Alytes_2014_31_1-2 - Amphibian Survival Alliance

Transcription

ISSN 0753-4973
VOLUME 31
|
ISSUE 1-2
|
SEPTEMBER 2014
VOLUME 31 (1-2)
SEPTEMBER 2014
Illustration. Milene Matos (Lissotriton boscai and Triturus marmoratus)
Published
by the
International
Society for
the Study and
Conservation
of Amphibians
(ISSCA)
In partnership
with Amphibian
Survival Alliance
(ASA) and
IUCN/SSC
Amphibian
Specialist Group
(ASG)
RESEARCH ARTICLE
2014 | VOLUME 31 | PAGES 3-12
Description of the larvae of Günther’s toad
Duttaphrynus hololius (Günther, 1876) (Anura:
Bufonidae) with notes on development and
oral ultra-structure
S. R. Chandramouli1*, Ayuthavel Kalaimani2
2.
1.
Centre for Ecological Sciences, Indian Institute of Science, Bangalore
No: 466/158/89, Murugan Koil Street,Thenimalai,Tiruvannamalai-606603,Tamil Nadu, India
Morphology of tadpoles of the little known, Peninsular Indian endemic bufonid Duttaphrynus hololius
(Günther, 1876) is described across different stages in detail, with observations on natural history. The
oral apparatus of these benthic feeding tadpoles was examined using scanning electron microscopy
and the ultra-structure is illustrated and described in detail. These larvae have a labial tooth row formula of 2(2)/2(2), with about 99 and 97 denticles on the anterior and posterior labia respectively. Novel information on the time taken for its ontogenetic development in the advanced stages is presented.
INTRODUCTION
Unlike any other terrestrial vertebrate groups, amphibians have complex life history traits and most of
them spend the early stages of their developmental cycle in the aquatic phase, as free-swimming larvae (Duellmann
& Trueb, 1994). Though there are exceptions in the form of direct developing taxa that hatch out from the eggs as
imagos of the adults (e.g. Rhacophoridae: Raorchestes), most species undergo this exhaustive transformation in
body form before attaining adulthood. This crucial phase of their life history remains poorly documented for most
of the amphibian species in India (Das & Dutta, 2007). Among them, genus Duttaphrynus is one of the poorly
known genera for which data on larval characteristics exist for a very few species (Das & Dutta, 2007). These
include the species Duttaphrynus melanostictus (Schneider, 1799), D. microtympanum (Boulenger, 1882), D.
scaber (Schneider, 1799), D. stomaticus (Lütken, 1864) and D. brevirostris (Rao, 1937) (see Das & Dutta, 2007).
Duttaphrynus hololius, one of the poorly known, endemic members of this genus, was reported from Hosur
very recently, with additional information on basic morphology (Chandramouli et al., 2011) and subsequently
recorded from a few other localities in the Eastern Ghats (Adimalliah et al., 2012; Kalaimani et al., 2012;
Srinivasulu et al., 2013). Preliminary data on the larvae and the habitat of this species were briefly presented in
Ganesh et al. (2013). Ganesh et al. (2013) provided notes on larval habitat of D. hololius and a brief description of
the overall morphology of tadpoles across Gosner stages 20-47. However, precise information on morphological
characteristics of the larvae belonging to each stage could not be elaborated since their study was entirely based
on live, uncollected samples.
The information provided by Ganesh et al. (2013) is supplemented here, based on the collection of new
samples and description of their morphology at different phases of metamorphosis. Also, the oral apparatus and
details of its ultra-structure are presented herein for the very first time.
Received 04 December 2013
*Corresponding author
Accepted 22 April 2014
Published Online 15 September 2014
© ISSCA and authors 2014
findthesnakeman@gmail.com
S. R. CHANDRAMOULI & AYUTHAVEL KALAIMANI
MATERIALS AND METHODS
This published work has been registered in ZooBank. The ZooBank LSIDs (Life Science Identifiers)
can be resolved and the associated information viewed through any standard web browser. The LSID for this
publication is: urn:lsid:zoobank.org:pub:0124C095-B8FE-4A6C-84B2-6ED204C13E28
While on a biodiversity inventory in the Eastern Ghats landscape near the inter state boundary of Karnataka
and Tamil Nadu, Southern India (12.38°N, 78.62°E, 677 m asl), bufonid tadpoles were observed in small puddles
amidst dry rocky hills. Site characteristics such as the habitat type, forest cover and the number and size of puddles
and their attributes such as size and depth were recorded. Four tadpole samples were collected and reared in
captive conditions. The tadpoles were reared in captivity and were allowed to feed on the same detritus material
collected from the native pool. They were retained for the following eight days before being fixed for preservation.
In captivity they were allowed to feed on the organic debris, which was collected from the native pool. In regular
intervals, tadpoles were euthanized and preserved in 10% formalin solution across different developmental stages.
The samples were fixed at different stages; i.e., 25 and 42, wherein, they were observed to show
considerable variation in the development of the oral apparatus. A piece of the tail muscle tissue was extracted
and stored in absolute ethanol from one tadpole prior to fixation. After fixation, the tadpoles were examined under
a Leica™ MZ75 microscope for observing oral morphological characteristics. Measurements were done using
the microscope. The collected larvae were unambiguously allocated to the species Duttaphrynus hololis based
on the morphological characters of the metamorphosed toadlets seen along with the tadpoles in the field, namely,
rudimentary toe webbing, smooth skin texture, indistinct parotoid glands and the absence of cephalic ridges. An
adult female (identified based on the absence of nuptial pads and vocal sac) of this species (identified based on
rounded parotoid glands, absence of cephalic ridges and rudimentary toe webbing) was examined, measured in
situ and released.
The above diagnostic morphological characters of this species were discussed in detail by Chandramouli
et al. (2011). Also, no other anuran species were encountered in the vicinity of the collection site. The specimens are deposited in the collection of the Centre for Ecological Sciences, Bangalore under the voucher number
CESF2354. The tadpoles were staged following the key provided by Gosner (1960) and were measured with a
Figure 1. A. Tadpoles of Duttaphrynus hololius seen in a rocky puddle;
B. freshly metamorphosed imagos of Duttaphrynus hololius; C. adult
female.
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ALYTES 2014 | 31
stereo microscope to the nearest 0.01 mm. Following measurements were recorded from the tadpoles – head-body
length (HBL), head-body width (HBW), head-body depth (HBD), total length (totL), eye diameter (ED), distance
between anterior eye margin and nostril (EN), anterior eye margin to snout tip (ES), tail length (TaL), inter orbital
distance (IO), inter narial distance (IN), caudal fin height (CFH), caudal muscle height (CMH), width of the posterior tooth row, width of the anterior tooth row, inter-tooth row distance, oral disc width, upper arm length (UAL),
lower arm length (LAL), palm length (PAL), femur length (FEL), tibia length (TBL), tarsus length (TAR) and foot
length (FOL). Eco-morphological classification of the larvae based on their microhabitat associations and feeding
habits follows Altig & Johnston (1989). Terminologies used in the description of oral apparatus follow McDiarmid
& Altig (1999). The adult toad and freshly metamorphosed toadlets encountered in the field were measured in situ
with a vernier caliper to the nearest 0.1 mm, followed by release in the place of capture.
Scanning electron microscopy
The oral disc of a stage 25 larva was dissected and kept in absolute alcohol overnight for dehydration. It
was then dried completely and mounted on a metallic stub with a thin layer of adhesive. The mounted sample was
then desiccated completely and coated with a thin film of gold. The gold coated sample was then subjected to exposure at different levels and under different degrees of magnification in the SEM equipment to study the detailed
ultrastructure of the oral apparatus.
RESULTS
Species identification
The tadpoles were allocated to the species Duttaphrynus hololius based on the developmental characters
of the larvae and morphology of the toadlets observed in the field along with the tadpoles. An intensive search in
the immediate surroundings of the pools with tadpoles was successfully able to uncover six freshly metamorphosed
imagos (SVL 10.3, 9.6, 11.3, 11.3, 10.3 and 11 mm; mean 10.63 mm) and an adult female of this toad from rock
crevices (fig. 1). These toadlets as well as the adult female showed the following diagnostic morphological traits:
absence of cephalic ridges, rudimentary toe webbing, rounded parotoid glands and skin without warts. Hence, they
were referred to this species.
Ecological notes
The tadpoles of Duttaphrynus hololius were observed on 1 June 2013, following a bout of sporadic rains
(in the pre-monsoon season), in ephemeral puddles formed by stagnant rain water on rocky hill tops (fig. 1.A).
