Taxonomic revision of the Jita snakes (Lamprophiidae: Boaedon) from São Tomé and Príncipe (Gulf of Guinea), with the description of a new species

Abstract

The taxonomic status of the São Tomé and Príncipe islands 'Cobras Jitas', genus Boaedon, has been a subject of confusion. Historically, these island populations have been referred to as part of either the Boaedon fuliginosus species complex or Boaedon capensis species complex, two of the most taxonomically challenging groups of African snakes, or considered a distinct taxonomic entity, B. bedriagae. Here we review the São Tomé and Príncipe populations through a combination of morphological and molecular data. Our results suggest that each island population represents a unique species. After a thorough review of the taxonomic and nomenclatural history of the group, we revalidate B. bedriagae, restricting the application of this name to the São Tomé population by the designation of a lectotype. We also describe the Príncipe population as a new species, Boaedon mendesi sp. nov. This description has implications to our understanding of the diversity and phylogeographic patterns of the Gulf of Guinea Oceanic Islands.

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African Journal of Herpetology
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Taxonomic revision of the Jita snakes
(Lamprophiidae: Boaedon) from São Tomé and
Príncipe (Gulf of Guinea), with the description of a
new species
Luis MP Ceríaco , Ana Lisette Arellano , Robert C Jadin , Mariana P Marques ,
Diogo Parrinha & Jakob Hallermann
To cite this article: Luis MP Ceríaco , Ana Lisette Arellano , Robert C Jadin , Mariana P Marques ,
Diogo Parrinha & Jakob Hallermann (2021): Taxonomic revision of the Jita snakes (Lamprophiidae:
Boaedon) from São Tomé and Príncipe (Gulf of Guinea), with the description of a new species,
African Journal of Herpetology, DOI: 10.1080/21564574.2020.1832152
To link to this article: https://doi.org/10.1080/21564574.2020.1832152
Published online: 15 Jan 2021.
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Taxonomic revision of the Jita snakes (Lamprophiidae:
Boaedon) from São Tomé and Príncipe (Gulf of Guinea), with
the description of a new species
Luis MP Ceríaco
a,b
, Ana Lisette Arellano
c
, Robert C Jadin
d
,
Mariana P Marques
b,e
, Diogo Parrinha
b
and Jakob Hallermann
f
a
Museu de História Natural e da Ciência da Universidade do Porto, Porto, Portugal;
b
Departamento de
Zoologia e Antropologia (Museu Bocage), Museu Nacional de História Natural e da Ciência, Universidade de
Lisboa, Lisboa, Portugal;
c
Golden Gate National Parks Conservancy, Fort Mason, San Francisco, USA;
d
Department of Biology and Museum of Natural History, University of Wisconsin Stevens Point, Stevens
Point, Wisconsin, USA;
e
Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), Rede de
Investigação em Biodiversidade e Biologia Evolutiva (InBIO), Universidade do Porto, Vairão, Portugal;
f
Center
of Natural History (CeNak), Zoologisches Museum, Universität Hamburg, Hamburg, Germany
ABSTRACT
The taxonomic status of the São Tomé and Príncipe islands ‘Cobras
Jitas’, genus Boaedon, has been a subject of confusion. Historically,
these island populations have been referred to as part of either the
Boaedon fuliginosus species complex or Boaedon capensis species
complex, two of the most taxonomically challenging groups of
African snakes, or considered a distinct taxonomic entity,
B. bedriagae. Here we review the São Tomé and Príncipe
populations through a combination of morphological and
molecular data. Our results suggest that each island population
represents a unique species. After a thorough review of the
taxonomic and nomenclatural history of the group, we revalidate
B. bedriagae, restricting the application of this name to the São
Tomé population by the designation of a lectotype. We also
describe the Príncipe population as a new species, Boaedon
mendesi sp. nov. This description has implications to our
understanding of the diversity and phylogeographic patterns of
the Gulf of Guinea Oceanic Islands.
ARTICLE HISTORY
Received 18 May 2020
Accepted 30 September
2020
ASSOCIATE EDITOR
E Stanley
KEY WORDS
Boaedon bedriagae;Boaedon
mendesi sp. nov.; Boaedon
fuliginosus complex;
endemism; island
biodiversity
Introduction
In just over a decade, several new vertebrate taxa have been described from the small island
nation of São Tomé and Príncipe in the Gulf of Guinea. These include the following new
species: a shrew, Crocidura fingui (Ceríaco et al. 2015); a gecko, Hemidactylus principensis
(Miller et al. 2012); four species of skinks, Trachylepis adamastor,T. principensis,
T. thomensis and Panaspis thomensis (Ceríaco 2015; Ceríaco et al. 2016; Soares et al.
2018, respectively); a cobra, Naja peroescobari (Ceríaco et al. 2017); a puddle frog;
African Journal of Herpetology is co-published by NISC Pty (Ltd) and Informa Limited (trading as Taylor & Francis Group)
CONTACT Luis MP Ceríaco luisceriaco@gmail.com © 2020 Herpetological Association of Africa
This article is registered in ZooBank under: urn:lsid:zoobank.org:pub:13E287C7-E644-4899-95B9-AF5B66931E89
This species is registered in ZooBank under: Boaedon mendesi: urn:lsid:zoobank.org:act:0936727A-A15F-4967-936E-
645A0EF97DCE
AFRICAN JOURNAL OF HERPETOLOGY
https://doi.org/10.1080/21564574.2020.1832152
Published online 15 Jan 2021

Phrynobatrachus leveleve (Uyeda et al. 2007); and a reed frog, Hyperolius drewesi (Bell
2016). These earlier works have considerably expanded our understanding of the diver-
sity, endemicity and biogeographic patterns of the herpetofauna of these two islands.
Five species of amphibians are currently known on São Tomé Island and three species
are present on Príncipe (Ceríaco et al. 2018). All of these amphibians are endemic to
their respective island. São Tomé Island hosts a total of 12 species of terrestrial reptiles,
of which six are island endemics, three are shared endemics with Príncipe Island, and
the remaining three also occur elsewhere in the continent (Ceríaco et al. 2018,2020). A
total of 14 species, including confirmed species and species groups with taxonomic
and nomenclatural uncertainties, have been recorded for Príncipe Island. Of these taxa,
eight are island endemics, three are shared endemics with São Tomé Island, and the
remaining three also occur on the continent (Ceríaco et al. 2018,2020).
Among the endemic reptiles of these islands, the African House Snakes of the genus
Boaedon Duméril, Bibron and Duméril, 1854, locally known as ‘Cobras Jitas’[=Jita snakes],
are probably some of the most well-known species in the country. Unlike the famous
‘Cobra-Preta’,Naja peroescobari,the‘Jitas’are usually appreciated by locals and recognised
as harmless to humans. Despite this general appreciation, these snakes are still poorly
studied, and their taxonomic allocation remains problematic. These island Boaedon have
one of the most complex and problematic taxonomic and nomenclatural histories of all
São Tomé and Príncipe reptiles. Indeed, Boaedon ranks amongst one of the most taxonomi-
cally challenging genera of African snakes (Greenbaum et al. 2015; Hallermann et al. 2020).
The taxonomic allocation of the São Tomé and Príncipe ‘Jitas’has been a matter of dis-
cussion and different interpretations by the authors who have recently dealt with them.
Historically the São Tomé and Príncipe Jitas have had an uncertain taxonomic allocation
and given a varied number of nomina (see below). Boulenger (1906) was the first to recog-
nise the São Tomé and Príncipe Jitas as a different species, Boaedon bedriagae. Sub-
sequently, various authors had different interpretations, both on the validity of the
taxon and its affinities. Angel (1920) and Roux-Estève and Guibé (1965) followed Boulen-
ger’s(1906) interpretation and considered bedriagae a valid species. Bogert (1940) con-
sidered that bedriagae was a subspecies of B. lineatus, a decision followed by Manaças
(1958) and Capocaccia (1961a,1961b). Roux-Estève and Guibé (1964), Manaças (1973),
Schätti and Loumont (1992), Chippaux (1999) and Hofer (2002), considered it as a subspe-
cies of Boaedon (or Lamprophis)fuliginosus. Chippaux (2006) considered bedriagae to be a
synonym of Boaedon fuliginosus, while Wallach et al. (2014) considered bedriagae to be a
synonym of Boaedon lineatus. Ceríaco et al. (2018) referred to the species as B. bedriagae,
noting that the Príncipe Jitas were in the process of being described as a new species.
Chippaux and Jackson (2019) considered it as a subspecies of B. fuliginosus.
One of the main issues regarding the taxonomic identity of the São Tomé and Príncipe
Jitas has roots in the contentious taxonomic and nomenclatural history of the remaining
species of the genus, especially B. fuliginosus and B. lineatus, the two taxa typically associ-
ated with this population. The taxonomy and nomenclature of B. fuliginosus and
B. lineatus has been a matter of debate for decades. Roux-Estève and Guibé (1964,
1965) presented an initial approach to the understanding of these two taxa.
Boaedon fuliginosus was described by Heinrich Boie (in Friedrich Boie 1827)asLycodon
fuliginosus. The original description states that the type locality of the species was Java,
which, as noted by Hallermann et al. (2020), represents an error. According to Brogersma
2L. M. P CERÍACO ET AL.

