Book review. Snakes of Central and Western Africa

Abstract

We analyze an important new opus on the snakes of West and Central Africa co-authored by Jean-Philippe Chippaux and Kate Jackson. We correct the identification of some of the illustrated snakes of the genera Dipsadoboa, Grayia, Limaformosa and Philothamnus. We provide more detailed localities for more than 30 photographs of snakes of the genera Atractaspis, Bitis, Boaedon, Bothrophthalmus, Causus, Dasypeltis, Dendrolycus, Eryx, Gonionotophis, Grayia, Hydraethiops, Leptotyphlops, Limaformosa, Mehelya, Myriopholis, Natriciteres, Philothamnus, Polemon, Python, Thelotornis, Tricheilostoma and Xenocalamus, from Botswana, Cameroon, Democratic Republic of the Congo, Ethiopia, Ghana, Ivory Coast, Liberia, Republic of the Congo and Tanzania. An interval of four years between the submission of the manuscript of the book and its publication explains the inaccuracy of many distribution maps, and the fact that recent taxonomic changes and numerous recently described species and genera were not included.

Full text

Herpetological Review 51(1), 2020
161BOOK REVIEWS 161
Herpetological Review, 2020, 51(1), 161–164.
© 2020 by Society for the Study of Amphibians and Reptiles
Snakes of Central and Western Africa
Jean-Philippe Chippaux and Kate Jackson. 2019. Johns Hopkins
University Press, Baltimore, Maryland (https://www.press.jhu.edu). x +
429 pp. Hardcover. US $84.95. ISBN: 978-1-4214-2719-5.
OLIVIER S. G. PAUWELS
Royal Belgian Institute of Natural Sciences
Rue Vautier 29, 1000 Brussels, Belgium
e-mail: osgpauwels@yahoo.fr
JONATHAN BRECKO
Royal Belgian Institute of Natural Sciences
Rue Vautier 29, 1000 Brussels, Belgium
and Royal Museum for Central Africa
Leuvensesteenweg 13, 3080 Tervuren, Belgium
The first edition of Jean-Philippe Chip-
paux’s Les serpents d’Afrique occidentale et
centrale was published two decades ago
in 1999, and was followed by two later edi-
tions, each bringing many additions and improvements, in 2001
and 2006, respectively (Akani 2002; Pauwels and Branch 2003). To
all herpetologists and field biologists dealing with snakes of West
and Central Africa, the ‘Chippaux’ was no longer just a technical
snake identification guide, it really became an ‘institution.’ Given
the numerous publications on the systematics and distribution
of African snakes during the last decade, it was definitely time
for a new, revised edition. The main surprise element for the new
‘Chippaux’ is, besides the shift to English, the addition of a second
author, although the general scope of the book stays the same.
The binding of the book is solid and the paper is of good qual-
ity. Although not a pocket guide, its size and weight (more than
one kg) allow bringing it in the field. The cover is nicely illustrated
by a photograph of a Dipsadoboa viridis, a welcome change from
the usual cobras or vipers on the cover of so many snake guides.
The first chapter of the book is dedicated to snake identification
(pp. 1–27) and presents the important diagnostic characters in
detail with the help of numerous photographs and drawings, as
well as keys to alethinophidian genera. It is followed by a chapter
on snake evolution (pp. 28–39) and chapters on the biogeography
of African snakes (pp. 40–48) and snakebite in sub-Saharan Africa
(pp. 49–52). Some of the introductory chapters would have ben-
efitted from more development, such as the chapter on snakebite,
because Chippaux is a leading authority in this domain. Of course,
field guides impose length constraints, and the authors provide
numerous references to publications for readers desiring to know
more (note, however, that there are several unused pages at the
beginning of the book and seven blank pages at the end of the
book).
The fifth chapter introduces the scolecophidians within just
a few pages (pp. 53–59). Similar to the former Chippauxs, scole-
cophidians receive a poor treatment relative to other snakes. The
authors state ‘We will deal with them [Scolecophidia] in less detail
than we devote to the alethinophidian snakes’ without justifica-
tion. The authors provide a key to the families and genera, a very
brief introduction to each genus, and a list of species with an im-
precisely written description of their distribution. They indicate
in their introduction on p. 53 that the scolecophidians are repre-
sented in the region by 49 species, but a few pages later they list 52
species, representing thus about 17% of the regional snake diver-
sity. Only five species are illustrated by a color photograph, one of
them not identified to the specific level. Knowing that this group
includes many species still to be described, we regret this decision
which helps to maintain a poor level of knowledge and popularity
of these fascinating, ecologically specialized snakes.
