The bees of the genus Centris Fabricius, 1804 described by Theodore Dru Alison Cockerell (Hymenoptera: Apidae)

In this paper the primary types of Centris bees described by the British entomologist Theodore Dru Alison Cockerell deposited in the Natural History Museum (London) and the Oxford University Museum of Natural History (Oxford) in the United Kingdom, as well as in the United States National Museum (Washington), American Museum of Natural History (New York), the Academy of Natural Sciences of Drexel University (Philadelphia), and in the California Academy of Sciences (San Francisco) in the United States were studied. To stabilize the application of the name C. lepeletieri (= C. haemorrhoidalis (Fabricius)), a lectotype is designated. The study of the primary types allow proposing the revalidation of C. cisnerosi nom. rev. from the synonymy of C. agilis Smith, C. nitida geminata nom. rev. from C. facialis Mocsáry, C. rufulina nom. rev. from C. varia (Erichson), C. semilabrosa nom. rev. from C. terminata Smith and C. triangulifera nom. rev. from C. labrosa Friese. Centris bakeri syn. nov., C. bimaculata carrikeri syn. nov., C. fusciventris matoensis syn. nov., C. heterodonta syn. nov. and C. elegans grenadensis syn. nov. are proposed as a new junior synonyms of C. varia, C. claripennis Friese nom. rev., C. caurensis, C. dentata Smith and C. elegans Smith, respectively. Centris ruae is withdrawn from the synonymy of C. transversa Pérez and proposed as a new junior synonym of C. nitida Smith. In addition, a lectotype for C. buchholzi Herbst (= C. wilmattae) is designated. Information on the repository of the lectotype of C. lepeletieri and images of most primary types studied here are also provided.


Introduction
Theodore Dru Alison Cockerell (1866Cockerell ( -1948 was a British entomologist, systematic biologist and one of the most important melittologist of all time (Engel & Davis 2012). His contributions to science began during the early years of his youth. Throughout his more than four decades of activity, Cockerell published more than 3900 articles (Zuparko 2007), becoming one of the most proliferous scientists of

Material and methods
All labels are here considered whitish and rectangular, and the data contained on them is black, handwritten or printed, unless otherwise indicated. The specific features of the labels, like coloration or type of writing, are given in squared brackets ([ ]). All yellow labels contain a barcode, which is part of the database of the United States National Museum collection (NMNH). Some information on the handwritten labels is difficult to decipher, in these cases the symbol ' [?]' was used. The backward slash (\) indicates different labels on the pin of the same specimen and two backward slashes (\\) indicate information on the back of the label. The copyright of the photographs belongs to the NMNH. The information related is provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10 th and Constitution Ave. N.W., Washington, DC 20560-0193 (https://www.nmnh.si.edu/).

Recognition of the type specimens
According to Cockerell (1949), he divided his collection of Neotropical bees in two parts; one was sent to the NMNH and the other to the AMNH. These two institutions house more than ⅔ of the primary types of Centris bees described by him. The rest of the type specimens are deposited, in decreasing order, in the collections of CAS, NHMUK, ANSP, OUMNH and ZMB. All primary types currently housed at CAS, except for C. hoffmanseggiae davidsoni, were collected by the American entomologist and curator of the Department of Entomology of the California Academy of Sciences Edward Payson Van Duzee   (MacFarland 1940). Probably, Cockerell studied these specimens at CAS during a visit to California. It is not clear whether the specimens currently housed at NHMUK already belonged to that collection or whether they were part of Cockerell's own collection and were sent there subsequently. Part of the type series of C. caesalpiniae and C. caesalpiniae rhodopus is currently housed at ANSP. Those specimens bear a printed label: "1980 Loan to USNM from Acad. of Nat. Sciences at Philadelphia". No additional information about this was found.
All Cockerell's primary types can be easily recognized by his handwriting style (see Horn & Kahle 1935: pl. 3, fig. 18). He labeled almost all the specimens he used in the original descriptions with the name of the species and the words "type", "cotype" or "co-type". Along with these labels, the data on collector, provenance and depository agreed with those mentioned by him in the original descriptions.
Considering this, I am confident that the specimens examined are true Cockerell's primary types and fully valid for the proposal of revalidations and new synonymies, as well as for the designation of lectotypes when necessary.

