Portugal: Coimbra: BMNH 1864.9.19.35–37 [syntypes].

Bocage (1864b) first described Chioglossa lusitanica in a paper that appeared on the Revue et Magasin de Zoologie Pure et Appliquée in August 1864, and a “duplicate” description was subsequently published in the Proceedings of the Zoological Society of London in November of the same year (Bocage 1864a). Although Bocage (1864a, 1864b) provided measurements for only one specimen, he based his description on an unspecified number of specimens from Coimbra, Portugal and promptly sent duplicates of his newly described species to other Portuguese and European museums (Bauer et al. 1993; Calado 2015). Even though they were not noted as types by Boulenger (1882a), the specimens presented to the British Museum were certainly part of Bocage’s type series. The three specimens were sent on 24 May 1864 via J. J. Monteiro, together with three manuscripts to be considered for publication in the Proceedings of the Zoological Society of London, one of which contained “la description, accompagnée d’une planche, du Salamandrien dont je vous adresse 3 individus, et que j’ai nommé Chioglossa lusitanica” (NHMA/DF/GüntherColl/16/1/102; Bocage 1864a). We here recognize these specimens as syntypes of Chioglossa lusitanica, which are in a poor state of preservation.

Angola: Caconda: BMNH 1887.3.23.1.

Bocage (1873a) described Hylambates anchietae based on a single specimen from “l’intérieur de Mossamedes”, and later recorded it from “Huilla, Caconda et Quindumbo” (Bocage 1895a). At that time, “l’intérieur de Mossamedes” corresponded to the inland areas of southwestern Angola, encompassing present day Namibe, Huíla and Cunene provinces. Perret (1976) examined the types in the Lisbon Museum and considered a juvenile specimen from Huíla (MB T. 13-233) to be the holotype. Even though Bocage (1873a) mentioned a single specimen in the original description, Perret (1976) referred additional material from Caconda to the type series: two juveniles and an adult male as “Cotype I” (MB T. 13-234) and an adult female as “Cotype II” (MB T. 236). It is possible that these “cotypes” were available to Bocage at the time of description, but the strict reference to a single individual in the description precludes them of being considered part of the type series. Although Boulenger (1882b) included the species in his Catalogue, there were no specimens in the British Museum at that time. Only a few years later Bocage sent a specimen to the British Museum, noted in a letter dated 19 March 1887 (NHMA/DF/ZOO/235/1/1/1/76) as “1. Un individu de l’Hylambates Anchietae de Caconda (Angola)”. Interestingly, a specimen at the Muséum d’Histoire naturelle, Genève (MHNG 953.11) has a label written by Bocage with the information “Hyl. Anchietae, Caconda (Angola), 1.” (LMPC pers. obs.). The specimen in question, collected by Frederick Creighton Wellman (1870–1960) at “Chiyaka District”, was exchanged from the British Museum (originally BMNH 1908.5.15.20), suggesting that the label was likely misplaced while both specimens were still in London. While Bocage (1873a) undoubtedly based his description on a single specimen, it is plausible to assume that the specimen from Caconda sent to the British Museum would have been part of what Perret (1976) considered as “cotypes”, even if this claim has no nomenclatural value.

Angola: “West Africa” [= Duque de Bragança]: BMNH 1875.5.22.3 (Fig. 6), BMNH 1875.5.22.2 [holotype; Figs 7, 8], Quissange: BMNH 1887.3.23.3, Caconda: BMNH 1887.3.23.2 (not examined) [syntype of Hylambates angolensis Bocage, 1893], “Angola”: BMNH 1896.2.28.2 (not examined).

Cystignathus bocagii was described by Günther (1865a), but the type material for the taxon is the subject of some confusion (Bocage 1866a, 1895a; Perret 1976; Marques et al. 2018; Frost 2024). In a letter dated 19 September 1864 (AHMB/CE/G75), Günther thanked Bocage for a shipment of reptiles from Duque de Bragança, noting that “several of the frogs appear to be new”. Günther stated that “I shall return all the specimens which you desire to keep, hoping that if you should receive duplicates at some future time, you will kindly communicate to us what you can spare”. In the original description Günther (1865a) acknowledged Bocage, “who has allowed me to examine the unique specimen brought from the province of Duque de Bragança (Angola) to the Lisbon Museum”. In a subsequent letter dated 24 July 1865 (AHMB/CE/G76) Günther classified the material sent earlier by Bocage, and although some specimens are noted as having been presented to the British Museum, the holotype of Cystignathus bocagii (identified as No. 11) is listed among the material to be returned to Lisbon (AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104; Fig. 5). In the following year, Bocage wrote back to Günther expressing doubts concerning the identity of two specimens returned by Günther as Cystignathus bocagii (No. 11, the holotype) and Leptopelis natalensis (No. 12), which were then sent back to London to be reexamined (AHMB/CE/G78, NHMA/DF/ZOO/200/1/184, NHMA/DF/GüntherColl/16/1/110). After reexamining the two specimens, Günther agreed with Bocage and confirmed that they both represented Cystignathus bocagii, asking to keep one as duplicate in the British Museum (AHMB/CE/G/79). In response, in a letter dated 10 July 1866, Bocage allowed Günther to keep one of the specimens but wished to keep the type in Lisbon (NHMA/DF/ZOO/200/1/185).

In the first issue of the Jornal de Sciencias Mathematicas, Physicas e Naturaes, published in November of the same year, Bocage (1866a) mentioned two specimens collected by Bayão from Duque de Bragança, stating that “Um d’elles é o typo da especie, o outro offerecemol-o ao Museu Britannico” [One of them is the type of the species, the other we offered to the British Museum]. Years later, in his Catalogue of the BatrachiaSalientia in the British Museum, Boulenger (1882b) mentioned two specimens from “W. Africa”, originally from the Lisbon Museum (Figs 6, 7). In his major work on Angolan herpetofauna, Bocage (1895a) stated “Deux individus, types de l’espèce, qui existent au Muséum Britannique et deux autres individus jeunes faisant partie de nos collections d’Angola sont les seuls exemplaires connus de cette espèce” [two specimens, types of the species, that exist in the British Museum and two other juvenile specimens that are part of our collections from Angola are the only known exemplars of this species]. When Perret (1976) examined the types in the Lisbon Museum he referred to a juvenile from Duque de Bragança as “Cotype I” (MB T. 15-232), and Frost (2024) stated that two specimens sent to the British Museum may be types.

We are confident that one of the two specimens from “West Africa” (BMNH 1875.5.22.2–3; Figs 6, 7) cited by Boulenger (1882b) corresponds to the holotype of Cystignathus bocagii. These specimens are accessioned in the register as Hylambates bocagei from “W. Africa” with a note stating they were “Received some years ago from the Lisbon Museum for examination”, suggesting that they were those initially sent in Bocage’s first shipment of specimens from Duque de Bragança and later sent again for reexamination (AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 110). While the type was supposed to be returned to Lisbon, we found no additional references that may explain why these specimens remained in London until they were accessioned in 1875. We speculate that Bocage received additional specimens in the meanwhile, leading him to present the specimens to Günther and mention two types in the British Museum (Bocage 1895a). The specimens in question were subsequently labeled as Leptopelis viridis, presumably by Parker (1936), who recognized this as the only taxon occurring in West Africa and was unaware of the true origin of the specimens labeled as “W. Africa”. Although the specimens still bear the original paper labels where the numbers mentioned in the letters would have been stated, these are completely faded and thus do not allow us to unambiguously match them with those cited in the letters. However, while both specimens generally fit Günther’s (1865a) description of Cystignathus bocagii, one of them more closely resembles the illustration of the holotype provided by Günther (1865a) even after 160 years of preservation (Fig. 8). The visible discrepancy between the specimen and Günther’s (1865a) figure is likely a result of the technique used to produce the illustration, resulting in a mirrored image of the specimen (Ceríaco and Bauer 2017). Furthermore, distinct pectoral glands are evident in BMNH 1875.5.22.3, a character not mentioned in the original description (Fig. 6). Based on this evidence, we recognize BMNH 1875.5.22.2 as the holotype of Leptopelis bocagii.

On 19 March 1887 Bocage sent additional Hylambates material to the British Museum (NHMA/DF/ZOO/235/1/1/1/76), including a specimen of Hylambates anchietae from Caconda (No. 1, see previous account), one specimen from Portuguese Guinea (No. 4, see Leptopelis viridis account) and two other specimens of a species that Bocage considered new: “2. Un individu d’une autre espèce de Hylambates, que je n’ai pu rapporter à aucune des espèces décrits dans votre Catalogue et que j’ai nommé provisoirement – H. angolensis, il vient de Caconda”, “3. Un autre individu, provenant d’un autre localité, qui me semble constituer à peine une variété interessante de l’ espèce précedente. Tous les individus que j’ai reçu de cette localité (Quissange) portent la grande tache noire sur les dos”. Some years later, Bocage (1893a) formally described Hylambates angolensis, and although he provided measurements for only an adult female, he mentioned material collected by Anchieta from “Caconda, (…) Quissange, Quibula, Quindumbo et Cahata, dans l’intérieur de Benguella”. Boulenger (1906 “1905”) referred Bocage’s Hylambates angolensis to the synonymy of bocagii, mentioning specimens sent by Bocage under that name, and was followed by subsequent authors (Parker 1936; Perret 1976).

