Echoes of a lost museum: Revision of the herpetological collections sent by Barbosa du Bocage from the Lisbon Museum to the British Museum of Natural History

353 Echoes of a lost museum: Revision of the herpetological collections sent by Barbosa du Bocage from the Lisbon Museum to the British Museum of Natural History Diogo Parrinha1,2,3, Francisco M. G. Calado4, Mariana P. Marques5, Aaron M. Bauer6, Luis M. P. Ceríaco1,2 1 BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485-661 Vairão, Portugal 2 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Portugal 3 Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, Rua do Campo Alegre 1021, 4169-007 Porto, Portugal 4 Aquário Vasco da Gama, R. Direita do Dafundo, 1495-718 Cruz Quebrada, Portugal 5 Section of Amphibians and Reptiles, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213-4080, USA 6 Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA https://zoobank.org/22DBAEFB-4690-47FD-9259-98013D7BF8CB Corresponding author: Diogo Parrinha ([email protected]) Academic editor Uwe Fritz | Received 26 August 2025 | Accepted 2 October 2025 | Published 22 October 2025 Citation: Parrinha D, Calado FMG, Marques MP, Bauer AM, Ceríaco LMP (2025) Echoes of a lost museum: Revision of the herpetological collections sent by Barbosa du Bocage from the Lisbon Museum to the British Museum of Natural History. Vertebrate Zoology 75: 353–404. https:// doi.org/10.3897/vz.75.e169790 Abstract As part of a nineteenth century scientific network, José Vicente Barbosa du Bocage regularly sent “duplicate” specimens from the zoological collections of the National Museum of Lisbon to natural history museums across Europe. These duplicates gained exceptional significance following the 1978 fire that destroyed the Lisbon Museum’s zoological collections, making them the last surviving representatives of its historical holdings. Despite their importance for taxonomic and nomenclatural stability, the full extent of Bocage’s duplicate specimens remains poorly documented. Here we present a comprehensive and integrative revision of the herpetological material sent by Bocage to the British Museum of Natural History. We assess its historical, taxonomic and nomenclatural value, providing an illustrated and annotated catalogue of type specimens. A total of 92 specimens representing 57 species were sent from Lisbon between 1864 and 1896, including 30 type specimens for 27 nominal taxa. We provide evidence for the correction of the type locality associated with the only surviving syntype of Agama anchietae, as well as the recognition of previously unknown types of Chioglossa lusitanica, Hylambates angolensis, Hylambates cynnamomeus, Cystignathus bocagii, Hyperolius insignis, Hyperolius huillensis, Hemidactylus cessacii and Ophirhina anchietae. Keywords Angola, Cabo Verde, Guinea-Bissau, history of science, nomenclature, taxonomy, type specimens Introduction The practice of exchanging specimens – often termed “duplicates” – with counterpart institutions was a common practice in most European and North American natural history museums in the nineteenth and early twentieth centuries (Cornish and Driver 2020; Nichols 2021; Kaiser 2022; Reese et al. 2024). This led to the diversification of collections and contributed to the dispersal of specific collections through several institutions, creating Vertebrate Zoology 75, 2025, 353–404 | DOI 10.3897/vz.75.e169790 Copyright Diogo Parrinha. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 354 important scientific networks and webs of knowledge. This practice had an unexpected but important outcome – it created “backup copies” of such collections deposited in several institutions, which become particularly relevant if the “original” collections were damaged or destroyed. Natural history museums are subject to a wide variety of catastrophic incidents that can lead to the total loss of their collections (Tyler et al. 2023). Unfortunately, such incidents continue to afflict museums in the present day, as exemplified in recent years by the destruction of collections by fire – Instituto Butantan (São Paulo, Brazil) in 2010, the National Museum of Natural History (New Delhi, India) in 2016 and the Museu Nacional (Rio de Janeiro, Brazil) in 2018 – or war – Assaray Al-Hamra Museums (Tripoli, Libya) in 2011. On 18 March 1978, a fire consumed Portugal’s National Museum of Natural History in Lisbon, especially affecting its Zoological Section – at the time known as Museu Bocage (present day Museu Nacional de História Natural e da Ciência) – completely destroying its collections (Fig. 1; Saldanha 1978; Ceríaco and Marques 2025). Among the lost specimens were the important series of material collected during the second half of the nineteenth century in the former Portuguese colonial territories in Africa, that have been used to document the fauna of those regions and describe dozens of new vertebrate taxa. Although any destruction of collections always represents an irreparable loss, these are particularly devastating when type specimens are amongst them. Due to the loss of type material, the taxonomic and nomenclatural status of many taxa described based on specimens from Museu Bocage remain problematic, especially those from Angola (see Marques et al. 2018). After the catastrophic fire, the specimens presented to other institutions in Portugal and abroad became the sole testament of these important collections that served as the basis for the descriptions of several taxa and provided important accounts of dozens of other species in poorly studied areas (Calado 2015). While specimens sent to other museums can help resolve these issues, the true extent of surviving material in foreign institutions remains unknown. The history and collections of Museu Bocage were deeply intertwined with that of its namesake. José Vicente Barbosa du Bocage (1823–1907; Fig. 2A) was a Portuguese zoologist who was appointed as director of the Zoological Section of the National Museum of Lisbon in 1858, and is considered as one of the country’s most influential zoologists (Madruga 2012, 2013, 2017; Gamito-Marques 2018, 2022; Ceríaco 2021). After inheriting the direction of a then moribund Museum which had been virtually abandoned during the late 1840s and 1850s, Bocage established a network of national and international collaborators which contributed to significantly expand its collections and regain scientific relevance among its European counterparts (Bocage 1862, 1865; Ceríaco 2021). Bocage meticulously studied and organized the zoological collections sent to the Lisbon Museum, publishing dozens of scientific papers and describing more than 200 species over nearly half a century of scientific activity (França 1908; Osório 1909; Madruga 2013; Calado 2015; Gamito-Marques 2018, 2022; Ceríaco 2021). Figure 1. The aftermath of the fire that destroyed Museu Bocage on 18 March 1978. Photo from Arquivo Histórico Museu Bocage. Vertebrate Zoology 75, 2025, 353–404 355 While Bocage’s works addressed several zoological groups (invertebrates, fishes, mammals, birds, amphibians and reptiles) and geographic contexts (Europe, Africa and Oceania), his major contributions were to African herpetology. Particularly, Bocage dedicated much of his career to the study of Angolan herpetofauna, receiving specimens collected by Portuguese officials and explorers such as Francisco António Pinheiro Bayão (1833–1883) and José Alberto de Oliveira Anchieta (1832–1897), that served as the basis for the descriptions of many new species and his magnum opus “Herpetologie d’Angola et du Congo” (Bocage 1895a). Despite the focus on Angolan herpetofauna, Bocage also extensively studied the herpetofauna (and overall biodiversity) of the Gulf of Guinea Islands and published several important works on the herpetofauna of Cabo Verde and other African countries (Bocage 1875, 1886a, 1886b, 1893b, 1893c, 1896a, 1903; Ceríaco 2021; Ceríaco et al. 2022). Due to the establishment of his network of collaborators, Bocage started receiving specimens of the African herpetofauna in the first years of his scientific career (Madruga 2012; Ceríaco 2021). Despite his already established scientific competency, at that time he was still unfamiliar with the African herpetofauna. To help him with the task of studying and organizing the collections he received from Africa, Bocage reached out to colleagues in other European museums with whom he exchanged opinions, literature and specimens. Bocage is known to have sent herpetological material to several European museums, most notably the Zoologisches Museum in Berlin (currently Museum für Naturkunde), the Muséum national d’Histoire naturelle in Paris, the Naturhistorisches Museum in Wien and the British Museum of Natural History (currently the Natural History Museum, London) (Günther 1912; Calado 2015; Ceríaco 2021). Amongst the herpetology curators of these museums, Bocage maintained particularly regular and cordial correspondence with Wilhelm Peters (1815–1883) – curator and director of the Zoologisches Museum from 1857 to 1883, Albert Günther (1830–1914; Fig. 2B) – assistant and keeper of zoology Figure 2. Portraits of correspondents from the Lisbon Museum and the British Museum. A José Vicente Barbosa du Bocage; B Albert Günther; C Georges Albert Boulenger. Photos from the Natural History Museum Archives and Library, London (A–B) and National Portrait Gallery, London (C). Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 356 at the British Museum from 1862 to1895 and Georges Albert Boulenger (1858–1937; Fig. 2C) – assistant and head of the reptile and fish section at the British Museum from 1882 to 1920. Type material of species described by Bocage has been identified in the collections of the Museum für Naturkunde (Bauer and Günther 1991, 1995; Bauer et al. 1996, 2003, 2006; Wallach et al. 2014; Tillack et al. 2021), the Muséum national d’Histoire naturelle (Brygoo 1985) and the British Museum (Broadley and Wallach 2009; Wallach et al. 2014; Marques et al. 2018; Hallerman et al. 2020). However, to date no systematic effort has been made to identify the full extent of herpetological specimens sent by Bocage to international museums, where additional type material may remain unaccounted for. Here we provide a comprehensive revision of the herpetological material presented by Bocage to the British Museum, including a commented and illustrated catalogue of type specimens. Materials and Methods Sources of data In order to identify the full extent of herpetological material sent by Bocage to the British Museum (BMNH), we examined four main sources of data (Fig. 3): 1) the herpetology department register at the British Museum (Fig. 3A); 2) specimens and their associated labels (Fig. 3B, C); 3) published literature; 4) unpublished documents deposited in the Arquivo Histórico Museu Bocage Figure 3. Sources of data examined. A Register noting specimens presented by Bocage; B Specimen jar with external label; C Original specimen label with locality and number mentioned in Bocage’s letter; D Extract of Bocage’s letter citing numbered specimens (NHMA/DF/ZOO/235/1/1/1/76). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 357 (AHMB) at the Museu Nacional de História Natural e da Ciência in Lisbon, Portugal and the Natural History Museum Archives and Library (NHMA) in London, United Kingdom. We first compiled all records accessioned in the herpetology department register that noted Bocage and/or the Lisbon Museum as the origin of the specimen. Next, we searched for all the identified records in the herpetology collections and examined the specimens to confirm identification and update nomenclature. All located specimens and associated labels were photographed. The compiled records were cross-referenced with relevant literature and archival documentation to screen for potential citations of specimens and additional data. Relevant material from other museum collections is also mentioned when appropriate, namely the Museum für Naturkunde, Belin (ZMB), the Muséum national d’Histoire naturelle, Paris (MNHN), the Muséum d’Histoire naturelle, Genève (MHNG), the Iziko South African Museum, Cape Town (SAM) and the lost collections of Museu Bocage, Lisbon (MB). Although the preferred institutional acronym for citing Natural History Museum specimens is currently NHMUK, we retain the historical acronym BMNH here for consistency with previous literature and documentation (Sabaj 2020). To identify the specific details of shipments to the British Museum, we examined documents relating to the shipment of specimens from the Lisbon Museum between 1865 and 1925, available at the AHMB. To further investigate the metadata associated with the specimens and resolve problems associated with the recognition of type material, we examined available correspondence exchanged between Bocage and his peers from the British Museum (Fig. 3C), namely John Edward Gray (1800– 1875), Albert Günther and Georges Albert Boulenger, as well as other relevant correspondents like Arthur O’Shaughnessy (1844–1881), Joachim John Monteiro (1833–1878) and Francisco Frederico Hopffer (1828– 1919), deposited in the AHMB and the NHMA. On the concept of type specimen A major difficulty when dealing with historical specimens such as these lies in the interpretations of the concept of “type” for the original authors and the subsequent reviewers of such material. Additional difficulties arise from the fact that it was not uniform practice for authors to explicitly designate specific specimens as types, leading to taxonomic and nomenclatural problems that persist until today (Bauer and Günther 1995; Dubois and Ohler 1996; Wüster and Tillack 2023; Bauer et al. 2024; Dubois et al. 2024). The concept of name-bearing type and the rules surrounding its designation and fixation have considerably evolved over the last 150 years, transforming personal interpretations into “codified” versions of the concept, such as that currently defined by the International Code of Zoological Nomenclature (hereafter “the Code”) and its subsequent amendments (ICZN 1999). Before 1999, the explicit fixation of the type material – either a single holotype, or two or more syntypes – was not a mandatory step for a new name to become nomenclaturally available (ICZN 1999). Article 72 of the Code defines a type series as “all the specimens on which the author established a nominal species-group taxon (with the exception of those excluded [Art. 72.4.1]); in the absence of holotype designation, or the designation of syntypes, or the subsequent designation of a lectotype, all are syntypes and collectively they constitute the name-bearing type” (Article 72.1.1.), consisting of “all the specimens included by the author in the new nominal taxon (whether directly or by bibliographic reference), except any that the author expressly excludes from the type series [Art. 72.4.6], or refers to as distinct variants (e.g., by name, letter or number), or doubtfully attributes to the taxon” (Article 72.4.1). However, as noted above, authors have not always unambiguously fixed the type series, complicating the work of subsequent researchers when endeavoring to locate these specimens in collections. To address these difficulties, the Code notes that “For a nominal species or subspecies established before 2000, any evidence, published or unpublished, may be taken into account to determine what specimens constitute the type series” (Article 72.4.1.1). A glimpse of Bocage’s interpretation of what constituted a “type” can be found on two papers, where he presented catalogues of the “exemplares typicos” of birds (Bocage 1896c) and mammals, amphibians and reptiles (Bocage 1897) that existed in the Lisbon Museum. Except for unique types, Bocage (1896c, 1897) did not state the precise number of specimens but mentioned only the localities and collectors, sometimes separately providing a list of additional specimens that he would have received afterwards from other localities. In both papers, it can be inferred that Bocage considered as “exemplares typicos” not only those specimens used in the original descriptions, but also other topotypical material not originally mentioned. Such a case is evident in his account of Helicops bicolor – described by Günther (1865b) based on two specimens presented by Bocage –, where Bocage (1897) mentioned the two “types” in the British Museum but still considered the specimens in the Lisbon Museum as “exemplares typicos”. Similarly, Bocage (1895a) mentioned two “types” of Hylambates bocagii in the British Museum and considered “exemplares typicos” in Lisbon as well, even though Günther (1865a) explicitly mentioned a unique specimen in the original description. In other instances, Bocage (1897) included among the “exemplares typicos” localities not mentioned in the original description, which was the case of Mabuia petersii as noted by Ceríaco et al. (2024), further demonstrating inconsistency and Bocage’s lack of a clear concept of “type”. Another difficulty, besides the cases where Bocage explicitly mentioned a single specimen (or that could implicitly be assumed from the text of the