A comprehensive survey of amphibians and reptiles in the … · Introduction The amphibians and...

IntroductionThe amphibians and reptiles of Madagascar are

extremely diverse and display a high level of endemism (Glaw and Vences, 2007). Although the amount of information regarding Malagasy herpetofauna has increased dramatically over the past decade further detailed surveys focused on Malagasy amphibians and reptiles are still required (D’Cruze et al., in press). The

majority of herpetological survey work in Madagascar has focused on long established protected areas (D’Cruze et al., in press). Current information regarding non-protected areas and secondary habitats is extremely

Herpetology Notes, volume 2: 31-44 (2009) (published online on 07 April 2009)

A comprehensive survey of amphibians and reptiles in the extreme north of Madagascar

Steven Megson1, Polly Mitchell1, Jörn Köhler2, Charles Marsh3, Michael Franzen4, Frank Glaw4, Neil D’Cruze*5

1 Frontier, The Society for Environmental Exploration, 50-52 Rivington Street, London, EC2A3QP, UK;

e-mail: [email protected] Hessisches Landesmuseum Darmstadt, Department of Natural

History – Zoology, Friedensplatz 1, 64283 Darmstadt, Ger-many; e-mail: [email protected]

3 Imperial College London, Silwood Park Campus, Buckhurst Road, Ascot, Berkshire, SL5 7PY, United Kingdom;

e-mail: [email protected] Zoologische Staatssammlung München, Münchhausenstr. 21,

81247 München, Germany; e-mail MF: [email protected]; e-mail FG: [email protected] The World Society for the Protection of Animals, 89 Albert

Embankment, London, WE1 7TP, United Kingdom; e-mail: [email protected] * corresponding author

Abstract. We surveyed the western dry forest and coastal forest of the Ampombofofo area in the extreme north of Madagascar for its herpetofauna. We recorded a total of nine amphibian and 46 reptile species via opportunistic searching and pitfall trapping. This is the first such survey and consequently providing new records for the area for all of the species found. For those species endemic to Madagascar, these records also represent the northernmost limit of their known range. We also surveyed the coastal forest of Orangea and recorded a total of three amphibian and 22 reptile species. Furthermore, we provide new additions to the species list for Montagne des Français. We discovered several species in this massif during our relatively short surveys that were not previously known from this site: Reptiles include Gehyra mutilata, Leioheterodon modestus, and Paracontias hildebrandti. Amphibians include an unidentified speces of Cophylinae and two additional species of Stumpffia. The results of this and other surveys in surrounding areas have now demonstrated that many of the species previously considered to be endemic to Montagne des Français are in fact regionally endemic to the north of Madagascar. However, the forested areas located in the extreme north are subject to numerous human induced environmental problems resulting in habitat destruction and remain a high conservation management priority. Therefore we strongly recommend that Montagne des Français should be granted full-protected status so that it is able to function as a biological refuge. Herein, we provide additional conservation and development recommendations for the other terrestrial ecosystems surveyed.

Keywords. Amphibia, Antsiranana region, Ampombofofo, conservation, Orangea, Madagascar, Montagne des Français, Reptilia.

Figure 1. Map of study area in northern Madagascar.

Steven Megson et al.32

scarce (Andreone et al., 2003; Kremen et al., 2008; D’Cruze et al., in press) and must be targeted in order to ensure that informative conservation decisions are made. This paper is part of a series of research articles (D’Cruze and Sabel, 2005; D’Cruze et al., 2006; D’Cruze et al., 2007, 2008) that contributes to the existing knowledge by highlighting the findings of an extensive biodiversity survey conducted in the north of Madagascar.

This and other fieldwork conducted in recent years has revealed the extraordinary importance of the extreme north as a biological center of herpetological diversity and endemism, which has included the discovery of a multitude of undescribed herpetological taxa (e.g. Pintak

and Böhme, 1988; Raselimanana et al., 2000; Glaw et al.,Raselimanana et al., 2000; Glaw et al., Glaw et al., 2001; 2005; 2006a, b, Raxworthy and Nussbaum 2006; Köhler et al., 2008) and resulted in significant geographic range extensions (e.g. Raxworthy and Nussbaum, 1994; Rakotondravony, 2006; D’Cruze et al., 2006, 2007). The herpetofauna in the extreme north of Madagascar is currently conserved by a network of protected areas consisting of Montagne D’Ambre National Park, Forêt d’Ambre Special Reserve, Analamera Special Reserve, Ankarana Special Reserve and Montagne des Français (recently awarded temporary protected status). Of these five biological refuges, three have been recently subjected to intensive survey work (Raxworthy and

SpeciesRelativeabundance

Ecologicaldistribution

Altitude (m a.s.l.)