The specimens sent by Bocage in 1887 were accessioned in the British Museum in the same order as listed in the letter, with the numbers BMNH 1887.3.23.1–4, where BMNH 1887.3.23.2 and 1887.3.23.3 are marked as types of Hylambates angolensis. Of these two putative types, we could only locate BMNH 1887.3.23.3 during our visits. Although the specimen bears the number “4” on the original label, the remaining data agree with Bocage’s description of No. 3 in the letter, i.e., “Hylambates angolensis var. ?” from “Quissange (Angola)”. However, it is unclear if the number was originally written by Bocage or subsequently added to the specimen label. Perret (1976) identified three syntypes of Hylambates angolensis in Lisbon: MB T. 14-244 from Quissange, MB. T. 14-242 from Caconda and MB T. 14-242 from Quindumbo. While the specimens in the British Museum were presented some years before the formal description was published, the contents of Bocage’s letter (NHMA/DF/ZOO/235/1/1/1/76) suggest that these were already included in his concept of Hylambates angolensis at the time. Although Bocage reconsidered the status of his specimens from Quissange and assigned them to Hylambates angolensis in the published description (Bocage 1893a, 1895a), the fact that he referred to specimen No. 3 (BMNH 1887.3.23.3) in his letter as a variety of his new taxon leads us to follow a conservative definition and not consider this specimen as part of the type series.

An additional specimen of Hylambates angolensis is mentioned in a letter dated 24 February 1896 (NHMA/DF/ZOO/235/1/1/1/76) and listed in the register with the number BMNH 1896.2.28.2 but could not be located. Neither the syntype of Hylambates angolensis (BMNH 1887.3.23.2) nor the last Hylambates specimen shipped from Lisbon (BMNH 1896.2.28.2) are recorded in the British Museum’s modern database and thus remain unaccounted for.

Guinea-Bissau: Bolama: BMNH 1887.3.23.4 [syntype of Hylambates cynnamomeus Bocage, 1893; Fig. 9].

In a letter detailing a shipment of Hylambates specimens (NHMA/DF/ZOO/235/1/1/1/76, see previous accounts), Bocage mentioned a specimen “4. Un individu d’une autre espèce de Hylambates, de Bolama (Guiné) peut-être voisin de H. rufus, mais qui me semble distinct de celui-ci”. Some years later Bocage (1893a) described Hylambates cynnamomeus, providing measurements for an adult male and mentioning an unspecified number of specimens from Quillenges in Angola and Bolama in Portuguese Guinea, noting also that the new species was similar to Hylambates viridis. In his catalogue of “types” in the Lisbon Museum, Bocage (1897) mentioned only Quillenges, seemingly restricting the type locality. Boulenger (1906 “1905”) mentioned a specimen sent by Bocage from Bolama under the name cynnamomeus, which he considered identical to viridis. These observations led Boulenger (1906 “1905”) to refer both Bocage’s angolensis and cynnamomeus, as well as Günther’s viridis, to the synonymy of Hylambates bocagei, while Parker (1936) regarded Hylambates viridis as a valid species from West Africa and considered cynnamomeus records from that region as referring to viridis. Upon examination of the types in the Lisbon Museum, Perret (1976) identified the “holotype” from Quillenges (MB T. 16-250) and four “paratypes” from Bolama (MB T. 16-248 and 16-249), confirming that the specimens from Bolama belonged to Leptopelis viridis. The specimen presented by Bocage to the British Museum in 1887 (BMNH 1887.3.23.4) is here recognized as a syntype of Hylambates cynnamomeus. While in his letter Bocage recognized it as a distinct, yet unnamed species (NHMA/DF/ZOO/235/1/1/1/76), the original label attached to the specimen bears Bocage’s yet unpublished name “Hyl. cinnamomeus n. sp.” (Fig. 9), thus representing compelling evidence that the specimen belonged to the original, composite, type series. Similarly to BMNH 1887.3.23.3, although the number on the specimen label (i.e., “2”) does not match, the remaining data agree with the description of specimen No. 4 in Bocage’s letter. It is unclear if the number was originally written by Bocage or subsequently added to the specimen label

Angola: Quissange: BMNH 1887.3.23.5.

This specimen was sent together with a shipment of Hylambates specimens on 19 March 1887, identified in both the letter and specimen label as No. 5 (NHMA/DF/ZOO/235/1/1/1/76). In his letter Bocage noted “5. Enfin un individu, de Quissange, que j’hésite à considérer comme étant le Breviceps gibbosus et qui me semble également distinct du B. mossambicus, dont je possède un individu identique à la fig de Peters”. Bocage (1873a) first assigned his Angolan material to Breviceps gibbosus and later to Breviceps mossambicus (Bocage 1895a). While we provisionally refer this specimen to the recently described Breviceps ombelanonga, further genetic sampling of historical localities is critical to identify populations of this morphologically conservative group (Nielsen et al. 2020).

Angola: Dombe (Benguella): BMNH 1947.2.21.3–4 [syntypes, originally BMNH 1896.2.28.3–4; Figs 10, 11].

Although Bocage (1895b) described Bufo dombensis based on 18 specimens collected by Anchieta at “Dombe (Benguella)”, Perret (1976) only identified two syntypes in the Lisbon Museum. The two syntypes (Figs 10, 11) in the British Museum were sent by Bocage on 24 February 1896 (NHMA/DF/ZOO/235/1/1/1/76) and were briefly cited by Boulenger (1898). Although not noted as types by Boulenger (1898), the specimens in the British Museum are generally regarded as syntypes (Marques et al. 2018; Baptista et al. 2023).

Genus Sclerophrys Tschudi, 1838

Angola: Benguela: BMNH 1872.2.15.3 [syntype of Bufo benguelensis Boulenger, 1882; Fig. 12], “W. Africa” [= Caconda]: BMNH 1883.7.26.27.

Bocage (1866b) described Bufo funereus based on a juvenile specimen from Duque de Bragança and later recorded additional specimens from Caconda (Bocage 1882b, 1895a). Even though Bocage (1895a) clearly referred to the specimen from Duque de Bragança as the type of the species, Perret (1976) considered two specimens from Caconda as “cotypes”, demonstrating that the author had a different and peculiar concept of “type”. Boulenger (1882b) described Bufo benguelensis based on three specimens, including one from Benguela sent by Bocage some years earlier (BMNH 1872.2.15.3). Even though it was formally described by Boulenger (1882b), the name was used by Bocage earlier in a letter from 24 May 1869 (NHMA/DF/ZOO/200/1/189), where he identified the specimen as “4. Bufo benguellensis nov. sp.” (Fig. 4). Furthermore, this was the name stated in the register when the specimen was accessioned a decade before Boulenger’s (1882b) publication (Fig. 3), and the label on the specimen jar identifying it as Bufo benguelensis attributes the nomen authorship to Bocage (DP pers. obs.). In an addendum to the same work, Boulenger (1882b) referred benguelensis to the synonymy of funereus based on an additional specimen sent by Bocage for comparison (BMNH 1883.7.26.27) (AHMB/CE/G93, NHMA/DF/ZOO/200/21/38). Although the second specimen bears only the locality “W. Africa”, it was most likely collected by Anchieta at Caconda considering that, in addition to the type of funereus from Duque de Bragança and the type of benguelensis sent to the British Museum, Bocage (1882b, 1895) only recorded the species from Caconda. Interestingly, Bocage (1866b, 1882b, 1895a, 1897) never directly reported the species from Benguela, even though he acknowledged Boulenger’s benguelensis as a synonym of funereus. Considering that no specific locality is stated in the letter where Bocage mentions the specimen, it is possible that Boulenger may have inferred the locality from Bocage’s name benguelensis, and the specimen actually originates from Caconda, at the time considered part of the “interior of Benguella”. While we assume that the type locality of Bufo benguelensis could be questionable, this scenario is merely speculative.

Angola: Duque de Bragança: BMNH 1864.10.28.1.

Bocage sent this specimen to the British Museum in 1864 along with several other specimens for Günther to examine, most of which were returned in the following year (AHMB/CE/G76). Although the original specimen label is completely faded, the specimen certainly corresponds to No. 19 of Bocage’s first shipment of Angolan material, which was noted as a duplicate specimen to be presented to the British Museum (NHMA/DF/GüntherColl/16/1/104). It was identified by Günther as Bufo pantherinus and later cited by Boulenger (1882b).

Angola: “W. Africa”: BMNH 1872.2.15.1.

