma Frota-Pessoa, 1954	BEL	Val (1982)
1,4Drosophila paramediostriata Townsend & Wheeler, 1955	BEL and PARP	MPEG	4
1,4Drosophila trapeza Heed & Wheeler, 1957	PARP	MPEG	1
IV subgroup
1,4Drosophila albicans Frota-Pessoa, 1954	PARP	MPEG	4
Without subgroup
3Drosophila argenteifrons Wheeler, 1954	CAS	Wheeler (1954)
Subgenus Phloridosa
1,4Drosophila lutzii Sturtevanti, 1916	BEL	MPEG	6
Subgenus Sophophora
melanogaster group
ananassae subgroup
Drosophila ananassae Doleschall, 1858	BEL, BRA, IGA, MEL, NIP, PAC, PRAP and POR	MPEG and Praxedes & Martins (2014)	107
Drosophila malerkotliana Parshad & Paika, 1964	ALM, BEL, BRA, BRE, IGA, JUR, MEL, NIP, PAC, PARG, PARP, POR, SAN, SAL and VIS	MPEG and Martins (2001)	533
melanogaster subgroup
Drosophila melanogaster Meigen, 1830	BEL, MEL and POR	MPEG and David et al. (2006)	15
Drosophila simulans Sturtevant, 1919	ALM, BEL, IGA, JUR, MEL, NIP and POR	MPEG and Dobzhansky & Pavan (1950)	44
montium subgroup
Drosophila kikkawai Burla, 1954	BEL, IGA, MEL, NIP, PAC, PARP and POR	MPEG and Praxedes & Martins (2014)	88
saltans group
cordata subgroup
Drosophila neocordata Magalhães, 1956	ALM, BEL, MEL, NIP and POR	MPEG and Praxedes & Martins (2014)	9
elliptica subgrupo
Drososphila neoelliptica Pavan & Magalhães in Pavan, 1950		MPEG and Praxedes & Martins (2014)	1
parasaltans subgroup
Drosophila parasaltans Magalhães, 1956	ALM, JUT, MEL and POR	MPEG and Praxedes & Martins (2014)	2
Drosophila subsaltans Magalhães, 1956	ALM, BEL, JUR, MEL, NIP, PAC and POR	MPEG and Magalhães (1956)	2
saltans subgroup
Drosophila austrosaltans Spassky, 1957	BEL, JUR, MEL and POR	MPEG and Praxedes & Martins (2014)	2
Drosophila prosaltans Duda, 1927	ALM, ALT, AVE, BEL, BRA, BRE, JUR, MEL, NIP, PAC, PARP and POR	MPEG and Dobzhansky & Streisinger (1944)	3
Drosophila pseudosaltans Magalhães, 1956	MEL and POR	MPEG and Bicudo (1973)	16
Drosophila saltans Sturtevant, 1916	ALM, MEL and POR	MPEG and De Toni et al. (2005)	1
sturtevanti subgroup
Drosophila dacunhai Mourão & Bicudo, 1967	MEL and POR	MPEG and Martins et at. (2008)	5
3Drosophila milleri Magalhães, 1962	MEL and POR	MPEG and Praxedes & Martins (2014)	5
Drosophila magalhaesi Mourão & Bicudo, 1967	ALM, BEL, JUR, MEL and POR	MPEG and Martins et at. (2008)	18
Drosophila sturtevanti Duda, 1927	ALM, AVE, BEL, BRE, IGA, ITA, JUR, MEL, NIP, PAC, PARP, POR and SAN	MEPG and Dobzhansky (1944)	132
willistoni group
bocainensis subgroup
Drosophila capricorni Dobzhansky & Pavan, 1943	BEL, MEL and POR	MPEG and Dobzhansky & Pavan (1950)	1
Drosophila fumipennis Duda, 1925	BEL, BRE, JUR, MEL and POR	MPEG and Pavan (1959)	35
Drosophila nebulosa Sturtevant, 1916	ALM, AVE, BEL, BRE, IGA, JUR, MEL, NIP, PAC, PARG, PARP, POR and SAL	MEPG and Dobzhansky & Pavan (1950)	63
willistoni subgroup
Drosophila equinoxialis Dobzhansky, 1946	ALM, BEL, BRE, IGA, MEL, NIP, PAC, PARP, POR and SAN	MPEG and Burla et al. (1949)	23
Drosophila paulistorum Dobzhansky & Pavan in Burla et al., 1949	ALM, BEL, BRE, IGA, JUR, MEL, NIP, PAC, PRAP, POR and SAN	MPEG and Townsend (1954)	105
Drosophila tropicalis Burla et al., 1949	BEL, MEL, POR, SAN and SAL	MPEG and Burla et al. (1949)	26
Drosophila willistoni Sturtevant, 1916	ALM, BEL, BRA, BRE, IGA, ITA, JUR, MEL, NIP, PAC, PARP, POR and SAN	MPEG and Dobzhansky & Mayr (1944)	104
Ungrouped species