Habitat type

Dry1

Dry2

Wet1

Wet2

CITESlisting

Regionallyendemic

Amphibia

Ranidae sensu lato

Hoplobatrachus tigerinus C S 10-100 F/A x x x x II

Ptychadena mascareniensis A T/S 10-100 F/A x x x x

Mantellidae

Aglyptodactylus securifer* SC T 10-100 F x x +

Blommersia cf. wittei* R S 10-100 F x

Boophis tephraeomystax C AB/T/S 10-100 F/A x x x x

Laliostoma labrosum SC T/S 10-100 F x x

Mantella viridis*** C T/S 10-100 F x x x x II +

Microhylidae

Stumpffia sp. 1 I T/S 10-100 F x x +

Stumpffia sp. 2 R T/S 10-100 F x +

Reptilia

Chamaeleonidae

Brookesia stumpffi SA AB/T 10-100 F x x x II

Furcifer oustaleti A AB/T 10-100 F/A x x x x II

Furcifer pardalis A AB/T 10-100 F/A x x x x II

Furcifer petteri SA AB/T 10-100 F/A x x x II

Gekkonidae

Blaesodactylus boivini A AB 10-100 F/A x x x x +

Ebenavia inunguis I AB/T 10-100 A x x

Geckolepis cf. maculata A AB 10-100 F/A x x x x

Geckolepis sp. I AB 10-100 F x x x x +

Hemidactylus frenatus I AB 10-100 A x x x

Hemidactylus mercatorius R AB 10-100 A x x

Lygodactylus heterurus trilineigularis R AB/T 10-100 F/A x +

Paroedura cf. karstophila R T 10-100 F x x +

Paroedura stumpffi C AB/T 10-100 F/A x x x x +

Phelsuma abbotti A AB 10-100 F/A x x x x II

Phelsuma grandis A AB 10-100 F/A x x x x II +

Uroplatus ebenaui I AB 10-100 F x x x x II

Uroplatus sp. nov. aff. henkeli C AB 10-100 F x x x x II +

Gerrhosauridae

Zonosaurus boettgeri I AB/T 10-100 F x x +

Scincidae

Amphiglossus ardouini R T 10-100 F x +

Cryptoblepharus boutonii cognatus C T 0-10 x x x x

Cryptoscincus minimus R T 10-100 F/A x x

Madascincus cf. intermedius 1 A T 10-100 F/A x x x x

Madascincus cf. intermedius 2 A T 10-100 F/A x x x x +

Trachylepis elegans A T 10-100 F/A x x x x

Trachylepis tavaratra C T 10-100 F/A x x x +

Table 1. Conservation status and distribution of the amphibian and reptile species found during surveys of Ampombofofo in northern Madagascar 2006-2007.

Amphibians and reptiles of northern Madagascar 33

SpeciesRelativeabundance

Ecologicaldistribution

Altitude (m a.s.l.)