Bocage (1864b) described Rana bragantina based on a specimen from Duque de Bragança but shortly thereafter referred it to the synonymy of Günther’s Rana occipitalis (Bocage 1866a, 1895a). The two authors exchanged letters in 1869 regarding the identity of two frogs that Bocage sent to the British Museum under the genus Rana (AHMB/CE/G85, NHMA/DF/ZOO/200/1/189, 190). One of the specimens was sent by Bocage as “No. 3. Rana plicigula nov. sp.” (Fig. 3) and was identified by Günther as Rana occipitalis. Although the original label with the number mentioned in the letter is missing, the specimen in question certainly corresponds to BMNH 1872.2.15.1, while the other specimen, identified with the number 5 both in the letters and the specimen label represents Amietia angolensis (see respective account). Both specimens were cited by Boulenger (1882b) as Rana occipitalis. The name Rana plicigula is stated in the exchanged letters, the register and the label on the jar of BMNH 1872.2.15.1, but it never appeared in a published form, as it was a working name for a species that Bocage intended to describe before knowing Günther’s opinion. Although there is no precise locality associated with the British Museum specimen, Bocage (1895a) knew the species only from Duque de Bragança, Dondo, Ambaca, Novo Redondo and Catumbela.

Angola: Duque de Bragança: BMNH 1864.10.28.10.

Bocage (1866a) recorded a single specimen of Hyperolius fulvovittatus from Duque de Bragança, later reported as Rappia fulvovittata (Bocage 1895a). Perret (1976) located the specimen in the Lisbon Museum and identified it as Afrixalus wittei. Although not mentioned by Bocage (1866a, 1895a), he had presented another specimen from Duque de Bragança to the British Museum in 1864 (AHMB/CE/G76), which was identified and cited as Rappia fulvovittata by Günther (1868b) and Boulenger (1882b). Since the specimen in the Lisbon Museum was destroyed by fire, BMNH 1864.10.28.10 is one of only two specimens of Afrixalus wittei known from Angola (Ceríaco et al. 2018).

Genus Hyperolius Rapp, 1842

Angola: Duque de Bragança: BMNH 1947.2.9.68 [lectotype, originally BMNH 1864.10.28.14; Fig. 13].

Among the material from Duque de Bragança sent by Bocage in 1864, Günther identified three specimens of a new species he described as Hyperolius nasutus (AHMB/CE/G76, Günther 1865a). In the short description, Günther (1865a) did not specify a type, but instead provided measurements for a single specimen and mentioned that “This species inhabits the province of Duque de Bragance, whence we have received it through M. Barboza du Bocage. Other specimens are in the Lisbon Museum”. In his letter to Bocage, Günther listed one specimen that was presented to the British Museum (No. 10) and two others to be returned to Lisbon (Nos 15 and 16), identifying specimen No. 15 as the “type” (AHMB/CE/G76; Fig. 5). Nevertheless, both Boulenger (1882b) and Bocage (1895a) referred to the specimen in the British Museum as the type. Perret (1976) did not note any type specimen referrable to this species in the Lisbon Museum, and subsequent authors continued to recognize BMNH 1947.2.9.68 as the holotype (Marques et al. 2018; Frost 2024).

This information leads to two conflicting interpretations: one in which Günther designated a “type” in communication to Bocage – corresponding to specimen No. 15, returned to Lisbon –, and another in which the specimen effectively measured and described by Günther is considered as the “type” – corresponding to specimen No. 10, presented to the British Museum. In any case, although the specimen in the British Museum is generally regarded as the holotype (Marques et al. 2018; Frost 2024), it is now clear that Günther examined three specimens, which collectively constitute the type series. Considering this conflicting information and potential interpretations, and the fact that the two specimens returned to Lisbon are now lost, we here designate BMNH 1947.2.9.68 (Fig. 13) as the lectotype of Hyperolius nasutus, which is still identified with the No. 10 in the original label and generally fits Günther’s (1865a) description. Hyperolius nasutus senso lato is a known species complex with at least 16 recognized species (Channing et al. 2013).

Angola: Duque de Bragança: BMNH 1864.10.28.11–13, Benguella: BMNH 1867.7.23.24 [syntype of Hyperolius insignis Bocage, 1867; Fig. 14], “W. Africa” [= Huíla]: BMNH 1872.2.15.4 [syntype of Hyperolius huillensis Bocage, 1873; Fig. 15], “W. Africa”: BMNH 1872.2.15.5.

The first specimens (BMNH 1864.10.28.11–13), collected by Bayão at Duque de Bragança, were presented to the British Museum in 1864 and identified by Günther as Hyperolius marmoratus (AHMB/CE/G76), leading Bocage (1866a) to report numerous specimens under that name. A subsequent specimen (BMNH 1867.7.23.24; Fig. 14) from Benguela was presented on 13 July 1867 as Hyperolius insignis (NHMA/DF/ZOO/200/1/195, NHMA/DF/GüntherColl/16/1/112), a species described by Bocage in the same year based on numerous specimens collected by Anchieta in Benguela (Bocage 1867b). This specimen, which generally agrees with that depicted by Bocage (1867b), was cited by Günther (1868b) as Rappia insignis and later by Boulenger (1882b) as Rappia marmorata, who noted it “as typical of Hyperolius insignis”. Although not noted elsewhere in the British Museum records, this specimen was certainly part of the type series and is here recognized as a syntype of Hyperolius insignis, adding to a surviving syntype in the Museum für Naturkunde (ZMB 6462) recently reported by Tillack et al. (2021). Although Perret (1976) reported two syntypes from São Salvador do Congo and Novo Redondo (currently M’banza-Kongo and Sumbe, respectively, both destroyed by fire), these were reported by Bocage (1887a) years after the original description and should not have been considered part of the type series. Both surviving syntypes in the British Museum and ZMB bear the locality “Benguella” noted in the original description.

According to the register, two additional specimens sent by Bocage were accessioned in 1872 as Hyperolius huillensis from “W. Africa”, a species described by Bocage in the following year based on four specimens collected by Anchieta at Huíla (Bocage 1873a). Boulenger (1882b) reported one specimen in the British Museum as Rappia marmorata, noting it “as typical of Hyperolius huillensis”. A list of amphibians sent to London is available on a letter from Bocage dated 24 May 1869 (NHMA/DF/ZOO/200/1/189), where two Hyperolius specimens are noted: “No. 1. Hyperolius huillensis nov. sp.” and “No. 2. Hyper. sp. ?” (Fig. 4). These specimens correspond to BMNH 1872.2.15.4 and 1872.2.15.5 (Fig. 15), which still bear the labels with the respective numbers mentioned in the letter (i.e., “1” and “2”). Although Perret (1976) identified four syntypes in the Lisbon Museum, the fact that Bocage sent a specimen to London identified as “Hyperolius huillensis nov. sp.” provides compelling evidence that it was used by him to conceptualize and describe the species, even though the formal description was only published some years later (Bocage 1873a), and we thus recognize BMNH 1872.2.15.4 as a syntype of Hyperolius huillensis. While we tentatively include BMNH 1872.2.15.5 in the material referred to Hyperolius cf. parallelus, neither Bocage nor Boulenger (1882b) reached a conclusive determination and the true identity of the specimen remains ambiguous. Since Bocage (1895a) regarded both insignis and huillensis as varieties of Rappia marmorata, he did not mention these taxa in his catalogue of type material in the Lisbon Museum (Bocage 1897). As noted by Marques et al. (2018) and Santos et al. (2021), the taxonomy of the Angolan members of the Hyperolius parallelus group is still unstable, and the proper identification of types of the numerous nominal taxa described and still associated with the group may prove critical for future taxonomic assessments.

Angola: Duque de Bragança: BMNH 1864.10.28.6–9.

The four specimens were collected by Bayão in 1864 and presented to the British Museum in the same year. Bocage tentatively referred them to the genus Stenorhynchus (NHMA/DF/GüntherColl/16/1/106), while Günther wrote that they “cannot be determined without other specimens” (AHMB/CE/G76). Although the specimen label attached to one of the specimens is completely faded, these most certainly correspond to No. 8 of Bocage’s first shipment of Angolan specimens (AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106) and were cited by Günther (1865a) and Boulenger (1882b).

Angola: Benguella: BMNH 1867.7.23.25.

The specimen in the British Museum, collected by Anchieta in “Benguella”, was likely among the first specimens of this species that Bocage examined. It was presented to the British Museum in 1867 as “Dactylethra Mulleri” (NHMA/DF/ZOO/200/1/195) and cited by Boulenger (1882b) as Xenopus muelleri, years before Bocage (1895a) described Xenopus petersii.

Angola: Duque de Bragança: BMNH 1864.10.28.2.

