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Catalogue of the amphibian and reptile type specimens of the Museu de História Natural da Universidade do Porto in Portugal, with some comments on problematic taxa


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Catalogue of the amphibian and reptile type specimens of the Museu de História Natural da Universidade do Porto in Portugal, with some comments on problematic taxa We present an annotated catalog of the type specimens of amphibians and reptiles in the collections of the Museu de História Natural da Universidade do Porto in Portugal. These specimens, all from present-day Angola, formed the basis of taxonomic descriptions by both José Júlio Bethencourt Ferreira and José Vicente Barbosa du Bocage in the latest 19th and early 20th century. We provide details for all type specimens and summarize the history and taxonomy for each species. Specimens of Rappia bocagei var. maculata and Typhlops bocagei could not be located during our survey, and we believe these to be lost. The collections at the University of Porto contain type specimens of one snake, Typhlops boulengeri, and eight frogs, Arthroleptis carquejai, Hylambates bocagei var. leucopunctata, Rappia platyceps var. angolensis, Rappia bivittata, Rappia fasciata, Rappia nobrei, Rappia osorioi, and Rappia seabrai. Of these, only two are currently recognized: Afrixalus osorioi and Arthroleptis carquejai.
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2014 | VOLUME 31 | PAGES 13-36
Catalogue of the amphibian and reptile type
specimens of the Museu de História Natural
da Universidade do Porto in Portugal,
with some comments on problematic taxa
*Corresponding author
1. Centro de Estudos de História e Filosoa da Ciência, Universidade de Évora,
Palácio do Vimioso, Largo Marquês de Marialva 8, 7000-554 Évora, Portugal
2. Museu Nacional de História Natural e da Ciência, Rua da Escola Politécnica 56-58, 1250-102 Lisboa, Portugal
3. Department of Vertebrate Zoology and Anthropology, California Academy of Sciences,
55 Music Concourse Drive, San Francisco, California 94118, USA
4. Departamento de Biologia, Universidade de Évora, Pólo da Mitra, Apartado 94, 7002-554 Évora, Portugal
Received 04 April 2014
Accepted 10 July 2014
Published Online 15 September 2014 © ISSCA and authors 2014
We present an annotated catalog of the type specimens of amphibians and reptiles in the collections
of the Museu de História Natural da Universidade do Porto in Portugal. These specimens, all from
present-day Angola, formed the basis of taxonomic descriptions by both José Júlio Bethencourt
Ferreira and José Vicente Barbosa du Bocage in the latest 19th and early 20th century. We provide
details for all type specimens and summarize the history and taxonomy for each species. Specimens
of Rappia bocagei var. maculata and Typhlops bocagei could not be located during our survey,
and we believe these to be lost. The collections at the University of Porto contain type specimens
of one snake, Typhlops boulengeri, and eight frogs, Arthroleptis carquejai, Hylambates bocagei
var. leucopunctata, Rappia platyceps var. angolensis, Rappia bivittata, Rappia fasciata, Rappia
nobrei, Rappia osorioi, and Rappia seabrai. Of these, only two are currently recognized: Afrixalus
osorioi and Arthroleptis carquejai.
Luis M. P. Ceríaco1,2*, David C. Blackburn3, Mariana P. Marques4,
Francisco M. Calado4
The Museu de História Natural da Universidade do Porto, Portugal, and its collections date back to the
second half of the nineteenth century. These collections are one of the largest zoological collections in Portugal,
probably only surpassed by the collections of the zoological section of the Museu da Ciência da Universidade de
Coimbra, in Coimbra and by the collections of the Centro de Zooglogia do Instituto de Investigação Cientíca
Tropical, in Lisbon. Included in the Porto collections are type specimens of several different taxa. During recent
studies of the late 19th and early 20th century collections, we identied type specimens of amphibian and reptile
species described by José Júlio Bethencourt Ferreira (1866-1936) in the beginning of the twentieth century. These
specimens received little attention subsequent to Ferreira’s original descriptive work though they are relevant to
several nomenclatural and taxonomic problems. In addition, the collection contains other specimens that appear to
be syntypes or paratypes of taxa described by José Vicente Barbosa du Bocage (1823-1927) in the second half of the
nineteenth century. Besides these herpetological specimens, we also located other mammalogical, ornithological,
and invertebrate type specimens. Here we present the complete catalogue of the extant herpetological type
specimens and contribute to clarifying several related taxonomic and nomenclatural issues. A brief note about the
history of the Museum and its collections is also presented.
The Universidade do Porto, created in 1911, is the direct heir of the defunct Academia Politécnica do Porto.
Formally created in December of 1837, the Academia Politécnica replaced the former Academia Real de Marinha
e Comércio. In comparison to other Portuguese scientic establishments, the history of the Academia Politécnica
do Porto has been almost forgotten. Consequently, particular parts of this institution, including the zoological
class and museum, are almost unknown. Since its beginning, the Academia Politécnica lectures included a class
dedicated to the study of natural history that focused especially in zoology, mineralogy, geognosy, mining, and
metallurgy (Basto, 1938). The “7th class”, as it was known, was divided in two years. For the zoological subjects,
it was focused on the study of comparative anatomy and physiology, to the classication of animals by “natural
families”, and to the description and study of economically useful species, always following Cuvier’s classication
scheme (Basto, 1938).
Giving the study plan of the 7th class, collections were needed for students to observe and compare different
animal groups. Unfortunately, there is little information on the collection and related facilities that were available
at the time of the Academia Politécnica. In 1845, under the direction of Professor José Carneiro da Silva (1791-
1853) who was in charge of the 7th chair beginning in 1840, the collections available for the class were small,
with the majority being loaned to the Academia by a local patron to whose family it later returned after his death
(Machado, 1937). In the following years, due to the initiative of the congressman José da Silva Passos, some funds
were allocated to improve the scientic collection of the Academia Politécnica. Some of those funds were spent
acquiring natural history material from Parisian dealers (Machado, 1937). However, the small teaching collection
remained impoverished during the late nineteenth century until Augusto Pereira Nobre (1865-1947) arrived at
the Academia Politécnica. Nobre was born in Porto and studied natural sciences at Universidade de Coimbra, but
later transferred to the Academia Politécnica do Porto to nish his studies. In Porto, Nobre began private studies
in malacology in the collections of the Museu Allen. His interest in maritime biology led him to transfer to Paris
where he studied with Edmond Perrier in the Sorbonne University and the Muséum national d’Histoire naturelle,
as well as with Armand Sabatier in the Station de Biologie Marine de Sète of Montpellier University (Almaça,
1997; Santos & Eiras, 2006). After returning to Portugal in 1891, Nobre was nominated as an assistant professor
of Botany in the Academia Politécnica do Porto in 1890 and as an assistant of the Zoology chair with the charge of
rearranging and cataloguing the collections (Almaça, 1997; Santos & Eiras, 2006). In the following years Nobre
published catalogues of the collections of the Polytechnical Academy (Nobre, 1892, 1893a, 1893b, 1895, 1897,
1899, 1903, 1904) and dedicated himself to enriching the collections. For that, Nobre acquired collections from
major European natural history dealers such as the Parisian house Deyrolle, the Hamburg house Umlauff, and the
house Schlütter from Halle in Germany. He also acquired specimens from local Portuguese dealers like António
F. F. Mendes, private collectors like Francisco Rodrigues Batalha or Braga Júnior (offered to the museum), and
scientic institutions like the Marine Biological Association of the United Kingdom from Plymouth, from which
the museum acquired a large collection of marine invertebrates in 1905. In addition, Nobre relied on contributions
from private individuals and other national and international natural history institutions, such as the Museu
Nacional de História Natural de Lisboa that regularly offered specimens from 1890 to the middle of the twentieth
century, or in the case of Siegfried Joffe from Germany.
More importantly, the collections grew by eld expeditions aimed at collecting specimens for the museum,
both in Portugal and in overseas territories. These expeditions not only provided large series of collections to
complete the museum and help in teaching, but also brought considerable novelties to zoological knowledge. The
most signicant of these expeditions was that of the Portuguese naturalist and explorer Francisco Newton (1864-
1909) to Angola. Newton, a personal friend of Augusto Nobre since childhood (Nobre, 1945), had explored and
collected specimens in Portuguese overseas territories, as São Tomé e Príncipe, Cape Verde, and Timor, under
the orders of Barbosa du Bocage. The Angolan expedition that Newton undertook from 1903 to 1905 for the
Academia Politécnica do Porto, resulted in a considerable collection of specimens from the most diverse groups:
insects, crustaceans, arachnids, shes, amphibians, reptiles, birds, and mammals. After arriving in Porto, part of
this collection was sent to Lisbon in 1904, where some of it was prepared and other portions sent to be studied by
specialists. That was the case of the collection of amphibians and reptiles made by Newton that was sent to José
Júlio Bethencourt Ferreira. At that time, Bethencourt Ferreira was the curator of the herpetological collections in
the zoological section of the Museu Nacional de História Natural de Lisboa, under the supervision of Barbosa du
Bocage. Nobre charged Ferreira with the task of identifying and classifying the collections made by Francisco
Newton in Angola and to return it to Porto after the work was completed (Ferreira, 1904, 1906). The rst shipment
sent by Nobre to Ferreira on 20th July 1904 contained the herpetological specimens collected by Newton in 1903
in the northern region of Kwanza river (Ferreira, 1904). It contained fourteen anuran taxa of which Ferreira
considered two to be new (Rappia nobrei, Hylambates bocagei var. leucopunctata), eleven snake species of which
one new taxon was described (Typhlops bocagei), and eight lizard species. In 1906, Ferreira described an additional
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Figure 1. Plate with specimen photography as presented by Ferreira (1906). Reproduced from the original.
seven new anuran taxa, including ve new species (Rappia bivittata, Rappia osorioi, Rappia seabrai, Rappia
fasciata, and Arthroleptis carquejai) and two new varieties (Rappia bocagei var. maculata and Rappia platyceps
var. angolensis) (Ferreira, 1906). Besides these new taxa, Ferreira (1906) provided records for species of another
seventeen frogs, fourteen snakes, and seven lizards. The paper was accompanied by a plate with photographs of
several of the newly described taxa (g. 1). Part of the collection made by Newton in Angola between 1903 and
1905 was only returned to Porto in November of 1908, following offers of other herpetological specimens from
the Museu Nacional de História Natural de Lisboa (also known since 1906 as Museu Bocage) to Porto, including
one rare specimen of the Cape Verde giant skink, Chioninia coctei (AHMB Div. 519-10; Ceríaco, in press). In
Lisbon, Antero de Seabra studied the collections of birds and mammals that had been sent there for preparation
(Seabra, 1905a, 1905b, 1906a, 1906b, 1906c, 1906d, 1906e, 1907). In March 1921, the Museum together with
the Estação de Biologia Marinha, the Laboratório de Entomologia Económica, and the general laboratory of the
Universidade do Porto were incorporated under the “Instituto de Zoologia” of Universidade do Porto. Following
the death of Augusto Nobre, who maintained the directorship of the Institute almost until his death, the museum
largely fell into a state of disrepair. Subsequent directors did not have the same dynamic inuence as Nobre, and
the collections and museum stagnated with time. Today the museum is closed to the public, though there are plans
for its reorganization and opening. The collections of the museum are displayed in two main rooms: the Portuguese
room and the General collections rooms maintaining almost the original display of the beginning of the twentieth
century (g. 2). This situation leads to the paradoxical question of how to preserve both the collections and the
museum itself. The early twentieth century display is not the best solution for preserving old and fragile specimens,
because of the excess light, fragile structure of the closets, and inability to control climate and pests effectively.