The next chapters present the Boidae and Pythonidae (pp.
60–71), viperids (pp. 72–103), elapids (pp. 104–135), lamprophiids
(pp. 136–287) and colubrids (pp. 288–384). Each of the 247 alethi-
nophidian species treated in the book is presented through a spe-
cific account. Species accounts generally follow an introduction to
the genus providing useful information and a key to the species.
The species accounts include a clear and informative text with a
general description of the geographic distribution, a morphologi-
cal description with emphasis on diagnostic characters, and occa-
sional taxonomic and other relevant comments. Maps are specific
or combine the distributions of several species. The authors chose
to use dotted maps, which would be very good if they included
at least all reliable published localities. Unfortunately, this is not
the case; in particular many recently published localities were not
included. For example, the map for Atheris hirsuta shows a single
dot in agreement with the species account that mentions that the
species is endemic to the Taï Forest in Ivory Coast, but in fact it
is more widely distributed (Penner et al. 2013). The map for Bitis
arietans on p. 83 shows no dot for Ghana, but the species is illus-
trated by a photograph of an individual taken in Ghana; the same
happens for Atractaspis aterrima on p. 143 and for Limaformosa
crossi (cited under Gonionotophis crossi) on pp. 211–212 (see Table
1). The map for Natriciteres variegata on p. 301 shows no dot for
Liberia, but the species is illustrated by a photograph taken in that
country; the only site in Liberia where the author of the photo-
graph, the late Bill Branch, worked is the Liberian part of Mount
Nimba (Rödel, pers. comm.). The map for Polemon bocourti on p.
176 shows a single dot for the Republic of Congo in the northern
part of the country, although the species is illustrated by a pho-
tograph taken in a coastal locality of that country (Table 1). Di-
etary observations made by Pauwels et al. (2004) on Psammophis
cf. phillipsii in Loango National Park, Gabon, were erroneously
included in the account and as a dot on the map for P. sibilans,
while the latter species does not occur in Gabon; the Gabonese
population actually belongs to P. mossambicus (Trape et al. 2019).
Additional localities for species of the northwestern part of the re-
gion covered by the book can be found in Trape and Mané (2006).
Another issue with the maps is their large size, about half a page
BOOK REVIEWSBOOK REVIEWSBOOK REVIEWS

Herpetological Review 51(1), 2020
162162 BOOK REVIEWS
each, and the fact that the geographic area covered by the book
represents less than half of the surface of each map. Reducing the
size of the maps and centering them on the region concerned by
the book would have saved many pages and enabled the photo-
graphs of snakes, generally much smaller than the maps, to be
enlarged.
In addition to the numerous excellent drawings illustrating the
diagnostic morphological characters in the general introduction
and in the introductions to various families, one finds 124 remark-
ably accurate snake head scalation drawings (each time a lateral,
dorsal and ventral view), nearly all performed by the talented hand
of Tuhin Giri who is to be congratulated. Each drawing caption
specifies the museum collection number of the specimen used,
which is precious information. It is therefore a pity that about 50
of these beautiful drawings were improperly typeset, thus cutting
off the bottom part of the illustration (e.g., Figs. 9.74, 10.21, 11.42).
The photographs are generally of good quality, although
sometimes too dark and too small. In total, 94 of the species (50
of them alethinophidians) and six of the genera treated were not
represented by at least one photograph or drawing. More than 80
of the photos were taken outside the region covered by the book or
show captive individuals of unknown origin. The photos of live in-
dividuals mention only the country where they were taken, with-
out more geographical precision, a real pity for all cases where
more detailed locality information was available, especially for
the rarely illustrated species. In order to not lose this important
information, we provide detailed localities for selected photos,
obtained directly from their respective authors (Table 1).