These larvae were found to be benthic feeders and were observed to feed actively on the algal growth at the bottom
surface of the pools. Many tadpoles (about 40 per pool) were observed in two adjacent stagnant water pools on an
open, rocky hilltop. The puddles were oval in shape, about 1-1.5 m in diameter (at the widest point) with a depth
of not more than 10-12 cm.
The immediate vicinity of the collection site was surrounded by rocky outcrops, with sparse vegetation,
predominantly with grass clumps and patches of moist soil. No other anuran species were observed in close
vicinity of these pools. The tadpoles in these pools were observed to be of different developmental stages,
suggesting that the observed individuals could probably belong to more than a single clutch that could have been
laid asynchronously with a short lag in between. An adult toad (SVL 27.1 mm) was also sighted in the immediate
vicinity. Four larvae were collected and reared in captivity. A brief note on their partial (advanced) ontogenetic
development is mentioned below.
Notes on metamorphosis and advanced ontogenetic development
Of the four samples collected on 1 June 2013, three were of stage 24 and one sample was in an advanced
stage 27. Upon collection, the tadpoles were uniform dark purplish brown above and pale white below with a
translucent skin. After a week of captive husbandry, two larvae were still found to be in stage 25 on 6 June 2013.
At this stage 25, they did not show any traces of limb buds while the other, older sample of a slightly advanced
stage, reached stage 31 with well developed hind limbs, which, by then, had emerged out of the body capsule. The
larvae of stage 25 were of the same overall dark purplish colouration but a feeble, inverted ‘V’ shaped marking
on the back had just started developing. There were no traces of forelimb buds at this stage. Upon further captive
5
Figure 2. Developmental stages of Duttaphrynus hololius (from top to bottom): larvae of stage 25, stage 31, stage 43 and a completely metamorphosed toadlet (all images of living specimens).
maintenance with the same feed, for two more days (i.e., 8 June 2013), the older larva of stage 31 reached stage
43, with the emergence of forelimbs from the body wall, and a slight reduction in the length of the caudal fin. As it
was raised further, the caudal fin started decreasing in both height and length, due to the absorption of nutrition for
Figure 3. Structure of the oral apparatus in a Duttaphrynus hololius
larva of stage 25 CESF2354 a.
6
Figure 4. SEM image of the oral disc of a stage 25 larva CESF2354
b of Duttaphrynus hololius, showing the characteristic features.
ALYTES 2014 | 31
Table 1. Measurements of Duttaphrynus hololius tadpoles.
CESF2354 a
(stage 25)
CESF2354 b
(stage 25)
Mean
CESF2354 c
(stage 43)
Tot L
18.9
19.33
19.11
16.36
Tail
12.26
12.57
12.41
8.48
HBL
6.35
6.76
6.55
8.11
HBW
3.41
2.86
3.13
3.21
HBDepth
2.28
-
2.28
-
ED
0.37
0.44
0.40
1.19
EN
0.6
-
0.6
0.7
ES
1.33
1.11
1.22
1.04
IN
0.86
0.9
0.88
0.83
IO
1.04
1.03
1.03
2.37
Caudal fin height
2.49
1.95
2.22
1.9
Caudal muscle height
1.44
1.5
1.47
1.54
oral disc width
1.84
1.62
1.73
2.3
post. tooth row width
0.05
0.05
0.05
-
ant. tooth row width
0.06
0.07
0.06
-
inter tooth row distance
0.1
0.11
0.10
-
UAL
-
-
-
1.49
LAL
-
-
-
1.53
PAL
-
-
-
1.87
FEL
-
-
-
2.8
TBL
-
-
-
2.29
TAR
-
-
-
1.43
FOL
-
-
-
1.95
metamorphosis. At this stage, the larvae had developed intensive pigmentation on the dorsal skin with the pale,
inverted ‘V’ shaped marking on the dorsum becoming more evident, and numerous orange colored spots on the
dorsum. The venter was more opaque than it was in the earlier stage. Freshly metamorphosed toadlet observed in
the field, with its tail completely absorbed, resembled the adult in colouration but with a greater density of orange
spots on the body. An evident, light colored inter-orbital band was observed in addition to the inverted ‘V’ marking
on the back in the developed toadlets (fig. 2). The larvae of stages 25 and 42 are illustrated and described below
in detail.
Description of larvae of Duttaphrynus hololius (Günther, 1875): (tab. 1)
Stage 25: CESF2354 a & b
Head-body ovoid, dorso-ventrally depressed; about twice as long as broad (HBW:HBL 0.47), slightly
longer than half the length of the tail (TaL:HBL 1.89). Snout blunt and rounded in dorsal view. Tail long and
tapering at the end, caudal fins better developed on the ventral than the dorsal part; caudal musculature intensive.
Overall colouration: dark purplish on the dorsum without any pattern; venter translucent and pale white. Gut
highly coiled and visible through the translucent skin in the posterior ventral part of the head-body.
Spiracle sinistral, vent-tube dextral; oriented posteriorly towards the right side in the ventral region right
at the junction of the tail beginning and the end of the posterior end of the head-body. Eyes small (0.4 mm
diameter); positioned dorsally; inter orbital space more than twice the eye diameter (IO:ED 2.5); nostrils slightly
closer to each other than the eyes (IO:IN 1.17); nostrils positioned dorsally, situated midway between the snout
tip and the eyes (EN:ES 0.49); snout rounded in dorsal and ventral views. Dorsal skin densely pigmented; overall
colouration dark purplish brown above without any distinct pattern and pale white ventrally. Oral sucker circular
7
Figure 5. Lateral, ventral and dorsal views of the Stage 25 larva CESF2354 a of Duttaphrynus hololius in preservation.
Figure 6. Scanning Electron Microscopic images of the upper (A) and lower labia (B) of the oral apparatus in a stage 25 larva CESF2354 b of
Duttaphrynus hololius.
and not emarginate; positioned antero-ventrally. Keratodont well developed; labial tooth row formula: 2(2)/2(2).
The upper (anterior) labium consists of a total of 99 denticles on the outer row, with 56 on the right half and 43
on the left half, demarcated by a short, interrupted, fleshy diastema at the centre of the left and right halves. The
interior row of the upper jaw has 97 denticles arranged in a single, continuous series. Interior tooth row of the
lower (posterior) labium separated into two halves, with 34 denticles on the right half and 38 on the left (fig. 3).
The exterior tooth row of the lower labium composed of a single, discontinuous row of 73 denticles. The upper
and lower jaw sheaths are indicated by a well developed, keratinized beak. The upper jaw sheaths indicated by a
8
ALYTES 2014 | 31
Figure 7. Scanning Electron Microscopic images of: A. the upper jaw sheath; B. keratodont structures in the upper jaw; C. lower labia in a
stage 25 larva of Duttaphrynus hololius.
continuous, undivided, serrated ridge.
Lower jaw sheath with an acute bend at the center. Lateral process absent on the upper jaw. Marginal
papillae located at the junction of the upper and lower labia arranged in two rows; beak on the upper labium formed
by a hard, keratinized ridge. Keratodont constituted by two concentric series, which consist of inwardly curved
tooth; those of the outer and inner rows of the upper and lower labia of the same size, made of elongate structures
with 6-7 pointed terminal ends (figs. 4-8).
9
Figure 8. Ventral view of the mouth of the stage 43 tadpole CESF2354 c of Duttaphrynus hololius showing disappearance of keratodont, and
initiation of development of the mouth.
Stage 43: CESF2354 c
Body elongate (SVL 8.11 mm) and moderately slender (BW 3.21 mm); habitus depressed; tail a little
shorter (TaL 8.36 mm) after a bit of absorption; fore and hind-limbs well developed with evident differentiation of
the digits. Oral apparatus transformed to mouth with the loss of keratodont and beak. Head differentiable from the
body and slightly broader (HW 3.88 mm) than the latter (BW 3.21 mm). Position of the eyes shifted from dorsal to
lateral; eye larger (ED 1.19 mm) than its distance to the nostril (EN 0.70 mm) and to the snout tip (ES 1.04 mm).
Eyes much separated from each other (IO 2.37 mm) than the nares (IN 0.83 mm).
Upper arms (UAL 1.49 mm), about as long as the lower arm (LAL 1.53), palm slightly longer (PAL
1.87 mm) than upper and lower arms. Thigh longer than the shank (FEL:TBL 1.22), tarsus shorter than the foot
(TAR:FOL 0.73). Toes basally webbed (fig. 9).