Yet, the display remains of historical value because it is a rare example of a natural history exhibition from the
beginning of the twentieth century. This collection is thus of unique importance to those interested in biodiversity
science as well as the history of science.
In general, many Portuguese collections require updated systematic organization. In addition, at least
one important collection was tragically destroyed the Bocage collections housed in Lisbon were lost in the
catastrophic re of 1978. Because of this, investigations of taxa described by Portuguese naturalists and the
relevant type specimens have been difcult over the past century. One of the few such studies based on Portuguese
collections is that of Perret (1976a), which was made before the catastrophic re and based on the amphibian
type specimens of Lisbon museum (Almaça & Neves, 1987). There are a variety of problems facing specimens
preserved in alcohol or formaldehyde in these collections. It is frequently difcult, if not impossible, to evaluate
the taxonomic status of older formalin- or alcohol-preserved specimens because they are often damaged, have
been dehydrated, or have faded colouration and patterns because of extensive exposure to light. However, their
study is important for addressing taxonomic, and nomenclatural queries. This is the situation with some amphibian
species described by Ferreira based on the Porto collections made by Francisco Newton (Ferreira, 1904, 1906).
During current investigations in the Museu de História Natural da Universidade do Porto, most of Ferreira’s anuran
types were located and identied. With this annotated type catalogue, we catalog the extant herpetological type
specimens present in the Porto collections, and comment on several outstanding taxonomic and nomenclatural
Figure 2. Picture of the General room of the Museu de História Natural da Universidade do Porto at the beginning of the twentieth century
(left) and actual picture of the same room (right). Credits: MHNFCP (left) and Luis Ceríaco (right).
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This published work has been registered in ZooBank. The ZooBank LSIDs (Life Science Identiers)
can be resolved and the associated information viewed through any standard web browser. The LSID for this
publication is:
The catalogue follows a similar structure to those published for the Institut für Systematische Zoologie
of the Museum für Naturkunde der Humboldt-Universitat zu Berlin (e.g., Bauer et al., 2006). For each taxon,
we provide the original name followed by the author, date of publication and page; the holotype or syntype
catalog numbers in the Museu de História Natural da Universidade do Porto (MHNFCP), along with information
derived from the jar labels, catalogue entries, and published descriptions about the collector, locality and date of
collections; the current name for the species; and other general remarks. For all of the holotypes or syntypes, we
present a colour photograph of the preserved specimen. In some cases, we referred to available correspondence in
the Arquivo Histórico do Museu Bocage (AHMB) at the Museu Nacional de História Natural e da Ciência, Lisboa
for clarifying particular details regarding specimens in the MHNFCP collections.
For the identication and clarication of problematic taxa, we conducted more detailed morphological
study. For these specimens, the following measurements were taken with a digital caliper following Bossuyt &
Dubois (2001): Snout-vent length (SVL), head width at the angle of jaws (HW), head length from the posterior
corner of mandible to tip of snout (HL), distance from posterior corner of mandible to the nostril (MN), distance
from the posterior corner of mandible to anterior corner of eye (MFE), distance from the posterior corner of
mandible to posterior corner of eye (MBE), distance between the anterior corner of eyes (IFE), distance between
the posterior corner of eyes (IBE), forelimb length from the elbow to base of outer palmar tubercle (FLL), hand
length from base of outer palmar tubercle to tip of third nger (HAL), inner toe length (ITL), third nger length
from base of rst subarticular tubercle (TFL), lower leg (crus, or tibiobula) length (TL), thigh (femur) length
from vent to knee (FL), foot length from base of inner metatarsal turbercle to tip of the fourth toe (FOL), fourth
toe length from base of rst subarticular tubercule (FTL), internarial distance (IN), distance from the anterior
corner of eye to nostril (EN), eye length (EL), distance from nostril to tip of snout (NS), maximum tympanum
diameter (TYD), and the distance between the tympanum and posterior corner of eye (TYE). Some specimens are
in poor condition, are still or were once preserved in formaldehyde, and/or are small and fragile, all of which inhibit
DNA-based or other detailed study. We compared the type material with specimens present in the collections of the
Locality in publication Corresponding taxa Current locality name Latitude Longitude Elevation
N’golla Bumba Rappia bivittata
Rappia platyceps var. angolensis
N’Golla Bumba
(Cuanza Norte Prov.) 09° 02'S 14° 36'E 750
"Quilombo" or
"Quilombo, Rio Luinha"
Rappia bivittata
Rappia fasciata
Rappia osorioi
Rappia platyceps var. angolensis
Rappia seabrai
(Cuanza Norte Prov.) 09° 20'S 14° 54'E 750-800
Rio Luinha Rappia bivittata Rio Luinha
(Cuanza Norte Prov.) 09° 16'S 14° 32'E 250
Cabiri Rappia nobrei Cabiri (Luanda Prov.) 08° 55'S 13° 40'E 60
Duque de Bragança Rappia seabrai Kalandula (Malanje Prov.) 09° 06'S 15° 57'E 1110
Gumba, Sa de Selles Hylambates bocagei var. leucopunctata Gumba
(Cuanza Sul Prov.) 11° 27’S 14° 29’E 1100
Cambondo Arthroleptis carquejai Cambondo
(Cuanza Norte Prov.) 09° 29'S 16° 38'E 1150
Quindumbo Typhlops boulengeri Quindumbo
(Benguela Prov.) 12° 28'S 14° 56'E 1350-1450
Golungo Alto Rappia bocagei var. maculata Golungo Alto
(Cuanza Norte Prov.) 09° 08'S 14° 46'E 630
Cacuaco (see account for
Rappia bocagei var. maculata)
(Luanda Prov.) 08° 47'S. 13° 22'E 40
Cabicula, Bom Jesus Typhlops bocagei Bom Jesus (Bengo Prov.) 09° 10'S 13° 34'E 87
Table 1. Localities for MHNFCP type specimens, with updated names (including province) and estimated latitude, longitude, and elevation.
California Academy of Sciences (CAS; San Francisco, USA), the Muséum national d’Histoire naturelle (MNHN;
Paris, France) and Zoologisches Museum für Hamburg (ZMH; Hamburg, Germany); for details on the specimens
examined, see Appendix 1. The results of these comparisons are presented in the general remarks for each taxon.
For identications, we relied especially on Schiøtz (1999), Noble (1924), Frétey et al. (2012), and Amiet (2012).
Geocoding of historical localities follows Cabral & Mesquitela (1989). Tab. 1 presents the list of type localities,
current locality names, and estimated latitude, longitude, and elevation.
Extant types
Rappia bivittata Ferreira, 1906
Original name. Rappia bivittata Ferreira, 1906: 161.
Type specimens. The three syntypes designated by Ferreira (1906) still exist in Porto. These are represented
by one specimen from “N’golla Bumba” (MHNFCP 017291, ♂, SVL 19.1 mm), one from “Quilombo”(MHNFCP
017296, Juv., SVL ≈ 14 mm), and one from “Rio Luinha” (MHNFCP 017302, ♂, SVL 19.7 mm), all of which are
from the Francisco Newton expedition to Angola in 1903-1905 (g. 3). Measurements of the type specimens are
presented in tab. 2 with exception of MHNFCP 017296 which is poorly preserved and so could not be measured
Present name. Hyperolius platyceps (Boulenger, 1900).
Figure 3. Dorsal and ventral views of the three syntypes of Rappia bivittata Ferreira, 1906. Top left: MHNFCP 017302 from Rio Luinha,
Angola. Top right: dorsal and ventral views of MHNFCP 017291 from N’golla Bumba, Angola. Below: Dorsal and ventral views of MHNFCP
017296, which is in poor condition because of past dehydration, from, Quilombo, Angola. Scale bar = 10 mm.
ALYTES 2014 | 31
Remarks. Ferreira’s (1906:161-162) description of Rappia bivittata was based on three syntype specimens.
Noble (1924) proposed the name Hyperolius ferreirai because the name Hyperolius bivittatus was preoccupied
by Hyperolius bivittatus Peters, 1865; note that Noble (1924) referred to this taxon as Rappia bivittatus and not
bivittata. Noble (1924) did not consult the specimens in Porto, but acknowledged that the species was only known
from the type specimens. The name Hyperolius ferreirai has been maintained to the present. Laurent (pers. comm.,
in Frost, 1985) suggested that Hyperolius ferreirai may be a synonym of Hyperolius platyceps Boulenger, 1900.
More recently, Frétey et al. (2011), following Laurent in Frost (1985) grouped Hyperolius ferreirai as a synonym
of Hyperolius platyceps.