The separation of the keys to genera and to species is very
handy. We tried the identification keys with fresh material from
Gabon and the Democratic Republic of the Congo. The first speci-
men we used, an Aparallactus modestus, did not fit anywhere in
the key to the genus, as it shows, similar to the individual illus-
trated on p. 165, a single supralabial (the 6th) in contact with the
parietal, while the key gave only two options: ‘5th and 6th supralabi-
als both in contact with parietal,’ or ‘the 5th but not 6th supralabial
in contact with parietal.’ We used other snakes, for which the keys
led to the right species and proved to be easy and practical. We
note that in the key to the genus Atheris on p. 74, A. hirsuta is said
to be distinguishable from A. hispida by its possession of only one
row of scales between the eye and the nasal, while in reality it has
two rows, as can be seen in the original description of A. hirsuta
and on the photos provided by Penner et al. (2013) and by Chip-
paux and Jackson themselves. In the same key, A. broadleyi is said
to be distinguishable from A. squamigera by the presence in the
former of a black stripe along the side of the head. Rather than a
black stripe (not visible indeed on Fig. 7.2), it is a ‘darker triangle’
as correctly mentioned in the species account for A. broadleyi, of-
ten poorly contrasted. In the key to Bitis species on p. 82, ‘A group
of several narrow, pointed scales between the two nostrils’ leads
to B. nasicornis, while ‘A single scale between the nostrils’ leads
to B. rhinoceros or B. gabonica, although it is impossible that the
nostrils are separated by a single scale. This means that a reader
with a poor knowledge of these snakes won’t identify a viper as
any of the latter two species. There are discrepancies between the
key and the species accounts for Polemon (p. 173). In the key, P.
notatus is said to have a ‘dark band across neck’ while in the spe-
cies account it is said to show ‘a broad pale band across the nape’;
in the key, P. gabonensis is said to have ‘no nuchal band’ but its
species account says it has ‘a pale band across the nape.’ Among
other discrepancies, it should be mentioned that in the key to Dip-
sadoboa, the character ‘subcaudals divided’ leads among others to
D. duchesnii, while its species account erroneously mentions that
it has single subcaudals. In the key to Telescopus on p. 362, ‘2 upper
labials in contact with the eye’ (versus 3) leads among others to T.
variegatus, while this species is illustrated by a drawing showing
three supralabials in contact with the eye. The last couplet of this
key gives an alternative between ‘19 dorsal scale rows’ for T. semi-
annulatus, whereas the species account says ‘19 (occasionally 17
or 21) oblique rows.’ For T. obtusus, the key says there are ‘21 dorsal
scale rows,’ whereas the species account says ‘23 (occasionally 21)
oblique rows.’
The individual shown on p. 217 to depict Gonionotophis poen-
sis was already used by Broadley et al. (2018) to illustrate the ge-
nus Limaformosa (Greenbaum, pers. comm., August 2019), thus
excluding an identification as Mehelya poensis; it rather seems
to fit with Limaformosa savorgnani, as it was initially tentatively
identified by Greenbaum (pers. comm.; Table 1). In the light of
the recent revision by Trape et al. (2019), the two individuals of
‘Psammophis phillipsii’ shown on Figs. 11.36 and 11.37 should be
re-identified as P. mossambicus, and the ‘P. sibilans’ on Fig. 11.46
as P. afroccidentalis. The ‘Philothamnus carinatus’ individual il-
lustrated by a photograph on p. 347 (see Table 1) was used in the
phylogenetic study of Engelbrecht et al. (2019) where it came out
as a P. heterodermus. Dipsadoboa underwoodi is illustrated by a
photograph taken in the Democratic Republic of the Congo. The
species account does not include this country in this species’ dis-
tribution, nor does the map. Based on the characters visible on
the photograph, in particular two supralabials in contact with the
orbit, and the color of the underside of the tail contrasting with the
lighter color of the belly, we believe this individual to be a Dipsa-
doboa weileri, a species well known from the Democratic Republic
of the Congo. The photograph of a juvenile ‘Grayia ornata’ shown
on pp. 8 and 382 actually represents a juvenile G. smithii, recog-
nizable by its laterodorsal pattern made of yellowish pyramids on
a dark brown background and its elongate posterior (7th) supra-
labial. This individual was sequenced and its identification as G.
smithii has been confirmed (Greenbaum, pers. comm.).
The chapters presenting the snake families are followed by a
bibliography (pp. 385–409) and an index (411–429), closing the
opus. The literature cited includes 614 references, among them
most of the original descriptions of the snakes of the region and
many other important historic references, which is very valuable.