Type data
This species was described based on a single female specimen collected by the English traveler, naturalist, ornithologist and entomologist William Frederick Henry Rosenberg (1868Rosenberg ( -1957 who made collecting trips to northern South America (Günter 1912

Type data
This species was described based on a single female specimen collected by Adan Rivera on February 9, 1948, at Zamorano, Honduras. Rivera and G. Cisneros were probably collectors who helped Cockerell during his stay in Honduras (Jesus Orozco, pers. com.). The holotype (Fig. 2)

Comments
The study of the type of Centris bakeri revealed that it corresponds to a male of C. varia with two yellow spots on the lateral surfaces of the second tergum. The coloration of the metasoma in both sexes of this species is very variable. Terga and sterna of specimens can be blackish, reddish brown or orange, with or without dark brown or yellowish spots or bands. Centris varia is one of the most widely distributed species of the subgenus C. (Centris). According to Moure et al. (2007), it occurs from Mexico to Argentina. Cockerell, 1937 Centris (Hemisia) nitens callospila Cockerell, 1937a: 1.

Type data
This species was described based on a single male specimen collected by John W. Ogilvie at Waranama, northeastern Guyana, on November 14, 1936. Ogilvie was a Scottish immigrant who went to South America to establish his fortune in gold prospection and exploitation of rubber tapping (Pourshariati 2017). Ogilvie, along with other people, made an expedition through the Amazon that begun in Manaus, Brazil, and ended in Georgetown, Guyana. The purpose of that expedition was mainly ethnological (Pourshariati 2017); however, biological collections were also made. At least some of the bees obtained during that trip were studied by Cockerell (1936Cockerell ( , 1937aCockerell ( , 1937b. Ogilvie also visited some Caribbean islands where he collected biological samples that were subsequently sent to other scientists for study (see Cockerell 1938Cockerell , 1939. During the collection of this new species, other Centris bees were also caught (Cockerell 1937b). Ogilvie also captured specimens of C. denudans Lepeletier, 1841, C. flavifrons clitelligera (Erichson, 1848) (= C. flavifrons (Fabricius, 1775)) and C. nitens Lepeletier, 1841

Comments
Although Cockerell labeled the type specimen as a new species, it was published as a variety of C. nitens. This species is only known from male specimens and could actually correspond to a metander male of C. nitens. Cockerell, 1922 Fig. 3 Centris disclusa Cockerell, 1922: 5-6.

Type data
This species was described based on two male specimens collected by Francisco Campos Ribadeneira in Posorja ("Posoya"), Ecuador. Campos (1878-1943 was an intellectual from Guayaquil and the first Ecuadorian entomologist (Barragán et al. 2009). He collected numerous insects and created the first entomological collection in that country (Moret 2005

Type data
This subspecies was described based on a single male specimen collected by the American naturalist Herbert Huntington Smith (1851Smith ( -1919

Type locality
Lesser Antilles: Grenada.

Comment
Centris elegans grenadensis only differs from C. elegans by the lighter hairs on hind legs and mesoscutellum ( Fig 3B). At NMNH, there were found three males of C. elegans, with dark hairs on legs and mesoscutellum, which according to their labels were collected along with the holotype of C. elegans grenadensis. Cockerell, 1912 Fig. 5 Centris inermis gualanensis Cockerell, 1912b: 568. Junior synonym of C. (Centris) varia (Moure 1969;Roig-Alsina 2000).