description), arises from the fact that Bocage did not provide any kind of catalog number that could unambiguously identify the specimens he was referring to. While in some cases Bocage stated how many specimens were available to him – e.g., 18 specimens of Bufo dombensis (Bocage 1895b) –, most often there was no mention of a precise Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 358 number, but instead a vague reference to “several specimens” from a given locality and collector. Such vague information requires further investigation to reach an idea of the number of specimens involved (sometimes through the analysis of the field notes and letters from the original collectors) and the subsequent history of dispersal of these specimens. A conservative interpretation of type series would imply that only those specimens explicitly mentioned in the original description would constitute name-bearing types. A more lenient interpretation, however, can include specimens that, although not explicitly mentioned in the original description, were available to Bocage prior to the publication and for which there is evidence that Bocage had already recognized as belonging to the new taxon. A practical example of these were some of the specimens shipped by Bocage to the British Museum, in which both the original specimen labels and/or the letters sent by Bocage already noted his acknowledgement that they represented distinct taxa. For example, in the cases of Chioglossa lusitanica, Hyperolius insignis or Hyperolius huillensis, Bocage presented specimens under his yet unpublished nomina, even if the description would only be published months or even years later (see Results for further comments; Fig. 4; Table 1). Even though Bocage did not explicitly mention these specimens in the original publications, we accept that they were used by Bocage to develop his concept and diagnoses of the new species and thus can be regarded as part of the type series. This approach, however, is solely based on Bocage’s conceptualization and not in subsequent interpretations. While Boulenger (1882b, 1885a, 1887, 1893, 1894b, 1896) referred to several specimens presented by Bocage to the British Museum as “one of the types” or Figure 4. Extract of letter from Bocage to Günther dated 24 May 1869 (NHMA/DF/ZOO/ 200/1/189) mentioning specimens sent via R. B. Sharpe, including “Hyperolius huillensis nov. sp.” and “Bufo benguellensis nov. sp.”, years before the descriptions were published. Vertebrate Zoology 75, 2025, 353–404 359 “as typical”, there seems to be no consistency or logical reasoning in what he considered to be a “type” or “typical” specimen. A particular case of permissive and inconsistent interpretations is that of Perret (1976), who examined the types of African amphibians in the Lisbon Museum. Perret identified several specimens as “cotypes”, a term that is not recognized by the Code, but has been traditionally used to refer either to syntypes or paratypes. In many cases, however, the specimens identified by Perret as “cotypes” were clearly not part of the original type series examined by Bocage, either because they represented localities not mentioned in the original publications and only reported by Bocage subsequent to the descriptions or because Bocage unambiguously mentioned a unique specimen in the original description. To address the lack of Bocage’s concept of “type” and the inconsistencies in subsequent interpretations, we here accept as being part of the type series those specimens for which there is compelling evidence suggesting that they were regarded by Bocage as novel species at the time of shipment, in cases where the putative types were sent before the description was published. We also accept as putative types those specimens presented to the British Museum after the original description was published, in the absence of compelling evidence suggesting that they were collected and first examined by Bocage posterior to publication. Results No shipments of reptiles or amphibians to the British Museum were found in the records of shipments from the Lisbon Museum at AHMB. Notwithstanding, several documents confirm the shipment of birds (AHMB/ Div. 471, 486, 493, 504, 509, 510, 532.7, 532.10), mammals (AHMB/Div. 509, 510, 513, 518, 522, 532.3, 532.8, 532.18) and shells (AHMB/Div. 472) from Lisbon to England, addressed directly to the British Museum or to particular researchers, namely Oldfield Thomas (1858– 1929; mammals), George Edward Dobson (1848–1895; mammals), Knud Christian Andersen (1867–1918; mammals), Richard Bowdler Sharpe (1847–1909; birds) and George Ernest Shelley (1840–1910; birds). However, despite several lists of herpetological material shipped to other European museums (Calado 2015), we found only a single reference to reptiles sent to the British Museum among these documents. The document (AHMB/Div. 500) is titled “Reptis remettidos em comunicaçao ao Dr. Günther – pelo vapor Malange – em 10 de Nov. 1883” [Reptiles sent in communication to Dr. Günther – by the steamer Malange – on 10 Nov. 1883] and lists four species of geckos from New Caledonia: “Rhacodactylus leachianus, Cuv”, “Rhacodactylus aubryanus, Boc.”, “Ceratolophus auriculatus, Bavay.” and “Lepidodactylus crepuscularis, Bavay”. These specimens were sent on a loan for examination and were not incorporated in the British Museum collections (Boulenger 1883, 1885a). On the other hand, detailed accounts of the herpetological specimens presented to the British Museum were found in the correspondence exchanged between Bocage and his colleagues in London. Bocage would often dispatch shipments through steamer or by an intermediate colleague like R. B. Sharpe or J. J. Monteiro, detailing the contents of each shipment in letters addressed to Günther or Boulenger. In his letters Bocage usually presented a list of numbered specimens, which in turn were accompanied by a specimen label with the corresponding number (Figs 3–5), allowing the unambiguous correspondence between the physical specimen and the citations in correspondence. The register lists a total of 92 herpetological specimens presented by Bocage to the British Museum, accessioned in 13 different occasions from 1864 to 1896. Through examination of the exchanged correspondence, we were able trace these specimens to a total of 12 shipments that occurred between 24 May 1864 and 2 June 1896 (Table 1). The first shipment, presented on 24 May 1864 (NHMA/ DF/GüntherColl/16/1/102), was sent via J. J. Mon teiro and contained specimens from the Portuguese fauna (BMNH 1864.9.19.35–39), including specimens of Bocage’s yet undescribed Chioglossa lusitanica, accompanied by a manuscript with the description to be considered for publication in the Proceedings of the Zoological Society of London (Bocage 1864a). A second shipment was sent in the same year, on 28 July 1864 (NHMA/DF/ GüntherColl/16/1/104), containing the first specimens collected by Bayão at Duque de Bragança (currently Kalandula, Malanje Province, Angola). This shipment contained specimens numbered 1–23 for Günther to identify, noting Nos 1–3 and 20–23 as snakes and Nos 4–19 as amphibians, allowing Günther to keep Nos 1–10 and 19 as duplicates for the British Museum collections. Günther described three taxa based on this material – Limnophis bicolor, Hyperolius nasutus and Leptopelis bocagii – and returned the remaining identified specimens to Lisbon (AHMB/CE/G76; Fig. 5), keeping in the British Museum those presented by Bocage (BMNH 1864.10.28.1–16). An additional shipment of specimens from Duque de Bragança followed on 25 May 1866 (NHMA/DF/GüntherColl/16/1/110), in which Bocage listed specimens numbered 1–6 (BMNH 1866.6.11.1–8), including species he believed to be new – Stellio angolensis and Ichnotropis bivittatus. In addition, this shipment contained three specimens from the same locality “envoyés en communication”, including two frogs previously examined by Günther (Nos 11 and 12; Fig. 5) and a young snake that Bocage could not identify (No. 7). Although these specimens were supposed to be returned to Lisbon, they were later accessioned in the British Museum with the note “received some years ago from the Lisbon Museum for examination” (BMNH 1875.5.22.2–4). Over the following years Bocage continued to send duplicates mostly from Angola, often including species he believed to be new (Table 1). While some of these shipments contained an assortment of duplicates of different taxa and geographic contexts, others were sent to exchange opinions on specific taxa, like those sent in preparation for