Among the material collected by Bayão at Duque de Bragança and sent to the British Museum in 1864, Bocage tentatively referred two specimens (No. 4) to “R. superciliaris ?” (NHMA/DF/GüntherColl/16/1/106), which Günther identified as Rana oxyrhyncha (AHMB/CE/G76), later reported by Boulenger (1882b). One of the specimens (originally BMNH 1864.10.28.3) was subsequently exchanged to the Iziko South African Museum in Cape Town, South Africa, where it remains deposited under the catalog number SAM ZR-002338.

Angola: Duque de Bragança: BMNH 1864.10.28.4–5.

Two additional Ptychadena specimens (No. 5) were included in the 1864 shipment from Duque de Bragança, tentatively identified as “R. Bibroni Hallowel ?” [sic] by Bocage (NHMA/DF/GüntherColl/16/1/106) and referred to Rana mascaraniensis by Günther (AHMB/CE/G76). Boulenger (1882b) cited three specimens of Rana mascaraniensis sent by Bocage, although we found no indication of additional Ptychadena specimens. The specimens were eventually associated with Rana mascaraniensis var. mossambica, until they were examined and identified as Ptychadena porosissima by John Poynton (1931–) in 1990. Although the species was described by Steindachner (1867) based on a specimen from “Angola” sent by Bocage to the Naturhistorische Museum in Wien, it most likely was part of the same material collected by Bayão at Duque de Bragança.

Angola: “W. Africa”: BMNH 1872.2.15.2, “Angola”: BMNH 1896.2.28.1.

Bocage (1866a, 1866b) described Rana angolensis based on two adult specimens collected by Bayão at Duque de Bragança. Perret (1976) mentioned only one specimen from Duque de Bragança as “holotype” in the Lisbon Museum, even though Bocage clearly mentioned two specimens in the description. In a letter from 24 May 1869, Bocage listed a specimen as “5. Rana sp. ? (Voisin de R. plicigula et de R. delalandii)” (NHMA/DF/ZOO/200/1/189; Fig. 4), which corresponds to BMNH 1872.2.15.2, still bearing a label with the number “5”. This specimen was accessioned as Rana plicigula together with BMNH 1872.2.15.1 (see Hoplobatrachus occipitalis account) and was later reported by Boulenger (1882b) as Rana angolensis. Bocage sent the second specimen (BMNH 1896.2.28.1) to Boulenger in 1896, asking him to compare it with Rana delalandii in a letter dated 24 February 1896 (NHMA/DF/ZOO/235/1/1/1/76). A specimen preserved at the Museum für Naturkunde (ZMB 10075) may represent the only surviving syntype of Amietia angolensis (Tillack pers. comm.).

Genus Tomopterna Duméril & Bibron, 1841

Angola: “W. Africa”: BMNH RR 1933.1.6.1 [reregistered, originally BMNH 1872.2.15.6].

Günther (1865a) described Pyxicephalus rugosus based on two specimens collected by Friedrich Welwitsch (1806–1872) at Pungo Andongo, but the name was preoccupied. Bocage sent one specimen to the British Museum in 1869 identified as “E/8. Pyxicephalus rugosus Gthr ?” (NHMA/DF/ZOO/200/1/189; Fig. 4), which was later reported by Boulenger (1882b) when he provided the replacement name Rana tuberculosa. Despite the lack of a precise locality, the specimen likely originates from southern Angola, as Bocage (1873a) recorded the species from “Huilla et d’autres localités dans l’intérieur de Mossamedes”. The specimen was reregistered in 1933 and is currently catalogued as BMNH RR 1933.1.6.1, although it certainly corresponds to BMNH 1872.2.15.6, the only specimen sent by Bocage as Pyxicephalus rugosus in the British Museum records and still identified with a label bearing the number mentioned in the 1869 letter (i.e., “E/8”).

Angola: Duque de Bragança: BMNH 1866.6.11.1.

The specimen was among the second batch of specimens collected by Bayão at Duque de Bragança, presented by Bocage to the British Museum in 1866 as “Nº 1 - Stellio angolensis Bocage nov. sp.” (NHMA/DF/GüntherColl/16/1/110). Bocage (1866a) recorded several specimens as Stellio nigricollis, stating that he had first thought they represented a new species he intended to name Stellio angolensis, but refrained from doing so after consulting Günther, who referred it to Agama nigricollis. Wagner et al. (2018) regarded angolensis to be a nomen nudum and suggested that Bocage’s reference to nigricollis was an error, also noted by Bocage himself (Bocage 1895a). We here confirm this observation, as in his letter dated 29 June 1866 Günther unambiguously wrote “Stellio angolensis = Agama atricollis”, and Bocage mentions also “Agama atricollis” in his response (AHMB/CE/G79, NHMA/DF/ZOO/200/1/185). The specimen presented to the British Museum was cited by Boulenger (1885a) as Agama atricollis in his Catalogue of Lizards and more recently as a newly described species by Marques et al. (2022).

Angola: Caconda: BMNH 1893.12.27.2.

This specimen was sent in 1893 as Agama atricollis and belongs to a recently described species known only from Angola and neighboring Namibia (Wagner et al. 2021). Bocage (1895a) mentioned specimens collected by Anchieta in Caconda.

Genus Agama Daudin, 1802

Angola: Caconda: BMNH 1893.12.27.5–6, “Hauts-Plateaux”: BMNH 1896.6.9.2–3.

Populations of ground agamas from the Angolan highlands have historically been associated with either Agama aculeata or Agama armata (Marques et al. 2018). Marques et al. (2018) referred Angolan populations to Agama aculeata, while Conradie et al. (2022) assigned new material to Agama armata based on a reticulated rather than striped gular pattern following Jacobsen (1992). Although the specimens sent by Bocage to the British Museum agree with this diagnosis, further research is needed to clarify taxonomic and geographic boundaries within the Agama aculeata complex (Leaché et al. 2014). Specimens BMNH 1893.12.27.5–6 were identified by Bocage as Agama armata on the specimen labels, while BMNH 1896.6.9.2–3 were referred to the same species in a letter dated 2 June 1896 (NHMA/DF/ZOO/235/1/1/1/76). Although Boulenger and Power (1921) cited a specimen from Caconda sent by Bocage (in error, see Agama anchietae account), the authors apparently did not cite any of the material presented by Bocage as belonging to the Angolan “plateau” species.

Mozambique: Lourenço Marques: BMNH 1896.6.9.1.

In May 1896, Bocage (1896a) recorded four specimens collected by Francisco Joaquim Dias Quintas (1851–date of death unknown) in Lourenço Marques (currently Maputo) as “Agama sp. ?”. In the same year, in preparation for a work on the ground agamas of Angola (Bocage 1896b), he wrote to Boulenger to exchange specimens and opinions (NHMA/DF/ZOO/235/1/1/1/76). In a letter dated 2 June 1896 Bocage discussed a shipment of Agama spp. to the British Museum, including a juvenile specimen from Mozambique (No. 3) that he tentatively identified as “Agama hispida ?”. The specimen was accessioned as Agama hispida with the number BMNH 1896.6.9.1 and a correction on the jar label indicates it was later reidentified as Agama armata. It was cited as Agama hispida var. armata by Boulenger and Power (1921). We provisionally treat the specimen as Agama cf. aculeata distanti based on geographic distribution and a striped gular pattern following Jacobsen (1992), although the status of the Agama aculeata complex requires further work to establish geographic and specific delimitations (Leaché et al. 2014).

Angola: Catumbella: BMNH 1893.12.27.7, “Région Littorale”: BMNH 1946.8.27.97 [syntype, originally BMNH 1896.6.9.4; Fig. 16].

Bocage (1867a) initially assigned material from coastal Angola (three specimens collected by Anchieta in “Dombe”, “Benguella” and “Catumbella”) to Agama aculeata. In the years that followed he received additional material from this region and started to note differences between these specimens and those from the Angolan hinterland, presenting a specimen from Catumbela to the British Museum in 1893 identified as “Agama sp. ?” (BMNH 1893.12.27.7). Bocage (1895a) recorded this distinction some years later in his major work on Angolan herpetofauna, noting specimens sent by Anchieta from “Benguella”, “Catumbella” and “Dombe”, and by Hermenegildo Capello (1841–1917) and Roberto Ivens (1850–1898) from “Mossamedes” to differ from those from the high plateaus of the Angolan hinterland, which he assigned to Agama armata (see Agama cf. aculeataaculeata account above). Although he noted the coastal specimens as distinct in a note following the Agama armata account, Bocage (1895a) refrained from naming this material at the time. On 3 April 1896, Bocage wrote to Boulenger asking for a specimen of Agama aculeata to compare with his material from coastal Angola (NHMA/DF/ZOO/235/1/1/1/76), to which Boulenger conceded by sending to Lisbon an adult male from Beaufort West, South Africa (AHMB/CE/B48). Shortly after, in a letter dated 2 June 1896, Bocage mentioned a shipment of ground agamas to the British Museum containing three samples, including an adult from Angola, “Région Littorale” (No. 2), that he believed to represent a new species (NHMA/DF/ZOO/235/1/1/1/76; Fig. 3). Later that year, Bocage (1896b) formally described the populations from coastal Angola as Agama anchietae, based on the previously mentioned material collected by Anchieta in “Benguella”, “Catumbella” and “Dombe”, and by Capello and Ivens in “Mossamedes”.

