Research Article |
Corresponding author: Favízia Freitas de Oliveira ( favosgyrl@gmail.com ) Academic editor: Thorleif Dörfel
© 2020 Favízia Freitas de Oliveira, Lívia Raquel de Sousa Silva, Fernando César Vieira Zanella, Caroline Tito Garcia, Heber Luiz Pereira, Claudia Quaglierini, Camila Magalhães Pigozzo.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
de Oliveira FF, de Sousa Silva LR, Zanella FCV, Garcia CT, Pereira HL, Quaglierini C, Pigozzo CM (2020) A new species of Ceratina (Ceratinula) Moure, 1941, with notes on the taxonomy and distribution of Ceratina (Ceratinula) manni Cockerell, 1912, and an identification key for species of this subgenus known from Brazil (Hymenoptera, Apidae, Ceratinini). ZooKeys 1006: 137-165. https://doi.org/10.3897/zookeys.1006.57599
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A new species of the small carpenter bee, genus Ceratina (Ceratinula) Moure, from the Cerrado Biome in midwestern Brazil is described and illustrated. Ceratina (Ceratinula) fioreseana Oliveira, sp. nov. is easily distinguished from its congeners by the size of the facial maculations and the honey-yellow color of the legs and antennal scape, which distinguish it especially from Ceratina (Ceratinula) manni Cockerell, 1912, the most similar species in terms of facial maculation patterns. The geographic records of C. manni, here interpreted as endemic to the semiarid Caatinga region in northeastern Brazil, are presented, with new records for the Brazilian states of Piauí, Ceará and Bahia. A morphological description of both species is provided, including a comparison with the type specimen of C. manni from the state of Paraíba (Guarabira, formerly named Independencia). An identification key is provided for the described species of Ceratina (Ceratinula) recorded for Brazil according to Moure’s Catalogue of Neotropical Bees.
Anthophila, biogeography, small carpenter bees, South American diagonal of open formations, taxonomy
Bees (Hymenoptera, Anthophila) are a diverse group of insects, with more than 20,000 described species and many more estimated to exist (
The small carpenter bees comprise a group of mainly solitary bees, with only a single genus, Ceratina Latreille 1802 (Ceratinini) (
The genus Ceratina is cosmopolitan and highly diverse, although in Australia the group is rare and limited in distribution (
Subgenera | Species number | Geographic distribution | |
---|---|---|---|
01 | Calloceratina Cockerell, 1924 | 15 | New World: Neotropical Region (Belize, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Panama, Paraguay, Trinidad and Tobago and Venezuela) and marginally in Nearctic Region (Mexico). |
02 | Catoceratina Vecht, 1952 | 2 | Old World: Burma, Thailand, and the Philippine Islands south to Sumatra, Java, and Borneo. |
03 | Ceratina s. str. Latreille, 1802 | 29 | Old World: France to Turkey, and south through Africa to South Africa, and east through Eurasia to Japan, and Thailand. |
04 | Ceratinidia Cockerell & Porter, 1899 | 50 | Old World: Sri Lanka and India throughout southeastern Asia, north through China to the maritime province of Siberia, also including all of Japan, Taiwan, the Philippines, and Indonesia east to the western tip of New Guinea. |
05 | Ceratinula Moure, 1941 | 37 | New World: Nearctic Region: United States of America (Florida, Georgia, Louisiana, North Carolina, Texas); Neotropical Region: Argentina (Misiones); Bahamas; Belize; Bolivia; Brazil (Amazonas, Ceará, Maranhão, Minas Gerais, Paraná, Paraíba, Pará, Rio de Janeiro, São Paulo); Costa Rica (Alajuela, San José); Cuba; Dominican Republic; Haiti; Honduras; Panama (Chiriquí, Coclé, Panamá); Paraguay; Peru (Loreto); Saint Vincent and the Grenadines; Trinidad and Tobago. |
06 | Chloroceratina Cockerell, 1918 | 2 | Old World: From northern Luzon, in the Philippine Islands. |
07 | Copoceratina Terzo & Pauly, 2001 | 2 | Old World: From Kenya to South Africa, Madagascar and Seychelles. |
08 | Crewella Cockerell, 1903 | 32 | New World: Neotropical Region: Argentina, Bolivia, Brazil, Colombia, French Guiana, Guyana, Panama, Paraguay, Uruguay, Venezuela. |
09 | Ctenoceratina Daly & Moure, 1988 | 10 | Old World: Africa, from Senegal to Ethiopia, south to South Africa. |
10 | Dalyatina Terzo, 2007 | 7 | Old World: From Spain, France, Croatia, Greece, Turkey, Turkmenistan to subsaharian Africa. |
11 | Euceratina Hirashima, Moure & Daly, 1971 | 38 | Old World: From Britain, Spain, and Morocco east through Europe and the Mediterranean basin to southern Russia, Pakistan, and Somalia. |