Habitat type

Dry1

Dry2

Wet1

Wet2

CITESlisting

Regionallyendemic

Boidae

Acrantophis madagascariensis** I T 10-100 F/A x x x x I

Sanzinia madagascariensis volontany** I AB/T 10-100 F/A x x x x I

Colubridae sensu lato

Dromicodryas quadrilineatus C T 10-100 F/A x x x

Heteroliodon fohy I T 10-100 F x x x +

Ithycyphus miniatus C AB/T 10-100 F/A x x x x

Langaha madagascariensis I AB/T 10-100 F x x

Leioheterodon madagascariensis A T 10-100 F/A x x x x

Leioheterodon modestus C T 10-100 F/A x x x x

Liophidium therezieni I T 10-100 F x x x x +

Liophidium torquatum C T 10-100 F/A x x x x

Madagascarophis colubrinus C AB/T 10-100 F/A x x x

Mimophis mahfalensis A AB/T 10-100 F/A x x x x

Pseudoxyrhopus quinquelineatus I T 10-100 F x x x

Stenophis granuliceps C AB/T 10-100 F/A x x x +

Stenophis inopinae I AB/T 10-100 F x x x +

Stenophis variabilis R AB/T 10-100 F x

Typhlopidae

Ramphotyphlops braminus I T 10-100 F x

Typhlops cf. microcephalus I T 10-100 F x x x x

Typhlops sp 2 R T 10-100 F x +

Xenotyphlops grandidieri R T 10-100 F x +

Pelomedusidae

Pelusios castanoides R S 10-100 F x

Table 1. continued

Relative Abundance: A = abundant, C = common, I = infrequent, R = rare. SA = seasonally abundant, SC = seasonally common. Eco-logical Distribution: AB = arboreal, T = terrestrial, S = semiaquatic. Habitat: F = Forest, A = anthropogenically disturbed. Seasonal Data: Wet = Wet season, Dry = Dry season, x = present. *Listed as Least Concern in the 2008 IUCN Red List of Threatened Species **Listed as vulnerable. ***Listed as Endangered. + = Regionally endemic

Figure 2. Photographic documentation of some of the habitat types surveyed during this study. A= Ampombofofo; B= Orangea; C= Montagne des Français; and D= Pitfall trap in Orangea.

Nussbaum, 1994; D’Cruze et al., 2007, 2008). Only the forest found within the Analamera Special Reserve and Ankarana Special Reserve have not been subject to in-depth biodiversity surveys that have resulted in detailed published species lists (but see Hawkins et. al., 1990).

The intention of this study was to survey a previously undocumented area of low altitude with large patcheslarge patches of relatively undisturbed western dry forest located western dry forest located outside of the protected areas in the extreme north of Madagascar. Firstly, this survey would serve to identify any species that do not currently receive protection within these biological refuges and provide researchers with the opportunity to assess the current conservation threats that they might face. Secondly, it was hoped

that the information gathered would contribute to the existing literature regarding Malagasy patterns of biodiversity by documenting the composition, geographical and ecological distribution of any species encountered. Crucially this information can then be used in the assessment of future conservation priorities.

Study Sites

The majority of our study was conducted by a team of researchers and volunteers in Ampombofofo in the Bobaomby region (Fig. 1), approximately 20 km north of the town of Antsiranana (Diego Suarez), the, the administrative capital of the Antsiranana province. The. The


Table 2. Location and description of pitfall lines used to capture amphibians and reptiles at Ampombofofo 2006-2007 (capture rate % is number of amphibians and reptiles caught per trap day).

Season Line Habitat Microhabitat Latitude (S)

Longitude(W)

Altitude (m)

Startdate

Finishdate

Days Trapdays

Capturerate %

Dry 1 1 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 18/10/2006 28/10/2006 11 121 3.31 Dry 1 2 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 18/10/2006 28/10/2006 11 121 0.83 Dry 1 3 Forest Primary Forest 12°05’58’’ 49°20’19’’ 19 18/10/2006 28/10/2006 11 121 1.65 Dry 1 4 Forest Steep hillside 12°03’12” 49°18’36” 24 01/11/2006 11/11/2006 11 121 0.83 Dry 1 5 Forest Steep hillside 12°03’11” 49°18’42” 30 01/11/2006 11/11/2006 11 121 0 Dry 1 6 Forest Steep hillside 12°03’10” 49°18’36” 58 01/11/2006 11/11/2006 11 121 0.83 Dry 1 7 Disturbed Open scrub 12°04’20” 49°18’19” 26 02/11/2006 12/11/2006 11 121 0Dry 1 8 Disturbed Open scrub 12°04’21” 49°18’24” 26 02/11/2006 12/11/2006 11 55 1.82 Dry 1 9 Forest Secondary

Forest 12°05’26’’ 49°19’50’’ 52 15/11/2006 25/11/2006 11 121 4.96

Dry 1 10 Forest Secondary Forest

12°05’26’’ 49°19’50’’ 52 15/11/2006 25/11/2006 11 121 3.31


12°05’25’’ 49°19’50’’ 50 15/11/2006 25/11/2006 11 121 3.31

Wet 1 12 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 20/01/2007 01/02/2007 11 121 5.79 Wet 1 13 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 20/01/2007 01/02/2007 11 121 8.26 Wet 1 14 Forest Primary Forest 12°05’58’’ 49°20’19’’ 19 20/01/2007 01/02/2007 11 121 20.7 Wet 1 15 Forest Secondary