In a revision of the Agama hispida and atra groups, Boulenger and Power (1921) recognized Agama anchieta as a valid species. The authors mentioned two specimens sent by Bocage to the British Museum, referring to a specimen from Caconda as “one of the types”. Marques et al. (2018) noted that this could not be a type, as Bocage (1895a, 1896b) never recorded the species from Caconda, and instead referred his material from that locality, in the highlands of the Angolan plateau, to Agama armata. Although it is impossible to state with certainty what led Boulenger and Power (1921) to record the type specimen as originating from Caconda, we are confident that this was done in error, as Bocage never mentioned this locality in his letters, nor is it stated in the original specimen label or the register. The locality “Caconda, Coast of Angola” appears only on the current label on the exterior of the specimen jar, which is a subsequent label presumably added when the specimen was reregistered as type in 1946, perpetuating Boulenger and Power’s (1921) error. While this is merely speculation, the source of the error may have been a misplaced label or confusion with other Agama specimens from Caconda sent by Bocage years earlier (see Agama cf. aculeataaculeata account). Considering this, we hereby correct the locality of the only surviving syntype of Agama anchietae (BMNH 1946.8.27.97; Fig. 16) to “Région Littorale, Angola”, corresponding to one of the coastal localities stated in the original description: Benguela, Catumbela, Dombe or Moçâmedes.

Angola: Duque de Bragança: BMNH 1866.6.11.2, Caconda: BMNH 1893.12.27.3, Quindumbo: BMNH 1893.12.27.4.

Two adult males (BMNH 1866.6.11.2, BMNH 1893.12.27.4) and one female (BMNH 1866.6.11.2) were sent from the Lisbon Museum. The specimen from Duque de Bragança was sent in 1866 as “Agama sp. ?” (NHMA/DF/GüntherColl/16/1/110) and was noted by Günther as “closely allied to Agama occipitalis, but has somewhat smaller scales” (AHMB/CE/G79), leading Bocage (1866a) to conservatively refer additional material collected by Bayão to “Agama occipitalis Gray. var. ?”. This specimen was later cited by Boulenger (1885a) as Agama planiceps. Butler (2020) validated the specific status of Agama schacki and identified cryptic diversity associated with this taxon. While the specimen from Quindumbo likely represents typical Agama schacki, those from Caconda and Duque de Bragança belong to two additional distinct lineages (Butler 2020).

Portugal: “Portugal”: BMNH 1864.9.19.38.

This specimen was among the first material presented to the British Museum in 1864, under the name Amphisbaena cinerea, and was cited by Boulenger (1885b) as Blanus cinereus. The genus Blanus is extremely conservative in external and internal morphology, and without a more precise locality it is, at present, impossible to determine to which of the two species present in Portugal this specimen belongs (Ceríaco and Bauer 2018; Villa et al. 2019).

Angola: Duque de Bragança: BMNH 1866.6.11.5–6.

Angolan material has been historically assigned to Chamaeleo gracilis and C. senegalensis (Marques et al. 2018). Bocage sent two specimens under the name Chamaeleo gracilis on 25 May 1866 (NHMA/DF/GüntherColl/16/1/110), and although Günther stated that “Chamaeleo gracilis is certainly not a distinct species” in a letter dated 29 June 1866 (AHMB/CE/G79), Bocage (1866a) referred his material from Duque de Bragança do this taxon, as did Boulenger (1887) when he cited the specimens presented to the British Museum.

Angola: Benguella: BMNH 1867.7.23.18.

Bocage (1867a) first recorded several specimens collected by Anchieta in Benguela, Catumbela and Dombe as Homodactylus bibroni, and presented a specimen to the British Museum on 13 July 1867 (NHMA/DF/ZOO/200/1/195). Boulenger (1885a) cited the specimen as Pachydactylus bibronii and was followed by Bocage (1895a).

Genus Hemidactylus Oken, 1817

Cabo Verde: “St. Iago”: BMNH 1875.4.26.10 (not examined) [putative syntype of Hemidactylus cessacii Bocage, 1873].

Bocage (1873a) described Hemidactylus cessacii based on three specimens (erroneously stated as a single specimen by Arnold et al. 2008) collected by Léon de Cessac (1841–1891) from Santiago Island in Cabo Verde. Bocage presented one specimen to the British Museum in 1875, together with additional duplicates of species described by him (NHMA/DF/ZOO/200/1/191). Boulenger (1885a) referred it to the synonymy of Hemidactylus bouvieri and cited the specimen sent by Bocage “as typical of H. cessacii”. In a first revised taxonomy of the endemic Hemidactylus from the archipelago, Arnold et al. (2008) described a population from Fogo Island as Hemidactylus lopezjuradoi and followed previous authors in assigning material from Santiago Island to Hemidactylus bouvieri (Boulenger 1885a; Loveridge 1947). Although Arnold et al. (2008) examined and photographed (see fig. 5E in Arnold et al. 2008) the specimen sent by Bocage to the British Museum in 1875, the authors did not fully assess the status of Hemidactylus cessacii and failed to recognize the possibility of surviving syntypes. In a subsequent revision, Vasconcelos et al. (2020) hypothesized that the Hemidactylus population from Santiago Island is conspecific with H. lopezjuradoi or represents a distinct species, although the lack of modern material from Santiago precludes further conclusions. Considering the pattern of local endemism observed on the reptile fauna from the Cabo Verde archipelago (Miralles et al. 2010; Vasconcelos et al. 2012, 2020), it is plausible that the Hemidactylus population from Santiago represents a distinct endemic species, for which the name cessacii would be available. On the other hand, if material from Santiago is conspecific with H. lopezjuradoi from neighboring Fogo Island, the nomen cessacii would represent a senior synonym and have priority. In any case, further research is needed to establish the status of Hemidactylus cessacii, and the specimen presented by Bocage to the British Museum may play a crucial role in this matter, as the only known surviving syntype. Although not recognized as a type by Arnold et al. (2008) or Vasconcelos et al. (2020), it is plausible to assume that the specimen was part of the type series, as no additional specimens were available to Bocage at that time, and it was presented in a shipment containing only duplicates of species described by Bocage (NHMA/DF/ZOO/200/1/191). Unfortunately, the specimen could not be located during our visits and remains unaccounted since last examined by Edwin Nicholas Arnold (1940–2023).

São Tomé and Príncipe: S. Tomé Island: BMNH 1893.12.27.1.

Hemidactylus greeffii was first described by Bocage in a paper written in Portuguese (Bocage 1886a), and a duplicate description written in French appeared immediately after in the same issue of the Jornal de Sciencias Mathematicas, Physicas e Naturaes (Bocage 1886b). The description was based on three specimens from “S. Tomé island” – one collected by Richard Greeff (1828–1892) and presented to the Lisbon Museum, and two from the Museu de Coimbra collected by Francisco Newton (1864–1909) (Bocage 1886a, 1886b). Although Bocage (1886a, 1886b) attributed the collection of the Coimbra specimens to Newton, this was most likely an error, as the now lost specimens were more likely to have been collected by Adolfo Moller (1842–1920) in 1885 (Lopes Vieira 1886). The specimen presented to the British Museum in 1893 was cited by Boulenger (1894a) and was likely a duplicate received at the Lisbon Museum after the original description, presumably collected by Newton, who explored the region from 1885 to 1895 (Ceríaco et al. 2022). Miller et al. (2012) reviewed its taxonomic status and designated a neotype from the California Academy of Sciences, San Francisco, USA.

Genus Lygodactylus Gray, 1864

Guinea-Bissau: Bissau: BMNH 1900386 [paralectotype, originally BMNH 1875.4.26.8; Fig. 17].

Bocage (1873a) described Hemidactylus gutturalis based on an unspecified number of specimens collected by Mr. Sá Nogueira (dates of birth and death unknown) at Bissau, sending a male to the ZMB in the same year (Bauer and Günther 1991) and another to the British Museum on 29 March 1875 (NHMA/DF/ZOO/200/1/191), cited by Boulenger (1885a) as “one of the types”. The specimen in Berlin (ZMB 7771) was recently designated as lectotype and the British Museum one as paralectotype (Lobón-Rovira et al. 2024).

Genus Pachydactylus Wiegmann, 1834

Angola: Benguella: BMNH 1867.7.23.17.

Bocage (1867a) reported several specimens of Pachydactylus ocellatus collected by Anchieta in Benguela. An adult male was presented to the British Museum in the same year (NHMA/DF/ZOO/200/1/195) and was later cited by Boulenger (1885a). Pachydactylus punctatus has long been recognized as a species complex with several distinct lineages occurring in Angola (Marques et al. 2018).