DISCUSSION
Of the nine species of Duttaphrynus known from peninsular India (Frost, 2013), larval traits have been
documented only in a very few species (Das & Dutta, 2007) among which none existed for the poorly known
D. hololius till very recently (Ganesh et al., 2013). However, even in the paper by Ganesh et al. (2013), only a
general description of larvae of stages 20-47 was presented briefly. Additionally, a stage-wise description of the
key morphological characters of the tadpoles could not be elaborated as their study was based only on observations
made on live individuals in the field. Hence, our novel data presented here fills in the gap in the existing knowledge
on the larval characters of this species by providing a detailed morphological description of the larvae at two major
transitional stages, based on vouchered material. Thus, this paper supplements the brief notes on the larvae of this
species provided by Ganesh et al. (2013).
Additionally, the data on the time taken for their advanced ontogenetic development is presented. The data
on breeding period and habitat associations of the larvae are consistent with those reported by Ganesh et al. (2013).
Morphologically, the tadpoles of D. hololius can be differentiated from the congeneric species described
as follows. In D. hololius, the tadpoles are dark purplish brown without any pattern in the early stages, and with
an inverted pale ‘V’ mark in the later stages (vs. black dorsal colour with shiny silvery spots on the body in D.
stomaticus; uniform black in colour in D. melanostictus; brownish tadpoles that are smaller than those of D.
melanostictus with relatively large nostrils in D. scaber) (Daniel, 1963).
It is interesting to note that no other anuran species were recorded to breed syntopically with D. hololius,
which has a peculiar preference to rocky pools for breeding and oviposition. This observation, again, corroborates
10
ALYTES 2014 | 31
Figure 9. Dorsal, ventral and lateral views of the stage 43 tadpole CESF2354 c of Duttaphrynus hololius.
that of Ganesh et al. (2013). Despite being a species found in the open rocky hills, a non mesic habitat, this species
has remained poorly known in terms of its biology for a very long time. Additional data on its distribution and
natural history being generated, would aid in assessing the conservation status of this taxon, which is currently
considered to be data deficient (Biju et al., 2004).
ACKNOWLEDGEMENTS
We are grateful to Mr. Sanjeev Kumar and the volunteers of the KANS (Anupama, Ashok, Karthikeyan,
Kesavan, Prem, Ranjith, Rohit, Sampatha and Tarsh) for their help extended in the field. Special thanks to Mr.
S.P. Vijayakumar and Dr. K.P. Dinesh for their suggestions and support; Dr. K. Shanker for the facilities provided
at the lab. We thank Mr. Govindaswamy and Ms. Vidya of the Department of Material Sciences, IISc for kindly
extending their technical expertise with scanning electron microscopic photography of the samples. Constructive
comments by two anonymous referees helped improve the manuscript.
11
REFERENCES
Adimalliah, D., Rao, V.V., Surender, G. (2012). Günther’s toad Duttaphrynus hololius (Günther, 1876) from
Nalgonda district, Andhra Pradesh. Cobra, 6(2): 8-11.
Altig, R., Johnston, G.F. (1989). Guilds of anuran larvae: Relationships among developmental modes, morphologies
and habitats. Herpetological monographs, 3: 81-109.
Biju, S.D., Dutta, S.K., Vasudevan, K., Vijayakumar, S.P., Srinivasulu, C., Bhuddhe, G.D. (2004). Duttaphrynus
hololius. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.1. Available from http://
www.iucnredlist.org (accessed September 2013).
Chandramouli, S.R., Ganesh, S.R., Baskaran, N. (2011). On recent sightings of a little-known south Indian toad,
Duttaphrynus hololius (Günther, 1876) with notes on its morphological characterization and ecology.
Herpetology Notes, 4: 271-274.
Daniel, J.C. (1963). Field guide to the amphibians of Western India. Part I. Journal of the Bombay Natural History
Society, 60(2): 415-438.
Das, I., Dutta, S.K. (2007). Sources of larval identities for amphibians of India. Hamadryad, 31(2): 152-181.
Duellman, W.E., Trueb, L. (1994). Biology of Amphibians. The Johns Hopkins University Press, Baltimore and
London.
Frost, D.R. (2013). Amphibian species of the World: an Online Reference. Version 5.6. American Museum of
Natural History, New York. Available from http://research.amnh.org/herpetology/amphibia/index.html
(accessed December 2013).
Ganesh, S.R., Kalaimani, A., Nath, A., Kumar, R.B. (2013). First observations on the larval characteristics of
Günther’s toad Duttaphrynus hololius (Günther, 1876). Herpetotropicos, 9: 5-8.
Gosner, K.L. (1960). A simplified table for staging anuran embryos and larvae with notes on identification.
Herpetologica, 16: 183-190.
Kalaimani, A., Nath, A., Kumar, R.B. (2012). A note on records of rare and endemic Duttaphrynus hololius
(Günther, 1876) from Tamil Nadu, Eastern Ghats, India. Frog Leg, 18: 27-30.
McDiarmid, R.W., Altig, R. (1999). Tadpoles: the biology of anuran larvae. University of Chicago Press, Chicago.
Srinivasulu, B., S.R. Ganesh & C. Srinivasulu (2013). New regional record and notes on historical specimens of
Günther’s Toad Duttaphrynus hololius with comments on other southeastern Indian congeners. Journal of
Threatened Taxa, 5(13): 4784-4790.
12
RESEARCH ARTICLE
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
Luis M. P. Ceríaco1,2*, David C. Blackburn3, Mariana P. Marques4,
Francisco M. Calado4
Centro de Estudos de História e Filosofia 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
1.
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.
INTRODUCTION
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 Científica
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 identified 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
Received 04 April 2014
Accepted 10 July 2014
luisceriaco@netcabo.pt
LUIS M. P. CERÍACO et al.
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 scientific 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 classification of animals by “natural
families”, and to the description and study of economically useful species, always following Cuvier’s classification
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 (17911853) 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 scientific 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 finish 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
scientific 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 field 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 significant of these expeditions was that of the Portuguese naturalist and explorer Francisco Newton (18641909) 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, fishes, 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 first 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.
15
seven new anuran taxa, including five 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 (fig. 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 influence 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 (fig. 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 fire of 1978. Because of this, investigations of taxa described by Portuguese naturalists and the
relevant type specimens have been difficult 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 fire 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 difficult, 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 identified. 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
problems.
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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MATERIALS AND METHODS
This published work has been registered in ZooBank. The ZooBank LSIDs (Life Science Identifiers)
can be resolved and the associated information viewed through any standard web browser. The LSID for this
publication is: urn:lsid:zoobank.org:pub:65ACCBB7-E69C-49EE-A866-EA2D122E302D
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 identification and clarification 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 finger (HAL), inner toe length (ITL), third finger length
from base of first subarticular tubercle (TFL), lower leg (crus, or tibiofibula) 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 first 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
Table 1. Localities for MHNFCP type specimens, with updated names (including province) and estimated latitude, longitude, and elevation.
Locality in publication
Corresponding taxa
Current locality name
Latitude
Longitude
Elevation
(m)
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
Gonguembo
09° 20'S
14° 54'E
750-800
Rio Luinha
Rappia bivittata
Rio Luinha
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
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
09° 08'S
14° 46'E
630
Cacuaco
(see account for
Rappia bocagei var. maculata)
Cacuaco
(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
17
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 identifications, 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.
SYSTEMATIC ACCOUNTS
Extant types
Amphibia
Hyperoliidae
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 (fig. 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
accurately.
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.
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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 %.
Rappia
bivittata
Rappia
fasciata
Rappia platyceps
var. angolensis
Hyperolius
platyceps
Hyperolius
cinnamomeoventris
(n = 12)
(n = 14)
MHNFCP
017291
MHNFCP
017302
MHNFCP
017294
MHNFCP
017303
SVL
19.05
19.65
25.13
26.21
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)
21.27-26.98 (23.98 ± 1.67) 21.46-29.29 (23.87 ± 2.18)
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
bivittata.
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
19
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 specific 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 (fig. 4).
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, confirms 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 identified specifically 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 flattened 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.
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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”
(fig. 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.
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.
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 confirms 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,
21
Table 3. Measures (in mm) of both syntypes
of Rappia nobrei Ferreira, 1904.
MHNFCP
017292
MHNFCP
017298
SVL
19.7
19.5
HW
6.46
5.91
HL
6.94
6.13
MN
5.76
5.23
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 difficult 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”
4.01
3.36
MFE
(MHNFCP 017307, ♂, 2 juveniles, SVL 26.3, 24.6, 17.6 mm
respectively) collected by Francisco Newton in Angola during the
2.32
1.44
MBE
1903-1905 expedition (fig. 6). This species is a patronym for Baltasar
3.10
2.42
IFE
Osório (1855-1926), ichthyologist, carcinologist, and director of the
4.41
4.34
IBE
Zoological section of the Museu Nacional de História Natural de
Lisboa (Museu Bocage).
3.95
4.02
FLL
Present name. Afrixalus osorioi (Ferreira, 1906).