in Roux-Estève and Guibé (1965), the type specimen has been lost. However, meristic scale
data and colouration unambiguously identify Boie’sB. fuliginosus as what most modern
authors recognise as the African Brown House Snake, Boaedon fuliginosus. It is probable
that the specimen was collected en route from Java to Europe. At the time the typical
voyages from the Dutch East Indies would have stopped at the Cape of Good Hope
(present-day Cape Town) and possibly in West Africa at one of the ports of the ‘Dutch Gold
Coast’, such as Elmina (present-day Ghana). Modern authors (Chippaux 2006;Trapeand
Mané 2006;Greenbaumetal.2015; Trape and Mediannikov 2016; Chippaux and Jackson
2019; Hallermann et al. 2020), tend to agree that although B. fuliginisous is still a very proble-
matic taxon, it corresponds to the dark-coloured Brown House Snake from Western Africa.
Boaedon lineatus was described by Duméril, Bibron and Duméril (1854) under the
French name Boedon Quatre-raises [ = four lines] based on two specimens collected in
‘Cote-D’or’, present-day Ghana. Later, Duméril (1861) established Boaedon quadrilineatum,
quadrilineatum literally meaning ‘four lines,’as an independent species and also mentions
that Günther (1858) considered Boaedon capense [=capensis] and Boaedon lineatus as
synonyms. B. fuliginosus and B. lineatus have sometimes been synonymised (Loveridge
1957; FitzSimons 1962; Roux-Estève and Guibé 1964; Broadley 1983), and much like
B. fuliginosus, B. lineatus has a complex taxonomic and nomenclatural history, with
several names currently placed under its synonymy (Wallach et al. 2014; Uetz et al.
2020). Modern authors (Chippaux 2006; Trape and Mané 2006; Greenbaum et al. 2015;
Trape and Mediannikov 2016; Chippaux and Jackson 2019) tend to agree that B. lineatus
is represented by the Striped House Snake from Western Africa described as uniformly
grey to reddish brown coloured, with two stripes on the side of head and two preoculars.
Recent authors tend to agree that both B. fuliginosus and B. lineatus represent species
complexes that need to be readdressed (Kelly et al. 2011; Wallach et al. 2014; Hallermann
et al. 2020), but so far this much needed taxonomic revision is still due. Given the con-
siderable number of names coined as subspecies of either fuliginosus or lineatus, the
number of names currently considered as synonyms of these two species and the impre-
cise data about the type locality of the nominotypical forms, this revision may prove to be
one of the most challenging taxonomic works of present-day African herpetology. These
species complexes have been the subject of recent studies (Greenbaum et al. 2015; Trape
and Mediannikov 2016; Hallermann et al. 2020) that have contributed to our understand-
ing of the diversity of the genus, but have not yet dealt with the root of the question,
namely the identity of B. fuliginosus,B. lineatus or even B. capensis.
This paper focuses on the São Tomé and Príncipe Boaedon populations. As part of an
ongoing revision of the herpetofauna of the islands, this paper presents a taxonomic and
nomenclatural review of the Jitas, including the description of a new species. We also
provide a taxonomic and nomenclatural review of the problematic Boaedon nigrum
Fischer, 1856.
Materials and methods
Material examined
For mensural and meristic comparisons, we examined specimens of Boaedon from
São Tomé and Príncipe. The specimens are deposited in the following institutions:
AFRICAN JOURNAL OF HERPETOLOGY 3

Muséum national d’Histoire naturelle (MNHN), Paris (France); Instituto de Investigação
Científica Tropical (IICT), Lisbon (Portugal); Museu Nacional de História Natural e da
Ciência (MUHNAC; formerly Museu Bocage - MB), Lisbon (Portugal); Center of Natural
History, Universität Hamburg, Zoologisches Museum (ZMH), Hamburg (Germany); Museo
Civico di Storia Naturale ‘Giacomo Doria’(MSNG), Genoa (Italy); University of Michigan
Museum of Zoology (UMMZ), Ann Arbor, MI (USA); California Academy of Sciences
(CAS), San Francisco, CA (USA); and Zoologisches Forschungsmuseum Alexander König
(ZFMK), Bonn (Germany). For a complete list of the specimens examined see the respect-
ive taxonomic accounts below. Locality geographic coordinates are reported in the form
of decimal degrees and use the WGS-84 map datum. Older (non-GPS) records were geor-
eferenced using the GEOLocate web application (https://www.geo-locate.org). All
elevations are reported as meters above sea level.
Molecular methods
DNA was extracted from 21 Boaedon tissue (Table 1) using a Qiagen DNeasy extraction
kit and protocol. We independently amplified two gene fragments (i.e. one mitochon-
drial [Cytochrome b(cyt b)], one nuclear [oocyte maturation factor mos (c-mos)]) using
Invitrogen Taq master mix by ThermoFisher Scientific(Waltham,MA,USA)withtwo
primer pairs: L14910 + H16064 (cyt b)andS77+S78(c-mos)fromEurofins Scientific
(Louisville, KY, USA). Cyt bwas amplified in 10 µl PCRs, with an initial denaturing temp-
erature of 94 °C for 3 minutes (min), followed by 40 cycles of denaturation at 94 °C for
30 seconds (s), annealing at 55°C for 30 s,and extension at 72°C for 90 s. C-mos was
amplified in 10 µl PCRs, with an initial denaturing temperature of 96°C for 1 min, fol-
lowed by 50 cycles of denaturation at 96°C for 10 s, annealing at 50°C for 30 s,and
extension at 60°C for 4 min (Slowinski and Lawson 2005). PCR amplicons were visual-
ised with 1% agarose gel with SYBRsafe gel stain and products were purified using
Table 1. Sequences and localities of individuals of Boaedon used generated in this study.
Species Catalog number cyt bc-mos Locality description
Boaedon bedriagae CAS218728 MW228125 MW228146 São Tomé Island
Boaedon bedriagae CAS218844 MW228126 MW228147 São Tomé Island
Boaedon bedriagae CAS218962 MW228127 –São Tomé Island
Boaedon bedriagae CAS219030 MW228128 –São Tomé Island
Boaedon bedriagae CAS219323 MW228129 MW228148 São Tomé Island
Boaedon bedriagae CAS219005 MW228130 –São Tomé Island
Boaedon bedriagae CAS219318 MW228131 MW228149 São Tomé Island
B. capensis b CAS168909 MW228132 –East Usambara Mtns, Tanzania
B. capensis b CAS168920 MW228133 –East Usambara Mtns, Tanzania
B. fuliginiosus a CAS201729 MW228134 –Byumba, Uganda
B. fuliginiosus a CAS201740 MW228135 –Byumba, Uganda
B. mendesi sp. nov. CAS233410 MW228136 MW228150 Príncipe Island
B. mendesi sp. nov. CAS238883 MW228137 –Príncipe Island
B. mendesi sp. nov. CAS238885 MW228138 MW228151 Príncipe Island
B. mendesi sp. nov. CAS238900 MW228139 MW228152 Príncipe Island
B. mendesi sp. nov. CAS238884 MW228140 MW228153 Príncipe Island
B. mendesi sp. nov. CAS238895 MW228141 MW228154 Príncipe Island
B. mendesi sp. nov. CAS238899 MW228142 MW228155 Príncipe Island
B. radfordi CAS201634 MW228143 –Bwindi, Uganda
B. radfordi CAS201635 MW228144 –Bwindi, Uganda
B. sp. 1 MVZ249808 MW228145 MW228156 Nkwanta, Volta region, Ghana
4L. M. P CERÍACO ET AL.