Type data
This subspecies was described based on male and female specimens collected by W.P. Cockerell on February 22-23, 1912at Gualán, Guatemala. Cockerell (1912b mentioned that the type specimen was a female, but in the type series housed at NMNH, two specimens of that sex were found. Both specimens agree with the information provided in the original description but one of them was labeled by Cockerell himself as "type", while the other bears a label with the word "cotype". The holotype (

Type data
This subspecies was described based on an apparently large series of males collected by Wilmatte Porter Cockerell and Adan Rivera at Zamorano and Tegucigalpa on flowers of Duranta plumieri Jacq. (Verbenaceae). Cockerell (1949) mentioned the type specimen was numbered "58887" but in the NMNH, two males with this number were found. One of them bears a label with the word "type" and the other is labeled "cotype". The holotype ( Fig. 6)

Centris lepeletieri
Type data Cockerell (1912c) recognized the existence of two species similar to each other, which would have been identified as C. haemorrhoidalis. According to him, the name C. haemorrhoidalis would be applied to Jamaican specimens where males have the first four terga and the anterior half of the fifth metallic blue, while females have the first four terga metallic blue and the rest reddish. Cockerell mentioned that the females from Santo Domingo, Dominican Republic identified by Lepeletier (1841) as C. haemorrhoidalis would correspond to a different species, since they have only the first three terga blue, proposing for them the name C. lepeletieri. Cockerell (1912c) was correct in indicating the existence of two similar species. However, C. lepeletieri was described based on specimens that correspond to the true C. haemorrhoidalis. The Jamaican species was described several decades later under the name C. dirrhoda Moure, 1960.
One of the females studied by Lepeletier (1841) was found at OUMNH and is here designated the lectotype of Centris lepeletieri. The specimen has the following data label: *Du [handwritten]\ Centris haemorrhoidalis Fabr Sto Domingo ♀.

Type data
This species was described based on a single male specimen collected by Cockerell's wife at Zamorano on February 8. The holotype (Fig. 7)

Comments
Cockerell considered describing C. robusta under the name "C. wilmattae", as an homage to his second wife. However, this name had already been used by Cockerell (1926a)

Comment
This species was cited as a junior synonym of C. varia (Moure et al. 2007), but it is actually a different species. Centris rufulina is only known from the type specimen and differs from C. varia, among other characters, by the shape of the third mandibular tooth (truncate in C. rufulina, acute in C. varia). Cockerell, 1938 Fig. 8 Centris versicolor vincentana Cockerell, 1938: 282-283.

Type data
This species was described based on an undetermined number of specimens of both sexes collected by J. Ogilvie at Barbados. Cockerell (1939) mentioned that the type of the species was a male. A male specimen belonging to the type series is currently housed at NHMUK and was labeled as type by Cockerell himself. Another male of the type series was found at NMNH and was labeled by Cockerell as cotype.
The female specimen mentioned in the original description was not found. the holotype or mentioned any data that could help to recognize it. However, Moure (1960) mentioned the type of this species was numbered 54853. In the NMNH, both specimens cited by Cockerell were found, with the number mentioned by Moure (1960), one bearing a label "type" and the other "cotype". The holotype (Fig. 8)

Type locality
Lesser Antilles: Saint Vincent and the Grenadines: Saint Vincent Island.

Type locality
Lesser Antilles: Barbados.

Type data
Cockerell described this subspecies apparently based on a single female specimen collected by the American entomologist and naturalist Charles Thomas Brues , specialist on Hymenoptera, Diptera and on insect diet (Salt 1955), during a trip to South America. Moure et al. (2007) indicated that it was described from syntypes. Nevertheless, only a single female with original Cockerell's label of "type" was found at NMNH. Due to the lack of evidence on the existence of more type specimens and the impossibility to deduce this fact from the original description, the female housed at NMNH is here interpreted as holotype, taking into account the information of the Cockerell's own label. The specimen ( Fig. 9) has the following data label:

Comment
This species was mistakenly proposed as a junior synonym of C. facialis Mocsáry, 1899(Rasmussen & Vivallo 2014), a species superficially similar. The study of large series of both species allowed to revert the proposed synonymy and to consider both as valid species. Females of C. geminata and C. facialis differ in the coloration of the prepygidial fimbriae, yellowish in the former and dark brown or blackish in the latter species. The males of both species differ in the size and pubescence of the hind femur (slightly larger than the hind tibia and covered by whitish pubescence on the dorsal surface in C. geminata; evidently larger than the hind tibia and covered completely by dark brown to black pubescence in C. facialis). Cockerell, 1936 Centris lanipes ogilviei Cockerell, 1936: 632.