4.33
4.33
HAL
Remarks. Ferreira’s (1906:162) description of Rappia osorioi
2.40
2.32
TFL
was based on three specimens, one male adult and two juveniles,
from “Quilombo”. The species is considered valid and is broadly
6.19
6.16
TL
distributed from northwestern Angola and across much of the Congo
7.64
8.73
FOL
Basin (Schiøtz, 1974; Perret, 1976b; Laurent, 1982; Channing, 2001),
5.50
6.59
FTL
and perhaps even Kenya (Köhler et al., 2005). Laurent (1972, 1982)
discussed patterns of morphological variation in A. osorioi and the
1.79
1.84
IN
minor phenotypic differences from two other Congo Basin species,
1.80
1.61
EN
A. equatorialis and A. leucostictus. In his discussion of the colour
pattern variation of A. osoroi, Laurent (1982) notes that “la phenotype
2.45
1.94
EL
représenté par l’holotype” is a somewhat rectangular elongate and
ITL
2.37
2.56
somber scapular spot. However, the citation provided in this discussion
FL
8.82
9.81
(Laurent, 1941), as well as his list of specimens examined, indicates
that this phenotype is that of the holotype of Megalixalus fornasinii
SL
2.23
1.94
congicus Laurent, 1941 and not type material of Afrixalus osoroi.
NS
0.67
0.75
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
conspecific with Megalixalus fornasinii congicus.
The existence of two different jars at MHNFCP containing specimens identified as possible types of
“osorioi” deserves 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 first 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 specifically 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
22
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Figure 6. Dorsal and ventral views of the syntypes of Rappia osorioi Ferreira, 1906 (catalogued as a lot, MHNFCP 017307). Scale bar = 10 mm.
Figure 7. Dorsal and ventral views of the syntype of Rappia platyceps var. angolensis Ferreira, 1906 (MHNFCP 017303). Scale bar = 10 mm.
23
Francisco Newton in Angola during the 1903-1905 expedition and still present in Porto (fig. 7). The other syntype
from “N’golla Bumba” was not located.
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
identification and delimitation (Frétey et al., 2011). Some hyperoliid species are characterized by remarkable
polymorphism (Hoffman & Blouin, 2000) that is in some cases reflected 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 confirm that the specimen is indeed a member of this species. More specifically, 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; fig.
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 first 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
Bocage).
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.
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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 fingers, 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
viridiflavus complex, there clearly is need for further study to resolve the systematics of these taxa.
Arthroleptidae
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 (fig. 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 conspecific 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 five 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 identification 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 bocagei” from 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 identified 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 fingers” (our translation). Based
25
Figure 9. Dorsal and ventral views of the holotype of Hylambates bocagei var. leucopunctata Ferreira, 1906 (MHNFCP 017324). Scale bar = 10 mm.
on our examination, the principal morphological feature suggesting that “var. leucopunctacta” is distinctive is the
expansion of the terminal disks of the fingers, 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 identification (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
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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 fire
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 (fig. 10). The species is a patronym for Bento
Carqueja (1860-1935), professor and naturalist in the
Academia Politécnica do Porto.
Table 4. Measures (in mm) of the holotype of Arthroleptis
Present name. Arthroleptis carquejai Ferreira, 1906.
carquejai Ferreira, 1904 (MHNFCP 018586) and the
Remarks. Since its initial description, this
specimens of A. carquejai from Zoologisches Museum
species
has
remained known only from the holotype
Hamburg.
and no subsequent authors have investigated its validity.
Our investigation suggests that Arthroleptis carquejai is
Holotype
specimens from ZMH
indeed a valid species and is likely part of the group of
species that includes Arthroleptis variabilis Matschie,
MHNFCP
ZMH A
ZMH A
1893, Arthroleptis perreti Blackburn, Gonwouo, Ernst,
018586
09493
09492
Rödel, 2009, and Arthroleptis palava Blackburn,
27.77
29.70
20.00
SVL
Gvoždik, Leaché, 2010, from Cameroon and other
9.68
10.66
6.76
HW
Central African countries. While the pigmentation of
the specimen has faded, Ferreira’s (1906) original plate
9.45
9.95
7.31
HL
(fig. 1) indicates that the gular and anterior venter were
8.26
6.79
6.70
MN
darkly pigmented with pale spots. Because the gular
region has pale spots but lacks a pale mid-line stripe,
6.72
6.49
5.44
MFE
the holotype of A. carquejai differs from A. variabilis,
3.25
3.19
2.50
MBE
in which there is a prominent and well-defined mid-line
3.88
4.31
3.61
IFE
gular stripe in females and juveniles, and pale markings
are generally lacking on the more uniformly pigmented
6.36
8.95
5.50
IBE
gular region of males (Blackburn et al., 2009). Darkly
5.85
7.79
5.10
FLL
pigmented gular and ventral surfaces are generally
uncommon in Arthroleptis but are characteristic of A.
8.64
9.39
5.98
HAL
variabilis and related species (Blackburn et al., 2009,
6.99
5.74
3.61
TFL
2010). During this work, we examined specimens from
14.20
15.68
10.10
Zoologisches Museum Hamburg (ZMH) collected during
FOL
the Hellmich expedition to Angola (Hellmich, 1957),
9.81
7.48
6.30
FTL
including several that are similar in external morphology,
2.09
2.99
1.91
IN
size, and proportions to the holotype of Arthroleptis
carquejai (tab. 4). Both specimens (ZMH A09492-93)
2.26
2.09
1.42
EN
were collected from “Roca Novo” (Roça Novo Mundo)
3.38
3.59
3.19
EL
not far from the type locality (approx. 135 km north)
of A. carquejai. These specimens exhibit the similar
3.87
4.55
2.69
ITL
dark ventral pigmentation and traces of a pale mid-line
FL
13.22
15.42
10.06
stripe (similar to A. palava; Blackburn et al., 2010) but
NS
1.37
1.82
1.03
it is not well-defined as it is in A. variabilis. In addition,
these ZMH collections demonstrate that A. carquejai is
EE
2.78
3.00
2.33
sympatric with Arthroleptis poecilonotus Peters, 1863,
TYE
0.99
1.14
0.64
a wide-ranging species complex in western and central
Africa, which was also collected at “Roca Novo” (ZMH
TYD
1.29
1.49
0.84
A09491, A09494-95).
27
Figure 10. Dorsal and ventral views of the holotype of Arthroleptis carquejai Ferreira, 1906 (MHNFCP 018586). Scale bar = 10 mm.
Reptilia
Typhlopidae
Typhlops boulengeri Bocage, 1893
Original name. Typhlops boulengeri Bocage, 1893: 117.
Type specimen. A syntype (MHNFCP 017434) from Angola (fig. 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 11. Dorsal and ventral views of the paratype of Typhlops boulengeri Bocage, 1893 (MHNFCP 017434). Scale bar = 10 mm.
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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
Amphibia
Hyperoliidae
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.
Reptilia
Typhlopidae
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.
DISCUSSION
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 fire), and a paratype of Typhlops
boulengeri Bocage, 1893, given by the Lisbon museum. In addition to the type specimens, our survey revealed
29
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 difficult to identify because of the lack of
labels or other identifying information.
Of the taxa described by Ferreira, we confirm 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 clarifications contribute to our understanding of anuran diversity in southwestern Africa and are a
modest first step towards refining 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 field studies will hopefully spark a new era for the Angolan biodiversity studies (Conradie et al.,
2012, 2013; Ernst et al., 2014). New field 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
description.
NOTE
After the acceptance and final 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.
ACKNOWLEDGEMENTS
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 scientific 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 first 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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ALYTES 2014 | 31
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APPENDIX 1
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,
1968-161A,1969-142.
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.
Arthroleptis carquejai: Zoologisches Museum für Hamburg: ZMH A09492-93.
Arthroleptis poecilonotus: Zoologisches Museum für Hamburg: ZMH A09491, A09494-95.
33
APPENDIX 2
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
AHMB CE G. 75
“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 fine portuguese fish: 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
AHMB CE G. 76
“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:
kindly presented to B. M.
34
1.
2.
3.
4.
6.
7.
8.
9.
10.
11.
Limnophis bicolour, g. & sp. n.
Causus rhombeatus
Lycophidium horstockii
& 5 I shall send the name at the later period end of this letter
has not arrived
Hyperolius marmoratus
cannot be determined without other specimens
Hyperolius fulvovittatus, Cope
- nasutus, sp. n.
Cystignathus bocagii, sp. n.
ALYTES 2014 | 31
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
Leptopelis natalensis
Hyperolius sp?
Hyperolius modestus
Hyperolius nasutus - type
Hyperolius sp? an nasutus, young.