Forest 12°05’26’’ 49°19’50’’ 52 02/02/2007 12/02/2007 11 121 9.09

Wet 1 16 Forest Secondary Forest

12°05’26’’ 49°19’50’’ 52 02/02/2007 12/02/2007 11 121 10.7


12°05’25’’ 49°19’50’’ 50 02/02/2007 12/02/2007 11 121 4.96

Wet 1 18 Disturbed Acacia ridge 12°05’57’’ 49°19’04’’ 81 14/02/2007 23/02/2007 10 111 17.1 Wet 1 19 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 14/02/2007 23/02/2007 10 111 15.3 Wet 1 20 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 14/02/2007 23/02/2007 10 111 9.91Wet 1 21 Disturbed Acacia ridge 12°05’57’’ 49°19’04’’ 81 09/04/2007 19/04/2007 11 121 5.79Wet 1 22 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 09/04/2007 19/04/2007 11 121 13.2 Wet 1 23 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 09/04/2007 19/04/2007 11 121 5.79Wet 2 24 Forest Secondary

Forest 12°05’26’’ 49°19’50’’ 52 15/05/2007 25/05/2007 11 121 5.79


12°05’26’’ 49°19’50’’ 52 15/05/2007 25/05/2007 11 121 2.48


12°05’25’’ 49°19’50’’ 50 15/05/2007 25/05/2007 11 121 1.65


12°05’19’’ 49°20’04’’ 38 21/04/2007 01/05/2007 11 121 1.65


12°05’19’’ 49°20’03’’ 36 21/04/2007 01/05/2007 11 121 6.61


12°05’19’’ 49°20’03’’ 35 21/04/2007 01/05/2007 11 121 4.96

Wet 2 30 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 04/05/2007 14/05/2007 11 121 2.48 Wet 2 31 Forest Primary Forest 12°05’58’’ 49°20’20’’ 19 04/05/2007 14/05/2007 11 121 1.65 Wet 2 32 Forest Primary Forest 12°05’58’’ 49°20’19’’ 19 04/05/2007 14/05/2007 11 121 0.83 Wet 2 33 Forest Secondary

Forest 12°05’23’’ 49°19’19’’ 28 28/05/2007 07/05/2007 11 121 4.13

Wet 2 34 Disturbed Open scrub 12°05’23’’ 49°19’19’’ 26 28/05/2007 07/05/2007 11 121 0Wet 2 35 Disturbed Open scrub 12°05’22’’ 49°19’19’’ 26 28/05/2007 07/05/2007 11 121 0.83 Dry 2 36 Forest Secondary

Forest 12°05’26’’ 49°19’50’’ 52 05/08/2007 15/08/2007 11 121 4.13


12°05’26’’ 49°19’50’’ 52 05/08/2007 15/08/2007 11 121 0.83


12°05’25’’ 49°19’50’’ 50 05/08/2007 15/08/2007 11 121 3.31

Dry 2 39 Disturbed Acacia ridge 12°05’57’’ 49°19’04’’ 81 17/08/2007 27/08/2007 11 121 1.65Dry 2 40 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 17/08/2007 27/08/2007 11 121 0.83Dry 2 41 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 17/08/2007 27/08/2007 11 121 0Dry 2 42 Disturbed Acacia ridge 12°05’57’’ 49°19’04’’ 81 11/10/2007 21/10/2007 11 121 1.65Dry 2 43 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 11/10/2007 21/10/2007 11 121 2.48Dry 2 44 Disturbed Acacia ridge 12°05’57’’ 49°19’03’’ 81 11/10/2007 21/10/2007 11 121 6.61Dry 2 45 Forest Secondary

Forest 12°05’26’’ 49°19’50’’ 52 26/11/2007 06/12/2007 11 121 2.48

Amphibians and reptiles of Northern Madagascar 35

Table 3. Amphibians and reptiles caught in the pitfall traps at Ampombofofo 2006-2007.