Bocage’s works Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 360 Table 1. Summary of specimens presented by Bocage to the British Museum. Shipment dates are inferred from Bocage’s letters and verbatim identifications are transcribed from the same letters or the original specimen labels. Type specimens are noted in bold and an asterisk (*) denotes material that could not be examined. Shipment date Original accession number [ current number] Verbatim identification Current identification Locality Published citations Archival citations 24 May 1864 BMNH 1864.9.19.35–37 “Chioglossa lusitanica”Chioglossa lusitanica Bocage, 1864 Coimbra, Portugal Boulenger (1882a: 6) NHMA/DF/GüntherColl/16/1/102, 103 BMNH 1864.9.19.38 “Amphisbaena cinerea Vandelli” Blanus sp. Portugal Boulenger (1885b: 434) NHMA/DF/GüntherColl/16/1/102 BMNH 1864.9.19.39* “Lacerta ocellata (?)” cf. Timon lepidus (Daudin, 1802) Portugal — NHMA/DF/GüntherColl/16/1/102, 104 28 July 1864 BMNH 1864.10.28.1 “batraciens” Sclerophrys regularis (Reuss, 1833) Duque de Bragança [= Kalandula], Angola Boulenger (1882b: 299) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104 BMNH 1864.10.28.2 “Rana superciliaris ?” Ptychadena oxyrhynchus (Smith, 1849) Duque de Bragança [= Kalandula], Angola Boulenger (1882b: 51 AHMB/CE/G76, NHMANHMA/ DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.3 [SAM ZR-002338] “Rana superciliaris ?” Ptychadena oxyrhynchus (Smith, 1849) Duque de Bragança [= Kalandula], Angola Boulenger (1882b: 51 AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.4–5 “Rana Bibroni Hallowel ?” [sic] Ptychadena porosissima (Steindachner, 1867) Duque de Bragança [= Kalandula], Angola Boulenger (1882b: 53) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.6–9 “Stenorhynchus ?” Phrynobatrachus natalensis (Smith, 1849) Duque de Bragança [= Kalandula], Angola Günther (1865: 481), Boulenger (1882b: 112) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.10 “batraciens” Afrixalus wittei (Laurent, 1941) Duque de Bragança [= Kalandula], Angola Günther (1868b: 479), Boulenger (1882b: 121) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104 BMNH 1864.10.28.11–13 “Hyperolius sp ?” Hyperolius cf. parallelus Günther, 1858 Duque de Bragança [= Kalandula], Angola —AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.14 [1947.2.9.68] “batraciens” Hyperolius nasutus Günther, 1865 Duque de Bragança [= Kalandula], Angola Günther (1865a: 482), Boulenger (1882b: 127), Marques et al. (2018: 106) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104, 106 BMNH 1864.10.28.15 “ophidiens” Lycophidion multimaculatum Boettger, 1888 Duque de Bragança [= Kalandula], Angola Boulenger (1893: 340), Broadley (1996: 19) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104 BMNH 1864.10.28.16 [1946.1.14.53–54]* “ophidiens” Limnophis bicolor Günther, 1865 Duque de Bragança [= Kalandula], Angola Günther (1865b: 96), Bocage (1866a: 47, 1895a: 77), Boulenger (1893: 275), Wallach et al. (2014: 384), Marques et al. (2018: 406), Conradie et al. (2020: 12) AHMB/CE/G76, NHMA/DF/GüntherColl/16/1/104 25 May 1866 BMNH 1866.6.11.1 “Nº 1 - Stellio angolensis Bocage nov. sp.” Acanthocercus ceriacoi Marques et al., 2022 Duque de Bragança [= Kalandula], Angola Boulenger (1885a: 359), Marques et al. (2022: 230) AHMB/CE/G79, NHMA/DF/ GüntherColl/16/1/110, NHMA/DF/ ZOO/200/1/185 BMNH 1866.6.11.2 “Nº 2 - Agama sp ?” Agama cf. schacki Mertens, 1938 Duque de Bragança [= Kalandula], Angola Boulenger (1885a: 358) AHMB/CE/G79, NHMA/DF/GüntherColl/16/1/110 BMNH 1866.6.11.3–4 [1946.9.3.47–48] “Nº 3 - Ichnotropis bivittatus Bocage” Ichnotropis bivittata bivittata Bocage, 1866 Duque de Bragança [= Kalandula], Angola Boulenger (1887: 79, 1921: 424) AHMB/CE/G79, NHMA/DF/ GüntherColl/16/1/110, NHMA/DF/ ZOO/200/1/185 BMNH 1866.6.11.5–6 “Nº 4 - Chamaeleo gracilis”Chamaeleo gracilis etiennei Schmidt, 1919 Duque de Bragança [= Kalandula], Angola Boulenger (1887: 449) AHMB/CE/G79, NHMA/DF/GüntherColl/16/1/110 BMNH 1866.6.11.7 [1946.8.15.27] “Nº 5 - Euprepes quinqueteniatus”Trachylepis bocagii (Boulenger, 1887) Duque de Bragança [= Kalandula], Angola Boulenger (1887: 204), Marques et al. (2018: 259), Ceríaco et al. (2024: 49) NHMA/DF/GüntherColl/16/1/110 BMNH 1866.6.11.8 [1946.8.19.13] “Nº 6 - Euprepes Gravenhorstii”Trachylepis bayonii (Bocage, 1872) Duque de Bragança [= Kalandula], Angola Boulenger (1887: 201), Bauer et al. (2003: 270), Marques et al. (2018: 256), Ceríaco et al. (2024: 37) NHMA/DF/GüntherColl/16/1/110 Vertebrate Zoology 75, 2025, 353–404 361 Shipment date Original accession number [ current number] Verbatim identification Current identification Locality Published citations Archival citations 13 July 1867 BMNH 1867.7.23.16 “Naja nigricollis”Naja nigricollis Reinhardt, 1843 Bissau, Guinea-Bissau Boulenger (1896: 379) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.17 “Pachydactylus ocellatus”Pachydactylus cf. punctatus Peters, 1854 Benguela, Angola Boulenger (1885a: 206) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.18 “Homodactylus Bibroni”Chondrodactylus pulitzerae (Schmidt, 1933) Benguela, Angola Boulenger (1885a: 202) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.19 “Mochlus (Eumeces) afer”Mochlus sundevallii (Smith, 1849) Benguela, Angola Boulenger (1887: 308) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.21 “Leptophis dorsalis Bocage” Philothamnus dorsalis Bocage (1866) Benguela, Angola Günther (1868a: 424), Boulenger (1894b: 101) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.22 “Psammophis elegans”Psammophis subtaeniatus Peters, 1882 Benguela, Angola Boulenger (1895: 212, 1896: 161), Broadley et al. (1977a: 13, 2002: 108), Marques et al. (2018: 352) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.23 “Alopecion variegatum Bocage n. sp.” Boaedon variegatus (Bocage, 1867) Benguela, Angola Boulenger (1893: 333), Hallerman et al. (2020: 22) NHMA/DF/ZOO/200/1/195, NHMA/DF/GüntherColl/16/1/112 BMNH 1867.7.23.24 “Hyperolius insignis Bocage” Hyperolius cf. parallelus Günther, 1858 Benguela, Angola Günther (1868b: 479), Boulenger (1882b: 122) NHMA/DF/ZOO/200/1/195, NHMA/DF/GüntherColl/16/1/112 BMNH 1867.7.23.25 “Dactylethra Mulleri”Xenopus petersii Bocage, 1895 Benguela, Angola Boulenger (1882b: 458) NHMA/DF/ZOO/200/1/195 BMNH 1867.7.23.26 [1946.8.15.37] “Euprepes binotatus Bocage n. sp.” Trachylepis binotata (Bocage, 1867) Benguela, Angola Boulenger (1887: 199), Bauer et al. (2003: 271), Marques et al. (2018: 257), Ceríaco et al. (2024: 42) NHMA/DF/ZOO/200/1/195 24 May 1869 BMNH 1872.2.15.1 “Nº 3. Rana plicigula Nov. sp.” Hoplobatrachus occipitalis (Günther, 1858) W. Africa [= Angola] Boulenger (1882b: 28) AHMB/CE/G85, NHMA/DF/ ZOO/200/1/189, 190 BMNH 1872.2.15.2 “Nº 5. Rana sp? (voisin de R. plicigula et de R. Delalandii)” Amietia angolensis (Bocage, 1866) W. Africa [= Angola] Boulenger (1882b: 51) NHMA/DF/ZOO/200/1/189 BMNH 1872.2.15.3 “Nº 4. Bufo benguellensis nov. sp.” Sclerophrys funerea (Bocage, 1866) Benguela, Angola Bocage (1867b: 845), Boulenger (1882b: 300), Marques et al. (2018: 70) NHMA/DF/ZOO/200/1/189 BMNH 1872.2.15.4 “Nº 1. Hyperolius huillensis nov. sp.” Hyperolius cf. parallelus Günther, 1858 W. Africa [= Huíla, Angola] Boulenger (1882b: 122) NHMA/DF/ZOO/200/1/189 BMNH 1872.2.15.5 “Nº 2. Hyper. sp ?” Hyperolius cf. parallelus Günther, 1858 W. Africa [=Huíla, Angola] — NHMA/DF/ZOO/200/1/189 BMNH 1872.2.15.6 [RR 1933.1.6.1] “E/8. Pyxicephalus rugosus Gthr ?” Tomopterna tuberculosa (Boulenger, 1882) W. Africa [= Angola] Boulenger (1882b: 30) NHMA/DF/ZOO/200/1/189 29 March 1875 BMNH 1875.4.26.8 [1900386] “Hemidactylus gutturalis Boc.” Lygodactylus gutturalis (Bocage, 1873) Bissau, Guinea-Bissau Boulenger (1885a: 161), Lobón-Rovira et al. (2024: 465), NHMA/DF/ZOO/200/1/191 BMNH 1875.4.26.9 [1946.8.18.43] “Euprepes Hopfferi Boc.” Chioninia stangeri (Gray, 1845) Ilhéu Raso, Cabo Verde Boulenger (1887: 158), Bauer et al. (2003: 273), Miralles et al. (2010: 17) NHMA/DF/ZOO/200/1/191 BMNH 1875.4.26.10* “Hemidactylus Cessaci Boc.” Hemidactylus cf. lopezjuradoi Arnold et al., 2008 Santiago, Cabo Verde Boulenger (1885a: 118), Arnold et al. (2008: 629; fig. 5E) NHMA/DF/ZOO/200/1/191 BMNH 1875.4.26.11* “Ascalabotes gigas Boc.” Tarentola gigas (Bocage, 1875) Ilhéu Raso, Cabo Verde Boulenger (1885a: 200) NHMA/DF/ZOO/200/1/191 Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 368 Figure 6. Specimen of Leptopelis bocagii mentioned in the 1866 letter (BMNH 1875.5.22.3), note presence of distinct pectoral glands. Photos by DP. tinct 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. 