One species, the parthenogenetic Ceratina dallatorreana Friese, is established in California, introduced by commerce. | |||
12 | Hirashima Terzo & Pauly, 2001 | 7 | Old World: From Tanzania and Nigeria south to South Africa, and Madagascar and Aldabra. |
13 | Lioceratina Vecht, 1952 | 8 | Old World: From India through Southeast Asia, Indonesia as far east as Bali, Sulawesi, and Philippines. |
14 | Malgatina Terzo & Pauly, 2001 | 1 | Old World: Madagascar. |
15 | Megaceratina Hirashima, 1971 | 1 | Old World: Africa, from Senegal to Zaire and east to Uganda. |
16 | Neoceratina Perkins, 1912 | 13 | Old World: From Turkey and Cyprus east through southwest and southern Asia and Indonesia to southern China, the Ryukyu Islands, Micronesia, Philippines, and south to the Bismarck Archipelago, the Solomon Islands, and eastern Australia as far as New South Wales. |
One species was introduced in Hawaii ( |
|||
17 | Neoclavicera Roig-Alsina, 2013 | 11 | New World: South America: from Argentina, Uruguay, Brazil, Paraguay, Bolivia and Peru. |
18 | Pithitis Klug, 1807 | 25 | Old World: Africa: from Senegal to Egypt south throughout Africa to Cape Province; Crete, eastward in Saudi Arabia, Yemen, Pakistan, India, and SriLanka, throughout southeast Asia to Philippines, Taiwan, Ryukyu Islands, southeast China, and through Indonesia east as far as Ambon. |
One species was introduced in Hawaii ( |
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19 | Protopithitis Hirashima, 1969 | 2 | Old World: Africa: Gabon, Congo, Angola, Zambia, Zaire, Tanzania, Mozambique, South Africa. |
20 | Rhysoceratina Roig-Alsina, 2013 | 10 | New World: South America: Argentina, Uruguay, Brazil, Paraguay, Colombia and Venezuela. |
21 | Simioceratina Daly & Moure, 1988 | 3 | Old World: Liberia to Kenya south to Namibia and Natal Province, South Africa. |
22 | Xanthoceratina Vecht, 1952 | 7 | Old World: Sri Lanka, Burma and Southeast Asia, including Indonesia as far east as Java, Philippines, and southern China. |
23 | Zadontomerus Ashmead, 1899 | 29 | New World: From Quebec and British Columbia in Canada to south throughout North and Central America until northern Colombia and Venezuela. |
Total of placed species | 338 | Several other species have not been yet placed into any subgenus. |
Some investigators have considered the genus Ceratina as a key taxon for understanding the transition from subsocial to social behavior, as they have a broad range of social behaviors, ranging from solitary, subsocial, and semisocial to eusocial colony organization (
Of the five subgenera with species recorded in Brazil, Ceratina (Ceratinula) Moure, 1941 is the most diverse, with 37 species, followed by: C. (Crewella) Cockerell, 1903 (32 species); C. (Calloceratina) Cockerell, 1924 (15 species); C. (Neoclavicera) Roig-Alsina, 2013 (11 species); and C. (Rhysoceratina) Michener, 2000 (10 species) (
Species of Ceratina (Ceratinula) Moure, 1941 recorded in Brazil according to “Catalogue of Bees (Hymenoptera, Apoidea) in the Neotropical Region – online version” (
Species | Type locality* and additional geographic records** | Domain | |
---|---|---|---|
1 | Ceratina (Ceratinula) augochloroides Ducke, 1910 | *Serra de Baturité (Ceará, Brazil) | Rain Forest highland enclave in Caatingas |
#Manaus (Amazonas, Brazil) | |||
2 | Ceratina (Ceratinula) biguttulata (Moure, 1941) | *Curitiba (Paraná, Brazil) | Atlantic Rain Forest |
#São Paulo (São Paulo, Brazil), Rio Grande do Sul (Brazil) | |||
3 | Ceratina (Ceratinula) combinata Friese, 1910 | *Belém (Pará, Brazil) | Amazonian |
4 | Ceratina (Ceratinula) fulvitarsis Friese, 1925 | *São Paulo (São Paulo, Brazil) | Atlantic Rain Forest |
5 | Ceratina (Ceratinula) immaculata Friese, 1910 | *Belém, Itaituba (Pará, Brazil) | Amazonian |
6 | Ceratina (Ceratinula) lucidula Smith, 1854 | *Santarém (Pará, Brazil) | Amazonian and Atlantic Rain Forest |
*(synonymous): São Paulo (São Paulo, Brazil) | |||
**Ceará, Minas Gerais, Pará, São Paulo (Brazil), Paraguay | |||
It is possible that different species are being considered as C. lucidula. | #Rio de Janeiro, Paraná, Rio Grande do Sul (Brazil) | ||
7 | Ceratina (Ceratinula) manni Cockerell, 1912 | *Guarabira (Paraíba, Brazil) | Caatingas |
#São Paulo (Brazil) | |||
8 | Ceratina (Ceratinula) melanochroa (Moure, 1941) | *Curitiba (Paraná, Brazil) | Atlantic Rain Forest |
#Rio Grande do Sul (Brazil) | |||
9 | Ceratina (Ceratinula) minima Friese, 1908 | *Trinidad | Caribe-Guajira Subequatorial and Amazonian |
**Trinidad and Tobago, Brazil | |||
# Merida (Venezuela) | |||
10 | Ceratina (Ceratinula) muelleri Friese, 1910 | *Belém, Óbidos (Pará, Brazil) | Amazonian and Atlantic Rain Forest |