The colouration patterns of two best preserved syntypes are similar. Both present a pair of pale dorsolateral
stripes extending from the anterior margin of the eye to the insertion of the leg, and some small dark spots
are still visible on the dorsum especially in specimen (MHNFCP 017302). Both H. platyceps and Hyperolius
cinnamomeoventris Bocage, 1866, exhibit this pattern, although H. cinnamomeonvetris usually presents a darker
band separating the dorsolateral colouration from that of the venter (Schiøtz, 1999; Amiet, 2012). While this
band is currently not visible in specimen MHNFCP 017291, Ferreira (1906) noted it in his description of Rappia
The morphology of the head also differs beween these two syntypes, with a more accuminate head (HW/HL
of 92%) in MHNFCP 017291 and a broader head (HW/HL of 109%) in MHNFCP 017302. Amiet (2012) used both
the wider head and shorter snout of H. platyceps as useful diagnostic features relative to H. cinnamomeoventris,
and these two syntypes of Rappia bivittata also differ in this way (tab. 2). Another important difference noted
by Amiet (2012) between these species is the presence of granulations on the palmar and plantar surfaces of H.
platyceps and their absence in H. cinnamomeoventris, which again is similar to the differences between MHNFCP
017302 and 017291, respectively. Based on our comparisons and study of both the best preserved syntypes, we
conclude that these represent two different species. The specimen from N’Golla Bumba (MHNFCP 017291)
seems to represent Hyperolius cinnamomeoventris Bocage, 1866, while the specimen from Rio Luinha (MHNFCP
Rappia platyceps
var. angolensis
017303 (n = 12) (n = 14)
SVL 19.05 19.65 25.13 26.21 21.27-26.98 (23.98 ± 1.67) 21.46-29.29 (23.87 ± 2.18)
HW 5.59 6.38 8.46 9.69 7.66-8.98 (8.24 ± 0.52) 6.33-8.88 (7.17 ± 0.68)
HL 6.07 5.85 8.44 7.68 7.44-9.06 (8.15 ± 0.57) 7.06-9.13 (7.82 ± 0.60)
HW/HL 92 109 100 126 98-105 (101 ± 2.14) 83-101 (92 ± 5.60)
EN 1.55 1.43 2.60 2.37 1.48-2.53 (2.13 ± 0.39) 1.98-3.00 (2.94 ± 0.33)
IFE 2.63 2.71 5.06 4.85 4.05-4.98 (4.53 ± 0.31) 3.53-5.09 (3.98 ± 0.42)
EN/IFE 58 52 51 48 36-62 (47 ± 7.84) 50-62 (58 ± 4.20)
IBE 3.44 5.39 7.95 8.06 6.92-8.27 (7.54 ± 0.43) 5.41-7.96 (6.64 ± 0.58)
FLL 4.22 3.85 5.57 5.46 4.44-5.80 (5.25 ± 0.49) 4.60-6.31 (5.07 ± 0.50)
HAL 3.40 4.39 6.23 6.74 6.00-7.46 (6.76 ± 0.51) 5.35-7.06 (6.28 ± 0.51)
ITL 2.07 1.52 2.55 3.26 2.72-3.76 (3.29 ± 0.30) 2.35-3.69 (2.98 ± 0.38)
TFL 1.45 2.80 5.42 4.79 3.69-4.95 (4.14 ± 0.39) 3.20-4.29 (3.95 ± 0.44)
FL 8.43 7.90 11.62 12.89 10.89-13.57 (12.25 ± 0.91) 8.75-11.66 (10.87 ± 1.18)
FOL 7.75 7.26 10.58 10.27 8.37-11.87 (10.40 ± 1.07) 8.26-16.73 (9.90 ± 1.97)
FTL 4.51 5.43 6.71 8.66 6.04-8.50 (7.19 ± 0.78) 4.74-7.20 (5.42 ± 0.83)
IN 1.19 1.34 1.98 2.58 1.69-2.43 (2.14 ± 0.22) 1.42-2.09 (1.77 ± 0.20)
EL 2.53 1.99 3.02 3.94 2.89-4.03 (3.50 ± 0.37) 2.36-3.39 (2.94 ± 0.33)
Table 2. General comparisons between Rappia bivittata, Rappia fasciata, and Rappia platyceps var. angolensis type specimens with Hyperolius
platyceps and Hyperolius cinnamomeoventris specimens (see Appendix 1). When n specimens > 1 measurements are provided as range,
followed by mean ± standard deviation. Measurements taken in mm; ratios in %.
017302) represents Hyperolius platyceps Boulenger, 1900.
According to the rules of the International Code of Zoological Nomenclature (ICZN, 1999), a nomen
cannot stand in two synonymies, except in the specic case in which holotypes represent hybrids. We designate
specimen MHNFCP 017302 as the lectotype of Rappia bivittata. Thus, Rappia bivittata and Hyperolius ferrerai
are unambiguously junior synonyms of Hyperolius platyceps Boulenger, 1900, as already suggested by Laurent
(pers. comm., in Frost, 1985), Schiøtz (1999), and Frétey et al. (2011). The two remaining syntypes, MHNFCP
017291 and 017296, should be considered paralectotypes.
Rappia fasciata Ferreira, 1906
Original name. Rappia fasciata Ferreira, 1906: 164.
Type specimen. A single specimen (MHNFCP 017294, ♀, SVL 25.1 mm) from “Quilombo”, captured by
Francisco Newton in Angola during the 1903-1905 expedition and still present in Porto (g. 4).
Present name. Hyperolius platyceps (Boulenger, 1900).
Remarks. Ferreira’s (1906:164) description of Rappia fasciata was based on a single specimen from
Francisco Newton’s Angola collection. To date, this taxon remains known only from the type locality. Laurent (pers.
comm. in Frost, 1985) suggested that Hyperolius fasciatus may be a synonym of Hyperolius platyceps Boulenger
1900, and this was followed recently by Frétey et al. (2011). Our comparison of the holotype with specimens of
both Hyperolius platyceps and H. cinnamomeoventris, a sometimes similar species, conrms the suspicions of
previous authors that this is a synonym of H. platyceps. These similarities are born out in measurements of relevant
specimens (tab. 2).
The general shape of the head and body of the holotype of Rappia fasciata is similar to that observed in H.
platyceps (Amiet, 2012). Similar to H. platyceps, HL is approximately equal to HW, and the relative proportions
of the snout (EN/IFE) are closer to the values observed in H. platyceps than in H. cinnamomeoventris.
The holotype of Rappia fasciata presents characters that allow it to be identied specically as the
pleurotaenia morph of H. platyceps (Frétey et al., 2011; Amiet 2012). These characters include its stout body, large
head with a rounded snout, and prominent dorsolateral stripes. Amiet (2012) used both the wider head and shorter
snout of H. platyceps as diagnostic features relative to H. cinnamomeoventris, and both characteristics are present
in the holotype of Rappia fasciata. As also indicated by Amiet (2012), the attened granulations on the palmar and
plantar surfaces also indicate that the holotype is more similar to H. platyceps than H. cinnamomeoventris.
Figure 4. Dorsal and ventral view of the holotype Rappia fasciata Ferreira, 1906 (MHNFCP 017294). Scale bar = 10 mm.
ALYTES 2014 | 31
Rappia nobrei Ferreira, 1904
Original name. Rappia nobrei Ferreira, 1904: 112.
Type specimens. Two syntypes (MHNFCP 017292, ♀, SVL 19.7 mm; 017298, ♀, SVL 19.5 mm) collected
by Francisco Newton in Angola during the 1903-1905 expedition and still in Porto, both from “Cabiri, Angola”
(g. 5). This species is a patronym for Augusto Nobre (1865-1946), director of the Museu de História Natural of
the Academia Politécnica do Porto.
Present name. Hyperolius cf. adspersus Peters, 1877.
Remarks. In part because of the long lack of study of these type specimens, Frost (2014) considers this
taxon as incertae sedis. Our study of the syntypes conrms that this is clearly a member of the genus Hyperolius.
Based on both geographical distribution and measurement data (tab. 3), it is plausible that this taxon is a junior
synonym of H. adspersus Peters, 1877, which was described based on a specimen also collected near the Atlantic
coast at Chinchoxo, Cabinda (Peters, 1877), approximately 455 km NW of Cabiri. Hyperolius adspersus is part
of the taxonomically problematic Hyperolius nasutus group, which currently contains 16 recognized species
(Channing et al., 2013). Of these 16 species, only three are known to occur near the type locality of Rappia nobrei:
Hyperolius adspersus Peter, 1877, Hyperolius nasutus Günther, 1865, and H. benguellensis (Bocage, 1893) (see
Channing et al., 2013). Both syntypes of R. nobrei fall within the range of SVL for H. adspersus (SVL 18.1-21.6
mm; Amiet, 2012) but also H. benguellensis (SVL 19-24 mm; Poynton & Broadley, 1987) and present a rather
large and blunt head, a character shared by both species. The pedal webbing of the R. nobrei syntypes is also
similar to both H. adspersus and H. benguellensis. While the two syntypes currently lack distinct pigmentation,
Figure 5. Dorsal and ventral view of the two syntypes of Rappia nobrei Ferreira, 1904 (Top specimen: MHNFCP 017292; lower specimen:
MHNFCP 017298). Scale bar = 10 mm.
the original description by Ferreira (1904) notes the presence of dark
pigmentation on the dorsum, especially on the head and middle of
the dorsum, with a whitish venter. Unfortunately, the available data
for H. adspersus and H. benguellensis as well as the preservation of
the syntypes make it difcult to determine to which species Rappia
nobrei should be referred.
Rappia osorioi Ferreira, 1906
Original name. Rappia osorioi Ferreira, 1906: 162.
Type specimens. Three syntypes from “Quilombo, Angola”
(MHNFCP 017307, ♂, 2 juveniles, SVL 26.3, 24.6, 17.6 mm
respectively) collected by Francisco Newton in Angola during the
1903-1905 expedition (g. 6). This species is a patronym for Baltasar
Osório (1855-1926), ichthyologist, carcinologist, and director of the
Zoological section of the Museu Nacional de História Natural de
Lisboa (Museu Bocage).
Present name. Afrixalus osorioi (Ferreira, 1906).
Remarks. Ferreira’s (1906:162) description of Rappia osorioi
was based on three specimens, one male adult and two juveniles,
from “Quilombo”. The species is considered valid and is broadly
distributed from northwestern Angola and across much of the Congo
Basin (Schiøtz, 1974; Perret, 1976b; Laurent, 1982; Channing, 2001),
and perhaps even Kenya (Köhler et al., 2005). Laurent (1972, 1982)
discussed patterns of morphological variation in A. osorioi and the
minor phenotypic differences from two other Congo Basin species,
A. equatorialis and A. leucostictus. In his discussion of the colour
pattern variation of A. osoroi, Laurent (1982) notes that “la phenotype
représenté par l’holotype” is a somewhat rectangular elongate and
somber scapular spot. However, the citation provided in this discussion
(Laurent, 1941), as well as his list of specimens examined, indicates
that this phenotype is that of the holotype of Megalixalus fornasinii
congicus Laurent, 1941 and not type material of Afrixalus osoroi.
Perret (1976b) lists three type specimens in Porto museum lacking
catalog numbers (one male holotype and two paratypes, a male and
a juvenile) and followed Laurent (1972) in recognizing this taxon as
conspecic with Megalixalus fornasinii congicus.
The existence of two different jars at MHNFCP containing specimens identied as possible types of
osorioideserves further comment. One jar (MHNFCP 017307) is labeled Rappia osorioi B. F. / Quilombo
Newton” and contains one adult and two juveniles. In addition, the jar also contains two paper notes. The rst is
written in pencil (most probably by Bethencourt Ferreira) and states “Rappia osorioi / n. sp. B. F. / Typo ♀ ado.”,
whereas the second note is written in ink with the same caligraphy but on the revserse of a label from the Lisbon
museum and states “Rappia osorioi BF / Typo ♀ N’Golla Bumba / Angola Newton”. Last, there is a third typed
note stating: Afrixalus osorioi osorioi (Ferreira) / Holotype: male / Paratypes: femelle et deux jeunes”. The second
jar (MHNFCP 017301) contains one adult specimen and the external label notes “Hyperolius platyceps fasciatus
(Ferreira) / F Newton”. However, inside the jar is a typed note stating “Paratype de Rappia osorioi Ferreira / est
en fait un Hyperolius platyceps fasciatus (Ferreira)”. Given that Laurent (1972) and Perret (1976b) examined
specimens in both jars, it is plausible that these typed notes were written by one of these authors.