Pif

all l

ine

num

ber

Agl

ypto

dact

ylus

sec

urif

er

Lal

iost

oma

labr

osum

Stum

pffi

a sp

. 1

Man

tella

vir

idis

Tot

al A

mph

ibia

Am

phig

loss

us a

rdou

ini

Mad

asci

ncus

cf.

inte

rmed

ius1

/2*

Bro

okes

ia s

tum

pffi

Fur

cife

r pe

tteri

Bla

esod

acty

lus

boiv

ini

Gec

kole

pis

cf. m

acul

ata

Lyg

odac

tylu

s he

teru

rus

Par

oedu

ra s

tum

pffi

Tra

chyl

epis

ele

gans

Tra

chyl

epis

tava

ratr

a

Typ

hlop

s m

icro

ceph

alus

Typ

hlop

s sp

. 2

Ram

phot

yphl

ops

bram

inus

Lio

phid

ium

torq

uatu

m

Lio

phid

ium

ther

ezie

ni

Mim

ophi

s m

ahfa

lens

is

Pse

udox

yrho

pus

quin

quel

inea

tus

Tot

al R

epti

lia

Tot

al s

peci

men

s

1 0 1 2 1 4 4

2 0 1 1 1

3 0 1 1 2 2

4 0 1 1 1

5 0 0 0

6 0 1 1 1

7 0 0 0

8 0 1 1 1

9 0 1 3 2 6 6

10 0 1 3 4 4

11 0 1 1 1 1 4 4

12 2 2 1 2 1 1 5 7

13 7 7 2 1 3 10

14 2 3 1 6 12 1 1 1 1 1 2 19 25

15 2 2 4 4 1 9 11

16 0 10 1 1 1 13 13

17 0 3 1 1 1 6 6

18 0 19 19 19

19 0 1 16 17 17

20 0 11 11 11

21 0 7 7 7

22 5 5 9 1 1 11 16

23 0 6 1 7 7

24 0 2 4 1 7 7

25 0 2 1 3 3

26 0 2 2 2

27 0 1 1 2 2

28 0 8 8 8

29 0 1 5 6 6

30 2 2 1 1 3

31 2 2 0 2

32 0 1 1 1

33 0 4 1 5 5

34 0 0 0

35 0 1 1 1

36 0 5 5 5

37 0 1 1 1

38 0 1 3 4 4

39 0 2 2 2

40 0 1 1 1

41 0 0 0

42 0 2 2 2

43 0 2 1 3 3

44 0 8 8 8

45 0 1 2 3 3

Total 2 14 3 7 26 1 128 1 2 2 1 1 4 57 6 3 1 2 4 1 1 1 216 242

close proximity to the coast and the sandstone geology has created a unique habitat matrix of large patches of primary western dry forest fringed by primary coastal forest. In addition, anthropogenically disturbed areas of habitat are also present throughout this region (Fig. 2 provides photographic documentation of the main habitat types surveyed during this study). Our study mainly concentrated on an area of forest surrounding the village of Ampombofofo with researchers operating from a base camp situated at 12°05’53’’S, 49°19’49’’E. Fieldwork in this region consisted of five survey periods,

which were conducted during both wet and dry seasons over a period of approximately 15 months. Each survey period lasted approximately 9 weeks in length and can be summarised as follows: Survey 1: 9 October to 14 December 2006; Survey 2: 4 January to 8 March 2007; Survey 3: 3 April to 13 June 2007; Survey 4: 4 July to 15 September 2007; and Survey 5: 8 October to 13 December 2007. Fieldwork was conducted throughout the full altitudinal range (0-100 m a.s.l.) within all habitat types.

A second team conducted fieldwork in the Andavakoera region of Montagne des Français (Fig. 1) (12°19’78’’S, 49°22’05’’E, 140 m a.s.l.), in January and February 2006 and again between 16 to 25 February 2008. The area has already been subject to a detailed survey effort (D’Cruze et al., 2007). The vegetation within the calcareous massif of Montagne des Français has been described as transitional mid-transitional mid-altitude rainforest and dry deciduous western forest (Ramanamanjato et al., 1999) (Fig. 2). A third teamA third team conducted fieldwork in an area of coastal scrub forest located within the area known as Orangea (Figures 1 and 2) (12°16’34’’S, 49°23’24’’E, ca. 25 m a.s.l.), in 22 February to 5 March 2007 and 19 to 28 February 2008.

All of the study sites are part of the Western Ecoregion defined by Cornet (1974), and the N Bemarivo centre of endemism identified by Wilmé et al. (2006). The region is characterized by a reliable wet season that runs from November to March during which the majority of rainfall occurs. This period is followed by a pronounced dry season that can last up to seven months during which rainfall is highly infrequent. However the mean annual precipitation of Montagne des Français is believed to be slightly higher than the 980 mm received by Antsiranana (Nicoll and Langrand, 1989).