It is possible that different species are being considered as C. muelleri. The diagnosis presented in the key is based on specimens collected in one of the type locality (Belém, Pará, Brazil), which fit perfectly with the original description of the species. | **Misiones (Argentina), Amazonas, Ceará, Maranhão, Minas Gerais, Paraná, Pará, Rio de Janeiro, São Paulo (Brazil) | ||
# Trinidad and Tobago, Rio Grande do Sul (Brazil) | |||
11 | Ceratina (Ceratinula) piracicabana Schrottky, 1910 | *Piracicaba (São Paulo, Brazil) | Atlantic Rain Forest |
#Brazil | |||
12 | Ceratina (Ceratinula) sclerops Schrottky, 1907 | * Encarnación (Itapúa, Paraguay) | Atlantic Rain Forest |
**Paraná, São Paulo (Brazil), Paraguay | |||
#Rio Grande do Sul (Brazil) | |||
13 | Ceratina (Ceratinula) turgida (Moure, 1941) | *Itatiaia (Rio de Janeiro, Brazil) | Atlantic Rain Forest |
#São Paulo, Paraná (Brazil) | |||
14 | Ceratina (Ceratinula) xanthocera (Moure, 1941) | *Mar de Espanha (Minas Gerais, Brazil) | Atlantic Rain Forest |
These are solitary bees, whose role in the pollination of plants is still not well studied, although their importance in pollination of the melon cactus Melocactus curvispinus Pfeiff. and the carnivorous corkscrew plant Genlisea violacea A.St.-Hil. has been reported (
Information on the geographic distribution of the described species of C. (Ceratinula) is still limited.
The present contribution describes and illustrates a distinctive new species of Ceratina (Ceratinula) Moure from midwestern Brazil (in the Cerrado Biome). Ceratina (C.) manni is redescribed, based on female and male individuals, and its geographical records updated. An identification key for the species of Ceratina (Ceratinula) so far recorded in Brazil is also provided, based on the list of
The specimens of the new species described here were collected in November 2018, during a rapid assessment for monitoring bee diversity, performed in the area surrounding a soybean field on the Nossa Senhora Aparecida farm, in Água Fria de Goiás, Goiás State, midwestern Brazil. The farm belongs to the Fiorese family (Oli Antonio Fiorese, Edileusa Fiorese, Henrique Gustavo Fiorese, Kaio Felipe Fiorese), who have adapted their production methods to meet the standards for environmental certification, and this is now considered a model farm.
Repository institutions of the specimens are: Entomological Collection of the Natural History Museum (MHNBA-MZUFBA) of the Biology Institute of the Federal University of Bahia, Ondina Campus, Salvador, Bahia, Brazil; Entomological Collection of the Latin American Institute of Life and Nature Sciences of the Federal University of Latin American Integration (CE-UNILA), Foz do Iguaçu, Paraná, Brazil; Reference Collection of the Laboratory of Bionomy, Biogeography and Insect Systematics (BIOSIS), a unit associated with the MHNBA, in Salvador, Bahia, Brazil. The syntype of C. (Ceratinula) manni Cockerell, 1912 is deposited in the Entomological Collection of the American Museum of Natural History (AMNH), New York, United States of America.
The description of C. (Ceratinula) fioreseana sp. nov. is based on the female holotype and male paratypes. The redescription of the female of C. (Ceratinula) manni Cockerell, 1912 is based on all specimens studied, checking the diagnostic characters presented in the original description (
Due to the COVID-19 pandemic, the identification key presented for Brazilian species of Ceratina (Ceratinula) was constructed based mainly on the original descriptions (as a first key attempt to aid identification, since the subgenus has not yet been revised), but was tested using a few specimens of seven different species that were accessible to us.
Specimens were studied and photographed using a Leica M165C stereomicroscope coupled to a Leica DFC295 digital camera, containing the program Leica Application Suite V4.1 Interactive Measurements, Montage. Measurements are given in millimeters and taken at the greatest width or length of structures. When we had access to more than one specimen, all were measured, and we report the average. The ocellocular distance is measured from the lateral ocellus, and the length of the anterior wing is measured from the costal sclerite. Abbreviations are:
DO diameter of the middle ocellus;
DS diameter of the scape;
DP diameter of a puncture;
F1, F2, F3 antennal flagellomeres 1, 2 and 3;
T metasomal tergum;
S metasomal sternum.
The classification follows
Ceratina lucidula Smith, 1854 by original designation.