Given the poor state of preservation of MHNFCP 017301, that Ferreira (1906) noted only three specimens
of Rappia osorioi, and that the only indication that MHNFCP 017301 is a paratype is based on a type-written note,
we choose to not consider it as a type specimen. As the original description mentions specically an adult male
and two juveniles from Quilombo, this additional “adult female type” from N’Golla Bumba cannot be considered
a syntype (ICZN, 1999).
Rappia platyceps var. angolensis Ferreira, 1906
Original name. Rappia platyceps var. angolensis Ferreira, 1906: 161.
Type specimens. One syntype from “Quilombo” (MHNFCP 017303, ♀, SVL 26.2 mm), collected by
SVL 19.7 19.5
HW 6.46 5.91
HL 6.94 6.13
MN 5.76 5.23
MFE 4.01 3.36
MBE 2.32 1.44
IFE 3.10 2.42
IBE 4.41 4.34
FLL 3.95 4.02
HAL 4.33 4.33
TFL 2.40 2.32
TL 6.19 6.16
FOL 7.64 8.73
FTL 5.50 6.59
IN 1.79 1.84
EN 1.80 1.61
EL 2.45 1.94
ITL 2.37 2.56
FL 8.82 9.81
SL 2.23 1.94
NS 0.67 0.75
Table 3. Measures (in mm) of both syntypes
of Rappia nobrei Ferreira, 1904.
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Figure 7. Dorsal and ventral views of the syntype of Rappia platyceps var. angolensis Ferreira, 1906 (MHNFCP 017303). Scale bar = 10 mm.
Figure 6. Dorsal and ventral views of the syntypes of Rappia osorioi Ferreira, 1906 (catalogued as a lot, MHNFCP 017307). Scale bar = 10 mm.
Francisco Newton in Angola during the 1903-1905 expedition and still present in Porto (g. 7). The other syntype
from “N’golla Bumba” was not located.
Present name. Hyperolius platyceps (Boulenger, 1900).
Remarks. The genus Hyperolius Rapp, 1842 contains more than 140 recognised species (Frost, 2014) that
display a wide breadth of phenotypic variation both within and between species that often complicates species
identication and delimitation (Frétey et al., 2011). Some hyperoliid species are characterized by remarkable
polymorphism (Hoffman & Blouin, 2000) that is in some cases reected by the number of nominal subspecies
(Wieczorek et al., 1998, 2000, 2001) and many junior synonyms. Many of the subspecies (and species) may in
fact represent colour variants of a single taxon (Kohler et al., 2005). However, in addition, genetic analyses have
revealed that many nominal taxa comprise two or more cryptic species (Rödel et al., 2002).
Ferreira’s (1906:161) description of Rappia (now Hyperolius) platyceps var. angolensis was based on
two specimens and Ferreira considered it rare. Ahl (1931) recognized this taxon as a species and also proposed
the replacement name Hyperolius angolanus because of the previously described Hyperolius marmoratus var.
angolensis Steindachner, 1862. More recently, Frétey et al. (2011), following Laurent (pers. comm., in Frost,
1985), recognized Hyperolius platyceps angolensis as a synonym of Hyperolius platyceps, although they made no
reference to examinations of the type specimens of H. platyceps angolensis. Our comparisons with other specimens
of H. platyceps conrm that the specimen is indeed a member of this species. More specically, it resembles the
“platyceps” morph because of the “hour-glass” pattern present on the dorsum.
Rappia seabrai Ferreira, 1906
Original name. Rappia seabrai Ferreira, 1906: 163.
Type specimens. One paratype from “Duque de Bragança” collected by “capitão Bayão” (Captain
Francisco António Pinheiro Bayão) in 1865 and still present in Porto (MHNFCP 018587, ♂, SVL 25.0 mm; g.
8). Ferreira (1906: 163) noted this additional specimen located in the Lisbon collections in his description and
thus we consider it a paratype. The Bayão collections, mostly from Duque de Brangança, were among the rst and
most important of Angolan herpetological collections as they served as the basis for taxonomic description of a
number of taxa by Bocage, Steindachner, and Günther. Ferreira apparently sent this specimen to the Porto museum
following his sending of the holotype. Unfortunately, the holotype from “Quilombo, Rio Luinha”, which was
collected by Francisco Newton in Angola during the 1903-1905 expedition, was not found during our survey and
we believe that it may be lost. The species is a patronym for Antero de Seabra (1883-1938), a mammalogist and
entomologist, and curator of the Zoological section of the Museu Nacional de História Natural de Lisboa (Museu
Present name. Hyperolius bocagei (Steindachner, 1867).
Figure 8. Dorsal and ventral views of the syntype of Rappia seabrai Ferreira, 1906 (MHNFCP 018587). Scale bar = 10 mm.
ALYTES 2014 | 31
Remarks. The species is currently recognized as valid (Frost, 2014), though no further data or records
of the species have been presented subsequent to its description. This species might be a synonym of Hyperolius
bocagei Steindachner, 1867 (Laurent pers. comm. in Frost, 1985; Frétey et al., 2011). The general morphological
characters noted by Ferreira (1906) agree with the original description of H. bocagei (Steindachner, 1867) and
subsequent descriptions (Bocage, 1895; Schiøtz, 1999), including granulations on the head, a truncated snout, the
presence of small terminal discs on the ngers, and a nearly complete pedal webbing. The SVL of the species,
21 mm in the holotype according to the original description, 25 mm in the the paratype, falls within the range of
the SVL H. bocagei (Schiøtz, 1999). Because of the poor condition of the available specimen, it is not possible
to discern its colouration and patterns. Based on the original description, Rappia seabrai presents a greyish
colouration on the back, with small darks spots dispersed dorsolaterally and on the limbs (especially the crus), a
yellowish venter, and a dark line beginning at the nostrils and extending to the insertion of the upper arm (Ferreira,
1906). This description generally agrees with the colouration of H. bocagei (Steindachner, 1867; Bocage, 1895;
Schiøtz, 1999). The type locality of Quilombo, currently Gonguembo, Cuanza Norte province, is approximately
140 km west of Kalandula falls at Malanje (formerly Duque de Brangança), which is both the type locality of
H. bocagei and the locality from which the paratype of R. seabrai was collected. Based on available evidence
we follow Frétey et al. (2011) by suggesting that R. seabrai is as a junior synonym of H. bocagei. However, as
indicated by Schiøtz & Von Daele’s (2003) suggestion that H. bocagei may be a junior synonym of the Hyperolius
viridiavus complex, there clearly is need for further study to resolve the systematics of these taxa.
Hylambates bocagei var. leucopunctata Ferreira, 1904
Original name. Hylambates bocagei var. leucopunctata Ferreira, 1904: 113.
Type specimen. Syntype (MHNFCP 017324, ♂?, SVL 25.1 mm ), from “Gumba, Sa [Serra] de Selles =
Angola” collected by Francisco Newton during the 1903-1905 expedition in Angola (g. 9). Ferreira (1904) notes
two syntypes, one adult and one juvenile from the same locality, collected by Francisco Newton in his expedition
to Angola. The second syntype may be lost (but see below).
Present name. Leptopelis bocagii (Günther, 1865).
Remarks. Leptopelis bocagii is a widespread species in southern and central Africa (Channing, 2001),
though whether all populations currently referred to this species are truly conspecic remains unclear (i.e., Amiet,
2012). In a review of the amphibians of the Zambezi region, Poynton & Broadley (1987) described a similar
species, Leptopelis parbocagii Poynton & Broadley, 1987 based on ve specimens collected at Mabwe on the
eastern shore of Lake Upemba, Zaire (currently the Democratic Republic of Congo; DRC). The distribution of
Leptopelis parbocagii overlaps that of L. bocagii which extends east from Angola and DRC to Malawi, Zambia,
and Mozambique, though doubts remain about the identication of these species (Schiøtz, 1999; Schiøtz & Von
Daele, 2003). Poynton & Broadley (1987) used the ratio of the interorbital distance to the nostril-tympanum
distance as a diagnosis for these two species: in L. bocagii this ratio is less than 36%, whereas in L. parbocagii it
is 36% or more. Because this ratio in MHNFCP 017324 is 33%, we refer it to L. bocagii rather than L. parbocagii.
Subsequent to the description of Hylambates bocagei var. leucopunctata, Ferreira (1906) referred to three
specimens of Hylambates bocageifrom Rio Luinha, Quilombo (currently Gonguembo), and N’golla Bumba.
Note that Ferreira (1904, 1906) mispelled the epithet bocagii, writing it always as bocagei; thus in citing Ferreira
(1904, 1906) directly we use his spelling of “bocagei”. In addition to the syntype located during this survey,
we found three additional specimens, one adult from “N’Golla Bumba” (MHNFCP 017323) and another adult
(MHNFCP 017325) for which the locality data is uncertain because of a damaged label. Ferreira (1906) notes two
adult specimens of Hylambates bocagei, one from “Rio Luinha, Quilombo” and the other from “N’Golla Bumba”,
which likely correspond to these two additional specimens. Among other specimens at MHNFCP identied as
Hylambates bocagei, there is a jar containing a juvenile specimen (MHNFCP 017326) bearing an original label
stating “Hylambates bocagei [torn area] … erreira.” It is possible that the missing portion would have stated “var.
leucopunctata Ferreira” and thus this might represent the missing juvenile syntype.
There are several of other records of Leptopelis bocagii from Angola. Laurent (1953) cited ten specimens
from Muita, Luembe and as well as later citing two others from Dundo (Laurent, 1954). Because Hylambates
angolensis, which was described by Bocage (1893) from a type series collected by José de Anchieta (1832-1897)
from the district of Benguela, is considered a synonym of L. bocagii (Frétey et al., 2011; Amiet, 2012), the known
distribution of L. bocagii is extensive. Ferreira (1904) considers his “var. leucopunctata” as “a transition between
H. viridis and H. bocagei” and distinguished this new taxon by the presence of a “very extensive and protruding
dorsal lateral fold” and by “the stronger development of the terminal disks of the ngers” (our translation). Based
on our examination, the principal morphological feature suggesting that “var. leucopunctacta” is distinctive is the
expansion of the terminal disks of the ngers, as noted by Ferreira (1904). While suggestive given the typically
non-expanded digits of Leptopelis bocagii, the absence of interdigital webbing clearly places the remaining syntype
with this species. In addition, the geographic distribution of Ferreira’s taxon occurs on the northwestern margin of
the wide geographic range of L. bocagii. Based on available evidence, we follow Amiet (2012) and Frétey et al.