ExoSAP-IT or gel excision. Sequencing was performed in both forward and reverse
directions using the PCR primers on an Applied Biosystens ABI3130xl Genetic Analyzer
at the California Academy of Sciences’Center for Comparative Genomics, and
sequence chromatographs were edited using Geneious 5.5. No internal stop codons
were found and gaps in alignments were treated as missing data. Novel sequences
from this study were deposited in GenBank (MW228125 - MW228156). These sequences
were combined with those from other Lamprophiidae utilised in Greenbaum et al. (2015)
(see Appendix).
Phylogenetic analyses
To determine the evolutionary history and distinction of Boaedon from the islands of
São Tomé and Príncipe, we conducted Bayesian mixed-model analyses performed in
MrBayes v.3.0b4 (Ronquist and Huelsenbeck 2003) using models selected based on
Akaike information criterion (AIC) conducted in MrModeltest 2.2 (Nylander 2004) run in
PAUP*v4.0b10 (Swofford 2002). Two simultaneous runs were conducted (three heated
and one cold chain with the default Markov chain Monte Carlo settings), for a total of
6×10
6
generations per run, sampling trees and parameters every 100 generations and
the first 1.5 × 10
6
generations from each run were discarded as burn-in. Trees were
rooted with outgroup taxa Amblyodipsas dimidiata,Bothrolycus ater,Bothrophthalmus
lineatus,Gonionotophis brussauxi, and Lycophidion capense included in Greenbaum
et al. (2015).
Finally, we computed pairwise comparisons of the cyt bgene fragment as a
measure of estimated genetic distances within and between species of Boaedon
using MEGA X (Kumar et al. 2018;Table 2). These analyses were conducted using
the Maximum Composite Likelihood model (Tamura et al. 2004)andinvolved76
informative nucleotide sites including all three codon positions. All ambiguous pos-
itions of the final 1 116 bp dataset were removed for each sequence pair (pairwise
deletion option).
Morphological methods
Specimens were measured with a flexible measuring tape for snout–vent length (SVL)
and tail length (TL) to a precision of one millimetre, and all other measurements were
recorded with a digital calliper or an ocular micrometer to a precision of a tenth of a
millimetre. Lepidosis was observed with the help of a stereomicroscope. Scale nomen-
clature, scales counts and measurements used in the descriptions follow Greenbaum
et al. (2015) and Hallermann et al. (2020). The following characters were measured:
snout-vent length (SVL), from the tip of the snout to the anterior edge of the
cloaca; tail length (TL), from the posterior edge of the cloaca to the tip of the tail.
For type material we also measured the head length (HL), from the tip of snout to
just behind the angle of the jaw; interocular distance and eye diameter. The following
scale counts were made: number of dorsal scale rows at midbody (MSR); number of
ventral scales (V), from the first scale broader than long to the cloacal plate (we
used the traditional method of ventral scale count to make our work comparable to
Greenbaum et al. (2015) and Hallermann et al. (2020)); number of subcaudal scales
AFRICAN JOURNAL OF HERPETOLOGY 5

Table 2. Pairwise sequence divergences within (diagonal in italics) and among (below diagonal) all Boaedon species as defined in this study. Sequences used were
only from the cyt bgene fragment and values are rounded to the nearest hundredth place. Values for B. mendesi sp. nov. are in bold.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16
1–B. bedriagae 0.01
2-B. capensis a 0.13 n/c
3-B. capensis b 0.13 0.10 0.06
4-B. capensis c 0.09 0.11 0.11 n/c
5-B. fradei 0.12 0.13 0.12 0.11 0.01
6-B. fulginosus a 0.11 0.13 0.12 0.13 0.11 0.01
7-B. fulginosus b 0.13 0.13 0.13 0.13 0.11 0.09 0.03
8-B. lineatus 0.12 0.13 0.12 0.12 0.14 0.11 0.11 n/c
9-B. mentalis 0.13 0.14 0.15 0.13 0.14 0.13 0.12 0.13 0.00
10 - B. olivaceus 0.12 0.12 0.12 0.12 0.12 0.09 0.10 0.10 0.11 0.01
11 - B. radfordi 0.11 0.12 0.11 0.12 0.11 0.09 0.07 0.11 0.11 0.10 0.01
12 - B. sp. 1 0.18 0.18 0.18 0.18 0.17 0.18 0.17 0.16 0.18 0.17 0.16 n/c
13 - B. sp. 2 0.10 0.12 0.12 0.12 0.10 0.08 0.06 0.11 0.11 0.09 0.06 0.16 n/c
14 - B. upembae 0.16 0.17 0.17 0.16 0.15 0.17 0.17 0.18 0.17 0.17 0.16 0.16 0.16 0.00
15 - B. virgatus 0.18 0.19 0.19 0.18 0.19 0.18 0.18 0.19 0.18 0.19 0.18 0.14 0.17 0.17 n/c
16 - B. mendesi sp. nov. 0.06 0.13 0.18 0.13 0.13 0.12 0.13 0.12 0.14 0.12 0.12 0.17 0.11 0.18 0.18 0.00
6L. M. P CERÍACO ET AL.

(SC); number of supralabials; number of infralabials; number of preoculars; number of
postoculars; number of supralabials touching the eye orbit; the ratio of V and Sc (V/SC)
was also recorded. The length and height of the loreal scale (LOR), as well as its ratio
(LOR-L/H) were recorded. Snout length is described in relation to the length of the par-
ietal shield (PAR): the PAR is either equal to the distance between the frontal and the
rostral scale or longer. Finally, colouration pattern was reported in preserved speci-
mens. A slash (/) represents characters from the right and left sides of the body,
always in that order.
Information on morphological characters of species and/or type material that could
not be examined, as well as supplemental data for all Central and West African
Boaedon was obtained from the relevant literature (e.g. Greenbaum et al. 2015;
Chippaux 2006; Trape and Mané 2006; Trape and Mediannikov 2016; Chippaux and
Jackson 2019; Hallermann et al. 2020). There has been a long history of confusion
regarding the identity of some of Boaedon taxa, and the allocation of certain names
to synonymy. This is especially true for Boaedon fuliginosus and B. lineatus. For these
reasons, we provide comparisons between the two São Tomé and Príncipe Boaedon
populations and all West and Central African congeners. From a taxonomy and nomen-
clature perspective, B. fuliginosus is still one of the most challenging groups within
Boedon. Our interpretation of ‘fuliginosus’includes only West African B. fuliginosus
(from Morocco in the north to Angola in the south), which comprise a genetic unit, fol-
lowing Hallermann et al. (2020). Morphologically similar fuliginosus-like species from East
and Central Africa are excluded from comparison. This is in accordance with the most
updated and informed interpretations of what ‘true’fuliginosus really is (Greenbaum
et al. 2015; Trape and Mediannikov 2016; Chippaux and Jackson 2019). The same com-
parisons apply to B. lineatus.B. capensis was not considered for these comparisons, as it
is an extralimital species.
Results
Phylogenetic analysis
We conducted a Bayesian phylogenetic analysis using mtDNA (cyt b, 1 116 bp) alone and
a second analysis that combined cyt band a nuclear (c-mos, 579 bp) gene fragment.
MrModeltest 2.2 selected the following partitioned nucleotide substitution models for
our dataset: cyt bcodon 1 (GTR+ Γ), codon 2 (HKY + I+Γ), codon 3 (GTR + I+Γ); c-mos
codon 1 (HKY), codon 2 (GTR+ Γ), codon 3 (HKY).
The phylogenies generated from these data and partitions produced nearly identi-
cal topologies, though less support in the combined analysis (Figure 1), with compar-
able relationships as found in recent studies (e.g. Kelly et al. 2011; Greenbaum et al.
2015). Similar to these studies we found strong support for a basal split within
Boaedon between a clade of B. upembae and B. virgatus and the remaining Boaedon,
though our analyses add a potential new species, B. sp. 1, that is sister to
B. virgatus. Furthermore, we concurred support for a split of B. mentalis from the
remaining Boaedon. Similar to previous studies, the majority of interspecies relation-
shipsinthiscladeofBoaedon were not supported. One consistency, however, was
support for the paraphyly of B. capensis and B. fulginosus, even after recognizing
AFRICAN JOURNAL OF HERPETOLOGY 7

Figure 1. Bayesian 50% majority-rule consensus phylogram of Boaedon inferred from a partitioned
dataset of the cyt b gene fragment. Posterior probability nodal support values of 0.95 and above
are displayed and considered highly supported, while values <0.95 are not displayed or represented
by **. Support values on the left result from the cyt b phylogeny, while values on the right are derived
from a separate Bayesian phylogeny utilizing a combined mtDNA (cyt b) and nuclear (c-mos) gene
fragment dataset (total of 1 695 bp). Species names follow those given in Table 1 and Appendix.
Photograph of Boaedon mendesi sp. nov. from Príncipe by LMPC.
8L. M. P CERÍACO ET AL.

some clades as distinct taxa (e.g. B. fradei,B. radfordi). More specifically, our cyt bphy-
logeny found strong support for a clade consisting of B. sp. 2, B. fuliginosus b, and
B. radfordi along with a sister relationship between two distinct island species of
Boaedon from Príncipe and São Tomé islands that are together sister to the
B. capensis c lineage found within a clade that includes sister taxa B. capensis aand
B. capensis b.
Our phylogenetic analyses found strong support that all our 16 designated species of
Boaedon belong to their own unique clade. Sequence divergences of cyt bamong these
taxa ranged from 6 to 19%, while divergences within species ranged between 0.0 and 6%
(Table 2). The sequence divergence within the island species from São Tomé (i.e.
B. bedriagae) was 0.532%, while the population from Príncipe was 0.394%. These
species are further supported by a 6% divergence, compared to the next nearest diver-
gence of 9% by B. bedriagae and B. capensis c. Based on these results we recognise the
population from Príncipe as being a distinct species from B. bedriagae and other
Boaedon under the General Lineage Concept of species (de Queiroz 1998) and therefore
distinguish it morphologically below.
Morphological analysis
Results of our morphological analyses are shown in Table 3. The populations
of São Tomé and Príncipe islands are very similar to each other, without major
morphological or meristic characters differences, although they can unambiguously
be diagnosed against each other by their colouration patterns (see diagnoses
below). As noted by Trape and Mediannikov (2016), colouration pattern is one of
the most reliable and useful characters for the genus. Both islands populations can
also be distinguished from the continental forms based on a combination of charac-
ters (see taxonomic accounts below). Based on these differences and the results of
our molecular analyses, we recognise these clades as new or revalidated species,
respectively.
Table 3. Comparison between B. bedriagae and B. mendesi sp. nov.
Boaedon bedriagae
(n= 37) Boaedon mendesi sp. nov. (n= 20)
Maximum SVL (cm) 107 122
Maximum TL (cm) 24.5 22
Midbody scale rows 25–31 24–29
Ventral scales 210–247 206–242
Subcaudal scales 65–95 61–81
Subcaudals rows Double Double
Preoculars 1 1 (rarely 2)
Postoculars 2 2
Supralabials
touching the eye
Usually 3rd, 4th and 5th, sometimes only 4th
and 5th
Usually 3rd, 4th and 5th, rarely 4th, 5th and 6th
Cloacal plate Entire Entire
Head stripes Present, broad Present, broad
Colouration
(dorsum)
Dark brown, with parallel mediodorsal and
dorsolateral bands running from the neck to
the tail
Dark brown, with a series of dorsolateral cream
blotches running from the neck to the
midbody
Colouration
(ventral)
Cream, heavily speckled with dark markings Whitish, without markings
AFRICAN JOURNAL OF HERPETOLOGY 9