Minute bees (3–6 mm long); usually metallic (rarely with the metasoma red); body elongated, with a slightly petiolate metasoma, the first segment as elongate-triangular; integument with extensive impunctate smooth areas, especially on head (paraocular area above antenna and on gena, sometimes on the whole head) and on mesonotum; second submarginal cell narrowed, sometimes converging to a point anteriorly, becoming almost triangular (
Among the species of Ceratina (Ceratinula) occurring in South America, C. (C.) fioreseana Oliveira, sp. nov. is similar to C. (C.) manni, especially in the pattern of facial maculation (Fig.
The male genitalia also differ widely between the species, especially in the structure and shape of the valves, gonostyle, S5, S6 and S7 (Figs
Both sexes: integument color tending more to greenish with golden metallic sheen. Female: five yellow maculations on face and one stripe on gena; median paraocular yellow maculation not filling the entire space between the eye and antennal socket, and not reaching the height of upper part of the epistomal suture (Figs
Female holotype of Ceratina (Ceratinula) fioreseana sp. nov., deposited at the Entomological Collection of the Natural History Museum of the Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil A head in frontal view B body in lateral view C head in lateral view D mesosoma in lateral view E labrum in frontal view.
Brazil, Goiás, Água Fria de Goiás, Fazenda Nossa Senhora Aparecida, Bayer Forward Farming, 14°49'25.946"S, 47°43'30.742"W (–14.823874, –47.725206), Cerrado vegetation (Savanna), alt. 1073 m a.s.l. (Fig.
♀: Structure (mm): total body length 4.9; forewing length 3.5; head width 1.37; eye length 0.83, width 0.47; gena width in profile 0.22; ocellocular distance 0.28; diameter of median ocellus 0.10; upper interorbital distance 0.86, median interorbital distance 0.75, lower interorbital distance 0.68; clypeus length 0.48, width 0.6; labrum length 0.17, width 0.36; scape length 0.3, width 0.08; F1 length 0.05; F2 length 0.05; F3 length 0.05; metatibia length 0.9, width 0.2; T2 width 1.25; T4 width 1.38. Antennal sockets located in deep depression (Fig.
♂: Structure (mm): total body length 4.5; forewing length 3.5; head width 1.37; eye length 0.87, width 0.52; gena width in profile 0.17; ocellocular distance 0.24; diameter of median ocellus 0.12; upper interorbital distance 0.85, median interorbital distance 0.64, lower interorbital distance 0.64; clypeus length 0.42, width 0.49; labrum length 0.18, width 0.33; scape length 0.24, width 0.08; F1 length 0.05; F2 length 0.065; F3 length 0.07; metatibia length 0.83, width 0.18; T2 width 1.15; T4 width 1.38. Antennal sockets located in deep depression (Fig.
Male paratypes of Ceratina (Ceratinula) fioreseana sp. nov., deposited at the Entomological Collection of the Natural History Museum of the Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil: Paratype A head in frontal view B body in lateral view C head in lateral view D mesosoma in lateral view. Paratype E labrum in frontal view F head in frontal view.
Male genitalia and associated metasomal sternum of paratype of Ceratina (Ceratinula) fioreseana sp. nov., deposited at the Entomological Collection of the Natural History Museum of the Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil A genital capsule in dorsal view B genital capsule in ventral view C genital capsule in lateral view D seventh sternum in dorsal view (E7) E sixth sternum in dorsal view (E6) F fifth sternum in dorsal view (E5) G seventh tergum in dorso-lateral view.
Some male specimens, such as the male paratype (Fig.
(1♀, 3♂) – Holotype ♀ (MHNBA) // Brazil, Goiás, Água Fria de Goiás, Fazenda Nossa Senhora Aparecida, Bayer Forward Farming, 14°49'25.946"S, 47°43'30.742"W (–14.823874, –47.725206), 29.XI.2018, 15:00–15:34 h, Cerrado Savanna, alt. 1073 m a.s.l. Paratype used for description ♂ (MHNBA) // Brazil, Goiás, Água Fria de Goiás, Fazenda Nossa Senhora Aparecida, Bayer Forward Farming, 14°49'25.946"S, 47°43'30.742"W (–14.823874, –47.725206), 30.XI.2018, 10:40–11:40 h, Cerrado Savanna, alt. 1073 m a.s.l. Paratypes: 2 ♂ (MHNBA) // Brazil, Goiás, Água Fria de Goiás, Fazenda Nossa Senhora Aparecida, Bayer Forward Farming, 14°49'25.946"S, 47°43'30.742"W (–14.823874, –47.725206), 29.XI.2018, 15:00–15:34 h, Cerrado Savanna, alt. 1073 m a.s.l. All specimens were collected with an entomological net on flowers of Mentha piperita L. (mint, family Lamiaceae), in planted vegetable garden.