(2011) by suggesting that Hylambates bocagei var. leucopunctata should remain a synonym of L. bocagii.
Doubts exist regarding the history of the type specimens of Leptopelis bocagii (Amiet, 2012) which
was described based on specimens from “Duque de Bragance” (now Malanje) in Angola. This uncertainty arises
because of apparent contradictions between the accounts of Günther (1864) and Bocage (1895), and demands
attention here. During the course of our work, we examined the correspondence between Günther and Bocage in an
attempt to reconstruct the history of the type specimen. Albert Günther (1864) described “Cystignathus bocagii
based on a single specimen from “Duque de Bragance (Angola)” loaned by Barbosa du Bocage (1923-1907),
director of the zoological section of the Museu Nacional de História Natural de Lisboa (Gray, 1864). In 1864,
Bocage loaned this specimen, surely collected by Francisco Pinheiro Bayão, who sent several collections from
the Duque de Bragança region to Bocage in 1863 (AHMB CN B19; AHMB CN B20; AHMB CN B21a; AHMB
CN B21b), to Günther to assist with its identication (AHMB CE G75, see Appendix 2). After his examination
of the material and published description of the species, based on a single specimen (AHMB CN B32), Günther
returned the specimen to Bocage in July 1865 (AHMB CE G76, See Appendix 2). Subsequent to the publication
of the species description, Bocage wrote a letter of thanks to Günther for naming the species after him (20 June
1865; AHMB CN B32).
On 25 May 1866, Bocage wrote again to Günther noting that he had a specimen of Leptopelis natalensis
that he thought could be “C. [Cystignathus] bocagii”, from another (unknown) location (AHMB CN B32). In the
same month Bocage sent a loan of the type and a second specimen of the species to Günther (AHMB CE G78, see
Appendix 2). After receiving the specimens, Günther wrote asking Bocage if he could retain the second specimen
in his collection (AHMB CE G79, see Appendix 2). Bocage apparently agreed to this since he later (Bocage,
1866) noted that the Zoological Section of the Museu Nacional de História Natural de Lisboa had two specimens
of “Cystignatus Bocagii” from the Duque de Bragança, both collected by Bayão. Bocage also indicated that one
of these was the type specimen, while the other was given to the British Museum. In 1895, Bocage again refered
to this species noting that the only known specimens of the species were “two types” in the British Museum and
that there were two other juvenile specimens in the Angolan collections of the Museu Nacional de História Natural
de Lisboa. The statement made by Bocage (1895) is problematic because it is unclear whether either of the two
specimens that Bocage sent to Günther in 1866, one of which was certainly the holotype, later returned to Lisbon
Figure 9. Dorsal and ventral views of the holotype of Hylambates bocagei var. leucopunctata Ferreira, 1906 (MHNFCP 017324). Scale bar = 10 mm.
ALYTES 2014 | 31
or if one or both instead remained in London. The timeframe of both collection and loan of the two other juveniles
mentioned by Bocage (1895) also remains unclear. The last African type catalog published by Bocage in 1897
stated only that a type of “Hylambates Bocagii” was in the Lisbon collections and noted the locality as “Angola:
Duque de Bragança, Bayão” (Bocage, 1897). Perret (1976a) claimed to have found the “cotype” of Hylambates
bocagii from “Duque de Bragança, Angola” (No. T. 15-232), a juvenile specimen. This specimen, lost in the re
that destroyed the museum in 1978, was surely one of the two juvenile specimens cited by Bocage (1895), but
it seems unlikely that this specimen was the holotype given the earlier the correspondence between Günther and
Bocage. Further investigations in the archives and collections of the Natural History Museum of London are
needed to address this problem.
Arthroleptis carquejai Ferreira, 1906
Original name. Arthroleptis carquejai Ferreira, 1906: 165.
Type specimen. Holotype (MHNFCP 018586, ♀ , SVL 27.8 mm) from “Cambondo, Angola” collected
by Francisco Newton during the 1903-1905 expedition in Angola (g. 10). The species is a patronym for Bento
Carqueja (1860-1935), professor and naturalist in the
Academia Politécnica do Porto.
Present name. Arthroleptis carquejai Ferreira, 1906.
Remarks. Since its initial description, this
species has remained known only from the holotype
and no subsequent authors have investigated its validity.
Our investigation suggests that Arthroleptis carquejai is
indeed a valid species and is likely part of the group of
species that includes Arthroleptis variabilis Matschie,
1893, Arthroleptis perreti Blackburn, Gonwouo, Ernst,
Rödel, 2009, and Arthroleptis palava Blackburn,
Gvoždik, Leaché, 2010, from Cameroon and other
Central African countries. While the pigmentation of
the specimen has faded, Ferreira’s (1906) original plate
(g. 1) indicates that the gular and anterior venter were
darkly pigmented with pale spots. Because the gular
region has pale spots but lacks a pale mid-line stripe,
the holotype of A. carquejai differs from A. variabilis,
in which there is a prominent and well-dened mid-line
gular stripe in females and juveniles, and pale markings
are generally lacking on the more uniformly pigmented
gular region of males (Blackburn et al., 2009). Darkly
pigmented gular and ventral surfaces are generally
uncommon in Arthroleptis but are characteristic of A.
variabilis and related species (Blackburn et al., 2009,
2010). During this work, we examined specimens from
Zoologisches Museum Hamburg (ZMH) collected during
the Hellmich expedition to Angola (Hellmich, 1957),
including several that are similar in external morphology,
size, and proportions to the holotype of Arthroleptis
carquejai (tab. 4). Both specimens (ZMH A09492-93)
were collected from “Roca Novo” (Roça Novo Mundo)
not far from the type locality (approx. 135 km north)
of A. carquejai. These specimens exhibit the similar
dark ventral pigmentation and traces of a pale mid-line
stripe (similar to A. palava; Blackburn et al., 2010) but
it is not well-dened as it is in A. variabilis. In addition,
these ZMH collections demonstrate that A. carquejai is
sympatric with Arthroleptis poecilonotus Peters, 1863,
a wide-ranging species complex in western and central
Africa, which was also collected at “Roca Novo” (ZMH
A09491, A09494-95).
Holotype specimens from ZMH
SVL 27.77 29.70 20.00
HW 9.68 10.66 6.76
HL 9.45 9.95 7.31
MN 8.26 6.79 6.70
MFE 6.72 6.49 5.44
MBE 3.25 3.19 2.50
IFE 3.88 4.31 3.61
IBE 6.36 8.95 5.50
FLL 5.85 7.79 5.10
HAL 8.64 9.39 5.98
TFL 6.99 5.74 3.61
FOL 14.20 15.68 10.10
FTL 9.81 7.48 6.30
IN 2.09 2.99 1.91
EN 2.26 2.09 1.42
EL 3.38 3.59 3.19
ITL 3.87 4.55 2.69
FL 13.22 15.42 10.06
NS 1.37 1.82 1.03
EE 2.78 3.00 2.33
TYE 0.99 1.14 0.64
TYD 1.29 1.49 0.84
Table 4. Measures (in mm) of the holotype of Arthroleptis
carquejai Ferreira, 1904 (MHNFCP 018586) and the
specimens of A. carquejai from Zoologisches Museum
Typhlops boulengeri Bocage, 1893
Original name. Typhlops boulengeri Bocage, 1893: 117.
Type specimen. A syntype (MHNFCP 017434) from Angola (g. 11) was donated to the Porto museum
from Lisbon. This is supported by the original label on the jar (“Typhlops boulengeri Boc. / Angola off. Mus.
Boc.”) and the fact that the catalog notes that the specimen MHNFCP 017434 was offered by Museu Bocage (L.
Sousa, pers. comm.). In his description of this species, Bocage (1893) noted that he received several specimens of
this species from Quindumbo, “dans l’intérieur de Benguella” that were collected by the explorer José de Anchieta.
Figure 10. Dorsal and ventral views of the holotype of Arthroleptis carquejai Ferreira, 1906 (MHNFCP 018586). Scale bar = 10 mm.
Figure 11. Dorsal and ventral views of the paratype of Typhlops boulengeri Bocage, 1893 (MHNFCP 017434). Scale bar = 10 mm.
ALYTES 2014 | 31
Present name. Afrotyphlops lineolatus (Jan, 1864).
Remarks. Though Laurent (1964) considered T. boulengeri to be a valid species, we follow the opinion
of Broadley & Wallach (2009) in referring Typhlops boulengeri to the synonymy of Afrotyphlops lineolatus. In
addition, Typhlops bocagei Ferreira, 1906, based on one specimen collected by Francisco Newton in Angola during
his 1903-1905 expedition, is also a synonym of this same species (see below).
Missing types
Rappia bocagei var. maculata Ferreira, 1906
Original name. Rappia bocagei var. maculata Ferreira, 1906: 160.
Present name. Hyperolius bocagei Steindachner, 1867.
Remarks. Ferreira described this new variety based on one specimen collected in “Golungo Alto” by
Francisco Newton during the 1903-1905 expedition to Angola. The description of this taxon by Ferreira (1906) was
based largely on the colour pattern of this single specimen. Despite thorough searches of the amphibian and reptile
collections in the Museu de História Natural da Universidade do Porto, we could not locate this specimen. There
are, however, three specimens noted as “Rappia bocagei” corresponding to a female and a male from “Cucuaco”
(MHNFCP 052189 – ♀ and MHNFCP 017306 – ♂) that might represent those from “Cacuaco” mentioned earlier
by Ferreira (1904: 112), and a third specimen noted only as deriving from “Angola” (MHNFCP 017308). In these
two works, Ferreira noted only three specimens of Rappia bocagei”, the two from “Cacuaco” (Ferreira, 1904)
and the “maculata” from “Golungo Alto” (Ferreira, 1906). Because it is unlikely that other specimens of this
species were catalogued in the Porto collections, it is conceivable that MHNFCP 017308 represents the holotype of
Rappia bocagei var. maculata. The loss of the original label may have resulted in the loss of the only information
relating this specimen to Ferreira’s work. However, despite the loss of the original label, the catalog notes that
MHNFCP 017308 was offered by Museu Bocage (L. Sousa, pers. comm.), which thus excludes the possibility of
this specimen being the holotype.
Typhlops bocagei Ferreira, 1904
Original name. Typhlops bocagei Ferreira, 1904: 114.
Present name. Afrotyphlops lineolatus (Jan, 1864).
Remarks. Ferreira (1904) described Typhlops bocagei based on two specimens from “Cabicula, Bom Jesus
(margens do Quanza)”, but we were unable to locate any specimens that plausibly represent these type specimens.