Type data
According to the original description, Cockerell proposed this subspecies based on two females collected by J.

Type data
Cockerell described this subspecies based on an undetermined number of male and female specimens collected on an adobe wall at San Francisco ranch near Zamorano, Honduras, on November 10. Although he mentioned that the type was from that locality, he failed to mention the sex of the specimen chosen. At NMNH, a male specimen was found with original labels of Cockerell, one with the name of the new subspecies and the word "type", and the other indicating that it was collected at an adobe wall in San Francisco, on November 3 not 10 as Cockerell mentioned explicitly. In the same collection, a female collected by Cockerell was found that bears an original label of Cockerell that agrees exactly with the information provided in the original description, but mentions in Cockerell's handwriting "cotype". Both specimens have subtle inconsistencies on the information labels and no additional specimens of the type series were found at NMNH or in any other collection that contains Cockerell's types of Centris bees (AMNH, ANSP, CAS, NHMUK, OUMNH and ZMB). Despite the differences cited above, I am interpreting the male specimen as the holotype of the species, following the information on its label. The holotype (Fig. 10)   , an former student and employee of the Escuela Agrícola Panamericana (Pitty 2008). It is not clear whether this specimen belonged to the type series, and unfortunately, it was not found during this research.

Type data
This species was described based on a single female specimen collected by the American limnologist and Amazon traveler Harriet Bell Merrill (1863-1915) during a research trip to South America. According to Rathbun (1911), she sold to the NMNH 500 specimens of insects collected during her expedition to Brazil, including probably the exemplar studied by Cockerell. The holotype (Fig. 11)

Type data
This species was described based on male and female specimens collected by J. Ogilvie at Barbados on April 25-26, 1938. Cockerell described both sexes of this species, but he did not indicate how many specimens of each sex he used. Apparently, the type series was composed of at least one male and two females, which were found at NMNH. The male was chosen as holotype. and it was labeled by Cockerell as such. The male holotype (Fig. 12)

Type locality
Lesser Antilles: Barbados.
Comment Moure et al. (2007) mentioned the sex of the holotype erroneously.

Type data
This species was described based on male and female specimens collected in Honduras. The female holotype was collected on March 30 at Escuela Agrícola Panamericana by Rua Williams, the wife of the American explorer and botanist Louis Otho Williams, who worked on the flora of Central America (Burger 1991). According to the original description, the type series was composed of another female specimen collected by Adan Rivera on April 8 and two males collected by Wilmatte P. Cockerell and Adan Rivera on January 28. The holotype (Fig. 13)

Type locality
Honduras: Francisco Morazán Department: Zamorano.  Snelling (1984) mentioned, he studied the type specimen of C. ruae, considering it conspecific with C. transversa Pérez, 1905. It is not clear whether he studied the type of the latter species to propose the mentioned synonymy, but according to the information he cited in his article, apparently he did not, assuming the differences between both species as a result of variation. The study of both type specimens allows recognizing C. transversa and C. ruae as different species, and proposing the transfer of the latter from the synonymy of the former species to the synonymy of C. nitida. Cockerell, 1949 Fig. 14 Centris rufomaculata Cockerell, 1949: 476. Junior synonym of C. trigonoides (Snelling 1984).

Type data
This species was described based on a single male specimen collected on February 7 at Zamorano, Honduras. The holotype (Fig. 14)
Comments Moure et al. (2007) were not able to find the depository of the holotype of this species. Centris bicornuta occurs from Mexico to Argentina (Snelling 1984;Roig-Alsina 2000). According to the accession book at the AMNH, the number 33337 was assigned to about 100 specimens that were sent from Cockerell's collection to the AMNH. This number is also present on the labels of the primary types of C. ceratocephala, C. euphenax, C. libertatis, C. maroniana and C. wilmattae (see below). Cockerell, 1949 Fig. 15 Centris (Melanocentris) durantae Cockerell, 1949: 474. Junior synonym of C. (Heterocentris) analis (Fabricius, 1804) (Michener 1954;Moure 1960).