Cabo Verde: Ilheo Raso: BMNH 1875.4.26.11 (not examined) [putative paralectotype].

Bocage (1875) described Ascalabotes gigas based on an unspecified number of specimens collected by Francisco Frederico Hopffer from Ilheo Raso (= Raso Islet), and in the same year presented a duplicate to the British Museum (NHMA/DF/ZOO/200/1/191). Boulenger (1885a) reported the specimen sent by Bocage, although he did not note it to be a type. Another duplicate was presented by Bocage to the ZMB and was designated lectotype (ZMB 8998) by Bauer and Günther (1991). Although not noted by Boulenger (1885a), it is plausible to assume that the specimen presented to the British Museum was part of the type series, considering that those collected by Hopffer in 1874 were the only specimens available to Bocage at the time (Bocage 1896a), and all specimens sent in the 1875 shipment are regarded as types. The putative paralectotype could not be located during our visits.

Angola: Duque de Bragança: BMNH 1946.9.3.47–48 [syntypes, originally BMNH 1866.6.11.3–4; Figs 18, 19].

Bocage received several specimens of this lacertid collected by Bayão from Duque de Bragança, two of which he sent to the British Museum under the name Ichnotropis bivittatus on 25 May 1866 (NHMA/DF/GüntherColl/16/1/110). In a letter dated 19 June 1866, Günther refers the specimens to “Algira (Tropidosaura) dumerilii” after comparing them to the type of that species (AHMB/CE/G79). Although Bocage did not agree with this decision (NHMA/DF/ZOO/200/1/185), he followed Günther’s opinion and referred his specimens to Ichnotropis dumerilii instead of describing a new species, although he still included the new name in his account (“Tropidosaura Dumerilii. Smith. Ichnotropis bivittatus. Nob. Mss.”, Bocage 1866a). Bocage also sent a specimen to Peters at the ZMB, who argued for the validity of Bocage’s bivittata (Peters 1882). Boulenger (1887) first cited the British Museum specimens as types of bivittata under the synonymy of Ichnotropis capensis, only later recognizing it as a valid species (Boulenger 1921). The two specimens in the British Museum and the one in Berlin (ZMB 5827) are the only surviving syntypes of Ichnotropis bivittata (Bauer and Günther 1995). Syntype BMNH 1946.9.3.48 is missing its right arm.

Cabo Verde: Ilheo Raso: BMNH 1946.8.18.43 [syntype of Euprepes hopfferi Bocage, 1875, originally BMNH 1875.4.26.9; Fig. 20].

Bocage wrote to Günther and O’Shaughnessy in 1875 asking for specimens of the recently described Euprepes fogoensis, leading to an exchange of specimens where Bocage sent a shipment with duplicates of species described by himself from Cabo Verde and Guinea Bissau (NHMA/DF/ZOO/235/1/1/1/76, NHMA/DF/ZOO/200/1/191, 194). Among these specimens was a syntype of Euprepes hopfferi, described in the same year based on several specimens collected by Hopffer at “Ilheo Raso” (Bocage 1875). This specimen was cited as “one of the types of Euprepes hopfferi” by Boulenger (1887) under the synonymy of Mabuia stangeri and was recently examined by Miralles et al. (2010). Another syntype (ZMB 8999) of Euprepes hopfferi exists in the ZMB (Bauer et al. 2003).

Genus Eumecia Bocage, 1870

Angola: Caconda: BMNH 1893.12.27.8.

Although not explicitly stated, Bocage (1870) seemingly described Eumecia anchietae based on a single specimen collected by Anchieta “sur le plateau de la Huilla”. Years later Bocage (1895a) reported additional material, all specimens having been collected by Anchieta at Huíla, Caconda and Galanga. One specimen from Caconda presented to the British Museum in 1893, reported by Boulenger (1894a) as Lygosoma anchietae, may be the only surviving specimen examined by Bocage. Curiously, the specimen bears an original label identifying it as Dumerilia bayonii [= Sepsina bayonii (Bocage, 1866)].

Genus Mochlus Günther, 1864

Angola: Benguella: BMNH 1867.7.23.19.

Bocage (1867a) first recorded this species as Mochlus afer and later as Lygosoma sundevallii (Bocage 1895a), noting specimens collected by Anchieta in Benguela and other localities. The specimen was presented to the British Museum in 1867 as “Euprepes (Eumeces) afer” (NHMA/DF/ZOO/200/1/195) and was erroneously accessioned in the register as Euprepes binotatus. The following specimen in the register (BMNH 1867.7.23.20) is listed as Mochlus afer and could not be located, suggesting that a cataloguing error resulted in a duplication and swapping of data, as only one specimen of Mochlus was presented by Bocage (NHMA/DF/ZOO/200/1/195) and cited by Boulenger (1887).

Genus Trachylepis Fitzinger, 1843

Angola: Duque de Bragança: BMNH 1946.8.19.13 [syntype, originally BMNH 1866.6.11.8; Fig. 21].

Bocage (1866a) first recorded this skink, collected by Bayão at Duque de Bragança, as Euprepes gravenhorstii based solely on Dúmeril and Bibron’s (1839) description of the species, and later sent specimens to Auguste Duméril (1812–1870) and Peters to compare with typical material of that species. Both authors agreed that Bocage’s specimens were specifically distinct, leading Bocage (1872) to describe them as Euprepes bayonii. Bocage confirmed these observations with Boulenger after examining specimens from Madagascar (Bocage 1895a). Boulenger (1887) cited the specimen presented to the British Museum but did not note it as a type. Nevertheless, this specimen and another presented by Bocage to the ZMB (ZMB 6477) have been recognized as syntypes of Euprepes bayonii (Bauer et al. 2003; Marques et al. 2018; Ceríaco et al. 2024). The specimen in the British Museum was presented on 25 May 1866 identified as Euprepes gravenhorstii (NHMA/DF/GüntherColl/16/1/110), and the one in Berlin was likely presented in the same period, although no date in the ZMB catalogue confirms this. While these specimens were presented years before Bocage (1872) formally described Euprepes bayonii, the clear reference to the material he had previously referred to Euprepes gravenhorstii (Bocage 1866a) in the description suggests that Bocage considered his observations of these specimens for the diagnosis of the new species and thus constitutes sufficient evidence to recognize them as part of the type series.

Angola: Benguella: BMNH 1946.8.15.37 [syntype, originally BMNH 1867.7.23.26; Fig. 22].

Bocage (1867a) first mentioned several specimens collected by Anchieta in Benguela, Dombe and Catumbela as Euprepes binotatus. A detailed description, however, was only provided in a subsequent paper in the same issue of the Jornal de Sciencias Mathematicas, Physicas e Naturaes, where Bocage mentioned only Benguela (Bocage 1867c). Nevertheless, all localities mentioned in the first paper are relatively close to each other in coastal Benguela Province and the specimens on which Bocage based his descriptions were most likely the same, hence most authors have considered Benguela, Dombe and Catumbela to be type localities (Marques et al. 2018; Ceríaco et al. 2024). Shortly before the publication of the formal description in August 1867, Bocage presented a specimen to the British Museum on 13 July as “Euprepes binotatus Bocage n. sp.” (NHMA/DF/ZOO/200/1/195). The specimen entry is crossed over in the register, most likely due to a cataloguing error (see Mochlus sundevallii account). Additional surviving syntypes from Catumbela and Benguela exist in the collections of the ZMB (ZMB 5830) and MNHN (MNHN 1462), respectively (Brygoo 1985; Bauer et al. 2003; Ceríaco et al. 2024). Boulenger (1887) did not recognize binotata as valid, citing the British Museum type under Mabuia quinquetaeniata.

Angola: Duque de Bragança: BMNH 1946.8.15.27 [syntype, originally BMNH 1866.6.11.7; Fig. 23].

Bocage (1866a) recorded several specimens collected by Bayão from Duque de Bragança as Euprepes quinquetaeniatus and promptly sent duplicates to several European museums (Brygoo 1985; Bauer et al. 2003; Marques et al. 2018; Ceríaco et al. 2024). After comparing his specimens with typical material of quinquetaeniatus, Bocage (1872) described the specimens from Duque de Bragança as Euprepes petersi. However, the name was preoccupied by Euprepes petersi Steindachner, 1867, so Boulenger (1887) provided Mabuia bocagii as a replacement name, including a new description and citing a specimen sent by Bocage to the British Museum, in addition to two other specimens collected by Welwitsch. The specimen was presented to the British Museum in 1866 as Euprepes quinquaetaeniatus (NHMA/DF/GüntherColl/16/1/110) and has been considered a potential syntype of Euprepes petersi, together with specimens presented to the ZMB (ZMB 6479) and MNHN (MNHN 1286 and 1286a) (Brygoo 1985; Bauer et al. 2003; Marques et al. 2018; Ceríaco et al. 2024). Although it was presented to the British Museum years before the formal description was published, Bocage (1872) explicitly mentioned the specimens he had previously identified as Euprepes quinquetaeniatus as representing the new species, thus providing compelling evidence that he considered them part of the type series.