Hyperolius sp?
Hyperolius modestus
Bufo panterinus
nos. 11-19 are returned in today’s parcel
Grayia triangularis
Ahaestulla irregularis
Boodon lineatus
nos. 20 - 22 have been returned through mr. Monteiro several months ago; I hope you have safely
received them.
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
Whitefriars
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
AHMB CE G78
“British museum
6. 5. 66
35
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 fingers hands, but such dilatations you find also in several Rana & other so called
oxydactyle frogs. Cystign. bocagii has no webb between the fingers & 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 confidence 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 printer’s 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 indefinitely 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
AHMB CE G79
“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 finger 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.
Lizards:
1. Stellio angolensis = Agama atricollis, A. Smith, Ill. Slep. Cpp p. 14. I have compared your specimen with the
type.
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”
36
SCIENTIFIC NOTE
2014 | VOLUME 31 | PAGES 37-39
Field observation of an adult Lesser treefrog
Dendropsophus minutus (Anura: Hylidae) being
consumed by a neotropical Lethocerus sp.
(Hemiptera: Belostomatidae) nymph
Ricardo Rocha1,2,3*, Thaís Almeida4, Adrià López-Baucells1,3,5
Centro de Biologia Ambiental, Departamento de Biologia Animal, Faculdade de Ciências da Universidade de Lisboa, Bloco C2, Campo
Grande, 1749-016 Lisboa, Portugal
2.
Metapopulation Research Group, Faculty of Biosciences, University of Helsinki, PO Box 65 (Viikinkaari 1), FI-00014 Helsinki, Finland
3.
Biological Dynamics of Forest Fragments Project, National Institute for Amazonian Research (INPA) and Smithsonian Tropical Research
Institute, C.P. 478, Manaus, AM 69011-970, Brazil
4.
Laboratório de Citotaxonomia e Insetos Aquáticos, Coordenação de Pesquisas em Entomologia – CPEN/Instituto Nacional de Pesquisas da
Amazônia – INPA, Av. André Araújo, 2936, Aleixo, CEP 69060-001, Manaus, AM, Brazil.
5.
Museu de Ciències Naturals de Granollers, Àrea Investigació en Quiròpters, Av. Francesc Macià 51, 08402 Granollers, Catalonia, Spain
1.
Amphibians constitute important items in the diet of many predators. Giant water bugs have been
reported to feed on several species of amphibians; however, there is still a poor understanding of the
complexity of their food webs. Here, we report the consumption of an adult Dendropsophus minutus
(Anura: Hylidae) by a Lethocerus sp. (Hemiptera: Belostomatidae) nymph, in Central Amazon, Brazil.
This represents the first observation of thropic interaction between Lethocerus sp. and D. minutus
and the first report of a neotropical Lethocerus sp. nymph feeding upon an adult vertebrate.
Giant water bugs of the family Belostomatidae are widely distributed throughout the world’s tropical and
temperate regions (Hungerford, 1919) and are rapid colonizers of newly formed temporary shallow water environments where they are often found in relatively high densities (Williams, 2006). These aquatic insects are known to
prey upon a great variety of taxa and have been suggested to significantly impact the ecological structure of their
prey communities (Ohba et al., 2008).
The cosmopolitan subfamily Lethocerinae presents the largest body size among belostomids and possesses raptorial forelegs which it uses to capture terrestrial and aquatic invertebrates, small fish, tadpoles, adult
anurans and, less frequently, snakes and turtles (Hirai & Hidaka, 2002; Mori & Ohba, 2004; Ohba et al., 2008;
Ohba, 2011). Lethocerinae nymphs are tadpole specialists, however, late stage nymphs are known to occasionally
prey upon post-metamorphic amphibians (Ohba et al., 2008).
Adult anurans are key elements of both terrestrial and aquatic food chains, featuring in the diets of vertebrates, invertebrates and even carnivorous plants (Toledo et al., 2007). In the neotropics, trophic interactions
between aquatic insects and frogs have been reported for several species, including for members of the genus
Lethocerus. However, although reports of adult Lethocerus spp. preying upon post-metamorphic anurans are not
uncommon (e.g. Toledo, 2005; Nenda et al., 2008; Pezzuti et al., 2008; de Andrade et al., 2010; Zaracho, 2012) no
trophic interaction between a neotropical Lethocerus nymph and adult amphibian has been reported.
The lesser treefrog Dendropsophus minutus (Peters, 1872) is a nocturnal arboreal hylid widespread
throughout South America, east of the Andes. Its reproduction period takes place in the rainy season from November to May and males usually call from emergent aquatic plants over or next to the water on permanent and temporary ponds (Lima et al., 2005). Adult D. minutus have been reported as prey items of the aquatic environment
Received 03 November 2013
Accepted 21 June 2014
ricardo.nature@gmail.com
RICARDO ROCHA et al.
Figure 1. (A) Third instar giant water bug Lethocerus sp. nymph and (B) the adult minute tree frog Dendropsophus minutus it was feeding
upon.
associated spiders of the family Pisauridae (fishing spiders) (Bernarde et al., 1999) and of adult individuals of the
family Belostomatidae (water bugs) (Bastos et al., 1994; Toledo, 2003). In this publication, we report a giant water
bug Lethocerus sp. nymph feeding upon an adult D. minutus in Central Amazon, Brazil. This constitutes the first
report of consumption of an adult vertebrate by a neotropical Lethocerus sp. nymph.
Our observation took place on 20 March 2013 in a temporary pond surrounded by mature secondary
regrowth, at the Porto Alegre reserve of the Biological Dynamics of Forest Fragments Project (BDFFP) (-2° 21’
26.64”, -59° 57’ 33.65”), Central Amazon, Brazil. The BDFFP harbors a rich and diverse amphibian fauna with
more than 42 described species (Zimmerman & Rodrigues, 1990); however, its belostomid fauna is poorly known.
At 21:32 pm a third instar Lethocerus sp. nymph (TL 31 mm) (fig. 1.A) was observed perching on the
pond vegetation and holding an adult D. minutus (SVL 24.3 mm) (fig. 1.B) with its raptorial forelimbs. The insect
had its proboscis inserted into the D. minutus’ vocal sac and upon disturbance stopped feeding and released the
frog. Since we did not observe the capture event itself we do not exclude the possibility of necrophagy but given
the frog’s small size we believe this indeed represented a case of opportunistic predation. Both specimens were
collected, preserved in 70% alcohol solutions and deposited at the herpetology and entomology collections of the
National Institute of Amazonian Research (INPA) in Manaus, Amazonas, Brazil (D. minutus catalogue number
INPA-H 32305). Unfortunately, at its current developmental stage the nymph lacks the anatomical features necessary for species level identification. However, it could be identified as Lethocerus sp. due to its characteristic three
segment antennae, the widely dilated anterior and posterior femurs and two large claws on the tarsi of the first pair
of legs (de Carlo, 1962).
Post-metamorphic anurans have been found to constitute a significant prey item for third-fifth instar
nymphs of Kirkaldyia (=Lethocerus) deyrolli in Japanese rice fields (Ohba et al., 2008). Along with the observation here reported, we witnessed several other adult and nymphal belostomids capturing tadpoles in the referred
temporary pond. However, although adult D. minutus were commonly observed in the pond, no other trophic
interaction between belostomids and adult frogs were observed.
The observation presented suggests that, when available, adult frogs especially of smaller species like D.
minutus, may constitute important components of the diet of late stage neotropical Lethocerus sp. nymphs. Rearing experiments would be valuable to unveil the contribution of main prey items to the specific growth rates of
different nymphal stages.
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ACKNOWLEDGEMENTS
We would like to thank Christoph F.J. Meyer, Paulo Bobrowiec and Jorge M. Palmeirim for their assistance during all phases of the project and José L.C. Camargo, Rosely Hipólito and Ary Jorge Ferreira for
logistic support. We also thank Sasha Vasconcelos for commenting on early drafts. Funding was provided by the
Portuguese Foundation for Science and Technology through the grants PTDC/BIA-BIC/111184/2009 and SFRH/
BD/80488/2011 and this is publication 638 in the Technical Series of the BDFPP.
LITERATURE CITED
Bastos, R.P., Oliveira, O.C., Pombal Jr, J.P. (1994). Hyla minuta (NCN). Predation. Herpetological Review, 25:
118.
Bernarde, P.S., Souza, M.B., Kokubum, M.C.N. (1999). Predation on Hyla minuta Peters, 1872 (Anura, Hylidae)
by ancylimetes spp. (Aranea, Pisauridae). Biociências, 7: 199-203.
de Andrade, C.A.F., Santana, D.J., de Carvalho-e-Silva, S.P. (2010). Predation on Scinax xsignatus (Anura: Hylidae) by the giant water bug Lethocerus annulipes (Hemiptera Belostomatidae) in a Brazilian restinga
habitat. Herpetology Notes, 3: 53-54.
de Carlo, J.M. (1962). Consideraciones sobre la biologia de Lethocerus mazzai (Hemiptera: Belostomatidae).