14242 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 Hy lambates 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. Vertebrate Zoology 75, 2025, 353–404 369 Figure 8. Preserved holotype of Leptopelis bocagii (left, BMNH 1875.5.22.2) and portion of Plate XXXIII adapted from Günther (1865a) depicting the same specimen (right). Plates often depict a mirrored image of the original illustration, thus the difference between the specimen and the plate. Figure 7. Holotype of Leptopelis bocagii (BMNH 1875.5.22.2). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 370 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. Leptopelis viridis (Günther, 1869)* Specimen. Guinea-Bissau: Bolama: BMNH 1887.3.23.4 [syntype of Hylambates cynnamomeus Bocage, 1893; Fig. 9]. Comments. 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 Figure 9. Syntype of Hylambates cynnamomeus (BMNH 1887.3.23.4), label not to scale. Photos by DP. Vertebrate Zoology 75, 2025, 353–404 371 Family Brevicipitidae Bonaparte, 1850 Genus Breviceps Merrem, 1820 Breviceps cf. ombelanonga Nielsen et al., 2020 Specimen. Angola: Quissange: BMNH 1887.3.23.5. Comments. 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). Family Bufonidae Gray, 1825 Genus Poyntonophrynus Frost et al., 2006 Poyntonophrynus dombensis (Bocage, 1895)* Specimens. Angola: Dombe (Benguella): BMNH 1947. 2.21.3–4 [syntypes, originally BMNH 1896.2.28.3–4; Figs 10, 11]. Comments. 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). Figure 10. Syntype of Poyntonophrynus dombensis (BMNH 1947.2.21.3). Photos by Frank Tillack. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 372 Genus Sclerophrys Tschudi, 1838 Sclerophrys funerea (Bocage, 1866)* Specimens. Angola: Benguela: BMNH 1872.2.15.3 [syntype of Bufo benguelensis Boulenger, 1882; Fig. 12], “W. Africa” [= Caconda]: BMNH 1883.7.26.27. Comments. 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. Figure 11. Syntype of Poyntonophrynus dombensis (BMNH 1947.2.21.4). Photos by Frank Tillack. Vertebrate Zoology 75, 2025, 353–404 373 Sclerophrys regularis (Reuss, 1833) Specimen. Angola: Duque de Bragança: BMNH 1864. 10.28.1. Comments. 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). Family Dicroglossidae Anderson, 1871 Genus Hoplobatrachus Peters, 1863 Hoplobatrachus occipitalis (Günther, 1858) Specimen. Angola: “W. Africa”: BMNH 1872.2.15.1. Comments. 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. Figure 12. Syntype of Bufo benguelensis (BMNH 1872.2.15.3). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 374 Family Hyperoliidae Laurent, 1943 Genus Afrixalus Laurent, 1944 Afrixalus wittei (Laurent, 1941) Specimen. Angola: Duque de Bragança: BMNH 1864. 10.28.10. Comments. 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 Hyperolius nasutus Günther, 1865* Specimen. Angola: Duque de Bragança: BMNH 1947. 2.9.68 [lectotype, originally BMNH 1864.10.28.14; Fig. 13]. Comments. 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). Hyperolius cf. parallelus Günther, 1858* Specimens. 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. Comments. 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 Vertebrate Zoology 75, 2025, 353–404 375 Figure 13. Lectotype of Hyerolius nasutus (BMNH 1947.2.9.68). Photos by DP. Figure 14. Syntype of Hyperolius insignis (BMNH 1867.7.23.24). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 376 (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. Family Phrynobatrachidae Laurent, 1941 Genus Phrynobatrachus Günther, 1862 Phrynobatrachus natalensis (Smith, 1849) Specimens. Angola: Duque de Bragança: BMNH 1864. 10.28.6–9. Comments. 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). Figure 15. Syntype of Hyperolius huillensis (BMNH 1872.2.15.4). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 377 Family Pipidae Gray, 1825 Genus Xenopus Wagler, 1827 Xenopus petersii Bocage, 1895 Specimen. Angola: Benguella: BMNH 1867.7.23.25. Comments. 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. Family Ptychadenidae Dubois, 1987 Genus Ptychadena Boulenger, 1917 Ptychadena oxyrhynchus (Smith, 1849) Specimen. Angola: Duque de Bragança: BMNH 1864. 10. 28.2. Comments. 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. Ptychadena porosissima (Steindachner, 1867) Specimens. Angola: Duque de Bragança: BMNH 1864. 10.28.4–5. Comments. 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. Family Pyxicephalidae Bonaparte, 1850 Genus Amietia Dubois, 1987 Amietia angolensis (Bocage, 1866) Specimens. Angola: “W. Africa”: BMNH 1872.2.15.2, “Angola”: BMNH 1896.2.28.1. Comments. 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 Tomopterna tuberculosa (Boulenger, 1882) Specimen. Angola: “W. Africa”: BMNH RR 1933.1.6.1 [reregistered, originally BMNH 1872.2.15.6]. Comments. 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”). Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 384 type (ZMB 8999) of Euprepes hopfferi exists in the ZMB (Bauer et al. 2003). Genus Eumecia Bocage, 1870 Eumecia anchietae Bocage, 1870 Specimen. Angola: Caconda: BMNH 1893.12.27.8. Comments. 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 Mochlus sundevallii (Smith, 1849) Specimen. Angola: Benguella: BMNH 1867.7.23.19. Comments. 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 Trachylepis bayonii (Bocage, 1872)* Specimen. Angola: Duque de Bragança: BMNH 1946.8.19.13 [syntype, originally BMNH 1866.6.11.8; Fig. 21]. Comments. 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 auFigure 20. Syntype of Euprepes hopfferi (BMNH 1946.8.18.43). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 385 thors 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. Trachylepis binotata (Bocage, 1867)* Specimen. Angola: Benguella: BMNH 1946.8.15.37 [syntype, originally BMNH 1867.7.23.26; Fig. 22]. Comments. 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. Figure 21. Syntype of Trachylepis bayonii (BMNH 1946.8.19.13). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 386 Trachylepis bocagii (Boulenger, 1887)* Specimen. Angola: Duque de Bragança: BMNH 1946. 8.15.27 [syntype, originally BMNH 1866.6.11.7; Fig. 23]. Comments. 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. Family Colubridae Oppel, 1811 Genus Philothamnus Smith, 1840 Philothamnus cf. angolensis Bocage, 1882 Specimen. Angola: Caconda: BMNH 1882.6.9.5 (not examined). Comments. 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 Figure 22. Syntype of Trachylepis binotata (BMNH 1946.8.15.37). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 387 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. Philothamnus dorsalis Bocage, 1866 Specimens. Angola: Benguella: BMNH 1867.7.23.21, “Angola” [= Luanda]: BMNH 1882.6.9.2. Comments. 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). Philothamnus girardi Bocage, 1893* Specimen. Equatorial Guinea: “Ile d’Anno-Bom, Golfe de Guinée”: BMNH 1946.1.6.3 [syntype, originally BMNH 1893.12.27.18; Fig. 24]. Comments. 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. Figure 23. Syntype of Trachylepis bocagii (BMNH 1946.8.15.27). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 388 Philothamnus heterolepidotus Günther, 1863 Specimen. Angola: “Angola” [= Duque de Bragança]: BMNH 1882.6.9.3. Comments. 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. Philothamnus semivariegatus (Smith, 1840)* Specimens. 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]. Comments. 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. Philothamnus thomensis Bocage, 1882* Specimens. 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. Comments. 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), Figure 24. Syntype of Philothamnus girardi (BMNH 1946.1.6.3). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 389 Figure 25. Syntype of Philothamnus smithii (BMNH 1946.1.5.98). Photos by DP. Figure 26. Syntype of Philothamnus thomensis (BMNH 1946.1.21.60). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 390 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 Zamenis scalaris (Schinz, 1822) Specimens. Portugal: Alfeite: BMNH 1893.12.27.14, Aldegallega [i.e., Aldeia Galega = Montijo]: BMNH 1893.12.27.15, Coimbra: BMNH 1893.12.27.16. Comments. 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). Family Lamprophiidae Fitzinger, 1843 Genus Boaedon Duméril, Bibron & Duméril, 1854 Boaedon variegatus (Bocage, 1867)* Specimen. Angola: Benguella: BMNH 1867.7.23.23 [paralectotype of Alopecion variegatum Bocage, 1867 and Boodon lineatus var. lineolata Bocage, 1895; Fig. 27]. Comments. 