Materials and MethodsDue to the lack of field information regarding the herpetofauna

of this location we used a wide range of sampling methods in order to collect data for as many species as possible. We also made a concerted effort to identify anthropogenic activities that represent threats to the conservation of this area. We used three 100 m pitfall trap-lines with drift fences as described in Raxworthy and Nussbaum (1994), which were left in place for a total of seven - 11 days. We conducted opportunistic searches (D’Cruze et al., 2007) across the full range of altitudes and habitats – night and day searching – in order to reveal the presence of species not captured by the other methods. We also made direct counts along line transects to identify species and assess their distribution and abundance. We specifically targeted calling anurans after rainfall.

A total of 45 trap lines were constructed at Ampombofofo during the survey period, the details of which are shown in table 2. A total of four trap lines were constructed in Orangea and six


Species IUCN Status CITES listing Regionally endemic

Amphibia

Ranidae sensu lato

Hoplobatrachus tigerinus LC II

Ptychadena mascareniensis LC

Hyperoliidae

Heterixalus andrakata LC

Reptilia

Chamaeleonidae

Furcifer oustaleti II

Furcifer pardalis II

Furcifer petteri II

Gekkonidae

Geckolepis sp. +

Hemidactylus mercatorius

Paroedura stumpffi

Phelsuma abbotti II

Uroplatus sp. nov. aff. henkeli II

Scincidae

Cryptoblepharus boutonii cognatus

Cryptoscincus minimus

Madascincus cf. intermedius 1 +

Madascincus cf. intermedius 2

Paracontias rothschildi +

Paracontias sp. nov. +

Trachylepis elegans


Dromicodryas quadrilineatus

Leioheterodon madagascariensis

Leioheterodon modestus

Liophidium therezieni +

Liophidium torquatum

Mimophis mahfalensis

Typhlopidae

Xenotyphlops grandidieri +

Table 4. Amphibian and reptile species found during surveys of Orangea in northern Madagascar 2007/2008 and their threat and conservation status.

in Montagne des Français (Tab. 5). We classified the species encountered during this survey using a system similar to that used by D’Cruze and Sabel (2005) that can be summarised as follows; Abundant (species encountered in large numbers regularly all year round), common (encountered on a regular basis all year round), infrequent (unpredictable and few individuals encountered), rare (rarely encountered), seasonally abundant (encountered in large numbers on a regular basis during a certain time of year but otherwise infrequent or rarely seen), or seasonally common (encountered on a regular basis during certain times of year but otherwise infrequent). These classifications are based on data collected using all survey techniques and refer to the total number of individuals encountered for each species. Finally, we also estimated the altitudinal range for all of the species currently recorded from the survey sites. We used the minimum and maximum elevations [combined data of our surveys and that of Raxworthy and Nussbaum (1994)] to calculate the elevational range of each species. This method makes the assumption that each species is distributed continuously through the minimum and maximum elevation recorded.

The following information was recorded for each collected specimen: date, latitude, longitude, habitat, microhabitat and altitude. We also made colour notes and took photographs. We took voucher specimens (preserved in 70% ethanol) and tissue samples for all voucher specimens. Reptile and amphibian specimens were deposited at the Université d’Antananarivo, Département de Biologie Animale (UADBA) and Zoologische Staatssammlung München (ZSM). A list of specimens can be found in Appendix I.

Results

Ampombofofo

A complete species list is given in table 1 along with the current CITES listing (CITES Species Database, Geneva, Switzerland) and IUCN status (IUCN Red List of Threatened Species 2008, Species Survival Commission, Gland, Switzerland) for each of the relevant species. Table 1 also includes ecological data for each species including habitat type (classified as either forest or anthropogenically disturbed habitat), ecological distribution and relative abundance.

During this survey, we recorded a total of nine amphibian and 46 reptile species in Ampombofofo region, giving a total herpetofaunal diversity of 55 species. It can now be assumed that the herpetofauna of the Ampombofofo region consists of at least one chelonian, 25 lizards, 20 snakes and nine species of anurans. We classify a total of 12 species (22%) as abundant, 13 species (24%) as common, 15 species (27%) as infrequent, 11 species (20%) as rare, two species (4%) as seasonably abundant and two species (4%) as seasonally common. Of the 55 species encountered we found 26 (47%) to be present during wet and dry season

sample periods, two species (4%) only in the dry season and eight species (15%) only during the wet season.