Physis, 23: 143-151.
Hirai, T., Hidaka, K. (2002). Anuran-dependent predation by the giant water bug, Lethocerus deyrollei (Hemiptera:
Belostomatidae), in rice field in Japan. Ecological Research, 17: 655-661.
Hungerford, H. (1919). The biology and zoology of aquatic and semi-aquatic Hemiptera. Kansas University Bulletin, 11: 1-341.
Lima, A.P., Magnusson, W.E., Menin, M., Erdtmann, L.K., Rodrigues, D.J., Keller, C., Hödl, W. (2005). Guide to
the frogs of Reserva Adolpho Ducke, Central Amazonia. Áttema Design Editorial, Manaus.
Mori, A., Ohba, S. (2004). Field observations of predation on snakes by the giant water bug. [In Japanese]. Bulletin
of the Herpetlogical Soecity of Japan, 2004: 78-81.
Nenda, S.J., Barrasso, D.A., Cajade, R. (2008). Physalaemus cuvieri Predation. Herpetological Review, 39: 210.
Ohba, S. (2011). Field observation of predation on a turtle by a giant water bug. Entomological Science, 14: 364365.
Odba, S., Miyasaka, H., Nakasuji, F. (2008). The role of amphibian prey in the diet and growth of giant water bug
in Japanese rice fields. Population Ecology, 50: 9-16.
Pezzuti, T.L., De Melo, A.L., Leite, F.S. (2008). Scinax eurydice Predation. Herpetological Review, 39: 341-342.
Toledo, L.F. (2003). Predation on seven South American anuran species by water bugs (Belostomatidae). Phyllomedusa, 2: 105-108.
Toledo, L.F. (2005). Predation of juvenile and adult amphibians by invertebrates: Current knowledge and perspectives. Herpetological Review, 36: 395-400.
Toledo, L.F., Silva, R.R., Haddad, C.F.B. (2007). Anurans as prey: an exploratory analysis and size relationships
between predators and their prey. Journal of Zoology, 271: 170-177.
Williams, D.D. (2006). The biology of temporary waters. Oxford University Press, New York.
Zaracho, V.H. (2012). Predation on Elachistocleis bicolor (Anura: Microhylidae) by Lethocerus annulipes (Heteroptera: Belostomatidae). Herpetology Notes, 5: 227-228.
Zimmerman, B.L., Rodrigues, M.T. (1990). Frogs, snakes and lizards of the INPA-WWF reserves near Manaus,
Brazil. In: Gentry A.H. (ed.) Four Neotropical rainforests, Yale University Press, New Haven: 426-456.
39
40
BIOGRAPHICAL NOTE
2014 | VOLUME 31 | PAGES 41-45
Compléments et rectificatifs à la biographie de
François Daudin
Roger Bour1*
1.
Reptiles et Amphibiens, UMR 7205 OSEB, Département Systématique et Évolution,
Muséum National d’Histoire Naturelle, CP 30, 25 rue Cuvier, 75005 Paris, France
This note completes the biography of François Marie Daudin (Bour, 2011), between 1634 and 1924.
Les parents du “premier” François Daudin
Jacques Daudain [sic] et Jeanne de Langlier se sont épousés le 2 février 1645, à Campeaux (Oise) ; lui
était de la paroisse de Campeaux, elle de celle de Saint-Arnoult, un village distant de 5 km. François Daudin (I)
fut baptisé le 15 février 1647 à Campeaux (Bour, 2011 : 5) ; son parrain était Guillaume Poller, sa marraine Denise
Langlier, de Saint-Arnoult (fig. 1).
(Source : Archives de l’état-civil du département de l’Oise).
Figure 1. Acte de baptême du “premier” François Daudin. Campaux (Oise), 15 février 1647 (Archives de l’état-civil du département de l’Oise).
Catherine de Regnonval (de Pouilly)
Catherine de Regnonval est née vers 1666, probablement à Beauvais. C’était la fille de Nicolas de
Regnonval (vers 1614-1683, Beauvais) et de Marguerite Foy (vers 1629-1679, Beauvais), petite-fille de Nicolas
de Regnonval (vers 1569-1652), juge consul, échevin, maire de Beauvais, et la sœur d’Anne de Regnonval (16531726) et de Nicolas de Regnonval, seigneur de Fabry a (1655-1716). Catherine de Regnonval ne doit pas être
confondue avec sa nièce Catherine de Regnonval de Saint Rémy (1683-1755), fille de son frère Nicolas, comme
nous l’avions relevé dans certaines généalogies, et admis dans notre étude (Bour, 2011 : 6, note 4).
Catherine de Regnonval s’est mariée le 14 septembre 1688 à Beauvais, église Saint-Sauveur, avec
a.
Fabry était un hameau de Pouilly, indiqué sur la carte de Cassini; aujourd’hui subsiste le bois de “Fabris”.
Received 18 December 2013
Accepted 27 January 2014
bour@mnhn.fr
ROGER BOUR
Louis Legras, chevalier, seigneur de Pouilly. Était notamment présent son beau-frère Louis Divery (1654-1727),
“conseiller au baillage du siège présidial de Beauvais”, époux de sa sœur Anne. Son beau-père Jacques Legras,
seigneur de Pouilly, mort quelques mois plus tôt, fut inhumé dans le chœur de l’église de Pouilly (dalle présente)
le 29 mars 1688. Un ancêtre, Jacques Legras, était déjà seigneur de Pouilly en 1598.
Louis Legras, seigneur de Pouilly, mourut à l’âge de 40 ans environ (né vers 1654 ; actes manquants) ;
il fut inhumé dans l’église de Pouilly le 23 décembre 1694. Veuve de Louis Legras, Catherine de Regnonval se
remaria rapidement : le 27 janvier 1695 elle épousa à Pouilly Jean Philip de Césan, né vers 1648, “colonel exempt
des gardes du corps du Roy”. Jean Philip de Sésan [selon sa signature ; variantes : Philippe, de Philippe…, de
Césan, de Cézan, de Cizeau…], décéda à l’âge de 85 ans et fut lui aussi inhumé en l’église de Pouilly (dalle
présente), le 11 septembre 1733. Il était “chevalier mestre de camp de cavalerie, capitaine exempt des gardes du
corps du Roy, chevalier de l’ordre royal et militaire de St. Louis”. Néanmoins il n’est pas cité, de même que les
Regnonval, dans les recueils des membres de l’ordre de Saint-Louis (d’Hozier, 1818).
Catherine de Regnonval, dame de Pouilly, décéda le 31 mars 1750 et fut inhumée le 1er Avril dans le
chœur de l’église de Pouilly (dalle présente, réunissant ses deux époux). Quatre petits neveux assistaient à la
cérémonie : Nicolas Pierre de Regnonval, seigneur de Fabry (né en 1728) ; Germer Raoul de Regnonval de
Courcelles, chanoine de l’Église de Beauvais (né en 1729) ; Jean Toussaint Le Caron de Troussures (né en 1709) ;
Claude Joseph Le Mareschal, maire de Beauvais, conseiller du Roy (né en 1707).
En novembre 1750 François Daudin (III), né à Campaux le 15 mai 1703, acheta aux héritiers de Catherine
de Regnonval le château et les terres de la seigneurie de Pouilly (acte passé devant Hainne, notaire, selon Troussures, 1895). Seigneur de Pouilly, Montoisel et autres lieux, il fut lui aussi inhumé, le 6 juin 1779, dans le chœur
de l’église de Pouilly (dalle présente).
En 1781, le 23 octobre, fut inhumé, toujours dans le chœur de l’église de Pouilly (dalle présente) Marie
Nicolas de Regnonval de Courcelles (né en 1729), “chevalier de l’ordre royal et militaire de St Louis, ancien
capitaine au régiment de Berry infanterie”, décédé la veille. Étaient présents Toussaint Stanislas de Regnonval [de
Martel], “écuyer, chevalier de l’ordre royal et militaire de St Louis, capitaine au régiment de Conty infanterie”
(né en 1736) ; Joseph François de Regnonval de Courcelles [de Hodan], “chanoine, diacre, de l’église cathédrale
de Beauvais” (né en 1738), “frères du défunt” ; Nicolas Pierre de Regnonval, “seigneur de Fabry, Mons, et autres
lieux” (né en 1728), cousin germain ; Louis Lucien Le Caron de Troussures (né en 1751).