Type data
This species was described based on a single male specimen collected by Wilmatte Cockerell on February 2, foraging on Duranta plumieri (Fabaceae). The holotype (Fig. 15)

Type data
This species was described based on a single male specimen collected by Rua Williams in January at Zamorano, Honduras, foraging on Petrea voluvilis L. (Verbenaceae). The specimen (Fig. 16)

Type locality
Ecuador.

Comment
Centris triangulifera nom. rev., C. labrosa Friese, 1899 and C. terminata Smith, 1874 belong to a complex internal lineage of C. (Heterocentris), which apparently also includes some undescribed species from Central and South America. Unfortunately, a taxonomic revision of this subgenus is not currently available, so the real identity of these species is unclear. Centris labrosa and C. triangulifera are different species, but the type specimen of the latter is almost identical with that of C. semilabrosa, which was proposed as a junior synonym of C. terminata (see Moure et al. 2007). Considering the complexity of the problem, I propose to remove C. triangulifera (see below) from the synonymy of C. labrosa -a synonymy proposed by Snelling (1984), who apparently did not study the type specimens of the latter species -and C. semilabrosa from the synonymy of C. terminata, at least until a taxonomic revision of this group is made that allows to distinguish the limits between each of these species. Cockerell, 1949 nom. rev. Fig. 17 Centris (Rhodocentris) triangulifera Cockerell, 1949: 477. Type data Cockerell (1949) did not mention the number of females he studied to describe this species. According to the label of the type specimen examined at NMNH, apparently he had only a single specimen collected by Adan Rivera on January 30 at Zamorano, Honduras. The holotype (Fig. 17)

Type data
As for Centris caurensis, the single male specimen used by Cockerell (1919) to describe this species was also collected by Melbourne Carriker in the same locality. The holotype (Fig. 19)

Comment
Cockerell (1919) cited a specimen also collected by M. Carriker at the type locality of C. fusciventris caurensis and C. fusciventris matoensis as a subvariety of the latter subspecies. The characters cited by him about the integument of the clypeus and the pubescence on the fourth tergum are within the variation range of C. caurensis. Cockerell, 1917 Centris (Melanocentris) maroniana Cockerell, 1917: 475.

Type data
This species was described based on a single male specimen collected in Maroni, French Guiana. According to the original description, the specimen belonged to the Queensland Museum and was sent to Cockerell by Le Moult. Eugène Le Moult (1882-1967) was a French naturalist and entomologist specialist on butterflies who lived in French Guiana (Lhoste 1987

Type locality
French Guiana: Maroni ("Guyane, Maroni"). It is not clear whether the type specimen was collected in Saint-Laurent-du-Maroni, one of the two sub-prefectures of French Guiana or near the Maroni River, a natural border between French Guiana and Suriname.

Comment
This is one of the most stunning species of the subgenus C. (Melanocentris). Unfortunately, nothing is known about its bionomy and distributional range.

Type data
This species was described based on a pair of at least two females collected by Geraldo Cisneros on November 3 and 5 at Zamorano, Honduras. The female collected on November 3 was found at NMNH and has the number 58888 in the catalog of types. According to the original description and the label that the found specimen bears, it was designated by Cockerell as the holotype. The specimen (Fig. 20
VIVALLO F., The Centris bees described by Theodore D.A. Cockerell Snelling (1984) proposed the synonymy of C. obsoleta Lepeletier, 1841, C. melanochlaena and C. zamoranensis, following the interpretation of Friese (1901) of the former species. Centris melanochlaena and C. zamoranensis are actually conspecific, but C. obsoleta is a different species restricted to South America. The synonymy proposed by Snelling (1984) was corrected by Moure et al. (2007), maintaining C. obsoleta and C. melanochlaena as different species, with C. zamoranensis as a junior synonym of the latter. Cockerell, 1935 Centris rufosuffusa Cockerell, 1935b: 12-13.

Type data
Although the original description does not mention the number of males that were studied by Cockerell (1935b) to describe this species, the collector's note mentioned by him refers to an aggregation of males in a nesting area, which leads to believe that he could have used more than one specimen. However, one specimen studied by Cockerell (1935b) is currently housed at NHMUK bearing a label that agrees with the information cited in the original description. Considering this, I interpret that specimen as the holotype. It has the following data label:  1975). During this period, he collected the specimen(s) used by Cockerell (1935b) to describe this species.