The specific epithet is a patronym honoring Oli Antonio Fiorese, Edileusa Fiorese, Henrique Gustavo Fiorese and Kaio Felipe Fiorese, owners of the Nossa Senhora Aparecida farm (located in Água Fria de Goiás, Goiás State, midwestern Brazil), where the type specimens of the new species were collected. We honor their recognition of bees as key pollinators important to sustainable production, adapting their production to meet the standards for environmental certification, and also adopting various pollinator-friendly measures through the Bayer Forward Farming Project. Because of these procedures, their property has been certified as a model farm by environmental agency Round Table on Responsible Soy Association (RTRS), being the only farm in Brazil within the Bayer Forward Farming Project, and the twenty-fifth in the world.
Ceratina manni
Cockerell, 1912: 47 [original description];
Ceratinula manni:
Ceratina (Ceratinula) manni:
Brazil, Paraíba, Guarabira (previously known as Independencia).
Both sexes: integument color tending more to greenish with golden metallic sheen. Female: five yellow maculations on face and one stripe on gena; median paraocular yellow maculation almost filling the entire space between the eye and antennal socket, and almost reaching the height of upper part of the epistomal suture (Fig.
Female syntype of Ceratina (Ceratinula) manni Cockerell, 1912 deposited at the Entomological Collection of the American Museum of Natural History (AMNH, New York, United States of America) A head in frontal view (Photograph: I_HHG3587) B body in dorsal view (Photograph: I_HHG3585) C body in lateral view (Photograph: I_HHG3586) D labels (Photograph: I_HHG3588). Ownership rights to these images and their copyright belong to AMNH and Hadel Go. Photographs by Copyright Hadel Go 2011-2016 downloaded from the Discover Life Website.
Female of Ceratina (Ceratinula) manni Cockerell, 1912 A head in frontal view B body in lateral view C head and mesosoma in lateral view D labrum in frontal view E median paraocular area and clypeus in detail F head in frontal view A, E specimen from Amélia Rodrigues, Bahia, deposited at the Reference Collection of the Laboratório de Bionomia, Biogeografia e Sistemática de Insetos (BIOSIS), Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil; Specimens B,C, F from Caracol, Piauí, and D Milagres, Bahia, deposited at the Entomological Collection of the Federal University of Integração Latino-Americana (CE-UNILA), in Foz do Iguaçu, Paraná, Brazil.
Male of Ceratina (Ceratinula) manni Cockerell, 1912, specimen from Crato, Ceará, deposited at the Entomological Collection of the Federal University of Integração Latino-Americana (CE-UNILA), in Foz do Iguaçu, Paraná, Brazil A head in frontal view B body in lateral view C labrum in frontal view D head and mesosoma in lateral view.
♀: Structure (mm): total body length 4.7; forewing length 3.2; head width 1.3; eye length 0.89, width 0.43; gena width in profile 0.26; ocellocular distance 0.33; diameter of median ocellus 0.12; upper interorbital distance 0.91, median interorbital distance 0.78, lower interorbital distance 0.67; clypeus length 0.43, width 0.61; labrum length 0.22, width 0.44; scape length 0.31, width 0.09; F1 length 0.09; F2 length 0.05; F3 length 0.05; metatibia length 0.54, width 0.16; T2 width 1.25; T4 width 1.45. Antennal sockets located in shallow depression (Fig.
Male genitalia and associated metasomal sternum of Ceratina (Ceratinula) manni Cockerell, 1912, specimen from Serra Negra do Norte, Rio Grande do Norte, Brazil, deposited at the Entomological Collection of the Federal University of Integração Latino-Americana (CE-UNILA), in Foz do Iguaçu, Paraná, Brazil A genital capsule in dorsal view (red arrow shows the bidentate apex) B genital capsule in ventral view C genital capsule in lateral view D seventh sternum in dorsal view (E7) E sixth sternum in dorsal view (E6) F fifth sternum in dorsal view (E5).
Head of Ceratina (Ceratinula) Moure, 1941 species in frontal view A (female holotype) B (male paratype) Specimens of Ceratina (Ceratinula) fioreseana sp. nov., deposited at the Entomological Collection of the Natural History Museum of the Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil C, D Ceratina (Ceratinula) manni Cockerell, 1912: C female specimen from Amélia Rodrigues, Bahia, deposited at the Reference Collection of the Laboratório de Bionomia, Biogeografia e Sistemática de Insetos (BIOSIS), Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil D male specimen from Crato, Ceará, deposited at the Entomological Collection of the Federal University of Integração Latino-Americana (CE-UNILA), in Foz do Iguaçu, Paraná, Brazil. Drawings by Luisa de Lima Ruschioni.
Head of Ceratina (Ceratinula) Moure, 1941 species in frontal view, showing the supraclypeal area and median paraocular area A, B female holotype of Ceratina (Ceratinula) fioreseana sp. nov., specimen deposited at the Entomological Collection of the Natural History Museum of the Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil C, D female specimen of Ceratina (Ceratinula) manni Cockerell, 1912 from Amélia Rodrigues, Bahia, deposited at the Reference Collection of the Laboratório de Bionomia, Biogeografia e Sistemática de Insetos (BIOSIS), Federal University of Bahia (MHNBA-MZUFBA), in Salvador, Bahia, Brazil. Details: pls = puncture line delimiting the supraclypeal plain raised area above; ues = upper part of the epistomal suture.