(Sources: Archives de l’état-civil du département de l’Oise ; église Saint-Lucien de Pouilly).
Marie Madeleine Louise Escallard de la Bellangerie
La mère de François Marie Daudin est née le 3 décembre 1754 à Sougé-le-Ganelon (Sarthe), et baptisée le
même jour. Le parrain était Louis Herbin, “chyrurgien juré”, la marraine Magdeleine Lambert, sa tante. Un frère,
Nicolas Pierre, est né au même lieu le 11 octobre 1756 ; parrain René François Herbin, marraine Marie Herbin. Le
père Nicolas Pierre “Lescalard de la Belleangerie” était officier de M. le prince de Bouillon (voir Bour, 2011 : 8,
note 8). Les parents se sont unis le 30 septembre 1749 à Sougé-le-Ganelon.
Nicolas Pierre Escalard, sieur de la Bellangerie (âgé de trente-deux ans environ), était le fils de “défunt
messire Pierre Escalar conseiller du Roi à l’élection de Berné [Bernay, dans l’Eure] et de défunte demoiselle
Anne Piot”. Magdeleine Fresnais (âgée alors d’environ dix-neuf ans) était la fille de “défunt maître René Fresnais
advocat au siège d’Assé-le-Boisne et de demoiselle Marie Magdeleine Lambert”. Parmi les témoins figuraient
Charles Herbin “advocat au siège royal de Fresnay” [Fresnay-sur-Sarthe], Charlotte et Elisabeth Fresnais, sœurs
de l’épouse, Marie Madeleine Lambert, sa tante. Sa mère Madeleine Renée Fresnais avait été baptisée le 23 juin
1731 (naissance le 22) à Sougé-le-Ganelon, fille de René Fresnais et Magdelainne Lambert. Le parrain était Maître
Urban [?] Lambert, notaire royal demeurant Saint-Léonard des Bois, la marraine demoiselle Françoise Fresnais,
épouse de maître Peltier notaire à Assé-le-Boisne.
(Source: Archives de l’état-civil du département de la Sarthe).
Adélaïde Geneviève de Grégoire de Saint-Sauveur
L’épouse de François Marie Daudin, Adélaïde Geneviève, née le 6 avril 1775 à Paris (Bour, 2011 : 13),
était une enfant “naturelle”, fille de Hyacinthe Philémon de Grégoire de Saint-Sauveur et de Jeanne Madeleine
Olivier. Père et mère l’ont reconnue lors du baptême à l’église Bonne-Nouvelle, à Paris. Elle avait une sœur aînée,
Adélaïde Jeanne, née le 9 novembre 1767.
Adélaïde Geneviève ne fut pas reconnue civilement par son père (décédé le 11 juillet 1784 à Bagnères-deBigorre), mais elle porta le nom de Grégoire de Saint-Sauveur grâce à une dérogation royale accordée en 1791, en
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réponse à une demande de légitimation signée de la mère et de la fille : “Elle ne conservera que précairement et
momentanément le nom de Grégoire de Saint-Sauveur, qui lui appartient ; ce nom ira se perdre dans celui de son
mari et sera totalement éteint après elle” (Levy, 1911). Dans cette note son année de naissance est fixée à 1771,
probablement par erreur lors de la copie de l’acte. Hyacinthe Philémon Grégoire de Saint-Sauveur était issu de
deux familles nobles du Gévaudan, les de Grégoire (d’Ispagnac) du côté paternel, les Chapelain d’Issenges du côté
maternel, dont les ascendants sont reconnus dès le XIVe siècle.
Le mariage religieux de François Daudin (alors domicilié 27 rue de Hauteville, ancien 9e arrondissement
de Paris, aujourd’hui dans le 10e) et d’Adélaïde Grégoire de Saint-Sauveur fut célébré le même jour que le mariage
civil, le 10 Décembre 1800, dans l’église Saint-Roch (dans l’ancien 2e arrondissement, aujourd’hui dans le 1er).
Vente des collections de Daudin
Le 4 pluviôse an XII (25 janvier 1804), Louis Dufresne, ami de Daudin (Bour, 2011 : 15) adressa à Lacepède (et non à Duméril, qui était alors son adjoint mais officieusement le dirigeant de la chaire d’ichtyologie et
d’herpétologie) le message suivant, qu’il recopia dans son journal :
“A M. de Lacepede
J’ai l’honneur de vous anoncer que j’ai achetté pour le museum / a la vente de feu Mr Daudin 140. bocaux contenant grenouille, Rainette / crapaud, Serpens lezards poissons, vers etc. tous conservé dans lesprit de /
vin ce lot a été adjugé a la Somme de 90. fr ce qui fait a peu pres / 12 s pour chaque bocal. compris la liqueur et
l’individue quil contient / plus vu un autre lot de 3 Serpens, un diodon et une belle espèce de Congre, / marbrée de
jaune Sur un fond brun cet individu je crois manque ala / Collection du museum, ce lot coute 5. francs Soixante
quinze Centimes, je / vous prie d’avoir la bonté den faire part al assemblée”.
Dans ce même journal, le mois précédent, Dufresne s’était adressé à de Fourcroy pour obtenir l’accord
de l’administration du Muséum, afin de récupérer dans les biens laissés par Daudin des dessins publiés dans les
Annales du Muséum, probablement l’œuvre de Barraband :
“Si vous jugez convenable de me donner une autorisation par / écrit a léffet de reclamer aupres des
heritiers de feu Mr / Daudin, quelques desseins originaux qui appartiennent aux / annales du museum, j’ai déjà
empeché d’en mettre un sur / l’inventaire de Ses effets, mais les officiers de Justice, ne peuvent / me le remettre,
Sans avoir eté préalablement autorisée par ladministration / du museum”.
(Source : Journal de Louis Dufresne conservé dans les archives du laboratoire de Zoologie, Mammifères
et Oiseaux, du Muséum de Paris).
Maurice Ernest Depouilly Daudin
Après avoir vécu à Pouilly l’arrière petit-fils de François Marie Daudin s’installa à Paris, avec sa mère, au
99 rue d’Amsterdam dans le 8e arrondissement (Bour, 2011 : 46). Étudiant, il habitait en 1891 16 rue Clapeyron ;
en 1908, avocat, il habitait 35 rue de Berne. Selon les registres des électeurs de Paris, en 1913, toujours avocat, et
fidèle au 8e arrondissement, il habitait 2 rue Corvetto ; il demeurait et exerçait encore en 1921. En 1916 sa mère,
alors âgée de 76 ans, était domiciliée à cette même adresse. Elle mourut quatre ans plus tard.
Ernest Depouilly Daudin décéda à Paris en 1924. Son testament a été partiellement publié (Anonyme,
1925a) ; les blancs sont dans le texte imprimé, mais certainement remplis dans la version olographe originale.
Nous reproduisons intégralement le texte publié :
Legs Depouilly-Daudin.
Aux termes de ses testament et codicille olographes, en date des 1er juillet 1913 et 11 novembre 1920, déposés chez Me André Charpentier, notaire à Paris, M. Maurice-Ernest Depouilly-Daudin, en son vivant célibataire
majeur, sans profession, demeurant à Paris, rue Corvetto, 2, décédé même ville, rue du Faubourg-Saint-Honoré,
208 a , où il se trouvait momentanément, le 11 septembre 1924, a fait des dispositions sujettes à l’autorisation
administrative dont la teneur suit :
1er juillet 1913.
« J’institue comme légataire universel le Muséum d’Histoire Naturelle à Paris, en mémoire de…..
a.
C’était l’adresse de l’hôpital Beaujon, transféré en 1935 à Clichy-sur-Seine.
43
ROGER BOUR
« La seule condition que j’y mette est la conservation par le Muséum des papiers et médailles, parchemins
et documents concernant la famille Daudin, ainsi que des ouvrages du naturaliste qu’on trouvera dans ma bibliothèque.
« Le Muséum devra également entretenir en bon état de réparation la sépulture de la famille Daudin
située dans le cimetière de Pouilly (Oise) dont…..
« Quant aux tableaux, porcelaines, objets d’art et mobiliers ils pourront être vendus.
« Le présent testament sera également valable dans le cas où je survivrais à ma mère sans avoir eu le
temps ni la possibilité d’en faire un nouveau.
« Il est le seul qui existe de moi à la présente date.
« Le légataire universel devra servir une rente viagère de…..
« Je nomme…..
« Je déclare formellement exclure tous collatéraux qui sont du reste à l’abri du besoin et n’ont aucun droit
sur ma fortune ou succession. »
11 novembre 1920.