Type locality
Trinidad and Tobago: Trinidad Island: Mayo-Rio Claro region: Point Radix.

Type data
This species was described based on one male and three females from Las Cruces, New Mexico, United States. According to the original description, Cockerell collected the type series on May 18, 1897, on Caesalpinia falcaria Fischer (Caesalpiniaceae). Two females were found at ZMB and ANSP bearing an original label of Cockerell that agrees with the data and the flower record cited by Cockerell. A female labeled "cotype" and a male labeled "Type" by Cockerell were found at NMNH and AMNH, respectively, but they don't have the date of collecting or the floral host. All these four specimens must compose the original type series studied by Cockerell. Although Cockerell (1897a) did not mention in the original description which specimen is the holotype, it can be inferred from the label of the male housed at AMNH. This interpretation is supported by the Articles 73.

Type locality
United States: New Mexico State: Las Cruces.

Type data
This species was described based on a single female specimen collected by Geraldo Cisneros on November 2, at Zamorano, central Honduras, foraging on Crotalaria L. sp. (Fabaceae). The type specimen (Fig. 21)
VIVALLO F., The Centris bees described by Theodore D.A. Cockerell

Comments
This species was proposed as a junior synonym of C. agilis Smith, 1874by Snelling (1984, but it is actually a different, albeit similar species. Centris cisnerosi has terga 3 and 4 brown (reddish brown in C. agilis), terga 2 to 4 covered with a light yellow pubescence (brownish in C. agilis), and the lower half of the clypeus areolate laterally (not areolate in C. agilis). According to the morphology of the mandible and the basitibial and pygidial plates, C. cisnerosi and C. agilis are related to one of the internal linages of the subgenus C. (Paracentris), not to C. (Aphemisia) as mentioned by Moure et al. (2007). No information about the person that caught the type specimen was found, but probably he was a collector who helped Cockerell during his stay in Honduras (Jesus Orozco, pers. comm.). Cockerell, 1923 Centris cockerelli resoluta Cockerell, 1923: 76-77.

Type data
This subspecies was described based on male and female specimens collected by the American entomologist Edward Payson Van Duzee at La Paz, Lower California, on June 3, 1921. Cockerell (1923) designated the female as the holotype which is currently housed at CAS and has the following data label:

Type data
This species was described based on a single male specimen collected by the American entomologist and dipterologist Charles Henry Typer Townsend (1863-1944) on March 27 during a trip to Peru. Townsend went there not to study bees, but cotton pests (Evenhuis 2013). He was a difficult man to work with and this no doubt led to his virtually isolating himself from other colleagues. Nevertheless, he and Cockerell had a cordial relationship and that could have facilitated the sending of some bees to be studied by this latter researcher. The male holotype, not female as mentioned in the original description, is housed at AMNH and has the following data label: Pachacayo.

Type data
This species was described based on two male specimens collected by Gordon Floyd Ferris on June 29, 1919, at La Paz, Mexico. Ferris (1893-1958 was an American entomologist specialist on 'small insects', like lice, parasitic flies and small blood-sucking Hemiptera (Usinger 1959). In 1919, he made an extensive field trip to Baja California with the American herpetologist Joseph Richard Slevin , where both males studied by Cockerell (1924) were collected. In the original description, Cockerell (1924) did not indicate which male was the holotype, and he also failed to indicate the sex of the exemplars, citing that they were two females. Snelling (1984) recognized the latter mistake and corrected the sex of the specimens. According to Cockerell (1924), Snelling (1984) and Moure et al. (2007), the type specimens are housed at CAS, but actually they are in the NMNH. One of the males bears a label of "type" and the other was labeled as "cotype". The holotype has the following data label:

Type locality
Mexico: Baja California Sur State: La Paz.