A total of 242 amphibians and reptiles were caught in pitfall traps, the details of which can be seen in table 3. These individuals were captured over 5349 trap days, giving an overall capture rate of 4.5% per trap day. The pitfall traps caught four amphibian, 11 lizard and seven snake species. Two fossorial species (Typhlops cf. microcephalus and Typhlops sp. 1) were only caught using pitfall traps. At leastAt least one amphibian and one reptile species represent what appear to be undescribed species: Stumpffia sp. 1, and Typhlops sp. 1. Of the 55 species encountered during this survey, one species, Cryptoblepharus boutonii cognatus, was not found throughout the entire altitudinal range of the Ampombofofo region.

With regards to primary habitat in the Ampombofofo region three species (6%) were found only in anthropogenically disturbed non-forest habitat, 24 species (44%) were found solely within relatively undisturbed forest habitat. 27 (49%) occurred in both undisturbed and disturbed habitat. One species (2%) , Cryptoblepharus boutonii cognatus, was only observed on rocks along the coast. 21 (38%) species were found solely in terrestrial situations, nine (16%) only in arboreal situations and 16 (29%) in both. Five species (9%) were found to be occurring in both terrestrial and semi aquatic habitat, three (6%) species solely in semi aquatic environment and one species (2%) occurred in all three habitats, terrestrial, semi aquatic and arboreal situations.

The majority of the species found in the Ampombofofo region are endemic to Madagascar (89%). Ptychadena mascareniensis and Leioheterodon madagascariensis are found on other Indian Ocean Islands. Pelusios castanoides occurs on the African mainland, Hoplobatrachus tigerinus is native to India, Cryptoblepharus boutonii occurs on many Indian Ocean Islands, the Australian region and African mainland, Hemidactylus frenatus has a wide global distribution and Hemidactylus mercatorius occurs on the African mainland and other Indian Ocean islands. We recorded 21 species (38%) that are regional endemics restricted to only a few places in north Madagascar (Table 1). One species is listed as endangered, two species are listed as vulnerable but need updating and seven species are listed as least concern on the 2007 IUCN Red List of Threatened Species. A total of 12 species (22%) are listed on the CITES appendices.


Orangea

A total of 34 reptile specimens were captured in the pitfall traps over a total of 381 trap days, giving an overall capture rate of 8.9%. The pitfalls captured five lizard and two snake species. During the survey a total of three amphibian and 22 reptile species were recorded in Orangea providing a total of 25 species. It can now be assumed that the herpetofauna of Orangea consists of at least three anurans, 15 lizards and seven snake species (Table 4).Twentytwo (88%) of the species encountered are endemic to Madagascar and six species (24%) are regionally endemic. Three species (12%) appear as Least Concern on the IUCN red list of endangered species and six (24%) appear in the CITES Appendices (Table 4).

Montagne des Français

A total of 23 amphibians and reptiles were captured in the pitfall traps. The 23 individuals were captured over 660 trap days resulting in a capture rate of 4%. The pitfalls set up in forest habitat yielded an average capture rate of 6%, where as the pitfalls placed in more anthropogenically disturbed areas only yielded an average capture rate of 1%.

At Montagne des Français a total of eight amphibian and 29 reptile species were recorded during the survey period. The majority of species found (95%) are endemic to Madagascar, with Leioheterodon madagascariensis occurring on other Indian Ocean Islands and Gehyra mutilata occurring widely across Southeast Asia. Of the species recorded, 19 (51%) are regionally endemic. Six (16%) of the species recorded were previously unknown

for the Massif: Cophylinae gen. sp., Gehyra mutilata, Leioheterodon modestus, Paracontias hildebrandti, Stumpffia sp. 3 and Stumpffia sp. 4. Three of which (8%) are locally endemic; Cophylinae gen. sp., Stumpffia sp. 3 and Stumpffia sp. 4. These new records bring the total number of herpetofaunal species in the area to 67.Two species (5%) appear as least concern on the IUCN red list of endangered species: Aglyptodactylus securifer, Mantidactylus bellyi and one species (3%) appears as Endangered: Mantella viridis. Seven species (19%) appear in the CITES appendices; Mantella viridis, Brookesia ebenaui, Brookesia sp. nov., Brookesia stumpffi, Furcifer oustaleti, Furcifer pardalis, Furcifer petteri and Uroplatus sp. nov. aff. henkeli.