Systematics
Boaedon bedriagae Boulenger, 1906
(Figures 2 to 5)
Boaedon quadrilineatus: Jan and Sordelli (1870: Plate II)
Boodon quadrilineatum: Bocage (1879: 87; 1887: 200)
Boodon capensis: Greeff(1884: 48)
Boodon capense: Bocage (1886: 69; 1889: 34); Vieira (1886: 237)
Boodon lineatus [part]: Bedriaga (1892: 904; 1893: 439); Bocage (1905: 93);
Henriques (1917: 81, 143); Themido (1941: 6).
Boodon bedriagae [part]: Boulenger (1906: 211).
Boodon bedriagae: Angel (1920: 199)
Boaedon lineatus bedriagae: Bogert (1940:6)
Boaedon lineatus bedriagai [part]: Manaças (1958: 188); Capocaccia (1961a: 63; 1961b: 288)
Boaedon fuliginosus bedriagae [part]: Roux-Estéve and Guibé (1964: 770); Chippaux and
Jackson (2019: 192)
Boaedon bedriagae [part]: Roux-Estéve and Guibé (1965: 400); Ceríaco et al. (2018: 103)
Boaedon fuliginosus bedriagai [part]: Manaças (1973: 225)
Lamprophis fuliginosus bedriagae [part]: Schätti and Loumont (1992: 30); Nill (1993: 72);
Hofer (2002: 89)
Lamprophis fuliginosus [part]: Chippaux (2006: 65)
Boaedon lineatus [part]: Wallach et al. (2014: 96)
The first available records referring to the species were published by Jan and Sordelli
(1870), who provide an illustration of a São Tomé specimen collected by the German
medical surgeon Carl Weiss (dates of birth and death unknown) (Figure 2). The specimen
depicted was identified as Boaedon quadrilineatum and is still present in the collections of
the ZMH (R01256; Figure 3). Subsequently, several authors provided additional records
from São Tomé island. Bocage (1879) notes a specimen of Boaedon quadrilineatum col-
lected by Custodio de Borja, a navy officer and General Secretary of the Government of
São Tomé e Príncipe. Interestingly, Bocage (1879) mentions that the local name was
already ‘Dgita’. The German naturalist Richard Greeff(1828–1892) who explored the
Gulf of Guinea islands between 1879 and 1880, published records of ‘Boodon capensis’
Figure 2. Head of Boaedon bedriagae identified as Boaedon quadrilineatus by Jan and Sordelli (1870).
Adapted from Jan and Sordelli (1870).
10 L. M. P CERÍACO ET AL.

from São Tomé (Greef 1884). Some of Greef’s original specimens are still extant in the col-
lections of ZMH and ZMB.
From May to September 1885, Adolfo F Moller (1842–1920), chief gardener of the
Coimbra Botanical Garden, collected plants and animals in São Tomé for the collections
of the University of Coimbra. A preliminary list of the material collected by Moller, including
a list of herpetological material identified by Bocage, was published by Vieira (1886). In this
list, Vieira (1886) cites the presence of ‘Boodon capense, Dum. & Bibr.’. These specimens
remained in Portugal in the collections of the Zoological Museum of the University of
Coimbra (currently Museu da Ciência da Universidade de Coimbra (MCUC)), Portugal
(Themido 1940). Notably, the specimens were not located in a recent visit to the MCUC
(LMPC pers. obs.). However, some specimens collected by Moller were sent to the
Russian naturalist Jacques von Bedriaga (1854–1906), who published a review of
São Tomé herpetofauna based on the specimens from Moller (Bedriaga 1892, 1893). In
this review, Bedriaga referred to the presence of ‘Boodon lineatus D.B.’and provided a
detailed description of the specimens at hand (Bedriaga 1892, 1893). To date, it is
unclear what happened to these specimens. According to the available biographies of Bed-
riaga (Böhme 1996;Bischoffand Böhme 2001), the Russian herpetologist never visited Por-
tugal. Bedriaga moved to Nice, France in 1881 and later moved to Florence, Italy, where he
died in 1906, never returning to Russia (Böhme 1996;Bischoffand Böhme 2001). It is prob-
able that these specimens were sent to Nice, but their whereabouts are currently unknown.
The Portuguese explorer Francisco Newton (1864–1909), hired by the Natural
History Museum of Lisbon for the zoological exploration of the Gulf of Guinea, col-
lected and sent several specimens of reptiles from São Tomé Island. Based on these
Figure 3. Specimen of Boaedon bedriagae (ZMH R01256) depicted in Jan and Sordelli (1870).
AFRICAN JOURNAL OF HERPETOLOGY 11

records Bocage (1886,1889,1905) published several records of São Tomé Jitas, either
identifying them as Boaedon capense (Bocage 1886, 1889) or B. lineatus (Bocage 1905).
Some years after Newton, the Italian naturalist Leonardo Fea (1852–1903) explored the
islands in the Gulf of Guinea between 1899 and 1900 and collected specimens for the
natural history museum of Genoa, Italy. Based on five specimens from São Tomé Island
(Vista Alegre and Ribeira Palma) and three specimens from Príncipe Island collected by
Fea, Boulenger (1906) described ‘Boodon bedriagae,’a new species named in honor of
Jacques von Bedriaga. According to Boulenger (1906), the main characters that differ-
entiate B. bedriagae from B. lineatus were 1) ‘the third upper labial constantly entering
the eye (only very exceptionally in B. lineatus)’and 2) ‘the markings of the head’(Boulenger
1906). The type series of B. bedriagae is currently extant in the collection of the Museo Civico
di Storia Naturale ‘Giacomo Doria’, in Genoa, Italy (Capocaccia 1961a, 1961b). Angel (1920)
presented a subsequent record of the newly described B. bedriagae and was the only author
until very recently to consider the taxon as a full species.
Bogert (1940) examined a single specimen from São Tomé collected by the Portuguese
naturalist José Correia (1881–1954) and he noted that São Tomé [and Príncipe] island Jitas
were not sufficiently distinct to be regarded as full species and that they were instead a sub-
species of Boaedon lineatus. According to Bogert (1940), the distinctive character presented
by Boulenger (1906)todistinguishB. bedriagae from B. lineatus, the contact between labials
and the eye, is ‘exceedingly variable in the typical form [lineatus], and thus can have no diag-
nostic significance in separating bedriagae’,but he argued that scale counts, size and colour-
ation warrant diagnostic differences between the two forms. Bogert (1940) also emphasised
the considerably larger size of the São Tomé forms compared to those from the continent.
Manaças (1958), following the taxonomic interpretation of Bogert (1940), reported three
specimens from the island. Cappocacia (1961a) reviewed the type series of Boodon bedriagae
and other specimens not studied by Boulenger (1906) present in the Genoa museum, noting
that São Tomé island population had characters that allowed differentiation from Príncipe
island population. Namely, the number of ‘longitudinal series of dorsal scales’was lower
in specimens from Príncipe than those from São Tomé. Due to the small amount of compara-
tive material at her disposal, Cappocacia (1961a) did not propose any taxonomic change. In a
review of the Boaedon genus, Roux-Estéve and Guibé (1964)consideredB. bedriagae asub-
species of B. fuliginosus, much like the authors considered B. lineatus a synonym of
B. fuliginosus. These authors refuted Boulenger’s(1906) diagnostic characters, noting the
main characters that differentiated bedriagae from the nominotypic form were a higher
number of subcaudal scales in both male and female bedriagae and a colouration pattern
more closely resembling B. quadrilineatus than B. fuliginosus. Without further explanations
Roux-EstéveandGuibé(1965) considered B. bedriagae a full species.
Subsequent authors provided more records of the São Tomé’s Jita and revisionary works
have presented different interpretations on the taxonomic identity. Manaças (1973)pre-
sented the taxon as Boaedon fuliginosus bedriagae; Schätti and Loumont (1992)andNill
(1993)asLamprophis fuliginosus bedriagae.Chippaux(2006) considered them a synonym
of B. fuliginosus. Wallach et al. (2014) synonymised the taxon to B. lineatus lineatus.
Ceríaco et al. (2018) considered it a full species, while Chippaux and Jackson (2019) con-
sidered it a subspecies of B. fuliginosus. In the past decade, the species has been collected
by teams of the California Academy of Sciences (CAS; Figure 4) and the Museu Nacional de
História Natural e da Ciência (MUHNAC; Figure 5) (this paper).
12 L. M. P CERÍACO ET AL.