The systematics of the African Typhlopidae has received considerable attention (Laurent, 1964; Roux-Estève, 1970,
1974; Broadley & Wallach, 2009; Segniagbeto et al., 2011; Hedges et al., 2014) and Typhlops bocagei remains
currently considered a synonym of A. lineolatus. Note that Typhlops boulengeri Bocage, 1893 is also considered a
synonym of A. lineolatus (see above). Some doubts remain regarding the taxonomy of Typhlops bocagei (Broadley
& Wallach, 2009) but without either the type specimens or new investigations in the region, little more can be said
on its status.
Of the 18 type specimens expected from Ferreira’s papers on Newton collections we located a total of
12. The two syntypes of Typhlops bocagei Ferreira, 1904 and the holotype of Rappia bocagei var. maculata
Ferreira, 1906 could not be located and are probably lost. One syntype of Rappia platyceps var. angolensis, one
syntype of Hylambates bocagei var. leucopunctata, and the holotype of Rappia seabrai were also not found.
However, two other type specimens were found during this survey: a paratype of Rappia seabrai Ferreira, 1906
from the Bayão collection of the Lisbon museum (and supposedly lost in the 1978 re), and a paratype of Typhlops
boulengeri Bocage, 1893, given by the Lisbon museum. In addition to the type specimens, our survey revealed
approximately 90 to 110 reptile specimens and 60 to 80 amphibians from the Newton Angolan collection; some
of these specimens are still in the original collecting jars and remain to be inventoried and catalogued. Most of
the non-type specimens cited by Ferreira (1904, 1906) are still present in the collections and in generally good
condition, though some are in need of immediate conservation measures. It is possible that some of the type
specimens thought to be missing are still present in the collection but difcult to identify because of the lack of
labels or other identifying information.
Of the taxa described by Ferreira, we conrm that two of these should be recognized today (Rappia osorioi,
now Afrixalus osorioi, and Arthroleptis carquejai). We argue above that Rappia nobrei should be considered a
junior synonym of Hyperolius adspersus. The problematic Rappia bivittata, Rappia fasciata, Rappia platyceps
var. angolensis should all be considered junior synonyms of Hyperolius platyceps. Pending further investigations,
we recommend that Hylambates bocagei var. leucopunctata be considered a synonym of Leptopelis bocagii. These
taxonomic clarications contribute to our understanding of anuran diversity in southwestern Africa and are a
modest rst step towards rening our knowledge of the species richness of Angola amphibians.
Angola is among the largest countries in Africa at 1246700 km2 and perhaps the only biodiverse country in
Africa that remains seriously lacking in surveys of vertebrate diversity. Angola presents a great variety of biomes
and habitats and represents an important puzzle piece for understanding biogeographic patterns across sub-Saharan
Africa. Unfortunately, there has been little study of Angolan amphibians and reptiles since the mid-1960s. Recent
studies have been prohibited by the civil war and resulting social instability that engulfed the country for nearly
three decades (1975-2002; or four decades of war, if considering the beginning of the independence war in 1962).
A few recent eld studies will hopefully spark a new era for the Angolan biodiversity studies (Conradie et al.,
2012, 2013; Ernst et al., 2014). New eld studies will surely deepen the knowledge of the Angolan herpetofauna,
and help to address many long-standing taxonomic problems for Angolan and, more generally, southern African
species. Museum studies, however, remain crucial for addressing taxonomic and nomenclatural issues, especially
given that many species reported by Ferreira (1904, 1906) and others have not been reported since their initial
After the acceptance and nal revision of this manuscript (July 2014), we located two specimens that may
represent one of the syntypes of Typhlops bocagei Ferreira, 1904 and the holotype of Rappia bocagei var. maculata
Ferreira, 1906. Further details on these specimens will be presented separately in a future study.
We thank N. Ferrand de Almeida, director of the Museu de História Natural da Universidade do Porto, for
granting free access to the collections and all documentation. We extend a special thanks to L. Sousa, curator of
the Porto collections for more than three decades, for her kind help with all parts of this museological work and her
dedication to the herculean task of preserving this rich scientic collection. We thank J. Hallermann for facilitating
research on the herpetological collections in the Zoologisches Museum Hamburg. Annemarie Ohler provided
access to the collections of the Muséum national d’Histoire naturelle Paris, and encouraged and provided very
useful comments to the rst author of the paper. Alain Dubois gave useful suggestions and comments regarding
some nomenclatural issues. The authors also want to thank J. Alves from the Museu Nacional de História Natural
e da Ciência, Lisbon for assistance with accessing the historical archive of Museu Bocage. Portions of this work
were funded by a Ph.D. dissertation grant to LMPC (SFRH/BD/66851/2009) funded by the Portuguese Foundation
for Science and Technology (FCT-MCTES).
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Specimens used in comparisons
Hyperolius platyceps: California Academy of Sciences: CAS 199349-51, 199354, 201956, 201958,
201963, 201966, 208544-46. Muséum national d’Histoire naturelle: MNHN 1968-160, 1968-160A,1968-161,
Hyperolius cinnamomeoventris: California Academy of Sciences: CAS 98164, 144780, 154511, 154519,
154746, 180133-35, 180141, 199355, 202322, 202508, 204508, 204512. Muséum national d’Histoire naturelle:
MNHN 01-584, 01-578, 01-579, 06-209, 21-17A, 21-17B, 21-134A, 21-164B, 21-164C, 21-164D, 46-149, 84-17,
96-571, 96-572.
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Arthroleptis poecilonotus: Zoologisches Museum für Hamburg: ZMH A09491, A09494-95.
kindly presented to B. M.
Letters between Albert Günther and Barbosa du Bocage concerning the type specimens
of “Cystignathus bocagii
All the letters are deposited in the Historical Archive of Museum Bocage (AHMB) in the Museu Nacional
de História Natural e da Ciência, Lisbon, Portugal.
Letter from Albert Günther to José Vicente Barbosa du Bocage: 19 September 1864
“British museum
19. Sept. 64
My dear sir
I have just returned from a journey which I undertook during my holidays, & hasten to thank you for your kindness
in sending me the reptiles from the province of the Duke de Bragance, & some very ne portuguese sh: all of
which arrived in perfect safety. I have not had time yet to examine the reptiles, but one of the snakes is unknown
to me, & several of the frogs appear to be new. I shall return all the specimens which you desire to keep, hoping
that if you should receive duplicates at some future time, you will kindly communicate to us what you can spare.
If you receive any other examples from the west coast of africa, which are doubtful to you, I should consider it as
a great kindness if you would allow me to examine them. Mr. Monteiro has delivered the reptiles, & the Trustees
of our museum will send you their acknowledgment. I am glad to hear from you that the specimens sent by me to
your museum, were of some use; I hope to be able to add others when I shall return your specimens.
Yours most Truly
A. Günther”
Letter from Albert Günther to José Vicente Barbosa du Bocage: 24 July 1865
“British museum
24. 7. 65.
My dear Sir,
Excuse me for having kept the specimens kindly lent to me, for so long a time. I send them off to your address
today, & hope that they will in safety reach you. I add the names:
1. Limnophis bicolour, g. & sp. n.
2. Causus rhombeatus
3. Lycophidium horstockii
4. & 5 I shall send the name at the later period end of this letter
6. has not arrived
7. Hyperolius marmoratus
8. cannot be determined without other specimens
9. Hyperolius fulvovittatus, Cope
10. - nasutus, sp. n.
11. Cystignathus bocagii, sp. n.
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12. Leptopelis natalensis
13. Hyperolius sp?
14. Hyperolius modestus
15. Hyperolius nasutus - type
16. Hyperolius sp? an nasutus, young.
17. Hyperolius sp?
18. Hyperolius modestus
19. Bufo panterinus
nos. 11-19 are returned in today’s parcel
20. Grayia triangularis
21. Ahaestulla irregularis
22. Boodon lineatus
nos. 20 - 22 have been returned through mr. Monteiro several months ago; I hope you have safely
received them.
23. Prosymna meleagris. - very rare, is returned today with the frogs.
I add in a second bottle some specimens as a present from me for your museum:
Xenopeltis unicolour, Siam / large brown snake /
Typhlops braminus, Siam
Vermicella occipitalis, Moreton Bay, (black & white snake)
Lialis punctulata, Sasan River
(Ophidioid Saurian).
Further I add some paper of mine. Many thanks for the loan of your valuable specimens!
The address of the manufacturers of bottles employed by the British museum, is
Messus Powel & Son
Glass works
London F. C.
The name of the frogs are
4. Rana oxyrhyncha
5. - mascareniensis
I am very sorry not to have a single specimen of Rana delalandii which is not entered into the register and catalogue
of the collection, & which, therefore, cannot could be given away. This species is not rare, & I have no doubt that
I can soon send you an example.
Thanking you again for your kindess
I remain
Yours must truly
A. Ghünther”
Letter from Albert Günther to José Vicente Barbosa du Bocage: 6 May 1866
“British museum
6. 5. 66
My dear sir
I do not consider the disks of Cystign. bocagii to be well enough developed to deserve the name of disks. It is
quite true the extremities of its ngers hands, but such dilatations you nd also in several Rana & other so called
oxydactyle frogs. Cystign. bocagii has no webb between the ngers & toes, & therefore it cannot be associated
with Leptolepis. I do not recollect the specimen of Leptolepis which I returned with C. bocagii, & cannot speak
with condence about its determination; but I should regard it as a great favour, if you would give me another
opportunity of examining these two specimens, so as to explain the question to your & to my own satisfaction.
I would return the specimens without detaining them for more than a week. I have the manuscript of the second
volume of the record in the printers hands, but have not been able to obtain your paper on arvicola. I shall retain
the proof-sheet for three other weeks, hoping to be able to insert it. I should be much obliged to you for a separate
copy of the paper, or for a short abstract of it. Have you now received the Zool. Soc. publications? The clerk
told me they had been sent to you some time back. The third part of the proceedings for 1865 will be out in the
course of next week, the print part of 1866 is published, but not without illustrations. I am afraid, Peter’s Fishes of
Mozambique will be indenitely delayed; as the [?] Prussian government wants too much money for [?], to spare
[?] a little for science.
Yours ever truly.
A Günther.”
Letter from Albert Günther to José Vicente Barbosa du Bocage: 29 June 1866
“British museum
29 June 1866
My dear Sir
I have to thank you much for your kindness in sending me your memory on arvicola, & the reptiles which I
received from Mr. Joanson. You are right that the two frogs are of the same species, vig. = Cystignath bocagii, they
differ widely from Hylambates Leptopelis which has broad disks [small drawing of a nger disk]. As you possess
two specimens of Cystign. boc., would you give us one of them for our collection? I should return the specimens
(with other things) [?] but I think it better to wait for your answer as regards this specimen of Cystign. The snake
is the young of some species of Psammophis; the [?] poster tooth is distinct on one side. But I would not determine
of what species it is the young, it may be new, but it is not advisable to describe a new species from such a young
specimen, from which no certain characters may can be taken.