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 Figure 27. Paralectotype of Boaedon variegatus and Boodon lineatus var. lineolata (BMNH 1867.7.23.23). Photos by DP. Vertebrate Zoology 75, 2025, 353–404 391 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 variegatums 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 Lycophidion multimaculatum Boettger, 1888 Specimens. Angola: Duque de Bragança: BMNH 1864. 10.28.15, Caconda: BMNH 1893.12.27.13. Comments. 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). Family Natricidae Bonaparte, 1838 Genus Limnophis Günther, 1865 Limnophis bicolor Günther, 1865* Specimens. Angola: Duque de Bragança: BMNH 1946. 1.14.53–54 (not examined) [syntypes, originally BMNH 1864.10.28.16; Figs 28, 29]. Comments. 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. Family Psammophiidae Bourgeois, 1968 Genus Psammophis Fitzinger, 1826 Psammophis leopardinus (Bocage, 1887) Specimen. Angola: “W. Africa” [= Duque de Bragança]: BMNH 1875.5.22.4. Comments. 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 lo- Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 392 Figure 29. Syntype of Limnophis bicolor (BMNH 1946.1.14.54). Photos by Veerappan Deepak. Figure 28. Syntype of Limnophis bicolor (BMNH 1946.1.14.53). Photos by Veerappan Deepak. Vertebrate Zoology 75, 2025, 353–404 393 cality (i.e., “West Africa”) stated on the British Museum records, precluding a confident identification in such a taxonomically challenging group. Psammophis subtaeniatus Peters, 1882* Specimen. Angola: Benguella: BMNH 1867.7.23.22 [holotype of Psammophis bocagii Boulenger, 1895; Fig. 30]. Comments. 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 Psammophylax ocellatus (Bocage, 1873) Specimen. Angola: Humbe: BMNH 1882.6.9.1. Comments. 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. Figure 30. Holotype of Psammophis bocagii (BMNH 1867.7.23.22). Photos by DP. Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 400 Conradie W, Keates C, Verburgt L, Baptista NL, Harvey J, Júlio T, Neef G (2022) Contributions to the herpetofauna of the Angolan Okavango-Cuando-Zambezi River drainages. Part 2: Lizards (Sauria), chelonians, and crocodiles. Amphibian & Reptile Conservation 16: 181–214. Cornish C, Driver F (2020) “Specimens distributed”: The circulation of objects from Kew’s Museum of Economic Botany, 1847–1914. Journal of the History of Collections 32: 327–340. https://doi. org/10.1093/jhc/fhz008 Dubois A, Denzer W, Entiauspe-Neto OM, Frétey T, Ohler A, Bauer AM, Pyron RA (2024) Nomenclatural problems raised by the recent description of a new anaconda species (Squamata, Serpentes, Boidae), with a nomenclatural review of the genus Eunectes. Zootaxa 37: 8–58. https://doi.org/10.11646/bionomina.37.1.2 Dubois A, Ohler A (1996) Early scientific names of Amphibia Anura. I. Introduction. Bulletin du Muséum national d’Histoire naturelle, Série 4, 18: 297–320. Duméril AMC, Bibron G (1839) Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Tome Cinquième, Contenant l’Histoire de Quatre-vingt-trois Genres et de Deux Cent Sept Espèces des Trois Dernières Familles de l’Ordre des Sauriens, Savoir: les Lacertiliens, les Chalcidiens et les Scincoïdiens. Librairie Encyclopédique de Roret, Paris, viii, 854 pp., 37 plates. Engel MS, Ceríaco LMP, Daniel GM, Dellapé PM, Löbl I, Marinov M, Reis RE, Young MT, Dubois A, Agarwal I, Lehmann A P, Alvarado M, Alvarez N, Andreone F, Araujo-Vieira K, Ascher JS, Baêta D, Baldo D, Bandeira SA, Barden P, Barrasso DA, Bendifallah L, Bockmann FA, Böhme W, Borkent A, Brandão CRF, Busack SD, Bybee SM, Channing A, Chatzimanolis S, Christenhusz MJM, Crisci JV, D’Elía G, Da Costa LM, Davis SR, De Lucena CAS, Deuve T, Elizalde SF, Faivovich J, Farooq H, Ferguson AW, Gippoliti S, Gonçalves FMP, Gonzalez VH, Greenbaum E, Hinojosa-Díaz IA, Ineich I, Jiang J, Kahono S, Kury AB, Lucinda PHF, Lynch JD, Malécot V, Marques MP, Marris JWM, McKellar RC, Mendes LF, Nihei SS, Nishikawa K, Ohler A, Orrico VGD, Ota H, Paiva J, Parrinha D, Pauwels OSG, Pereyra MO, Pestana JB, Pinheiro PDP, Prendini L, Prokop J, Rasmussen C, Rödel M-O, Rodrigues MT, Rodríguez SM, Salatnaya H, Sampaio I, Sánchez-García A, Shebl MA, Santos BS, Solórzano-Kraemer MM, Sousa ACA, Stoev P, Teta P, Trape J-F, Van-Dúnem Dos Santos C, Vasudevan K, Vink CJ, Vogel G, Wagner P, Wappler T, Ware JL, Wedmann S, Kusamba Zacharie C (2021) The taxonomic impediment: A shortage of taxonomists, not the lack of technical approaches. Zoological Journal of the Linnean Society 193: 381–387. https://doi.org/10.1093/zoolinnean/zlab072 França C (1908) Le Professeur Barbosa du Bocage, 1823–1907: éloge historique prononcé à la séance solennelle du 2 mai 1908. Bulletin de la Société Portugaise des Sciences Naturelles 2: 141–161. Frost DR (2024) Amphibian Species of the World: An Online Reference. Version 6.2. American Museum of Natural History, New York, NY, https://amphibiansoftheworld.amnh.org/index.php [accessed 8 May 2025]. Gamito-Marques D (2018) A space of one’s own: Barbosa du Bocage, the foundation of the National Museum of Lisbon, and the construction of a career in zoology (1851–1907). Journal of the History of Biology 51: 223–257. https://doi.org/10.1007/s10739-017-9487-6 Gamito-Marques D (2022) The golden age (1862–1910) of the Zoological Section of the Museu Nacional de Lisboa (National Museum of Lisbon), Portugal. Archives of Natural History 59: 160–174. https:// doi.org/10.3366/anh.2022.0765 Goodwin ZA, Harris DJ, Filer D, Wood JRI, Scotland RW (2015) Widespread mistaken identity in tropical plant collections. Current Biology 25: R1066–R10697. https://doi.org/10.1016/j.cub.2015.10.002 Günther ACLG (1865a) Descriptions of new species of batrachians from West Africa. Proceedings of the Zoological Society of London 1864: 479–482. Günther ACLG (1865b) Fourth account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, Series 3, 15: 89–98. Günther ACLG (1868a) Sixth account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, Series 4, 1: 413–429. Günther ACLG (1868b) First account of species of tailless batrachians added to the collection of the British Museum. Proceedings of the Zoological Society of London 1868: 478–490. Günther ACLG (1912) The History of the Collections Contained in the Natural History Departments of the British Museum. Vol II. Appendix. General History of the Department of Zoology from 1856 to 1895. British Museum, London, 109 pp. Hallermann J, Ceríaco LMP, Schmitz A, Ernst A, Conradie W, Verburgt L, Marques MP, Bauer AM (2020) A review of the Angolan house snakes, genus Boaedon Duméril, Bibron and Duméril (1854) (Serpentes: Lamprophiidae), with description of three new species in the Boaedon fuliginosus (Boie, 1827) species complex. African Journal of Herpetology 69: 29–78. https://doi.org/10.1080/21564574.2020. 1777470 Heumann I, MacKinney AG, Buschmann R (2022) Introduction: The issue of duplicates. The British Journal for the History of Science 55: 257–278. https://doi.org/10.1017/S0007087422000267 Hughes B (1985) Progress on a taxonomic revision of African green tree snakes (Philothamnus spp.). In: Schuchmann KL (Ed.) Proceedings of the International Symposium on African Vertebrates. Zoological Research Museum A. Koenig, Bonn, 511–530. Hughes B, Wade E (2002) On the African leopard whip snake, Psammophis leopardinus Bocage, 1887 (Serpentes, Colubridae), with the description of a new species from Zambia. Bulletin of the Natural History Museum 68: 75–81. International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, XXIX + 306 pp. Jacobsen NHG (1992) The status of Agama aculeata armata Peters 1854 (Reptilia: Agamidae). Journal of the Herpetological Association of Africa 41: 30–34. Kaiser K (2022) Duplicate networks: The Berlin botanical institutions as a “clearing house” for colonial plant material, 1891–1920. British Journal for the History of Science 55: 279–296. https://doi. org/10.1017/S0007087422000139 Kapun M, Schwentner M, Haring E, Akkari N, Kroh A, Kruckenhauser A, Vohland K (2025) Museomics, the extended specimen and collectomics – How to frame and name the diversity of information linked to specimens in natural history collections. Natural History Collections and Museomics 2: 1–21. https://doi.org/10.3897/ nhcm.2.161331 Kellner A (2024) Biological collections in danger? Anais da Academia Brasileira de Ciências 96: e2024961. https://doi.org/10.1590/0001376520242024961 Leaché AD, Wagner P, Linkem CW, Böhme W, Papenfuss TJ, Chong RA, Lavin BR, Bauer AM, Nielsen SV, Greenbaum E, Rödel M-O, Schmitz A, LeBreton M, Ineich I, Chirio L, Ofori-Boateng C, Vertebrate Zoology 75, 2025, 353–404 401 Eniang