According to our results, São Tomé and Príncipe islands populations are not con-
specific. Since B. bedriagae was described based on specimens from both São Tomé
and Príncipe, the type series is currently composed by homonyms. Therefore, following
Article 74.1 from the International Code of Zoological Nomenclature (ICZN 1999), it is
mandatory to designate a lectotype to B. bedriagae from the available syntypes. For
Figure 4. Live photo of Boaedon bedriagae. Photo by Andrew Stanbridge.
Figure 5. Specimen of Boaedon bedriagae (MUHNAC/MB03–978) from São Tomé Island.
AFRICAN JOURNAL OF HERPETOLOGY 13

this, we chose specimen MSNG/CE 30987b, from Vista Alegre, São Tomé island, part of the
original type series, as a lectotype of B. bedriagae.
Lectotype —MSNG/CE 30987b, an adult from Vista Alegre [0.318941°, 6.676295°, 336 m],
Mé-Zochi district, São Tomé Island, collected by Leonardo Fea on 1900.
Additional material —SÃO TOMÉ ISLAND: Água Grande region: MUHNAC/MB03-
000727, São Marçal [0.32487°, 6.73906°, 9 m] collected by Manuel Sacramento and
Afonso A Junior on 7 July 1984; MUHNAC/MB03-000978, São Tomé [0.333333°,
6.733333°, 9 m] collected by an unknown collector on 7 November 1984. Cantagalo
region: IICT/R 51/1954, Água Izé [0.218168°, 6.725793°, 26 m] collected by Fernando
Frade on 9 December 1954; MUHNAC/MB03-00719, 00721, Água Izé [0.217896°,
6.725°, 38 m] collected by Ruben Soares, Teresa Pité and Artur Serrano on 14 and 18
June 1984, respectively; CAS 218844-218847, Caxueira, along Água Péte Péte, [0.298°,
6.73036°, 49 m] collected by Robert C Drewes, Ricka E Stoelting and Jens V Vindum
on 5 April 2001; CAS 258789, Roça Água Izé, shade plantation along road to Bernardo
Faro [0.227529°, 6.726019°, 33 m] collected by Rayna C Bell, Maria A Jerónimo, Andrew B
Stanbridge and Luis Mendes on 39 Setember 2015. Caué region: IICT/R 8/1954, Roça
Porto Alegre [0.033333°, 6.533333°, 16 m] collected by Fernando Frade on 11 October
1954; IICT/R 2-1967, Angra Toldo [0.15770°, 6.67070°, 19 m] collected by an unknown col-
lector on 1 June 1967; CAS 219311, Cruzeiro [0.288278°, 6.681194°, 303 m] collected by
Ricka E Stoelting on 5 May 2001; CAS 258812, inland from Praia Micondo [0.173444°,
6.67625°, 72 m] collected by Rayna C Bell, Maria A Jerónimo and Andrew B Stanbridge
on 30 September 2015; UMMZ 187961-187966, Pousada, area 850 m [0.231722°,
6.597921°, 749 m] collected by Ronald A Nussbaum and Michael E Pfrender on 4 and 7
June 1988; UMMZ 187969-187970, Colónia Açoreana 60 m [0.183333°, 6.683333°, 125 m]
collected by Ronald A Nussbaum and Michael E Pfrender on 10 June 1988. Lembá
region: IICT/R STP12, Ponta-Figo [0.339465°, 6.54286°, 151 m] collected by an unknown col-
lector on 24 May 1958; CAS 259001Contador Valley, road to aqueduct/reservoir system past
Ponta Figo [0.316417°, 6.550278°, 639 m] collected by Rayna C Bell, Maria A Jerónimo and
Andrew B Stanbridge on 13 October 2015. Lobata region: IICT/R 17/1967, Roça Boa Entrada
[0.35°, 6.666667°, 180 m] collected by an unknown collector in 20 August 1967; MUHNAC/
MB03-000724, Guadalupe [0.379342°, 6.638327°, 14 m] collected by Ruben Soares, Teresa
Pité and Artur Serrano on 4 July 1984; CAS 219030, Rio do Ouro below Agustinho Neto
[0.365472°, 6.644917°, 160 m] collected by Robert C Drewes, Ricka E Stoelting and Jens V
Vindum on 13 April 2001; CAS 218962, base of Muquinqui [0.381194°, 6.648778°, 137 m]
on 11 April 2001; CAS 218728, coast road S of Lagoa Azul [0.408083°, 6.58675°, 0 m] on
30 March 2001; CAS 258724, Obo National Park, Monte Carmo Roca, outside of Malanza
village, EMOLVE plantation (Agripalma) [0.105194°, 6.602583°, 79 m] collected by Rayna C
Bell, Lauren A Scheinberg, Andrew B Stanbridge and Ricardo F de Lima. Mé-Zóchi
region: IICT/R 4/1966, 18/1967, 21/1967, Roça Potó-Correia [0.29685°, 6.680288°, 291 m]
collected by an unknown collector on 20 and 23 August 1967; MUHNAC/MB03-000720,
000722, Nova Moca [0.283333°, 6.633333°, 771 m] collected by Ruben Soares, Teresa Pité
and Artur Serrano on 15 June 1984; MUHNAC/MB03-000725, Monte Café [0.299976°,
6.640108°, 694 m] collected by Ruben Soares, Teresa Pité and Artur Serrano on 24 June
1984; MB03-000726, Potó Correia a 7 km da Madalena [0.29685°, 6.680288°, 291 m]
14 L. M. P CERÍACO ET AL.

collected by Ruben Soares, Teresa Pité and Artur Serrano on 20 June 1984; MUHNAC/MB03-
000978 (Figure 5), Jardim Botânico Bom Sucesso [0.27427°, 6.58581°, 1 209 m] collected by
Luis MP Ceríaco and Mariana P Marques on 15 February 2015; CAS 219005, Java [0.261083°,
6.650889°, 588 m] collected by Robert C Drewes, Ricka E Stoelting and Jens J Vindum on 11
April 2001; CAS 219277, Bom Sucesso [0.288861°, 6.612417°, 1 155 m] collected by Robert C
Drewes and Ricka E Stoelting on 26 April 2001; CAS 219318, road from Bom Sucesso to radio
tower [0.275778°, 6.610417°, 1 234 m] collected by Ricka E Stoelting on 5 May 2011; CAS
219322, W of radio antenna above Bom Sucesso [0.275556°, 6.605194°, 1 324 m] collected
by Ricka E Stoelting on 6 May 2011; CAS 219323, Bom Sucesso [0.288667°, 6.612444°,
1 151 m] collected by Ricka E Stoelting on 7 May 2011; CAS 252830, forest between Bom
Successo and Lagoa Amelia [0.284811°, 6.602397°, 1 312 m] collected by M Jirku on 31
October 2011; CAS 258725, 258726 Roça Bombaim [0.245861°, 6.632278°, 462 m] collected
by Rayna C Bell, Lauren A Scheinberg, Maria A Jerónimo and Andrew B Stanbridge on 23
September 2015; CAS 258858 ca 0.9 km E of Madalena, along ES4 [0.327028°, 6.673222°,
232 m] collected by Rayna C Bell, Lauren A Scheinberg, Maria A Jerónimo and Andrew B
Stanbridge on 2 October2015; CAS 258969, Bom Sucesso [0.2875°, 6.611639°, 1 147 m] col-
lected by Rayna C Bell, Lauren A Scheinberg, Maria A Jerónimo, Andrew B Stanbridge,
Robert C Drewes and KB Lim on 12 October 2015; ZFMK 82964 Sao Nicolao [0.27972°,
6.62556°, 968 m] collected by Daniel Hofer on October 2003. Unknown locality: IICT/R
3/1966, São Tomé Island collected by unknown collector on 20 August 1966; IICT/R 19/
1967, São Tomé Island, collected by unknown collector on 24 August 1967; IICT/R STP13,
São Tomé Island, collected by unknown collector on 7 November 1984; MNHN 1920.10-
14, Sans localité precise (= without precise locality), 13 February 1920; ZMH R01256
(Figure 3), without precise locality (St. Thomas), collected by Carl Weiss in 1858.
Diagnosis —Boaedon bedriagae (Figures 2 to 5) can be distinguished from all of its
Central and West African congeners by the following combination of characters: dark
brown colouration with parallel mediodorsal and lateral bands running from the neck
to the tail; two broad, cream-coloured, dark brown-bordered bands running on the
side of the head; the upper band starting on the nasals and extending above the eye
to the posterior part of the head, the lower band usually starting on the posterior
lower part of the eye and extending to the corner of the mouth; venter cream-coloured
speckled with dark brown markings; the 3rd, 4th and 5th supralabials contacting the eye,
one preocular; 25 to 31 midbody scales rows, 210 to 247 ventral scales and 65 to 95 sub-
caudals, and a maximum snout–vent length of 107 cm.
It can be distinguished from B. fuliginosus by its higher number of ventral and sub-
caudal scales (210 to 247 and 65 to 95, respectively, in B. bedriagae versus 194 to 229
and 46 to 71 in W-African B. fuliginosus fide Hallermann et al. (2020), by having three
supralabials touching the eye (versus only two in B. fuliginosus fide Hallermann et al.
2020), by having two broad cream, dark-bordered bands running on the side of the
head (versus two thin white lines in B. fuliginosus), and by having dark brown colour-
ation with parallel mediodorsal and lateral bands running from the neck to the tail
(versus a homogeneous dark brown colouration in B. fuliginosus); it can be distin-
guished from B. lineatus by dark brown colouration with parallel mediodorsal and
lateral bands running from the neck to the tail (versus a uniformly grey to reddish
brown colouration in B. lineatus) and by having only one preocular (versus usually
AFRICAN JOURNAL OF HERPETOLOGY 15