Drosophila impudica Duda, 1927	BEL, JUR, NIP, PAC and PARP	MPEG, MZSP and Pavan (1959)	8
Drosophila tuchaua Pavan, 1950	BEL, MEL and POR	MPEG and Pavan (1950)	55
Subgenus Siphlodora
1,4Drosophila flexa Loew, 1866	NIP	MPEG	2
Hirtodrosophila
Ungrouped species
3Hirtodrosophila pictiventris Duda, 1925	Localities not mentioned	Vilela & Bächli (2004)
Hirtodrosophila subflavohalterata Burla, 1956	MEL and POR	Robe et al. 2014
Genus Mycodrosophila
3Mycodrosophila brunnescens Wheeler & Takada, 1963	BEL	Wheeler &Takada (1963)
3Mycodrosophila elegans Wheeler & Takada, 1963	CAS	Wheeler &Takada (1963)
Mycodrosphila projetans Wheeler & Takada, 1963	MEL and POR	Robe et al.(2014)
Genus Neotanygastrella
3Neotanygastrella chymomyzoides Duda, 1927	BEL	Hendel (1936)
Neotanygastrella tricoloripes Duda, 1925	SAN	Hendel (1936)
Genus Paraliodrosophila
Paraliodrosophila antennata Wheeler, 1957		Wheeler (1954)
latifasciaeformis group
Scaptodrosophila latifasciaeformis (Duda, 1940)	BEL, BRE, IGA, JUR, MEL, NIP, PAC, PARP and POR	MPEG and Pavan (1959)	50
Genus Zaprionus
vittiger group
Zaprionus indianus Gupta, 1970	ALM, BEL, MEL, NIP, PAC, PARP, POR and SAN	MEPG and David et al. (2006)	8
Genus Zygothrica
dispar group
aldrichii subgroup
3Zygothrica aldrichi Sturtevant, 1920	BEL	Hendel (1936)
3Zygothrica mediovitta Grimaldi, 1987	SAN	Grimaldi (1987)
Zygothrica microeristes Grimaldi, 1987	ALM, BEL, ITA, OBI and ORI	MZSP and Grimaldi (1987)
Zygothrica paraldrichi Burla, 1956	ALM, BEL, ITA, MAR, OBI and SAN	MZSP and Grimaldi (1987)
Zygothrica pilipes Hendel, 1936	ITA, OBI and SAN	MZSP and Hendel (1936)
3Zygothrica radialis Grimaldi, 1987	ALM, BEL and ITA	MZSP and Grimaldi (1987)
3Zygothrica somatia Grimaldi, 1987	ALM, OBI and SAN	MZSP and Grimaldi (1987)
Zygothrica zygia Grimaldi, 1987	OBI and SAN	Grimaldi (1987)
caudata subgroup
Zygothrica caudata (Hendel, 1913)	SAN	Hendel (1936)
dispar subgroup
Zygothrica dispar (Wiedmann, 1830)	SAN	MZUSP and Hendel (1936)
Zygothrica prodispar Duda, 1925	BEL, ITA, MAR and SAN	MPEG and Grimaldi (1987)	3
atriangula group
Zygothrica atriangula Duda, 1927	SAN	Burla (1956)
atriangulata group
Zygothrica virgatinigra Burla, 1956	MEL and POR	Robe et al. (2014)
bilineata group
Zygothrica bilineata (Williston, 1896)	ITA	Grimaldi (1990)
orbitalis group
Zygothrica orbitalis (Sturtevant, 1916)	SAN	Hendel (1936)
vittinubila group
1,4Zygothrica vittinubila Burla, 1956	NIP	MPEG and MZUSP	1
Ungrouped species
3Zygothrica laevifrons Duda, 1927	SAN	Hendel (1936)
Subfamily Steganinae
Genus Leucophenga
3Leucophenga argenteofasciata Kahl, 1917	SAN	Duda (1927)
Genus Rhinoleucophenga
1,4Rhinoleucophenga punctulata Duda, 1929	CAS	MPEG	5

• ¹
  New records for the state of Pará; ²New records for Brazil; ³Species that are recorded in Brazil only in the state of Pará; ⁴New records for Amazon

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, 1927) only from Costa Rica; and Drosophila neochracea (Wheeler, 1959) from Bolivia and Ecuador (Bächli 2015). The vittimaculosa group of Zygothrica (Z. vittinubilaBurla, 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 pallidipennisDobzhansky & Pavan, 1943, 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). 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). 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.

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