The incompleteness of some maps is explainable for the same
reason for which recent revisions and species descriptions are not
included—while the book was published in 2019, the manuscript
was submitted for publication four years earlier, on 27 May 2015
(Chippaux, pers. comm.). After that only some publications could
be integrated (only six references published after 2015 are listed
in the literature cited). It is a pity that the revision of the Naja
melanoleuca complex by Wüster et al. (2018) leading to the rec-
ognition at species level of N. subfulva and to the description of N.
guineensis and N. savannula, all occurring in the area covered by
the book, could not be taken into account, as well as the descrip-
tion a year before of Naja peroescobari from São Tomé (Ceríaco et
al. 2017). The revision of Gonionotophis by Broadley et al. (2018)
led among other results to the description of two new genera
found in the area covered by the book, the revalidation of Mehelya,
and the confirmation of the validity of Limaformosa savorgnani
as a separate species. Limaformosa chanleri, known among other
countries from the Democratic Republic of the Congo, Rwanda
and Burundi (Wallach et al. 2014), has been omitted in the book.
Another major work that could not be included was the revision
of Boaedon by Trape and Mediannikov (2016) that led to the de-

Herpetological Review 51(1), 2020
163BOOK REVIEWS 163
table 1. Localities for selected photographs presented by Chippaux and Jackson (2019), provided by the respective photographers (pers. comm. to
OSGP, 2019). Dept. = Department; DRC = Democratic Republic of the Congo; Prov. = Province; RC = Republic of the Congo.
Species Fig. numbers and Locality
photographer
Leptotyphlops scutifrons 1.27 & 5.4, S. Spawls Otse, South-East District, Botswana
Myriopholis macrorhyncha 5.5, S. Spawls Resort on the Awash River, Sodere, Ethiopia
Tricheilostoma bicolor 5.6, S. Spawls Wa, Upper West Region, Ghana
Eryx colubrinus 6.6, S. Spawls Northern Tanzania
Python natalensis 6.11, S. Spawls Otse, South-East District, Botswana
Bitis arietans 7.15, M.-O. Rödel Comoé National Park, north-eastern Ivory Coast
Causus maculatus 7.30, S. Spawls Wa, Upper West Region, Ghana
Atractaspis dahomeyensis 9.19, M.-O. Rödel Ananda, eastern-central Ivory Coast
Polemon bocourti 9.61, M. Burger 2.5 km E of Lombo, about 9 km E of Hinda, Kouilou Dept., RC
Polemon collaris 9.66, M. Burger About 24 km SE of Tsinguidi, Niari Dept., RC
Polemon notatus 9.781, M. Burger 1 km SE of Tchitobo, Kouilou Dept., RC
Xenocalamus michelli 9.87, E. Greenbaum Near Manono airport (7°16’39.8”S, 27°23’23.2”E; alt. 627 m asl), Tanganyika
Prov., DRC (field nr ELI355)
Boaedon radfordi 10.9, E. Greenbaum Shatuma-Abis village (2°00’47.3”N, 30°50’26.4”E; alt. 2000 m asl), Lendu
Plateau, Ituri Prov., DRC (paratype UTEP 20996)
Bothrophthalmus brunneus2 10.17, M. Burger Near Banjoko village, about 3 km SSW of Mayoko Poste, Niari Dept., RC
Bothrophthalmus lineatus 10.18, M.-O. Rödel Forêt Classée de la Haute Dodo, south-western Ivory Coast
Bothrophthalmus brunneus2 10.19, M.-O. Rödel Near Dja Faunal Reserve, south-eastern Cameroon
Dendrolycus elapoides 10.26, K. Mebert Uma (0°32’4.75”N, 25°54’3.36”E), Tshopo Prov., DRC (individual registered as
RBINS 18729)
Gonionotophis brussauxi 10.30, M. Burger 3.4 km SSW of village Bandjoko, 8.5 km SSW of Mayoko Poste, Niari Dept., RC
Limaformosa crossi3 10.35, S. Spawls Wa, Upper West Region, Ghana
Limaformosa cf. savorgnani4 10.43, E. Greenbaum Mayimbili (1°23’48.8”N 28°35’09.2”E; alt. 742 m asl), Ituri Prov., DRC (field nr
EBG 2637)
Mehelya stenophthalmus3 10.44, M. Burger 2 km SE of Chiboula village, about 10 km NW of Mandingo-Kayes, Kouilou
Dept., RC
Hydraethiops melanogaster 12.12, E. Greenbaum Near Epulu River (1°23’49.0”N 28°35’09.1”E; alt. 753 m asl), Ituri Prov., DRC
(field nr EBG 2612)