two in B. lineatus); it can be distinguished from B. olivaceus,B. poensis and B. radfordi by
having the subcaudals divided (versus single in B. olivaceus,B. poensis and B. radfordi); it
can be distinguished from B. virgatus by having two broad cream, dark-bordered bands
running on the side of the head (versus two thin white lines in B. virgatus), by a higher
number of midbody scales rows (25 to 31 in B. bedriagae versus 23 to 25 in B. virgatus),
and a higher number of ventral and subcaudal scales (210 to 247 and 65 to 95, respect-
ively, in B. bedriagae versus 186 to 223 and 42 to 64 in B. virgatus); it can be distin-
guished from B. upembae by having more ventral scales (210 to 247 in B. bedriagae
versus 175 to 197 in B. upembae), and a higher number of midbody scale rows (25 to
31 in B. bedriagae versus 21 to 23 in B. upembae). It can be distinguished from
B. perisilvestris by the 3rd, 4th and 5th supralabial in contact with the eye (versus
usually the 4th and 5th in B. perisilvestris), by having two broad cream, dark-bordered
bands running on the side of the head (versus no lines in B. perisilvestris), and by having
dark brown colouration with parallel mediodorsal and lateral bands running from the
neck to the tail (versus a homogeneous dark brown colouration in B. perisilvestris); it can
be distinguished from B. paralineatus by the 3rd, 4th and 5th supralabial in contact with
the eye (versus only the 4th and 5th in B. paralineatus), by its lower number of midbody
scale rows (25 to 31 in B. bedriagae versus 31 to 35 in B. paralineatus); it can be distin-
guished from B. subflavus by the 3rd, 4th and 5th supralabial in contact with the eye
(versus only the 4th and 5th in B. subflavus); having two broad, cream-coloured and
dark-bordered bands running on the sideofthehead(versusonlyonelinein
B. subflavus), and by having dark brown colouration with parallel mediodorsal and
lateral bands running from the neck to the tail (versus a homogeneous yellow to
light brown colouration in B. subflavus); it can be distinguished from B. longilineatus
by having a higher number of subcaudals (65 to 95 versus 42 to 63 in
B. longilineatus) and by having two broad coloured and dark-bordered bands
running on the side of the head (versus only one, continuous from the snout to the
tail, in B. longilineatus); it can be distinguished from B. littoralis by having dark
brown colouration with parallel mediodorsal and lateral bands running from the
neck to the tail (versus homogeneous light brown colouration in B. littoralis).
Distribution —The species is endemic to São Tomé island and Rolas islet, Gulf of Guinea.
The species has been recorded from the northern, central and south-eastern areas of the
island (Figure 6), and is likely distributed throughout.
Habitat and natural history notes —Not much is known about the natural history of
this species. Manaças (1973) noted that one female collected in July 1967 had 13 eggs,
each around 26 mm high by 12 mm wide. Another female, collected in August 1967, had
a total of five eggs, the largest measuring around 63 mm high by 14 mm wide and the
smallest measuring 28 mm high by 12.5 mm wide. The species is commonly found in
forested areas, but also in slightly disturbed habitats. Nill (1993) noted that in human
settlements the species could rely on Hemidactylus mabouia and Panaspis thomensis
as prey items. The same author found juveniles hunting Hyperolius and Phrynobatrachus
at night.
16 L. M. P CERÍACO ET AL.

Boaedon mendesi sp. nov.
(Figures 7 to 8)
Boodon geometricum ?: Bocage (1887: 199)
Boodon lineatus [part]: Bocage (1903: 54); Henriques (1917: 81)
Boodon bedriagae [part]: Boulenger (1906: 211)
Boaedon lineatus bedriagai Boulenger (Capocaccia 1961a: 63; 1961b: 288; Manaças 1958:
188)
Boaedon fuliginosus bedriagae [part]: Roux-Estéve and Guibé (1964: 770); Chippaux and
Jackson (2019: 192)
Boaedon bedriagae [part]: Roux-Estéve and Guibé (1965: 400); Ceríaco et al. (2018: 103)
Boaedon fuliginosus bedriagai: Manaças (1973: 225)
Lamprophis fuliginosus: Chippaux (2006: 65)
Figure 6. Confirmed records of Boaedon bedriagae in São Tomé Island.
AFRICAN JOURNAL OF HERPETOLOGY 17

Figure 7. Live photo of the holotype of Boaedon mendesi sp. nov. (MUHNAC/MB03–977).
Figure 8. Holotype of Boaedon mendesi sp. nov. (MUHNAC/MB03–977).
18 L. M. P CERÍACO ET AL.

Boaedon lineatus [part]: Wallach et al. (2014: 96).
Fewer historical records of the Príncipe Jita exist compared to the São Tomé Jita. Bocage
(1887) provided the first record of the taxon from Príncipe island based on a specimen
collected by Francisco Newton and Bocage identified it as a putative Boodon
geometricum. Subsequently, Bocage (1903) cited additional specimens from Príncipe,
this time identifying them as Boodon lineatus, and noting the local common name
‘Cobra Cabussam’. As noted in the account of Boaedon bedriagae, Boulenger (1906)
used three specimens from Príncipe island as part of the syntypes of Boodon bedriagae.
The taxonomic identity of the Príncipe population faced the same uncertainties as
B. bedriagae, since at the time they were considered the same taxonomic entity.
Different authors have argued that the Príncipe population is either a subspecies of
B. fuliginosus (Roux-Estéve and Guibé 1964; Manaças 1973; Hofer 2002; Chippaux and
Jackson 2019), a subspecies of B. lineatus (Manaças 1958; Capocaccia 1961a, 1961b;), a
valid species (Roux-Estéve and Guibé 1965), a synonym B. fuliginosus (Chippaux 2006)
or a synonym of B. lineatus (Wallach et al. 2014). Capocaccia (1961a) was the first to
denote some morphological differences between the São Tomé and Príncipe populations,
although she refrained from making any taxonomic decision, due to the small number of
individuals at her disposal. Ceríaco et al. (2018) noted that the Príncipe population was
different from the one from São Tomé and referenced a description elsewhere (this
paper). According to our molecular and morphological results, the Príncipe population
is not conspecific with B. bedriagae from São Tomé. As there are no available names to
the Príncipe population, we describe it here as a new species, Boaedon mendesi sp. nov.
Holotype —MUHNAC/MB03-977 (Figures 7 to 8), an adult male collected in the vicinities of
Santo Cristo (1.63633°, 7.42690°, 80 m), Príncipe Island, Republic of São Tomé e Príncipe, col-
lected by Luis MP Ceríaco, Mariana P Marques, and Pedro NP Ceríaco on 10 February 2015.
Paratypes —All specimens from the Island of Príncipe, Republic of São Tomé e Príncipe.
Three specimens: CAS 233410, Príncipe Island [1.659306°, 7.395389°, 178 m] collected by
Robert C Drewes, Joseph Uyeda and Jens V Vindum on 11 May 2006; IICT/R 52-56/1957,
Principe Island, collected by Décio Passos on March 1955; MB03-001027, road to Porto
Real [1.62617°, 7.41344°, 38 m] collected by Luis MP Ceríaco, Mariana P Marques, Ana C
Sousa and Ostilino C Rocha on 17 February 2016.
Additional material. CAS 238883, on road near Ponta do Sol [1.654889°, 7.381278°, 200 m]
collected by Robert C Drewes on 29 April 2008; CAS 238884, near Ponta do Sol [1.655889°,
7.380944°, 200 m] collected by Robert CDrewes on 29 April 2008; CAS238885, 238895, Bom
Bom, East slope [1.698833°, 7.403667°, 53 m] collected by Robert C Drewes on 29 April and 2
May 2008, respectively; CAS 238899, 238900, Bom Bom [1.698167°, 7.402667°, 58 m] col-
lected by Robert C Drewes on 3 May 2008; CAS 251572, base of João Dias Filho, João
Dias Pai [1.600917°, 7.374278°, 215 m] collected by Robert C Drewes on 27 May 2012;
CAS 261030, trail from Pico Mesa camp to Baia das Agulhas [1.588181°, 7.357781°, 293 m]
Rayna C Bell, Lauren A Esposito, Felipe Spina, and Ostilino C Rocha on 5 December 2016;
IICT/R 26-1954, Roça Sundy [1.668775°, 7.383353°, 169 m] collected by Fernando Frade
on 17 November 1954; IICT/R 113, 114-1956, Roça Sundy [1.668775°, 7.383353°, 169 m]
AFRICAN JOURNAL OF HERPETOLOGY 19