Type data
On May 18, 1896, at Mesilla Valley, Cockerell collected, on flowers of Caesalpinia falcaria, one male and seven females that were used to describe this species. The male of the type series is currently Centris hoffmanseggiae davidsoni Cockerell, 1904 Centris hoffmanseggiae subsp. davidsoni Cockerell, 1904a: 160. Junior synonym of C. (Paracentris) hoffmanseggiae (Snelling 1956).

Type data
This subspecies was described based on a single male specimen collected by dr. Davidson, probably the British geodesist, astronomer and geographer George Davidson (1825Davidson ( -1911, who worked in California State since 1850 (Davenport 1937
housed at NMNH and was designated by Fox (1899) as the lectotype. The specimen (Fig. 22)

Type locality
United States: New Mexico State: Las Cruces. Cockerell (1897a) mentioned that the type series was collected at "College Farm" in Mesilla Valley. Contrary to this information, the male studied by him bears a label of provenance of Las Cruces.

Comments
In the description of C. caesalpiniae, Cockerell (1897a) also mentioned a female collected by S. Steel on May 17, but it is not clear whether it belonged to the type series of the species. However, that specimen was not found in the collections visited during this research. Moure et al. (2007) mentioned the existence of a type series of C. morsei, but this cannot be inferred from the original description or from the label of the specimen housed at NMNH. In this collection, another male was found with the following data label: . Although this specimen bears a red label of type, it is not part of the type series because the date of collecting is different from that cited in the original description, and it also has a flower record not mentioned by Cockerell (1897b). Cockerell, 1923 Centris pallida callognatha Cockerell, 1923: 78. Junior synonym of C. (Paracentris) pallida Fox, 1899(Snelling 1956.

Type data
This species was described based on a single metander male collected by the American entomologist specialist in Orthoptera Latreille, 1793 Albert Pitts Morse

Type locality
United States: New Mexico State: Mesilla ("bed of the Rio Grande").

Type data
This subspecies was described based on a single female specimen collected by Edward P. Van Duzee on April 11, 1921, at Guaymas, northwestern

Type data
This species was described based on a single male specimen collected by E.P. Van Duzee at Punta Willard, Isla Tiburón, on July 3, 1921. The holotype is housed at CAS and has the following data label:

Comment
The type specimen of Centris rhodoleuca looks very different from males of C. rhodopus, but it is actually conspecific, corresponding to a metander male. Cockerell, 1897 Centris caesalpiniae var. rhodopus Cockerell, 1897a: 395.

Type data
This species was described based on two males and two females collected by Cockerell in Mesilla Valley, United States, foraging on Caesalpinia falcaria. One of those females was found at ANSP and one male at NMNH. Both specimens bear a label "cotype" and are in good condition. According to the treatment that Cockerell gave to his type specimens, both these exemplars must be paratypes.

Type data
This species was described based on a single male specimen collected at Mesilla Park, New Mexico, on June 24. The holotype (Fig. 24)

Type locality
United States: New Mexico State: Mesilla Park.

Comment
Centris rhodopus pulchrior is slightly larger than normal males of C. rhodopus, the hairs on the hind legs are lighter and the whitish bands on the terga are more defined. Cockerell, 1923 Centris tiburonensis Cockerell, 1923: 78.

Type data
This species was described based on a single female specimen collected by Wilmatte P. . The current condition and depository of the female paralectotype are unknown. The fixation of the lectotype allows confirmation of the synonymy between this species and C. wilmattae proposed by Zanella (2002).

Type data
This species was described based on a single female specimen collected in the central highlands of Peru. The holotype (Fig. 25)

Comment
Centris chlorura has only been recorded from its type locality. Snelling (1984) erroneously proposed this species as a junior synonym of C. festiva Smith, 1854. This mistake was emended by Moure et al. (2007).

Type data
It is not clear from the original description how many females were used by Cockerell (1919) to describe this subspecies. However, it was apparently based on a single specimen collected by Melbourne A. Carriker in Aroa city, northwestern Venezuela. The holotype (Fig. 26)

Comments
Centris claripennis was erroneously described from São Paulo, southeastern Brazil. Its distribution range in South America includes Colombia, Venezuela, and Trinidad and Tobago (unpubl. data). This species was erroneously proposed as a junior synonym of C. fuscata Lepeletier, 1841(Moure et al. 2007). The study of the type specimens of both species allows removing the former species from the synonymy of the latter, proposing its revalidation. Both sexes of C. claripennis can easily be identified by the yellowish coloration of the wings, being dark brown on the distal half. Centris heterodonta Cockerell, 1912 Centris (Ptilotopus) heterodonta Cockerell, 1912a: 43-44.