Dasypeltis palmarum 13.19, M. Burger 6 km W of Hinda, Kouilou Dept., RC
Philothamnus heterodermus5 13.64, M. Burger 5 km NE of Mengo, about 16 km SW of Hinda, Kouilou Dept., RC
Philothamnus dorsalis 13.66, M. Burger 1 km E of Carriere Mambanziba, 14 km E of Hinda, Kouilou Dept., RC
Thelotornis kirtlandii 13.95, M.-O. Rödel Forêt Classée de la Haute Dodo, south-western Ivory Coast
Grayia ornata 13.113, M. Burger About 5 km NE of Tchiboula village, Kouilou Dept., RC
Grayia ornata 13.114, M. Burger Vemba River, 1 km SW of Youbi village, Kouilou Dept., RC
Grayia smithii6 13.115, E. Greenbaum Vicinity of Epulu (1°24’05.0”N 28°34’12.0”E; alt. 747 m asl), Ituri Prov., DRC
(field nr EBG 2634)
Grayia smithii 13.117, M. Burger 2.8 km E of Hinda, Kouilou Dept., RC
1 The captions for figures 9.78 and 9.79 are inadvertently switched in the book. 2 Treated as ‘B. lineatus, unicolored form’ by Chippaux and Jackson. 3 Included in Goni-
onotophis by Chippaux and Jackson. 4 Erroneously identified as Gonionotophis poensis. 5 Erroneously identified as P. carinatus. 6 Erroneously identified as G. ornata.

Herpetological Review 51(1), 2020
164164 BOOK REVIEWS
scription of five species occurring in the area under consideration:
B. littoralis, B. longilineatus, B. paralineatus, B. perisilvestris and
B. subflavus. Snakes described in 2018 but not integrated in the
book also include Letheobia akagerae Dehling et al. (Typhlopidae)
from Rwanda, and Echis romani Trape (Viperidae) from Nigeria to
the Central African Republic. Inevitably, several snakes described
in 2019 could not be included: Myriopholis occipitalis Trape and
Chirio from the Central African Republic and Chad, Tricheilosto-
ma kongoensis Trape from the Democratic Republic of the Congo,
Afrotyphlops chirioi Trape from the Central African Republic, A.
rouxestevae Trape from Cameroon, Letheobia logonensis Trape
from Chad, Atractaspis branchi Rödel et al. from Guinea and Li-
beria, Polemon ater Portillo et al. from the Democratic Republic
of Congo, and Psammophis afroccidentalis Trape et al. from West
Africa. This renders the Naja, Boaedon, Psammophis and Goni-
onotophis sections of the book already obsolete.
French is an official language in 17 of the 23 countries covered
by the book. Some researchers will find the use of English chal-
lenging, but this new book will be a most useful tool for the nu-
merous non-French speaking persons interested in the snakes of
the region. The price of the book is much beyond what most re-
searchers from the area covered by the book (and beyond) can af-
ford. However, thanks to the useful identification keys, the numer-
ous photographs and high-quality drawings, the information-rich
text, and the abundant and pertinent literature cited, it remains an
excellent guide, and we strongly encourage local and internation-
al scientific libraries and all naturalists and herpetologists working
on African snakes to purchase this important new tool.
Acknowledgments.—We thank Jean-Philippe Chippaux for use-
ful discussions, and Marius Burger, Eli Greenbaum, Konrad Mebert,
Mark-Oliver Rödel, and Steve Spawls for information on photographs
they took.
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The Field Herping Guide: Finding Amphibians
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ISBN:978-0-820-35458-3.
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Department of Biology
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3300a Kindschi Hall of Science
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e-mail: moorejen@gvsu.edu
For many of us, our first forays into
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a salamander, snake, or frog, and staring
in wonder at its bright yellow spots,
feeling the roughness of its keeled scales,
or smelling the pungent twinge of musk.
Amphibians and reptiles appeal to young