1. Stellio angolensis = Agama atricollis, A. Smith, Ill. Slep. Cpp p. 14. I have compared your specimen with the
2. is very closely allied to Agama occipitalis, but has somewhat smaller scales.
3. Ichnotropis bivittatus = Algiru (Tropidosaura) dumerilii, Smith, App. p. 7 I have compared your specimen with
the type.
4. Chamaeleo gracilis is certainly not a distinct species; see Gray’s paper in Proc. Zool. Soc. 1864
5. & 6. are well determined.
I remain yours very truly
A. Günther”
... We focused mainly on records archived in MHNC-UP, in the Arquivo Histórico Ultramarino (AHU) and in Arquivo Histórico de Angola (AHA). As for bibliography, we consulted the published papers resulting from the study of the Newton's Angolan expedition collections (Ferreira, J.B. 1904(Ferreira, J.B. , 1906Nobre, 1905b;Seabra, 1905aSeabra, , b, c, 1906aSeabra, , b, c, 1907Seabra, , 1909 as well as recent reviews done by Ceríaco et al. (2014) and Marques et al. (2018). The MHNC-UP collection specimens were examined, and all present data was cross-referenced with the internal museum databases, documentation, and the resulting papers. ...
... The newly described taxa resulting from Newton's Angola expedition are presented in Table 2. The herpetological specimens have recently been addressed by Ceríaco et al. (2014) and Santos et al. (in press). Ceríaco et al. (2014) presented an updated catalogue of the amphibian and reptile type specimens extant today in MHNC-UP and located 12 of the 18 type specimens published by Ferreira, J.B. (1904Ferreira, J.B. ( , 1906. ...
... The herpetological specimens have recently been addressed by Ceríaco et al. (2014) and Santos et al. (in press). Ceríaco et al. (2014) presented an updated catalogue of the amphibian and reptile type specimens extant today in MHNC-UP and located 12 of the 18 type specimens published by Ferreira, J.B. (1904Ferreira, J.B. ( , 1906. In a recent review of the species of the genus Boaedon from Angola, Hallerman et al. (2020) used three specimens collected by Newton (MHNC-UP/REP 166-168) as paratypes in the description of Boaedon bocagei Hallermann, Ceríaco, Schmitz, Ernst, Conradie, Verburgt, Marques and Bauer 2020. ...
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Francisco Newton (1864–1909) was an important Portuguese naturalist who conducted several expeditions for more than 25 years during the late nineteenth and early twentieth century and who greatly contributed to the study of Portuguese overseas colonies. In 1902, Newton was nominated to undertake a zoological survey of Angola, commissioned by the Academia Politécnica do Porto (APP; precursor of University of Porto), an institution that had recently started to develop a zoological museum. Between 1903 and 1906, Newton collected several hundred zoological specimens that resulted in the description of 27 new taxa. Here we present a reconstruction of the expedition, providing georeferenced collecting localities and other historical data.
... Despite their size, small collections can have disproportionate importance (Winker & Withrow 2013). This is certainly the case of the MHNCUP collections, whose number of type specimens is considerable for its size (see Ceríaco et al. 2014), but also harbour interesting and rare specimens, such as the one used as the holotype of the newly described species. Within the MHNCUP collections, the specimens collected by Francisco Newton during his 1903-1905 expedition to Angola are of special relevance for the study of the herpetofauna of the whole region. ...
... Within the MHNCUP collections, the specimens collected by Francisco Newton during his 1903-1905 expedition to Angola are of special relevance for the study of the herpetofauna of the whole region. This collection was studied in the past by Ferreira (1904Ferreira ( , 1906 but was then subsequently abandoned and forgotten by the scientific community until Ceríaco et al. (2014) published its type catalog. The collection has recently been reviewed and a detailed and commented list of its specimens published (Santos et al. in press). ...
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Three species of Phrynomantis Peters, 1867, have been historically recorded for Angola: P. affinis, P. annectens and P. bifasciatus. As noted by all authors who have dealt with specimens of P. bifasciatus from the country, the Angolan population is characterized by an odd coloration pattern for the species, which led Boulenger to consider it a different variety. A revision of the extant specimens of Angolan Phrynomantis available in natural history collections, specimens collected in recent field surveys, as well as recent sightings and photographs allows the recognition of the Angolan population of P. cf. bifasciatus as a new species, endemic to the coastal lowlands of western Angola. The new taxon is described solely based on its coloration pattern and morphology, and it is separated from nominotypic P. bifasciatus by more than one thousand kilometers. The revision of these historical specimens also allowed us to confirm a second record of P. affinis in the country and to contribute to an overall better understanding of the distribution of the species of the genus on the continent.
... According to [1], over 8524 amphibian species have been identified worldwide and occupy virtually all habitats, with the exception of the coldest and driest regions, as well as the most remote oceanic islands [2]. Amphibians play critical roles in the functioning of ecosystems [3] and they are increasingly being used in pollution studies as bioindicator organisms [4]. ...
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The aim of the current study was to provide a description of the species composition and habitat association of anuran species found in the Keffa area of southwest Ethiopia, among the least studied natural areas in Ethiopia. Data were gathered in both the rainy and dry seasons. Opportunistic observations, audio, and visual encounters were used to perform the survey. To capture different frog species, drift fence and pitfall trap techniques were used. Three habitat types-wetlands, agricultural land, and riverine forests-were explored for the frog species. A total of 3672-person field hours were spent and 5678 individuals of 26 different anuran species from eight different families were recorded. Of these, six species were new records for the Keffa area. Species determination was assessed using morphology and validated with genetic barcoding. The mitochondrial 16S rRNA and COI gene sequence analysis have been used in molecular systematic analyses of the frog species. The Leptopelis ragazzii that was discovered to be present in the earlier study was actually Leptopelis vannutellii, as evidenced by DNA sequencing. The Hyperolius individuals were initially categorized as H. nasutus, but after conducting DNA barcoding, they were found to be H. microps. At least three potential new candidate species (Leptopelis sp. 1, Leptopelis sp. 2, and Phrynobatrachus sp. 1) were discovered. Hyperoliidae was the most abundant family from the recorded anurans, followed by Phrynobatrachidae, while Conrauidae was the least abundant. Most species were found in the wetland habitat, followed by riverine forest, while agricultural land had the least. Wetlands had the highest values for the Shannon diversity index (2.131), followed by agricultural land (1.58) and riverine forests (1.459). The environmental variables revealed that temperature had a considerable negative impact, while precipitation had a non-significantly positive relationship with anuran abundance. The Keffa area is rich in frog species, accounting for 35% of the country's batrachofauna. However, the habitats have been fragmented and ruined by agricultural expansion, water drainage from the wetlands, and the conversion of horticulture to field crops. We recommend that non-governmental organizations, districts, regional officials, and local people in and around Keffa actively engage in wetland and forest conservation and protection.
... Amphibians are distributed all over the world, except in the coldest and driest regions, and most remote oceanic islands (Ceríaco et al., 2014;Foden and Stuart, 2009). They are increasingly used as bioindicator organisms in pollution studies due to their bi-phasic nature ( Jongsma et al., 2014;McMenamin et al., 2008;Saber et al., 2017). ...
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The present study was carried out to describe the species composition, diversity, distribution, and species richness of endemic frog species in the Keffa area, southwest Ethiopia. Data were collected during the wet and dry seasons from 2018 to 2021. The survey was conducted through visual encounters, opportunistic observations, and acoustic encounters. Drift fence and pitfall trap methods were employed to capture frog species. Endemic anurans were investigated in three habitat types: wetland, agricultural land (enset vegetation), and riverine forest. From a total of 3672 person field hours, 2951 individuals belonging to 13 species such as Leptopelis susanae, Leptopelis ragazzii, Leptopelis vannutellii, Hemisus microscaphus, Afrixalus clarkei, Afrixalus enseticola, Paracassina obscura, Phrynobatrachus inexpectatus, Phrynobatrachus minutus, Xenopus largeni, Ptychadena beka, Ptychadena erlangeri, and Ptychadena neumanni were captured. Among these, Leptopelis susanae and Xenopus largeni were recorded for the first time in the study area. Arthroleptidae, Hyperoliidae, and Ptychadenidae families were the most represented, with three species each. The family Pipidae represented the least number comprising only one species. Agricultural land had the highest values for Shannon index (1.401), followed by riverine forest (1.182) and wetland (1.14) although species richness is highest in the wetlands. More endemic frog species were recorded in agricultural habitats, so local communities should continue to cultivate crops like enset plantations. The Keffa area is rich in endemic frog species, accounting for 34% of the country's frog endemism. Five of the thirteen identified endemic species have been identified as threatened by IUCN Redlist status. As a result, we propose that local communities, district and regional officials, and non-governmental organizations working in and around Keffa focus on wetland and forest conservation and protection.
... The present type catalogue follows the structure of similar recent publications (see Ceríaco et al., 2014b;Conradie et al., 2015Conradie et al., , 2019. A list of all extant material in the MD collections is presented. ...
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The herpetological collections of the Museu do Dundo in Lunda Norte Province, northeast Angola, are among the most important in southern Africa and represent one of the largest collections of Angolan amphibians and reptiles in the world. The collection comprises more than 2,750 preserved specimens, including type specimens of taxa described by Raymond F. Laurent during the 1950s and 1960s, when he was affiliated with the Musee royal de ´ l’Afrique centrale (RMCA) in Tervuren, Belgium, and the Museum of Comparative Zoology (MCZ) at Harvard University, Cambridge, Massachusetts, where portions of these type series were also deposited. We provide details for all type specimens and summarize the history and taxonomy for each species represented in the type collection. The collections contain type specimens of 28 amphibian and reptile species, including seven snakes: Typhlops praeocularis lundensis, Dispholidus typus punctatus, Lycodonomorphus subtaeniatus, Lycophidion hellmichi, Gonionotophis brussauxi prigoginei, Prosymna ambıgua brevis ´ , and Elapsoidea decosteri huilensis; 13 lizards: Rhoptropus boultoni montanus, Rhoptropus taeniostictus, Hemidactylus nzingae, Lygodactylus tchokwe, Cordylus vittifer machadoi, Chamaesaura anguina oligopholis, Gerrhosaurus bulsi, Nucras scalaris, Ichnotropis bivittata pallida, Mabuya bayonii huilensis, Mabuya ivensii septemlineata, Trachylepis raymondlaurenti, and Eumecia anchietae major; one amphisbaenian: Monopeltis vanderysti vilhenai; and seven frogs: Hyperolius machadoi, Hyperolius marmoratus alborufus, Hyperolius vilhenai, Ptychadena grandisonae, Ptychadena loveridgei, Ptychadena perplicata, and Ptychadena upembae machadoi. The type specimens of the snake Xenocalamus bicolor machadoi were not found in the collections. A brief history of the museum and remarks on the overall herpetological collections are also provided. We also note additional information about the related type material of these taxa at the MCZ.