« Les dispositions ci-incluses sont confirmées. »
Pour la période qui nous intéresse ici les comptes rendus des assemblées des professeurs (une dizaine par
an) sont conservés sous formes de minutes aux Archives nationales et de procès-verbaux au Muséum. Quant à la
correspondance, le courrier au départ est archivé au Muséum et le courrier à l’arrivée aux Archives nationales.
Louis Alexandre Mangin étant alors Directeur, l’assemblée fut informée du testament olographe de Maurice Ernest Depouilly Daudin le 19 février 1925, par une lettre adressée par Me Louis Georges Robert Violette, notaire
à Méru. L’acceptation du legs fut votée à l’unanimité par l’assemblée des professeurs du 19 mars. Dès le mois
de juillet 1925 les biens mobiliers étaient vendus et l’appartement avait un acquéreur, M. Théret, peut-être Jean
Marie Théret (il a alors 80 ans), le notaire qui avait déjà acheté le château des Daudin à Pouilly à la fin du XIXe
siècle. “Par décret en date du 22 novembre 1925, rendu sur le rapport du ministre de l’instruction publique et des
beaux-arts, le directeur du Muséum d’histoire naturelle est autorisé à accepter, sous bénéfice d’inventaire, au nom
de cet établissement, le legs universel fait par M. Maurice-Ernest Depouilly-Daudin” (Anonyme, 1925b). La suite
est complexe, du fait de la demande d’indemnisation de certains parents, du dépôt d’une main levée liée à la rente
viagère, empêchant la vente, et d’une mauvaise volonté montrée par certains protagonistes…
Les minutes des assemblées mentionnent un dossier particulier concernant le legs Depouilly Daudin, mais
il n’est pas localisé pour l’instant. Ce qui est le plus surprenant, pour ne pas dire le plus regrettable à nos yeux, c’est
que dans les documents du Muséum examinés pas la moindre allusion n’a rappelé la mémoire de François Marie
Daudin. Selon Pascale Heurtel et Antoine Monaque, la Bibliothèque centrale ne possède ni manuscrits ni médailles
provenant de ce legs, et d’éventuels ouvrages provenant de la bibliothèque du naturaliste n’y sont pas référencés.
Quant à la sépulture de la famille Daudin, à Pouilly, il est probable que le Muséum l’a oubliée depuis longtemps.
Comme l’avait auguré Hyacinthe Daudin, le temps déchire et disperse les feuillets de la vie. Les recherches sur
la famille Daudin restent donc inachevées. Il est néanmoins établi qu’Ernest Depouilly Daudin était le dernier
descendant direct de François Marie Daudin, et qu’il mourut célibataire à l’âge de 59 ans, sans postérité.
ENGLISH SUMMARY
Additions and corrections to the biography of François Daudin.
Parents of “first” François Daudin. Jacques Daudain [sic] from Campeaux and Jeanne de Langlier from
Saint-Arnoult were married on 2 February 1645 in Campeaux. François Daudin (I) was christened in this village
on 15 February 1647.
Catherine de Regnonval (de Pouilly). Catherine de Regnonval was born about 1666, probably in Beauvais. She married on 14 September 1688 in Beauvais Louis Legras, knight, seigneur of Pouilly, then on 27 January
1695, in Pouilly, Jean Philip de Césan. She died on 31 mars 1750 and was buried in the choir of the church of
Pouilly. On November 1750 François Daudin (III) bought from the heirs of Catherine de Regnonval the castle and
grounds of the seigneury of Pouilly.
Marie Madeleine Louise Escallard de la Bellangerie. The mother of François Marie Daudin was born
on 3 December 1754 in Sougé-le-Ganelon (Sarthe). Her parents were married in that town on 30 Septembre 1749.
Nicolas Pierre Escalard de la Bellangerie, about 32 years old, was the son of Pierre Escalar(d), councillor of the
King in Bernay (Eure), and Magdeleine Fresnais, about 19 years old, was the daughter of Maître René Fresnais,
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lawyer in Assé-le-Boisne (Sarthe).
Adélaïde Geneviève de Grégoire de Saint-Sauveur. The wife of François Marie Daudin, Adélaïde Geneviève, born on 6 April 1775 in Paris, was a natural child, daughter of Hyacinthe Philémon de Grégoire de
Saint-Sauveur and Jeanne Madeleine Olivier. Adélaïde Geneviève was allowed to bear the name of Grégoire de
Saint-Sauveur thanks to a royal exemption given in 1791. Hyacinthe Philémon Grégoire de Saint-Sauveur was
issued from two noble families of Gévaudan (Lozère), de Grégoire (d’Ispagnac) on the father’s side, Chapelain
d’Issenges on the mother’s side, both attested since the XIVth century.
Sale of the collections of Daudin. On 25 January 1804, Louis Dufresne, friend of Daudin, wrote to Lacepède that he bought for the Museum from Daudin’s auction 140 jars including frogs, tree frogs, toads, snakes,
lizards, fish, and worms, all in spirit, adjudicated at 90 francs.
Maurice Ernest Depouilly Daudin. The great grand-son of Daudin died in Paris on 11 September 1924.
He chose the Museum as his sole legatee. “The only condition that I put is the conservation by the Museum of
papers and medals, manuscripts and documents concerning the Daudin family, as well as works of the naturalist
kept in my library. The Museum will also maintain in good repair the tomb of the family Daudin located in the
churchyard of Pouilly (Oise)”. However these various objects are presently unlocated, and the grave is fortunately
maintained by the town council of Pouilly. Ernest Depouilly Daudin lived as bachelor with his mother, died in
1920, and had no posterity.
REMERCIEMENTS
Nous remercions Monsieur le Maire de Pouilly et son épouse pour leur cordial accueil. Au Muséum nous
remercions Antoine Monaque et Pascale Heurtel, de la Bibliothèque centrale, pour leurs recherches sur le legs Depouilly Daudin, et Marie Portas, du Laboratoire des Mammifères et Oiseaux, qui nous a communiqué les passages
du journal de Dufresne mentionnant les collections de Daudin.
LITTÉRATURE CITÉE
Anonyme, (1916). Les ambulances au front. Transport rapide des blessés. Le Gaulois, 14044: 3 (27 mars 1916).
Anonyme, (1925a). Direction du Contentieux (3e bureau). Dons et legs. Legs Depouilly-Daudin. Recueil des actes
administratifs de l’année 1925, bulletin officiel d’information de la Préfecture de Paris et de la Préfecture
de la Seine, Partie municipale, Deuxième section, 57(278): 11403 (27 novembre 1925).
Anonyme, (1925b). Muséum d’histoire naturelle. Journal officiel de la République française, 57, 278: 11403 (27
novembre 1925).
Bour, R. (2011). François Marie Daudin (29 août 1776-30 novembre 1803), auteur de l’Histoire naturelle, générale
et particulière, des Reptiles. Alytes, 28(1/2): 1-76.
d’Hozier, J. F. L. (1818). Recueil de tous les membres composant l’ordre royal et militaire de Saint-Louis, depuis
l’année 1693, époque de sa fondation ; précédé des édits de création et autres relatifs au dit ordre. Ouvrage
posthume. Volume 2. Paris, au bureau général du Bon Français: 1-448.
Levy, É. (1911). Légitimation des enfants naturels. Les lettres de légitimation à la fin de l’ancien régime (17891791). Bulletin de la Société d’études législatives. Rapports et comptes-rendus des séances, travaux relatifs aux questions étudiées par la Société, 10: 399-414.
Troussures, comte de [L. L. Le Caron], (1895). Armorial des familles du Beauvaisis dont les membres ont pris part
aux assemblées de la Noblesse pour l’élection des députés aux États généraux en 1789. Mémoires de la
Société académique d’Archéologie, Sciences & Arts du département de l’Oise, 16(1): 592-642.
45
CONTENTS
Research Articles
S. R. Chandramouli, Ayuthavel Kalaimani
Description of the larvae of Günther’s toad Duttaphrynus hololius (Günther, 1876) (Anura: Bufonidae)
with notes on development and oral ultra-structure..............................................................................
3-12
Luis M. P. Ceríaco, David C. Blackburn, Mariana P. Marques, Francisco M. Calado
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....................................
13-36
Scientific Note
Ricardo Rocha, Thaís Almeida, Adrià López-Baucells
Field observation of an adult Lesser treefrog Dendropsophus minutus (Anura: Hylidae) being
consumed by a neotropical Lethocerus sp. (Hemiptera: Belostomatidae) nymph................................
37-39
Biographical Note
Roger Bour
Compléments et rectificatifs à la biographie de François Daudin........................................................
This journal is available online at www.amphibians.org/alytes
Cover photo: Duttaphrynus hololius © S. R. Chandramouli
41-45

Alytes_2014_31_1-2 - Amphibian Survival Alliance

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