EA, El Din B, Lemmon AR, Burbrink FT (2014) A hybrid phylogenetic-phylogenomic approach for species tree estimation in African Agama lizards with applications to biogeography, character evolution, and diversification. Molecular Phylogenetics and Evolution 79: 215–230. https://doi.org/10.1016/j.ympev.2014.06.013 Letsch H, Greve C, Hundsdoerfer AK, Irisarri I, Moore JM, Espeland M, Wanke S, Arifin U, Blom MPK, Corrales C, Donath A, Fritz U, Köhler G, Kück P, Lemer S, Mengual X, Salas NM, Meusemann K, Palandačić A, Printzen C, Sigwart JD, Silva-Brandão KL, Simões M, Stange M, Suh A, Szucsich N, Tilic E, Töpfer T, Böhne A, Janke A, Pauls SU (2025) Type genomics: A framework for integrating genomic data into biodiversity and taxonomic research. Systematic Biology: syaf040. https://doi.org/10.1093/sysbio/syaf040 Lobón-Rovira J, Bauer AM, Vaz Pinto P, Trape J-F, Conradie W, Kusamba C, Júlio T, Cael G, Stanley EL, Hughes DF, Behangana M, Masudi FM, Pauwels OSG, Greenbaum E (2024) Integrative revision of the Lygodactylus gutturalis (Bocage, 1873) complex unveils extensive cryptic diversity and traces its evolutionary history. Zoological Journal of the Linnean Society 201: 447–492. https://doi. org/10.1093/zoolinnean/zlad123 Lopes Vieira AX (1886) Notícia acerca dos produtos zoológicos trazidos da ilha de S. Thomé para o Museu Zoológico da Universidade de Coimbra, pelo Sr. Adolpho F. Moller, em 1885. O Instituto, Série 2, 34: 235–241. Loveridge A (1947) Revision of the African lizards of the family Gekkonidae. Bulletin of the Museum of Comparative Zoology 98: 1–469, 7 plates. Loveridge A (1958) Revision of five African snake genera. Bulletin of the Museum of Comparative Zoology 119: 1–198. Madruga C (2012) The zoological collections of the Museu de Lisboa and the networks of scientific correspondence and exchange (1858–1898). In: Roca-Rosell A (Ed.) The Circulation of Science and Technology: Proceedings of the 4th International Conference of the European Society for the History of Science, Barcelona (Spain), 18–20 November 2010. Societat Catalana d’Història de la Ciència i de la Tècnica, Barcelona, 928–933. Madruga C (2013) José Vicente Barbosa du Bocage (1823–1907): A construção de uma persona científica. MSc Thesis, Universidade de Lisboa, Lisboa, 109 pp. Madruga C (2017) Expert at a distance: Barbosa du Bocage and the production of scientific knowledge on Africa. Journal of History of Science and Technology 11: 57–74. https://doi.org/10.1515/host2017-0004 Margócsy D (2022) Malinowski and malacology: Global value systems and the issue of duplicates. British Journal for the History of Science 55: 389–409. https://doi.org/10.1017/S0007087422000255 Marques MP, Ceríaco LMP, Blackburn DC, Bauer AM (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840–2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement II): 1–501. Marques MP, Parrinha D, Lopes-Lima M, Tiutenko A, Bauer AM, Ceríaco LMP (2024) An island in a sea of sand: A first checklist of the herpetofauna of the Serra da Neve inselberg, southwestern Angola. ZooKeys 1201: 167–217. https://doi.org/10.3897/zookeys.1201.120750 Marques MP, Parrinha D, Santos BS, Bandeira S, Butler BO, Sousa ACA, Bauer AM, Wagner P (2022) All in all it’s just another branch in the tree: A new species of Acanthocercus Fitzinger, 1843 (Squamata: Agamidae), from Angola. Zootaxa 5099: 221–243. https://doi. org/10.11646/zootaxa.5099.2.4 Mecke S, Mader F, Kieckbusch M, Kaiser H, Böhme W, Ernst R (2016) Tracking a syntype of the Australian skink Anomalopus leuckartii (Weinland, 1862): “Lost” treasures in the Senckenberg Natural History Collections Dresden highlight the importance of reassessing and safeguarding natural history collections. Vertebrate Zoology 66: 169–177. https://doi.org/10.3897/vz.66.e31545 Miller EC, Sellas AB, Drewes RC (2012) A new species of Hemidactylus (Squamata: Gekkonidae) from Príncipe Island, Gulf of Guinea, West Africa with comments on the African-Atlantic clade of Hemidactylus geckos. African Journal of Herpetology 61: 40–57. https:// doi.org/10.1080/21564574.2012.666680 Miralles A, Vasconcelos R, Perera A, Harris DJ, Carranza S (2010) An integrative taxonomic revision of the Cape Verdean skinks (Squamata, Scincidae). Zoologica Scripta 40: 16–44. https://doi. org/10.1111/j.1463-6409.2010.00453.x Nelson G, Ellis S (2018) The history and impact of digitization and digital data mobilization on biodiversity research. Philosophical transactions of the Royal Society B 374: 20170391. https://doi. org/10.1098/rstb.2017.0391 Nichols CA (2021) Exchanging Objects: Nineteenth-Century Museum Anthropology at the Smithsonian Institution. Museums and Collections 12. Berghahn Books, New York, NY, and Oxford, 268 pp. Nielsen SV, Conradie W, Ceríaco LMP, Bauer AM, Heinicke MP, Stanley EL, Blackburn DC (2020) A new species of rain frog (Brevicipitidae, Breviceps) endemic to Angola. ZooKeys 979: 133–160. https:// doi.org/10.3897/zookeys.979.56863 Osório B (1909) Elogio historico do illustre naturalista e professor J. V. Barbosa du Bocage. Memorias do Museu Bocage 1: i–xlii. Padial JM, Miralles A, De la Riva I, Vences M (2010) The integrative future of taxonomy. Frontiers in Zoology 7: 16. https://doi. org/10.1186/1742-9994-7-16 Parker HW (1936) Amphibians from Liberia and the Gold Coast. Zoologische Mededelingen, Leiden 19: 87–102. Parrinha D, Marques MP, Gonçalves FMP, Tiutenko A, Bauer AM, Ceríaco LMP (2025a) The genus Holaspis (Squamata: Lacertidae) in Angola: A tale of forgotten specimens and disappearing forests, with the description of a new species. Salamandra 61: 53–69. Parrinha D, Marques MP, Sousa ACA, Bauer AM, Ceríaco LMP (2025b) Buried in the sands of time: A new species of Sepsina Bocage, 1866 from Angola (Squamata: Scincidae). Annals of Carnegie Museum 90: 183–193. Parrinha D, Soares LB, Cartaxana A, Alves MJ (2024) The zoological collections of Portuguese oceanographic campaigns in former colonial territories. Natural History Collections and Museomics 1: 1–34. https://doi.org/10.3897/nhcm.1.136860 Perret J-L (1976) Révision des amphibiens africains et principalement des types, conservés au Musée Bocage de Lisbonne. Arquivos do Museu Bocage, Série 2, 6: 15–34. Peters WCH (1882) Naturwissenschaftliche Reise nach Mossambique auf Befehl seiner Majestät des Königs Friedrich Wilhelm IV in den Jahren 1842 bis 1848 ausgeführt. Zoologie, III, Amphibien. G. Reimer, Berlin, xv, 191 pp., 33 plates. Reese H, Finney V, Ville S (2024) Value, knowledge and reputation: Zoological exchange by Australian museums, 1870–1900. British Journal for the History of Science 58: 81–100. https://doi.org/10.1017/ S0007087424000700 Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 402 Sabaj MH (2020) Codes for natural history collections in ichthyology and herpetology. Copeia 108: 593–669. https://doi.org/10.1643/ ASIHCODONS2020 Saldanha L (1978) Museu Bocage. Copeia 1978: 739–740. Santos BS, Marques MP, Bauer AM, Ceríaco LMP (2021) Herpetological results of Francisco Newton’s Zoological Expedition to Angola (1903–1906): A taxonomic revision and new records of a forgotten collection. Zootaxa 5028: 1–80. https://doi.org/10.11646/zootaxa.5028.1.1 Santos BS, Marques MP, Ceríaco LMP (2024) Lack of country-wide systematic herpetology collections in Portugal jeopardizes future research and conservation. Anais da Academia Brasileira de Ciências 96: e20230622. https://doi.org/10.1590/0001-3765202420230622 Sigwart JD, Schleuning M, Brandt A, Pfenninger M, Saeedi H, Borsch T, Häffner E, Lücking R, Güntsch A, Trischler H, Töpfer T, Wesche K, Consortium C (2025) Collectomics – towards a new framework to integrate museum collections to address global challenges. Natural History Collections and Museomics 2: 1–20. https://doi. org/10.3897/nhcm.2.148855 Steindachner F (1867) Reise der österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Theil. I. Amphibien. Kaiserlich-Königliche Hofund Staatsdruckerei, Wien, 70 pp., 5 plates. Tillack F, de Ruiter R, Rödel M-O (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum für Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution 97: 407– 450 https://doi.org/10.3897/zse.97.68000 Trape J-F, Crochet P-A, Broadley DG, Sourouille P, Mané Y, Burger M, Böhme W, Saleh M, Karan A, Lanza B, Mediannikov O (2019) On the Psammophis sibilans group (Serpentes, Lamprophiidae, Psammophiinae) north of 12°S, with the description of a new species from West Africa. Bonn Zoological Bulletin 68: 61–91. https://doi. org/10.20363/BZB-2019.68.1.061 Tyler MJ, Fucsko LA, Roberts JD (2023) Calamities causing loss of museum collections: A