Angola: Caconda: BMNH 1882.6.9.5 (not examined).

Bocage and Günther exchanged letters in 1882 (NHMA/DF/ZOO/200/21/39, 41, 42), when Bocage was preparing his revision of the genus Philothamnus (Bocage 1882a). In this context, Bocage sent a shipment of Philothamnus specimens to the British Museum, including a specimen from Caconda under the name Philothamnus irregularis (NHMA/DF/ZOO/200/21/42). Although Bocage (1882a) did not cite specimens from Caconda in his generic revision, he later mentioned material collected by Anchieta at this locality (Bocage 1895a). The specimen presented to the British Museum was later reported by Boulenger (1894b) as Chlorophis irregularis, although subsequent works suggest that historical records of Philothamnus irregularis from Angola are referrable to either P. angolensis or P. hoplogaster (Hughes 1985; Marques et al. 2018). The specimen presented by Bocage could not be located during our visits, thus precluding confirmation of its identity.

Angola: Benguella: BMNH 1867.7.23.21, “Angola” [= Luanda]: BMNH 1882.6.9.2.

Bocage (1866b) described Leptophis dorsalis based on one specimen collected by Bayão at Duque de Bragança and another by Anchieta at Molembo, although Bocage (1882a) later designated the latter as the type and did not mention the specimen from Duque de Bragança again (Bocage 1882a, 1895a). Bocage later reported additional material, including specimens collected by Anchieta at Benguela and by João Osmundo Toulson (1832–1873) at Loanda (Bocage 1867a, 1882a). Although specimen BMNH 1882.6.9.2 is accessioned and cited only with the locality “Angola” (Günther 1868a; Boulenger 1894b), the original specimen label stating the name of the collector – Toulson – allowed us to infer Luanda as the specific locality (Bocage 1867a, 1895a). It is unclear how many specimens Toulson collected in Luanda, but the fact that Bocage (1882a) did not mention this material suggests that a single specimen was sent by Toulson in 1867 and subsequently presented to the British Museum (Bocage 1867a). The specimens sent to the British Museum were cited as Ahaethulla dorsalis by Günther (1868a) and Philothamnus dorsalis by Boulenger (1894b).

Equatorial Guinea: “Ile d’Anno-Bom, Golfe de Guinée”: BMNH 1946.1.6.3 [syntype, originally BMNH 1893.12.27.18; Fig. 24].

Bocage (1893b, 1893c) briefly described Philothamnus girardi based on eight specimens collected in 1892 by Francisco Newton on Anno-Bom Island and promptly presented a duplicate to the British Museum in the same year. This specimen was cited by Boulenger (1894b) as “one of the types” and by Loveridge (1958) as a “cotype”. Hughes (1985) suggested it was one of three surviving syntypes “and should be considered a neotype”. Although Wallach et al. (2014) considered this specimen to be a lectotype, Hughes’ (1985) statement does not constitute a lectotype designation compliant with Article 74.5 of the Code, and therefore BMNH 1946.1.6.3 should retain the status of syntype. It is unclear if additional syntypes were sent to other European museums, making the specimen in the British Museum the only known surviving type.

Angola: “Angola” [= Duque de Bragança]: BMNH 1882.6.9.3.

In preparation for a work on the Philothamnus in the Lisbon Museum, Bocage presented several specimens of this genus to the British Museum (NHMA/DF/ZOO/200/21/42). Although the only Philothamnus heterolepidotus mentioned in the letter was a specimen to be returned to the British Museum that Bocage had requested for examination, a specimen from “Angola” was accessioned with the remaining Philothamnus presented in this shipment. The specific locality Duque de Bragança can be inferred from the original specimen label that states the date and name of the collector – 1864, Bayão –, as recorded by Bocage (1882a). This specimen was cited by Boulenger (1894b) as Chlorophis heterolepidotus.

Mozambique: “Moçambique”: BMNH 1882.6.9.7, Angola: Humbe: BMNH 1946.1.5.98 [syntype of Philothamnus smithii Bocage, 1882, originally BMNH 1882.6.9.6; Fig. 25].

In his revision of the genus, Bocage (1882a) described Philothamnus smithii based on several specimens from Angola and Portuguese Guinea, including a specimen collected by Anchieta in 1876 at Humbe. A specimen from Humbe was presented to the British Museum in the same year under the name Philothamnus smithii (NHMA/DF/ZOO/200/21/42) and cited “as one of the types” by Boulenger (1984b). In the same shipment, Bocage presented a specimen from Mozambique under the name Philothamnus punctatus (NHMA/DF/ZOO/200/21/42). Bocage (1882a) noted specimens from Mozambique collected in 1868 by Canto and Valdez, and in 1869 by Cabral (dates of birth and death unknown), although without providing specific collecting localities. Specimen BMNH 1882.6.9.7 bears only the locality “Moçambique” and the year 1868, suggesting it was collected by Canto and Valdez. Both specimens presented to the British Museum in 1882 were cited by Boulenger (1894b) as Philothamnus semivariegatus.

S. Tomé and Príncipe: “Isle de S. Thomé”: BMNH 1946.1.21.60 [syntype, originally BMNH 1882.6.9.4; Fig. 26], BMNH 1893.12.27.17.

Bocage (1882a, 1882b) described Philothamnus thomensis based on five specimens from S. Tomé Island. One specimen was presented to the British Museum in the same year (NHMA/DF/ZOO/200/21/42), and another in 1893. Boulenger (1894b) cited the two specimens sent by Bocage, noting one to be “one of the types”. Hughes (1985) argued that BMNH 1946.1.21.60 corresponded to one of the adults mentioned by Bocage (1882a, 1882b) and “it would be wise” to designate it as lectotype. Although Wallach et al. (2014) regarded this statement as a lectotype designation, Hughes (1985) did not take explicit action complying with Article 74.5 of the Code, and therefore BMNH 1946.1.21.60 should retain the status of syntype rather than lectotype. This may be the only surviving type, although it remains unclear if additional specimens exist in other collections.

Genus Zamenis Wagler, 1830

Portugal: Alfeite: BMNH 1893.12.27.14, Aldegallega [i.e., Aldeia Galega = Montijo]: BMNH 1893.12.27.15, Coimbra: BMNH 1893.12.27.16.

In preparation for his second volume of the Catalogue of Snakes (Boulenger 1894b), Boulenger wrote to Bocage on 6 November 1893 (AHMB/CE/B43) asking for specimens of Rhinechis scalaris. Bocage presented three specimens from Portugal in the same year (NHMA/DF/ZOO/235/1/1/1/76), which were cited as Coluber scalaris by Boulenger (1894b). Notably, no specimens from Alfeite or Montijo exist today in Portuguese collections (Santos et al. 2024).

Angola: Benguella: BMNH 1867.7.23.23 [paralectotype of Alopecion variegatum Bocage, 1867 and Boodon lineatus var. lineolata Bocage, 1895; Fig. 27].

Bocage (1867c) described Alopecion variegatum based on three specimens from Benguela collected by Anchieta and one from Novo Redondo collected by J. A. Botelho (dates of birth and death unknown) (Bocage 1895a), and in the same year presented a specimen from Benguela to the British Museum (AHMB/CE/G81, NHMA/DF/ZOO/200/1/195, NHMA/DF/GüntherColl/16/1/112). Günther (1868a) likely did not consider this a valid species, as he did not include it in the new additions to the collection. Boulenger (1893) cited the specimen sent to the British Museum as “one of the types” of Alopecion variegatum, under the synonymy of Boodon lineatus, and Bocage (1895a) later referred his specimens from coastal Angola to a new variety Boodon lineatus var. lineolata. In a recent revision of the Boaedon fuliginosus complex in Angola, Hallermann et al. (2020) validated the specific status of Boaedon variegatus and designated the British Museum specimen as paralectotype of both Alopecion variegatum and Boodon lineatus var. lineolata. Another specimen from the original type series survives in the ZMB (ZMB 6469) and was designated lectotype (Hallerman et al. 2020).

Genus Lycophidion Fitzinger, 1843

Angola: Duque de Bragança: BMNH 1864.10.28.15, Caconda: BMNH 1893.12.27.13.

Among the specimens collected by Bayão at Duque de Bragança and sent by Bocage to the British Museum in 1864, Günther identified No. 3 as Lycophidion horstockii (AHMB/CE/G76). This specimen was presented to the British Museum and is still identified with the No. 3 in the original label. Bocage (1966a) followed Günther’s opinion and recorded seven specimens collected by Bayão at Duque de Bragança as Lycophidion horstockii var. A. Boulenger (1893) referred Bocage’s specimen from Duque de Bragança to Lycophidion capense var. B and was followed by Bocage (1895a). A second specimen, collected by Anchieta in Caconda, was presented to the British Museum in 1893 identified as Lycophidion capense var. multimaculata (Bocage 1895a; Boulenger 1896). Donald Broadley (1932–2016) examined the specimens in London and Lisbon in 1968 and cited them in his revision of the genus Lycophidion (Broadley 1996).