♂: Structure (mm): total body length 3.9; forewing length 3.06; head width 1.3; eye length 0.76, width 0.46; gena width in profile 0.23; ocellocular distance 0.24; diameter of median ocellus 0.12; upper interorbital distance 0.84, medium interorbital distance 0.61, lower interorbital distance 0.57; clypeus length 0.49, width 0.57; labrum length 0.21, width 0.33; scape length 0.23, width 0.08; F1 length 0.05; F2 length 0.04; F3 length 0.06; metatibia length 0.5, width 0.14; metasomal width 1.19 (measured on T4). Antennal sockets located in shallow depression (Fig.
In females, the large elliptical yellow longitudinal maculation on the disc of the clypeus is sometimes enlarged apically, as observed in some specimens from Piauí State; these also have a small translucent brown oval maculation in the middle of the disc (Fig.
(new geographical records indicated by*). Ceratina manni is endemic to northeastern Brazil and occurs mainly within the limits of the Caatinga region (Fig.
Geographic distribution map of Ceratina (Ceratinula) fioreseana sp. nov. and Ceratina (Ceratinula) manni Cockerell, 1912 in and Brazil: (1) Goiás, Água Fria de Goiás, Fazenda Nossa Senhora Aparecida, Bayer Forward Farming (fioreseana Type locality); (2) Bahia, Maracás; (3) Bahia, Milagres; (4) Bahia, Amélia Rodrigues; (5) Piauí, Caracol; (6) Ceará, Crato; (7) Rio Grande do Norte, Serra Negra do Norte, Estação Ecológica do Seridó; (8) Paraíba, Cacimba de Dentro, Fazenda Cachoeira da Capivara; (9) Paraíba, Mamanguape; (10) Paraiba, João Pessoa, Periurban Area – Sítio ponta de Gramame; (11) Paraíba, Conde, Granja Pitumirim; (12) Paraíba, Alhandra, Assentamento Tapuiu – Sítio Olho D’água; (13) Paraíba, Guarabira (manni Type locality).
Brazil: *Piauí State: Caracol. *Ceará state: Crato. Rio Grande do Norte state: Serra Negra do Norte (
(11♀, 2♂). 1 ♂ // (CE-UNILA HYAP 6072) // Brasil, Rio Grande do Norte, Serra Negra do Norte, Estação Ecológica do Seridó, 17.xii.1994, Zanella FCV and Moura ON leg.; 1 ♀ // (CE-UNILA HYAP 6073) // Brasil, Rio Grande do Norte, Serra Negra do Norte, Estação Ecológica do Seridó, 30.xii.1994, Zanella FCV and Moura ON leg.; 1 ♀ // (CE-UNILA HYAP 3350) // Brasil, Paraíba, Cacimba de Dentro, Fazenda Cachoeira da Capivara, 25.x.2003, F. Zanella leg.; 1 ♀ // (CE-UNILA HYAP 1109) // Brasil, Piauí, Caracol, 518 m, 13.xii.2010, F. Zanella and A. Carvalho leg.; 1 ♀ // (CE-UNILA HYAP 1110) // Brasil, Piauí, Caracol, 518 m, 13.xii.2010, F. Zanella and A. Carvalho leg.; 1 ♂ // (CE-UNILA HYAP 1120) // Brasil, Ceará, Crato, Estrada para Exú, Encosta, 07.ii.2011, F. Zanella leg.; 1 ♀ // (CE-UNILA HYAP 1117) // Brasil, Bahia, Maracás, 13°26'33.8"S, 40°20'42.6"W, 945 m, 13.iii.2012, Zanella FCV. leg.; 1 ♀ // (CE-UNILA HYAP 1118) // Brasil, Bahia, Maracás, 13°26'33.8"S, 40°20'42.6"W, 945 m, 13.iii.2012, Zanella FCV. leg.; 1 ♀ // (CE-UNILA HYAP 1112) // Brasil, Bahia, Milagres, 12°54'19.2"S, 39°50'46.5"W, 758 m, 16.iii.2012, Zanella FCV. leg.; 1 ♀ // (CE-UNILA HYAP 1115) // Brasil, Bahia, Milagres, 12°54'19.2"S, 39°50'46.5"W, 758 m, 16.iii.2012, Zanella FCV. leg.; 1 ♀ // (BIOSIS-UFBA, Favízia 06380) // Brasil, Bahia, Amélia Rodrigues, 12°22'31.70"S, 38°46'05.82"W, 21.i.2017, Hora: 13:10 P. 11, n° 404, on flowers of Stemodia foliosa Benth. Silva, Anjos and Melo leg.; 1 ♀ // (BIOSIS-UFBA, Favízia 06381) // Brasil, Bahia, Amélia Rodrigues, 12°22'31.70"S, 38°46'5.82"W, 21.i.2017, Hora: 13:10 P. 11, n° 405, on flowers of Stemodia foliosa Benth. Silva, Anjos and Melo leg.; 1 ♀ // (BIOSIS-UFBA, Favízia 06382) // Brasil, Bahia, Amélia Rodrigues, 12°22'31.70"S, 38°46'5.82"W, 11.iii.2017, Hora: 10:00 P. 11, on flowers of Stemodia foliosa Benth. Silva, Anjos and Melo leg. (Fig.