historical and global perspective. Zootaxa 5230: 153–178. https://doi.org/10.11646/zootaxa.5230.2.2 Vasconcelos R, Köhler G, Geniez P, Crochet P-A (2020) A new endemic species of Hemidactylus (Squamata: Gekkonidae) from São Nicolau Island, Cabo Verde. Zootaxa 4878: 501–522. https://doi. org/10.11646/zootaxa.4878.3.4 Vasconcelos R, Perera A, Geniez P, Harris DJ, Carranza S (2012) An integrative taxonomic revision of the Tarentola geckos (Squamata, Phyllodactylidae) of the Cape Verde Islands. Zoological Journal of the Linnean Society 164: 328–360. https://doi.org/10.1111/j.10963642.2011.00768.x Villa A, Kirchner M, Alba DM, Bernardini F, Bolet A, Luján AH, Fortuny J, Hipsley CA, Müller J, Sindaco R, Tuniz C, Delfino M (2019) Comparative cranial osteology of Blanus (Squamata: Amphisbaenia), Zoological Journal of the Linnean Society 185: 693–716. https://doi.org/10.1093/zoolinnean/zly082 Wagner P, Butler BO, Ceríaco LMP, Bauer AM (2021) A new species of the Acanthocercus atricollis complex (Squamata: Agamidae). Salamandra 57: 449–463. Wagner P, Greenbaum E, Bauer AM, Kusamba C, Leaché AD (2018) Lifting the blue-headed veil – Integrative taxonomy of the Acanthocercus atricollis species complex (Squamata: Agamidae). Journal of Natural History 52: 771–817. https://doi.org/10.1080/00222933.20 18.1435833 Wallach V, Williams KL, Boundy J (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, FL, 1227 pp. Wüster W, Tillack F (2023) On the importance of types and the perils of “en passant” taxonomy: A brief history of the typification of Coluber naja Linnaeus, 1758 (Serpentes: Elapidae) and its implications, with the designation of a lectotype. Zootaxa 5346: 403–419. https://doi. org/10.11646/zootaxa.5346.4.3 Zamudio KR, Kellner A, Serejo C, de Britto MR, Castro CB, Buckup PA, Pires DO, Couri M, Kury AB, Cardoso IA, Monné ML, Pombal Jr. J, Patiu CM, Padula V, Pimenta AD, Ventura CRR, Hajdu E, Zanol J, Bruna EM, Fitzpatrick J, Rocha LA (2018) Lack of science support fails Brazil. Science 361: 1322–1323. https://doi.org/10.1126/ science.aav3296 Appendix 1 Archival documents Arquivo Histórico Museu Bocage Div. 471, Diversos, 30 May 1855, “Lista das aves qe. forão pa. Inglaterra em 30 de Maio de 1855”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 472, Diversos, 30 May 1855, “Lista das conchas que forão para Inglaterra em 30 de Maio de 1855”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 486, Diversos, August 1874, “Aves remettidas em communicação ao Sharpe em Agosto de 1874”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 493, Diversos, May 1879, “Exemplares remettidos (em communicação) a Mr. Shelley – 6 Tenente est. Londres em maio de 1879”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 500, Diversos, 10 November 1883, “Reptis remettidos em communicação ao Dr. Günther – pelo vapor Malange – em 10 de Nov. 1883”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 504, Diversos, undated, “Aves de Angola remettidas a Mr. Shelley – de Londres”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 509, Diversos, November 1888, “Aves e mammiferos offerecidos ao Museu de Londres em Novembro de 1888”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Vertebrate Zoology 75, 2025, 353–404 403 Arquivo Histórico Museu Bocage Div. 510, Diversos, November 1888, “Aves escolhidas pa. o Museu de Londres – Nov. de 1888”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 513, Diversos, undated, “Enviado em communicação a Old. Thomas”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 518, Diversos, March 1897, “Exemplares enviados a M. Oldfield Thomas em Março de 1897”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 522, Diversos, 17 March 1904, “Exemplares enviados para Londres em 17 Março 1904”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 532.3, Diversos, 4 January, “Foi p. Londres (Andersen) no dia 4 de Janeiro e para o Nobre tambem”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 532.7, Diversos, “Aves remetidas pa. R. B. Sharpe”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 532.8, Diversos, undated, “Mammiferos remetidos em consulta a Mr. Dobson”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage Div. 532.10, Diversos, undated, “Aves remetidas a Sharpe”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico do Museu Bocage AHMB/Div. 532.18, Diversos, undated, “List of rodents received from the Lisbon Museum”. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G75, Correspondence, 19 September 1864. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G76, Correspondence, 25 July 1865. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G78, Correspondence, 6 May 1866. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G79, Correspondence, 29 June 1866. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G81, Correspondence, 19 July 1867. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G85, Correspondence, 26 June 1869. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G93, Correspondence, 16 March 1882. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/G94, Correspondence, 10 June 1882. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/B43, Correspondence, 6 November 1893. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/B44, Correspondence, 29 March 1894. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Arquivo Histórico Museu Bocage CE/B48, Correspondence, 14 April 1896. Museu Nacional de História Natural e da Ciência, Universidade de Lisboa. Natural History Museum Archives DF/GüntherColl/16/1/102, The Günther Collection, Letters to Albert and R. W. T. Günther, 24 May 1864. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/GüntherColl/16/1/104, The Günther Collection, Letters to Albert and R. W. T. Günther, 28 July 1864. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/GüntherColl/16/1/106, The Günther Collection, Letters to Albert and R. W. T. Günther, 20 June 1865. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/GüntherColl/16/1/110, The Günther Collection, Letters to Albert and R. W. T. Günther, 25 May 1866. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/GüntherColl/16/1/112, The Günther Collection, Letters to Albert and R. W. T. Günther, 13 July 1867. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/184, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 11 April 1866. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/185, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 10 July 1866. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/189, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 24 May 1869. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/190, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 16 June 1869. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/191, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 29 March 1875. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/194, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 10 June 1875. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/1/195, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 13 July 1867. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/21/38, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 2 March 1882. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/21/39, Department of Zoology, Keeper of Zoology’s Correspondence and Parrinha D et al.: Herpetological collections sent by Bocage to the British Museum 404 Files, 11 March 1882. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/21/41, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 5 April 1882. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/200/21/42, Department of Zoology, Keeper of Zoology’s Correspondence and Files, 20 May 1882. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/235/1/1/1/76, Department of Zoology, Reptile Section Correspondence and Papers, 19 March 1887. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/235/1/1/1/76, Department of Zoology, Reptile Section Correspondence and Papers, 5 December 1893. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/235/1/1/1/76, Department of Zoology, Reptile Section Correspondence and Papers, 24 February 1896. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/235/1/1/1/76, Department of Zoology, Reptile Section Correspondence and Papers, 2 June 1896. Library and Archives collections of the Natural History Museum, London. Natural History Museum Archives DF/ZOO/235/1/1/1/76, Department of Zoology, Reptile Section Correspondence and Papers, 3 April 1896. Library and Archives collections of the Natural History Museum, London.