collected by João Tendeiro on 17 November 1954; IICT/R 8/1967, Roça Sundy [1.668775°,
7.383353°, 169 m] collected by an unknown collector on 4 July 1967; IICT/R 63-1955, Prin-
cipe Airport [1.662259°, 7.41235°, 180 m] collected by Décio Passos on 6 September 1955;
IICT/R 109/1955, Santo António [1.636944°, 7.419444°, 12 m] collected by João Tendeiro on
28 November 1955; MB03-000977, trail to Santo Cristo [1.63427°, 7.42802°, 134 m] collected
by Luis MP Ceríaco, Mariana P Marques, Pedro NP Ceríaco and Ostilino C Rocha on 10
February 2015; UMMZ 187968, Ribeira Bacharel [1.614117°, 7.401889°, 19 m] collected by
Ronald A Nussbaum and Michael E Pfrender on 26 June 1988. Unknown locality: IICT/R
58-59/1955, Principe Island, collected by Décio Passos on April 1955.
Diagnosis —Boaedon mendesi sp. nov. (Figures 7 to 8) can be distinguished from all of its
Central and West African congeners by the following combination of characters: brown to
dark brown colouration with a series of parallel dark brown blotches extending from neck
to about midbody; two broad, cream-coloured, dark brown-bordered bands running on
the side of the head; the upper band starting on the nasals and extending above the
eye to the posterior part of the head, the lower band usually starting on the posterior
lower part of the eye and extending to the corner of the mouth; whitish to cream venter
colouration without markings; the 3rd, 4th and 5th (but sometimes the 4th, 5th and 6th)
supralabials contacting the eye, one preocular; 24 to 29 midbody scales rows, 206 to 242
ventral scales and 61 to 81 subcaudals, and a maximum snout–vent length of 122 cm.
The newly described species can be distinguished from B. bedriagae by its brown to
dark brown colouration with a series of parallel dark blotches extending from the neck
to about midbody (versus dark brown colouration with parallel mediodorsal and lateral
bands running from the neck to the tail in B. bedriagae) and a whitish venter (versus
cream, heavily speckled with dark markings in B. bedriagae). It can be distinguished
from B. fuliginosus by its higher number of ventral and subcaudal scales (206 to 242
and 61 to 81, respectively, in B. mendesi sp. nov. versus 194 to 229 and 46 to 71 in
West African B. fuliginosus fide Hallermann et al. (2020), by having three supralabials
touching the eye (versus only two in B. fuliginosus), by having two broad cream-coloured,
dark-bordered bands running on the side of the head (versus two thin white lines in
B. fuliginosus), and by having brown to dark brown colouration with a series of parallel
dark blotches extending from the neck to about midbody (versus homogeneous dark
brown colouration in B. fuliginosus). B. mendesi sp. nov. can be distinguished from
B. lineatus by its brown to dark brown colouration with a series of parallel dark blotches
extending from the neck to about midbody (versus a uniformly grey to reddish brown
colouration in B. lineatus) and by having only one preocular (versus usually two in
B. lineatus). B. mendesi sp. nov. can be distinguished from B. olivaceus,B. poensis and
B. radfordi by having the subcaudals divided (versus single in B. olivaceus,B. poensis
and B. radfordi). B. mendesi sp. nov. can be distinguished from B. virgatus by having
two broad cream-coloured, dark-bordered bands running on the side of the head
(versus two thin white lines in B. virgatus), and a higher number of ventral and subcaudal
scales (206 to 242 and 61 to 81, respectively, in B. mendesi sp. nov. versus 186 to 223 and
42 to 64 in B. virgatus). It can be distinguished from B. upembae by having more ventral
scales (206 to 242 in B. mendesi sp. nov. versus 175 to 197 in B. upembae), and a higher
number of midbody scale rows (24 to 29 in B. mendesi sp. nov. versus 21 to 23 in
B. upembae). It can be distinguished from B. perisilvestris by the 3rd, 4th and 5th
20 L. M. P CERÍACO ET AL.

supralabial in contact with the eye (versus usually the 4th and 5th in B. perisilvestris), by
having two broad cream-coloured, dark-bordered bands running on the side of the
head (versus no lines in B. perisilvestris), and by having brown to dark-brown colouration
with a series of parallel dark blotches extending from the neck to about midbody (versus
homogeneous dark brown colouration in B. perisilvestris). It can be distinguished from
B. paralineatus by the 3rd, 4th and 5th supralabial in contact with the eye (versus only
the 4th and 5th in B. paralineatus) and by its lower number of midbody scale rows (24
to 29 in B. mendesi sp. nov. versus 31 to 35 in B. paralineatus). It can be distinguished
from B. subflavus by the 3rd, 4th and 5th supralabial in contact with the eye (versus
only the 4th and 5th in B. subflavus), having two broad cream-coloured, dark-bordered
bands running on the side of the head (versus only one line in B. subflavus), and by
having brown to dark brown colouration with a series of parallel dark blotches extending
from the neck to about midbody (versus homogeneous yellow to light brown colouration
in B. subflavus). It can be distinguished from B. longilineatus by having a higher number of
subcaudals (61 to 81 in B. mendesi sp. nov. versus 42 to 63 in B. longilineatus) and by
having two broad cream-coloured, dark-bordered bands running on the side of the
head (versus a single, continuous line from the snout to the tail, in B. longilineatus). It
can be distinguished from B. littoralis by having brown to dark brown colouration with
a series of parallel dark blotches extending from the neck to about midbody (versus
homogeneous light brown colouration in B. littoralis).
Description of holotype —Adult male (Figures 7 to 8), 68.5 cm SVL; head subtriangu-
lar, slightly distinct from the neck, HL 3.5% of SVL (2.43 cm); pupil elliptical, eye diam-
eter 0.37 cm; loreal rectangular, almost twice as long (0.27 cm) as high (0.16 cm); body
cylindrical; tail moderately short (21.6% of SVL). Supralabials 8/8, 3rd, 4th and 5th on
both sides touching the eye; infralabials 9/9, first on each side in contact behind
mental, first three on both sides in contact with anterior chin shields and 4th on
both sides in contact with posterior chin shields; one preocular on both sides not
touching frontal; 2 postoculars, the lower one in contact with 5th and 6th supralabials
(both sides of head); temporals 1 + 2+3 on both sides; two internasals; nasal divided;
frontal longer than wide; dorsal scales smooth, 22 MSR one head-length posterior to
Table 4: Morphometric and meristic data on Boaedon mendesi sp. nov. types series. Measurements are
in cm. All abbreviations follow the Materials and methods
MUHNAC/MB03-977
Holotype
MUHNAC/MB03-1027
Paratype
IICT 52-1955
Paratype
CAS 233410
Paratype
Sex Male Unsexed Unsexed Female
SVL 69.5 47 122 46.4
Tail length 15 10.2 19 88
SVL/TL
MDSR 26 27 27 26
Ventral scales (V) 212 208 232 232
Subcaudals (SC) 69 68 62 66
V/SC
Subralabials (entering eye) 3rd, 4th, 5th 4th, 5th, 6th 3rd, 4th, 5th 3rd, 4th, 5th
Loreal length 2.7 2.6 3.5 2.4
Loreal height 1.6 1.1 1.8 1.0
Loreal L/H ratio
Preocular 1 1 1 1
Postocular 2 2 2 2
AFRICAN JOURNAL OF HERPETOLOGY 21

jaw rictus, 26 MSR at midbody; ventrals 212; cloacal plate entire; 69 subcaudals, all
paired.
Colouration of holotype in life —The colouration is brown with a series of parallel dark
blotches extending from the neck to about midbody (Figure 7). Two broad cream-
coloured, dark-bordered bands running on the side of the head; the upper band starting
on the nasals and extending above the eye to the posterior part of the head, the lower
band starting on the posterior lower part of the eye and extending to the corner of the
mouth (Figure 7). Venter whitish. Colouration in preservative present as described
above, but blotches are less evident (Figure 8).
Variation —Variation in measurements and scalation of the paratypes of B. mendesi sp. nov.
is presented in Table 4. All paratypes agree entirely with the holotype, with the exception
Figure 9. Confirmed records of Boaedon mendesi sp. nov. in Príncipe Island.
22 L. M. P CERÍACO ET AL.