Angola: Duque de Bragança: BMNH 1946.1.14.53–54 (not examined) [syntypes, originally BMNH 1864.10.28.16; Figs 28, 29].

Among the first shipment of specimens collected by Bayão at Duque de Bragança and subsequently sent by Bocage to the British Museum for identification, two specimens were used by Günther (1865b) to describe a new genus and species, Limnophis bicolor. Since Bocage had additional specimens sent by Bayão (Bocage 1866a), he presented the types to the British Museum (Boulenger 1893; Bocage 1895; AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104). Although the types were recently examined and photographed for a revision of the genus (Conradie et al. 2020), we were unable to locate and examine them during our visits.

Angola: “W. Africa” [= Duque de Bragança]: BMNH 1875.5.22.4.

On 25 May 1866 Bocage sent a young snake that he could not identify, listed as “No. 7 – Coronella ? jeune ? – Duque de Bragança” (NHMA/DF/GüntherColl/16/1/110). In response, Günther identified it as “the young of some species of Psammophis”, adding that “it may be new, but it is not advisable to describe a new species from such a young specimen” (AHMB/CE/G79). Although the specimen was “envoyés en communication” and should have been returned to Lisbon, it was later accessioned in 1875 as “Coluber” from “W. Africa” with the note “received some years ago from the Lisbon Museum for examination”, and still bears the original label with the number 7. Although Bocage (1895a) never cited any Psammophis from Duque de Bragança, this may have been because neither he nor Günther could initially identify the specimen to specific level, and it had already been presented to the British Museum when Bocage received additional comparative material that allowed him to describe new varieties to allocate his specimens (Bocage 1887b, 1895a). Bocage (1887b) described Psammophis sibilans var. leopardinus based on a juvenile collected by Capello and Ivens from “l’intérieur de Mossamedes” and an adult collected by Anchieta in Catumbela, and later reassigned this material to what he called Psammophis sibilans var. C (Bocage 1895a). While Boulenger (1896) considered Bocage’s var. leopardina to be a synonym of Psammophis sibilans, he referred the specimen sent years earlier by Bocage to Psammophis brevirostris, a determination that remained associated with the specimen until our revision. The specimen fits the description of Psammophis leopardinus and could only be confused with Psammophis zambiensis Hughes & Wade, 2002, which is also known from Angola and could be expected to occur in the region of Duque de Bragança (Broadley 2002; Hughes and Wade 2002; Marques et al. 2018; Trape et al. 2019). However, it can be readily distinguished from P. zambiensis based on color pattern, with pale crossbars on the back of the head, dorsum with a “chain-like” pattern on its anterior portion and uniform brown posteriorly, and a uniform cream ventrum (versus a greenish brown dorsum with more extensive “chain-like” pattern, and ventral scales irregularly edged with black on their free margin in P. zambiensis). This record represents an extension of about 300 km from the northernmost confirmed records of the species into inland Angola, defying the assumption that Psammophis leopardinus is restricted to arid savannas and semi-desert (Broadley 2002; Trape et al. 2019). The fact that none of the authors who dealt with the Psammophis sibilans complex cited this specimen (Broadley 2002; Hughes and Wade 2002; Trape et al. 2019) could probably be explained by the vague locality (i.e., “West Africa”) stated on the British Museum records, precluding a confident identification in such a taxonomically challenging group.

Angola: Benguella: BMNH 1867.7.23.22 [holotype of Psammophis bocagii Boulenger, 1895; Fig. 30].

This specimen was collected by Anchieta in Benguela, which Bocage (1867a) identified as Psammophis elegans and presented to the British Museum in 1867 under that name (NHMA/DF/ZOO/200/1/195). It was later used by Boulenger to describe Psammophis bocagii, first introduced with a brief description and only the locality “Angola” in a key to the genus Psammophis (Boulenger 1895). Boulenger (1896) later provided a detailed description of Psammophis bocagii, citing the specimen from Benguela sent by Bocage. Although ranges of ventrals and subcaudals were provided by Boulenger (1895, 1896), these were derived from the data provided by Bocage (1895a) for his Psammophis sibilans var. A, as listed in Boulenger’s (1896) chresonymy. The specimen was examined by Broadley (1977a, 2002).

Genus Psammophylax Fitzinger, 1843

Angola: Humbe: BMNH 1882.6.9.1.

Bocage (1873a) described Psammophylax ocellatus based on a single specimen collected by Anchieta from “l’intérieur de Mossamedes (Gambos)”, and later reported additional specimens collected by Anchieta in Humbe but referred this material to Psammphylax rhombeatus (Bocage 1895a). One of these specimens was presented to the British Museum in 1882 as Psammophylax ocellatus (NHMA/DF/ZOO/200/1/195). Even though Günther thought that “Psammophylax ocellatus seems to be an excellent species” (AHMB/CE/G94), Boulenger (1896) reported the specimen as Trimerorhinus rhombeatus. Broadley examined the specimen in 1968 and recovered ocellatus as a valid subspecies (Broadley 1977b). While recent material collected in the Humpata Plateau validate the specific status of Psammophylax ocellatus (Branch et al. 2019), the specimen sent by Bocage to the British Museum remains among the few historical records of the species from Cunene Province.

Angola: Caconda: BMNH 1893.12.27.19 [putative syntype of Ophirhina anchietae Bocage, 1882; Fig. 31].

Bocage (1882b) described Ophirhina anchietae based on an unspecified number of specimens collected by Anchieta in Caconda, although implying that more than one specimen was available. A juvenile specimen was presented to the British Museum in 1893, and Boulenger wrote to Bocage on 29 March 1894 stating that he thought Ophirhina anchietae to be a synonym of Pseudaspis cana (AHMB/CE/B44; Boulenger 1896). Bocage agreed with Boulenger’s opinion on a letter dated 3 April 1894 (NHMA/DF/ZOO/235/1/1/1/76) and later referred his material to this species (Bocage 1895a). Although not noted by Boulenger (1896) as a type, the type series included at least one juvenile (Bocage 1882b, 1895a) and the specimen presented to the British Museum is topotypical, raising the possibility that it was part of the type series of Ophirhina anchietae.

Guinea-Bissau: Bissao: BMNH 1867.7.23.16.

Bocage (1895a) considered three varieties of Naja nigricollis and coined var. occidentalis for some of his specimens from Angola and Guinea-Bissau. Bocage (1895a) mentioned only a juvenile collected by Barahona (presumably Henrique César da Silva Barahona e Costa, dates of birth and death unknown) in Bissau, even though he had presented an adult to the British Museum years earlier under the name Naja nigricollis (NHMA/DF/ZOO/200/1/195), that Boulenger (1896) assigned to the “forma typica”.

Angola: Quindumbo: BMNH 1893.12.27.11.

Bocage (1873b) described Onychocephalus anomalus based on three specimens collected by Anchieta in Huíla and one sent by Joaquim José Graça (1825–1889) without stating a specific locality, but later wrote that the original description had been made based on specimens from Bibala and Huíla (Bocage 1895a). Additional material received subsequently included specimens from Quindumbo, one of which was presented to the British Museum in 1893 (Boulenger 1896).

Angola: Quindumbo: BMNH 1946.1.11.18 [lectotype of Typhlops boulengeri Bocage, 1893, originally BMNH 1893.12.27.12; Fig. 32].

Bocage (1893a) described Typhlops boulengeri based on an unspecified number of specimens collected by Anchieta at Quindumbo, one of which he presented to the British Museum in the same year (Boulenger 1896). Broadley and Wallach (2009) designated the British Museum specimen as lectotype of Typhlops boulengeri, which the authors considered a synonym of Afrotyphlops lineolatus.

Angola: Quissange: BMNH 1893.12.27.9–10.

Bocage (1886c) described Typhlops (Onychocephalus) humbo based on two specimens collected by Anchieta in Quissange and later mentioned additional material from the same locality and collector sent in 1890 (Bocage 1895a). Bocage presented two specimens of Typhlops from Quissange to the British Museum in 1893, one of which was reported by Boulenger (1896) as Typhlops humbo and the other as Thyphlops mucruso. None of the specimens were considered by Boulenger (1896) to be types, suggesting they were part of the material sent by Anchieta in 1890. Broadley and Wallach (2009) referred Bocage’s humbo to the synonym of Afrotyphlops schlegellii, including the two specimens from Quissange presented by Bocage to the British Museum. Although further research is needed to stabilize the taxonomy of the Afrotyphlops schlegellii species complex, some authors recognize petersii as a subspecies from Angola and Namibia (Marques et al. 2024).