... Oliveira et al. 2016;Branch et al. 2017aBranch et al. , 2019bHeinicke et al. 2017;Agarwal et al. 2017;Gonçalves et al. 2019;Vaz Pinto et al. 2019). Other works related to the study of the Angolan herpetofauna include the revision of historical collections (Ceríaco et al. 2014b) and the publication of an atlas on the diversity and distribution of the amphibians and reptiles of the country ) and subsequent abridged syntheses of its overall diversity (Branch 2018;Baptista et al. 2019b;Branch et al. 2019c). Ongoing field surveys and several taxonomical reviews are under way, which will yield further additions to the knowledge of the herpetofauna of the country. ...
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An integrative taxonomic review of the genus Boaedon in Angola is provided. A molecular phylogeny, based on 99 genetic samples for which the mitochondrial markers 16S rRNA have been sequenced, reveals 23 monophyletic species-level groups in Africa and indicates the presence of nine species in Angola. Based on both phylogenetic and morphological data, we revalidate and designate a neotype for B. angolensis, describe three new species for Angola (e.g. B. bocagei sp. nov., B. branchi sp. nov., and B. fradei sp. nov.), revalidate B. variegatus from its synonymy with B. lineatus and designate a lectotype for this taxon, and identify B. lineatus var. lineolatus as a junior synonym of B. variegatus. The taxonomic status of the recently described B. paralineatus from Central Africa is discussed with respect to the more inclusive B. lineatus group. Moreover, we report on a new country record for Angola, namely B. mentalis, which we elevate here to full species and discuss the taxonomic status of this species in southern Africa. Finally, we provide an identification key and updated distribution maps for all Boaedon species occurring in Angola, including the Cabinda enclave.
... Very few papers were published during this period (Poynton and Haacke 1993;Ruas 1996Ruas , 2002. After the war, numerous researchers initiated several independent research projects and a constant flow of papers, reporting faunistic data (Ceríaco et al. 2014(Ceríaco et al. , 2016a(Ceríaco et al. , b, 2018aConradie et al. 2016;Baptista et al. 2019b;Butler et al. 2019), new country records (Ernst et al. 2014(Ernst et al. , 2015Branch and Conradie 2015), and new species descriptions Stanley et al. 2016;Ceríaco et al. 2018bCeríaco et al. , 2020aBranch et al. 2019a;Marques et al. 2019a, b) has been published. An updated atlas for the Angolan herpetofauna was recently published by Marques et al. (2018), and, together with Branch et al. (2019b) and Baptista et al. (2019a), these sources can now be used as a general reference and overview that is constantly being updated as new discoveries are published. ...
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We systematically assess the herpetofaunal diversity of the Serra do Pingano Forest Ecosystem (SPFE) and additional localities throughout the northern Angolan province of Uíge during four independent Rapid Assessment (RA) field campaigns held between 2013 and 2019. These assessments represent the first systematic surveys of amphibians and reptiles from the province, and thus we provide the first province-wide species list. We collected data on the status and current threats to amphibians and reptiles in the proposed Serra do Pingano Rainforest National Park and were able to document 33 species of reptiles from Uíge province. Of the 33 species recorded from the province, 10 species are exclusively found in the SPFE. Amphibian surveys yielded 47 amphibian species from the province. These include 14 new country records and additional records that may represent undescribed species. This raises the amphibian count for Angola to at least 133 species, which includes 18 species exclusively found within the SPFE. Species-richness estimators indicate that more species should be detected if survey efforts are intensified. The species composition in the SPFE is unique and consists of a high proportion of forest specialists with restricted ranges and species found nowhere else in the country. This emphasizes today's paramount importance of the SPFE, which is threatened by increasing agricultural encroachment and uncontrolled timber extraction and charcoal production. These principal factors need to be controlled and/or abandoned in already impacted areas. Conservation strategies should particularly consider the strict protection of remaining intact forests and both lentic and lotic aquatic systems. They are not only crucial for safeguarding a significant number of species that depend on these habitats for reproduction; they also provide key ecosystem services to the local population. Angola, and Uíge province in particular, is at a crossroads concerning decisions and trade-offs among utilization, conservation, and preservation of its forests and, thus, substantial parts of the country's biodiversity. The establishment of a National Protected Area in the Serra do Pingano Ecosystem is therefore a necessary and urgently needed first step towards protecting Angola's national biodiversity heritage.
... Rela -de -bocage Hyperolius bocagei (Steindachner, 1867) Laurent, , 1954Ceríaco et al., 2014b Laurent, 1954Ceríaco et al., 2016c, no prelo Rela -comprida -de--dartevelle ...
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Os anfíbios de Angola têm sido estudados desde os meados do século xix por exploradores e cientistas de todo o Ocidente, com colecções depositadas em cerca de 20 museus e instituições da Europa, América do Norte e África. Este estudo sofreu uma interrupção significativa durante as quase quatro décadas da luta de libertação e guerra civil de Angola e, como consequência, o conhecimento sobre a biodiversidade do país tornou‑se obsoleto, com lacunas críticas. Em 2009 começou uma nova era nos estudos da biodiversidade angolana, com expedições no Sudoeste, Nordeste, Sueste e Noroeste de Angola, e levando a descobertas empolgantes, como novos registos para o país, descrições de novas espécies, aumento de áreas de distribuição e actualizações taxonómicas. Actualmente, encontram‑se registadas 111 espécies de anfíbios neste país (das quais 21 são endémicas), mas este número é uma subestimativa e as diversas questões taxonómicas ainda não resolvidas desafiam o estudo de todos os outros aspectos relativos a este grupo. A fauna de anfíbios de Angola continua a ser uma das mais mal conhecidas de África e ainda há muito por fazer para compreender a sua diversidade, evolução e necessidades de conservação. Este capítulo apresenta uma panorâmica geral do conhecimento existente relativo aos anfíbios de Angola, incluindo uma lista de espécies actualizada, comentários sobre grupos problemáticos, espécies endémicas, biogeografia, descobertas recentes e tópicos de investigação prioritários.
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With alarming frequency, significant collections in natural history museums have been destroyed or damaged through insurrections, cyclones, wars, fires, floods, or earthquakes particularly in the nineteenth century but continuing into the twentieth century with World War II bombings, fires, and earthquakes being the primary causes of loss in fifty-seven institutions across thirty countries. We review the loss or damage of museum collections globally, and their varied causes. We emphasize the benefits of dispersal of a sample of paratypes across institutions as an essential feature of taxonomic practice. We argue that museums do not own type material but are acting as perpetual custodians of type material on behalf of science and society in general, and that museums, therefore, have an obligation to minimize the risk to their collections. The significance of the loss of type material would be ameliorated if, when there are numerous paratypes or syntypes, members of a type series were distributed among several institutions. This is currently common practice but historically this was not always the case and might not be possible if only a single holotype is available. We also reaffirm the need for scientists around the globe to develop specific protocols to protect collections of biological and cultural materials from loss or damage from natural and human-created disasters now and into the future. We comment on recent moves to modify the Zoological Code of Nomenclature to allow the use of images as "type" material when describing new species with the image serving as a substitute for "physical" specimens deposited in museum collections. Although our focus is on herpetological collections, our particular interest and area of expertise, our observations apply broadly to all collections, including those of animals, plants, and anthropological or ethnographic material.
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Amphibian faunas of the seven Central Afncan States and of Angola sum up to 359 species in 53 généra and 10 families, among which two are Gymnophiona and eight Anura. Taxonomie composition at the spécifie level of these faunas are mostly rclated to the geographical position of the country. Oceanic îslands in the Biafra Gulf (Sào Tomé, Principe) display a high endemism and lack of biogeographic unity. Respectively, Hypcroliidae (15 g. ; 121 sp.) and Ranidae (13 g. ; 108 sp.) add up to 53% and 64% of ail généra and species. Their two richest généra Hyperolius and Pluynobatrachus include 99 species (28% of the total fauna). Central Afnca and Angola harbour 128 endemic species, of them occuring in Cameroon and Congo-Kinshasa, two countries with high up to 4000 and 5000 m mountains and relatively the most surveyed. KEYWORDS Amphibians, Central Africa, Angola, Biogeography, Insulanty RESUME-La faune des Amphibiens des sept pays d'Afrique Centrale et d'Angola compte 359 espèces regroupées en 53 genres et 10 familles, dont 2 de Gymnophiones et 8 d'Anoures. L'analyse biogéographique de ce nche peuplement montre que sa composition spécifique, dans chaque pays, est fortement liée à la position géographique de celui-ci. Les îles océaniques du Golfe du Biafra (Sào Tomé, Principe) manifestent une forte originalité et une absence d'unité biogéographique. Les deux familles des Hypéroliidés, avec 15 genres et 121 espèces, et des Ranidés, avec 13 genres et 108 espèces, réunissent 53% des genres et 64% des espèces ; leurs deux genres les plus diversifiés, Hyperolius et Pluynobatrachus, totalisent 99 espèces, soit 28% du peuplement. L'entité géographique considérée renferme 128 espèces endémiques, présentes surtout au Cameroun et au Congo-Kinshasa, pays possédant des systèmes montagneux de 4 à 5000 m d'altitude et relativement mieux inventoriés.
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All eastern and southern African species of Typhlopidae (excluding Letheobia) are reviewed with synonymies, diagnoses, and distribution maps. The 27 species from this region are arranged as follows: three species remain in Typhlops (with T. cuneirostris calabresii elevated to full species status), 16 species are transferred from Typhlops to a new genus endemic to Africa (including a new species described from Kenya), three species remain in Rhinotyphlops, four species are transferred from Rhinotyphlops to a new genus endemic to Africa (with R. schlegelii brevis and R. s. mucruso elevated to full species status), and one invasive species of Ramphotyphlops is included, which has been established in a few areas. Keys to the included species and all six African typhlopid genera are provided. A current taxonomic synopsis is provided for extralimital African and Southwest Asian typhlopid species.
This paper reviews the problematic taxonomy of the superspecies Hyperolius viridiflavus (the senior synonym of H. marmoratus) (Anura: Hyperoliidae). The classifications proposed by various authors for the complex are summarized and compared. Significant differences in classification have resulted in the complex being divided into as few as three species (with numerous subspecies) or as many as 18 species (with a number of subspecies). The review shows that the taxonomy of this complex has not been resolved, as a consequence of dependence on the use of dorsal colour patterns as diagnostic characters. Colour pattern is polymorphic within taxa, and shows both geographical as well as non-geographical variation. Recent work has shown molecular techniques to be a valuable means of distinguishing taxa previously considered to be subspecies. It is proposed that the use of molecular techniques and phylogenetic analysis of these data is the most suitable means of resolving the taxonomy of the members of this complex.