Species so far recorded for Brazil, according to the “Catalogue of Bees (Hymenoptera, Apoidea) in the Neotropical Region” (
The identification key presented above is probably not exhaustive, as there are likely to be many species that have not yet been described or recorded in Brazil, but are known from neighboring countries. Therefore, we recommend that the characters be verified in the original descriptions and with a reference collection or type material, in order to more confidently assign species names.
1 | Face with yellow-pigmented maculations | 2 |
– | Face without yellow-pigmented maculations | 11 |
2 | Yellow maculations present on clypeus and on paraocular region; with yellow genal stripe; yellowish or whitish pronotal lobe | 3 |
– | Yellow maculations restricted to either clypeus or to paraocular region, if on both, then only in lower paraocular area; lacking yellow genal stripe; pronotal lobe variable | 5 |
3 | Color tending more to greenish with golden metallic sheen; scape color variable. Facial pattern consisting of five yellow maculations: one large elliptical longitudinal maculation in median paraocular area, extending upward and downward from level of antennal socket, almost reaching or not the height of upper part of the epistomal suture; one smaller rounded maculation on lower paraocular area near tentorial pit; one median, longitudinal subtriangular maculation occupying large area of clypeus; dorsal face of supraclypeal raised surface subtriangular or subpentagonal; frons level variable | 4 |
– | Color tending more to brownish, with strong blue-violet sheen on head, mesosutum an metasoma; scape reddish brown. Facial pattern consisting of three yellow maculations: one large elliptical longitudinal maculation in median paraocular area, extending upward and downward from level of antennal socket, not reaching the height of upper part of the epistomal suture (maculation width ca. 1.5DS, length 4DS, ending at a height ca. 1.2DS from epistomal suture – scape maximum width); and one median, longitudinal subtriangular maculation occupying large area of clypeus; supraclypeal plain raised surface subrectangular; frons below the rest of the head |
Ceratina (Ceratinula) muelleri Friese, 1910 (Brazilian States: Amazonas, Pará, Maranhão, Ceará, Minas Gerais, Rio de Janeiro, São Paulo, Paraná; Argentina) |
4 | Yellow maculation on median paraocular area almost reaching the height of upper part of the epistomal suture (maculation width ca. 1.4DS, length 2.4DS, ending at a height ca. 0.5DS from epistomal suture – scape maximum width – Figs |
Ceratina (Ceratinula) manni Cockerell, 1912 (Brazilian States: Piauí, Ceará, Rio Grande do Norte, Paraíba, Bahia – Caatinga biome and ecotone zone) |
– | Yellow maculation on median paraocular area not reaching the height of upper part of the epistomal suture (maculation width ca. 1DS, length 2.3DS, ending at a height ca. 1.25DS from epistomal suture – scape maximum width – Figs |
Ceratina (Ceratinula) fioreseana Oliveira, sp. nov. (Brazil: Goiás State – Cerrado biome) |
5 | Facial maculations restricted to median paraocular region, close to antennae | 6 |
– | Facial maculations restricted to lower paraocular region and clypeus | 8 |
6 | Larger in size (5 mm to larger); small facial maculations, width smaller than scape diameter | 7 |
– | Smaller in size (4–4,5 mm); large facial maculations, width larger than scape diameter | Ceratina (Ceratinula) biguttulata (Moure, 1941) (Brazil: Paraná State) |
7 | Yellow maculations very small, almost imperceptible; anterior part of scape and meso and metatarsi, yellowish brown; very sparse punctation (3–4DP); metatibia strongly thickened | Ceratina (Ceratinula) melanochroa (Moure, 1941) (Brazil: Paraná State) |
– | Yellow maculations small, but distinct, elongated; antennae dark brown with underside of flagellum rust-brown; meso and metatarsi dark brown; punctation less sparse (2–3DP); metatibia normal | Ceratina (Ceratinula) sclerops Schrottky, 1907 (Brazilian states: São Paulo, Paraná; Paraguay) |
8 | Variable in size; antennal scape and meso and metatibiae blackened; other characters, variable | 9 |
– | Small in size (around 4 mm); antennal scape, tibiae and tarsus, tegulae and wing venation honey-yellow; small elongated maculation on clypeus; pronotal lobes with yellow maculation | Ceratina (Ceratinula) xanthocera (Moure, 1941) (Brazil: Minas Gerais State) |
9 | Normal face, with shallow sutures; head nearly glabrous, with extensive impunctate areas; protibia and tarsus and metatarsus and other characters, variable | 10 |
– | Swollen face, with deep sutures; head with punctation distinct, most abundant; elongated maculation in middle of clypeus, narrowing upward, not dilated apically; protibia and tarsus and meta tarsus yellowish; microreticulate area in basal area of the propodeum narrowest, microreticulate sculpture more uniform, regular; puncture line delimiting the supraclypeal plain raised area above, with lateral branches divergent basally | Ceratina (Ceratinula) turgida (Moure, 1941) (Brazil: Rio de Janeiro State) |