Type data
As for Centris bakeri, the female specimen studied by Cockerell

Comments
The study of the type specimens of C. dentata and C. heterodonta allows proposing the synonymy between these both species. Despite the distribution range mentioned by Moure et al. (2007), this species occurs only in South America (unpubl. data). Cockerell, 1919 Fig. 27 Centris pachysoma Cockerell, 1919: 189. Type data Cockerell (1919) described this species based on a single female specimen collected by William Rosenberg, the same collector of the type specimen of C. calloxantha (see below). The holotype (Fig. 27)

Type data
This species was described based on a single female specimen collected in Cacao, northern Guatemala, by two renowned entomologists, the German-American Eugene Amandus Schwarz (1844Schwarz ( -1928 (Howard 1928) and the American Herbert Spencer Barber (1882Barber ( -1950. Both researchers were skillful collectors specialized in the study of beetles (Caudell 1907). The holotype (Fig. 28)

Type data
This species was described based on a couple of female and male specimens collected in Arequipa, southern Peru. Both specimens were collected by Edmundo Escomel (1880-1959), a Peruvian doctor who dedicated his free time as a meticulous observer of the flora, fauna and the natural environment (Velarde 2006). The male holotype (Fig. 29)

Comment
This species was correctly interpreted as a valid species by Moure et al. (2007) and Vivallo (2013).

Type data
Cockerell (1919) did not specify how many males he studied to describe this species. According to the labels of the type specimen housed at NMNH, it was based on a single specimen collected by William Rosenberg in Chanchamayo, Peru. The holotype (Fig. 30)

Comment
The species described by Cockerell only differs from normal males of C. bicolor by the presence of dark brown hairs on the mesoscutum.

Type data
The type specimen of this species was collected in 1911 during the Stanford Expedition by the embryologist Harold Heath

Discussion
As is generally the case with particularly diverse lineages, the taxonomy at specific level can be quite complex, not only because of the large number of species described, but also because of the large number of the synonymies proposed throughout the taxonomic history of the group. This is observed in Centris, where approximately one third of the proposed names are currently considered synonymous (see Moure et al. 2007).
The access to the type material, along with its careful study, allows evaluating the status of each proposed name, as well as, in some cases, to solve misidentifications or misinterpretations that originated in the past and have remained unnoticed until now (see, for example, Vivallo 2019Vivallo , 2020. It is logical to think that in Centris, as well as in other especially diverse groups of bees, the real number of species will change in the future, once the primary types of all the proposed species are studied. This will allow not only to correct erroneous identifications of specimens deposited in collections and museums, but also to recognize and describe new species and, above all, to correctly associate the bionomic data and ecological relationships that are being continuously published, with the real identity of the species that are being studied.
The detailed record of the primary types, their labels and the confirmation of their repository are fundamental steps to solve current taxonomic problems and avoid future problems related to the identity of the species. This also allows other taxonomists to have appropriate tools to locate and recognize which specimens were specifically studied by the authors of each species at the time they described them, facilitating their re-study or reinterpretation if necessary.
Hogan (OUMNH), Seán Brady (NMNH), and Michael Ohl (ZMB) for allowing me to visit their institutions to study the specimens under their care. I also thank Robert Zuparko, Norman Penny, Vince Lee (CAS), Stefan Cover (MCZ), Lars Vilhelmsen (NHMD), Jesús Orozco (Departamento de Ciencia y Producción Agropecuaria, Escuela Agrícola Panamericana El Zamorano, Tegucigalpa, Honduras) and the anonymous reviewers for their help. Financial support was provided by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, grant 444320/2014-8), Brazil. This paper is part of the SIGMA project Nº21565 MN/UFRJ and contribution number 36 from the HYMN.