of MUHNAC/MB03-1027, in which the 4th, 5th and 6th supralabials are in contact with
the eye.
Distribution —The species is endemic to Príncipe Island, Gulf of Guinea. Currently the
species has only been recorded from the northern, central and southwestern areas of
the island (Figure 9), although it is most likely widely distributed.
Habitat and natural history notes —Not much is known about the natural history of
this species. The holotype was found basking in a tree during day (Figure 10) in an aban-
doned coffee and cocoa plantation (Figure 11). The species is commonly found in more
forested areas, but also in slightly disturbed habitats.
Etymology —The specific epithet is a patronym in the masculine genitive singular
named after the Portuguese entomologist Luís Fernando Marques Mendes (Lisbon,
1946 –), former director of the Centro de Zoologia of the former Instituto de Investigação
Científica Tropical (Lisbon), from 1998 to 2014, and who has dedicated part of his career to
the study of the biodiversity of São Tomé and Príncipe.
Figure 10. Live photo of the holotype of Boaedon mendesi sp. nov. (MUHNAC/MB03–977), basking in a
tree during daytime. Photo by Luis MP Ceríaco.
AFRICAN JOURNAL OF HERPETOLOGY 23

The identity of Boodon nigrum Fischer, 1856
The first herpetological specimens from São Tomé and Príncipe known to have reached
European collections were those collected by the German medical surgeon Carl Weiss
and then deposited in the collections of the Hamburg museum. Weiss sailed on the
ship ‘Adolph’to São Tomé on 20 August 1847, and in September, he continued to the
Gold Coast with H Bartsch. He then returned to São Tomé, via Príncipe where he
served as a doctor. He then sent specimens from São Tomé to Hamburg in December
1847, 28 February 1848 and on 7 January 1849 (Weidner 1993). As an example of the
importance of Weiss collections, eight bird species from São Tomé, including the
endemic São Tomé Weaver, Ploceus sanctithomae (Hartlaub, 1848) and the Príncipe
Golden Weaver, Ploceus princeps (Bonaparte 1851), were described based on the material
collected by Weiss.
In addition to the specimen studied and published by Jan and Sordelli (1870;Figures 2
to 3), Fischer (1856) described another snake species, Boodon nigrum, based on two other
specimens from ‘St. Thomé (West-Afrika)’collected in 1847 by Weiss. A figure of one of the
syntypes accompanied the original description (Figure 12). However, Fischer’s description
was neglected by the subsequent authors who studied and published on the São Tomé
Jitas. Recently Hallermann (2006) rediscovered one of the remaining syntypes and ident-
ified the specimen as a B. virgatus. This interpretation was followed by Wallach et al.
(2014), who considered nigrum as a junior synonym of virgatus. Our analysis of the
extant syntype of B. nigrum (ZMH R08378; Figure 13) confirms the identity proposed by
Hallermann (2006). The specimen has 22 midbody scale rows, a head with two thin,
whitish lines on both sides, and homogeneous brownish body colouration without any
Figure 11. Type locality of Boaedon mendesi sp. nov. in the vicinities of Santo Cristo, Príncipe Island.
Photo by Pedro NP Ceríaco.
24 L. M. P CERÍACO ET AL.

stripes, typical of B. virgatus (see B. bedriagae taxonomic account above). Boaedon virgatus
occurs in continental West Africa, from Guinea to the Republic of the Congo (Wallach et al.
2014; Chippaux and Jackson 2019); it is not known to occur in the Gulf of Guinea Islands.
The locality data confusion may be the result of mislabelling and it is likely that Weiss may
have unintentionally mixed Gold Coast specimens with those from São Tomé. This scen-
ario is consistent with other similar mistakes recently documented from specimens col-
lected by Weiss and Fischer (Ceríaco et al. 2018). Therefore, we agree with the
decisions of Hallermann (2006) and Wallach et al. (2014) and consider B. nigrum a
junior synonym of B. virgatus, which is therefore unavailable to be applied to the
São Tomé Island population of Jita.
Figure 12. Illustration of one the syntypes of Boaedon nigrum by Fischer (1856). Adapted from Fischer
(1856).
AFRICAN JOURNAL OF HERPETOLOGY 25

Discussion
Our results support the recognition of Jitas from the São Tomé and Príncipe islands as
two distinct species, Boaedon bedriagae, endemic to São Tomé island and Rolas islet,
and Boaedon mendesi sp. nov. endemic to Príncipe island. In the light of the recent
reviews of the herpetofauna of these islands, this result is not surprising. Species
long considered to be conspecific among the two islands have been shown to rep-
resent distinct taxa, sometimes closely related (Jesus et al. 2006; Miller et al. 2012;
Bell et al. 2015; Bell 2016; Soares et al. 2018), but in other cases not even being the
respective sister species pairs (Uyeda et al. 2007; Ceríaco et al. 2015,2016). Using an
integrative approach, combining molecular and morphological data and a thorough
revision of the nomenclatural history of the genus in São Tomé and Príncipe, we con-
tribute to a better understanding of the diversity and endemicity rates of these islands.
With the recognition of Boaedon bedriagae as a valid species and Boaedon mendesi sp. nov.
as a new endemic species from Príncipe island, our current understanding of the herpe-
tofauna of São Tomé and Príncipe increase the percentage of island endemics. Currently
13 reptile species are known to occur in Príncipe island: two are introduced (Hemidactylus
longicephalus and H. mabouia, see Ceríaco et al. 2020); one is shared with the continent
(Trachylepis affinis, see Ceríaco et al. 2016); two species are shared with São Tomé, but
endemic to the country (Letheobia feae and L. newtoni, see Ceríaco et al. 2018); and
eight are island endemics (Afrotypholps elegans,Boaedon mendesi sp. nov., Feylinia
polylepis,Hapsidophrys principis,Hemidactylus principensis,Lygodactylus delicatus,
Panaspis africana, and Trachylepis adamastor, see Ceríaco et al. 2018, unpubl. data). For
São Tomé island, 11 species are currently recorded: two are introduced (Hemidactylus
longicephalus and H. mabouia, see Ceríaco et al. 2020); one is shared with the continent
(Pelusios castaneus, see Fritz et al. 2010); two species are shared with Príncipe, but
Figure 13. Syntype of Boaedon nigrum (ZMH R08378).
26 L. M. P CERÍACO ET AL.

endemic to the country (Letheobia feae and L. newtoni, see Ceríaco et al. 2018); and seven
are island endemics (Boaedon bedriagae,Hemidactylus greeffii,Lygodactylus thomensis,
Panaspis thomensis,Philothamnus thomensis,Trachylepis thomensis, and Naja peroescobari,
see Ceríaco et al. 2018). These numbers are impressive, especially when taking into
account the small area of Príncipe and São Tomé Island (136 and 855 km
2
, respectively).
Boaedon mendesi sp. nov. represents the 10th species of the genus described in the
past five years (Greenbaum et al. 2015; Trape and Mediannikov 2016; Hallermann et al.
2020). Despite the obvious taxonomic and nomenclatural challenges presented by the
group, our knowledge on the genus is currently in a state of flux, with several other
species being described from across Africa (JH pers. obs.). Although the most recent
descriptions are all based on both morphological and molecular data, our current under-
standing of several taxa is still deficient. Further studies are required to test the phylogeo-
graphic affinities of all known species of the genus, as well as to stabilise the problematic
taxonomy of B. fuliginosus and B. lineatus.
Acknowledgments
We thank several entities and persons that contributed to the current study. From São Tomé and
Príncipe, special thanks are owed to Arlindo Ceita Carvalho, director of the General Office for the
Environment from the Ministry of Environment and Natural Resources, to José Cassandra, President
of the Regional Government of Príncipe; to Daniel Ramos, director of Príncipe Obo National Park; and
particularly to Ostelino da Conceição Rocha (aka Balô) for his valuable help during field work. Pedro
Nuno Pires Ceríaco and Ana Carolina Sousa are also thanked for their support during fieldwork. We
thank the curators for loan of material and hosting during visits: Wolfgang Böhme (ZFMK), MO Rödel
& Frank Tillak (ZMB), Jens Vindum and Lauren Scheinberg (CAS), Ivan Ineich (MNHN), Carol Spencer
and Ted Papenfuss (MVZ). Anna Sellas, Beth Moore, and Anthea Carmichael for their support with
sequencing at CAS Center for Comparative Genomics. Giulliano Doria took photos of the lectotype
of Boaedon bedriagae housed in the collections of the Museo Civico di Storia Naturale ‘Giacomo
Doria’. Although the patronym of this species honor Luis Mendes, we want also to dedicate this
paper to António Bívar de Sousa, for his constant support and friendship. Robert “Bob”Drewes is
also thanked for his constant support and enthusiasm regarding the study of São Tomé and Principe
biodiversity. Mariana P Marques is currently supported by FCT, contract SFRH/BD/129924/2017.
Lisette Arellano was supported by the California Academy of Sciences’Summer Systematics Institute
(SSI) National Science Foundation Research Experiences for Undergraduates initiative and the CAS
Robert T Wallace endowment.
ORCID
Luis MP Ceríaco http://orcid.org/0000-0002-0591-9978
Ana Lisette Arellano http://orcid.org/0000-0001-9604-745X
Robert C Jadin http://orcid.org/0000-0003-3028-3315
Mariana P Marques http://orcid.org/0000-0002-1712-2632
Diogo Parrinha http://orcid.org/0000-0002-1302-025X
Jakob Hallermann http://orcid.org/0000-0002-8835-9303
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