10 | Color metallic dark green; large elongated yellow maculation on disc of clypeus; clypeus integument polished, smooth |
Ceratina (Ceratinula) minima Friese, 1908 (Brazil: northern region; Trinidad and Tobago) |
– | Color metallic olive-green; yellow band on apical border of clypeus; clypeus integument slight microreticulate | Ceratina (Ceratinula) piracicabana Schrottky, 1910 (Brazil: São Paulo State) |
11 | Clypeus with wide light-brown stripe on apical margin (at least 1/6 of clypeus width) | 12 |
– | Clypeus with uniform color, without yellow band, sometimes with only the extreme edge lighter | 13 |
12 | Body metallic green; face uniformly green, without maculation; legs brownish | Ceratina (Ceratinula) augochloroides Ducke, 1910 (Brazil: Ceará State) |
– | Body metallic olive-green; apical margin of clypeus, mandibles, labrum, basal antenna joint, tegula and legs honey-yellow | Ceratina (Ceratinula) lucidula Smith, 1854 (Brazil States: Pará, Ceará, Minas Gerais, São Paulo; Paraguay) |
13 | Head with greenish metallic sheen above clypeus; lower area of gena, next to mandibles, pale brownish white; tarsi yellowish brown | Ceratina (Ceratinula) fulvitarsis Friese, 1925 (Brazil: São Paulo State) |
– | Head with bluish metallic sheen above clypeus; lower area of gena near mandibles, of same color as rest of gena; meso- and metarsi dark brown | Ceratina (Ceratinula) immaculata Friese, 1910 (Brazil: Pará State) |
As stressed by
With the exception of C. (C.) combinata and C. (C.) minima, which were described from and known only from male specimens, the Brazilian fauna of Ceratina (Ceratinula) has been described based on females, with shape and presence or absence of these yellow maculations explained in the original descriptions.
The identification key presented above is probably not exhaustive, as there are likely to be many species that have not yet been described or recorded in Brazil, but are known from neighboring countries. Therefore, we recommend that the characters be verified in the original descriptions and with a reference collection or type material, in order to more confidently assign species names.
Several unidentified specimens of Ceratina (Ceratinula) have been reported in local inventories of bee faunas (some of them cited in the Introduction). Although we have not yet had access to the specimens collected in these surveys, this suggests that the species richness is higher than presently recorded, and that much more collection effort and taxonomic work is necessary to gain a comprehensive knowledge of the diversity and distribution patterns within the subgenus.
Apifaunistic surveys are particularly important sources of floral records for bees on their host plants (
The authors are indebted to Bayer Crop Science, through its Bee Care Brazil Initiative, for the financial support for this publication, and to the Bayer Forward Farming Project for the financial support for the collecting expedition to the Nossa Senhora Aparecida Farm (Água Fria de Goiás, state of Goiás) where the specimens of Ceratina (Ceratinula) fioreseana Oliveira, sp. nov. were collected. The authors graciously thank Oli Antonio Fiorese, Edileusa Fiorese, Henrique Gustavo Fiorese, and Kaio Felipe Fiorese (the esteemed Fiorese family) for kindly receiving us on their property and supporting the research that made possible the description of the new species presented here. The authors would also like to thank the Universidade Federal da Bahia (UFBA) and the Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB) for financial support provided to the BIOSIS Laboratory, where we performed the taxonomic part of this study; to Sra. Sônia Guimarães, owner of the Guimarães Farm (Amélia Rodrigues, Bahia) where Ceratina (Ceratinula) manni was collected; and also the Centro Universitário Jorge Amado (UNIJORGE) for providing the laboratory infrastructure where part of the material was prepared by the second author. Thanks are due also Christian Maus (Entomology & Pollination Science Lead, Bayer AG, Global Regulatory Scientific Affairs, Crop Science, Monheim, Germany) for his support in translating some Ceratina (Ceratinula) original descriptions from German to English, and to Natali Pereira do Santos (Inteligência Tropical, CEAT, Bayer), for her assistance in constructing the geographic distribution map. The specimens of Ceratina (Ceratinula) manni Cockerell, 1912 recorded at Milagres and Maracás in Bahia state, Crato in Ceará, and Caracol in Piauí studied in this paper and deposited at CE-UNILA were a contribution of the Semiarid Biodiversity Research Program, approved by the Brazilian National Council for Scientific and Technological Development (CNPq) (PPBIO Caatinga, Invertebrados project; see