Discussion

Pitfall Trapping

The pitfall traps set up in forest habitat of Montagne des Français yielded a higher average capture rate then those deployed in the more anthropogenically disturbed areas. It is difficult to determine the specific causal factors responsible for this from the results of our particular study. However, it seems that our data supports the contention that anthropogenic habitat alteration (loss, degradation or fragmentation) can have a strong negative influence on tropical herpetofaunal communities (Vallan, 2003). We recommend that more research should be conducted out to investigate this further. Unfortunately we can not directly compare pitfall trapping between the various study sites because this survey method was not kept constant e.g. pitfall trapping differed greatly in terms of duration and seasonal implementation.


Site,Line

Season Habitat Microhabitat Latitude Longitude Altitude (m a.s.l.)

Startdate

Finishdate

Days Trapdays

Capturerate %

O1 Wet 1 Forest Coastal scrub forest

12°14'43’’ 49°22'53’’ 14 22/02/2007 05/03/2007 12 120 1.6


12°16’34’’ 49°23’24’’ 25 19/02/2008 28/02/2008 10 74 18.9


12°16’37’’ 49°23’34’’ 28 19/02/2008 28/02/2008 10 110 10


12°16’32’’ 49°23’28’’ 27 20/02/2008 26/02/2008 7 77 9.1

MDF1 Wet 3 Forest Valley 12°20’24’’ 49°21’27’’ 170 16/02/2008 25/02/2008 10 110 7.27

MDF2 Wet 3 Forest Slight hillside 12°20’27’’ 49°21’23’’ 185 16/02/2008 25/02/2008 10 110 5.45

MDF3 Wet 3 Forest Slight hillside 12°20’27’’ 49°21’18’’ 185 16/02/2008 25/02/2008 10 110 6.36

MDF4 Wet 3 Disturbed Canyon 12°19’59’' 49°20’60'’ 232 16/02/2008 25/02/2008 10 110 0.91

MDF5 Wet 3 Disturbed Canyon 12°19’64'’ 49°20’59'’ 262 16/02/2008 25/02/2008 10 110 1.81

MDF6 Wet 3 Disturbed Canyon 12°19’63'’ 49°20’53’' 288 16/02/2008 25/02/2008 10 110 0.91

Table 5. Location and description of pitfall lines used to capture amphibians and reptiles at Orangea (O1-O4) and Montagne des Français (MDF1- MDF6) 2007-2008 (capture rate % is number of amphibians and reptiles caught per trap day).

Range extensions: Ampombofofo

All of the species discovered in this survey are new records for Ampombofofo. It is important to note that for all of these species this locality data represent the northernmost limit of their known range. Many species encountered during this study require special mention as their occurrence in Ampombofofo significantly contributes to the current information regarding their distribution in Madagascar.

Until recently, Mantella viridis was considered as critically endangered (now changed to endangered by Andreone et al., 2008a) and was thought to be restricted to only a few localities in the extreme north of Madagascar (D’Cruze et al., 2007; Glaw and Vences, 2007). However a recent study conducted by Mercurio and Andreone (2008) revealed the presence of this species in a total of 42 different localities. Our study revealed that M. viridis was locally abundant in the Ampombofofo region, which supports the suggestion that the status of this species needs to be re-evaluated (Mecurio and Andreone, 2008).

The presence of Liophidium therezieni at this locality is noteworthy because (although its conservation status has yet to be assessed) prior to this survey this terrestrial snake was only known from two locations; Montagne des Français and Anatelo (Franzen et al., in press). Its presence in the Ampombofofo region and at Orangea represents only the third and fourth localities for this species. Our findings indicate that it is a relatively adaptable species that is not restricted to limestone habitat. The specimen ZSM 1612/2008 was found at 11:00 approximately 30-40 cm under the ground when digging holes for the construction of basecamp. The individual was unearthed in an area of secondary western dry forest (12°05’52”S, 049°19’48”E). During the survey in Ampombofofo six other individuals were recorded. Individuals were encountered throughout a variety of different habitat types. L. therezieni was observed both on the steep slopes of the primary western dry forest at the Pteropus rufus roost and in the primary western dry forest surrounding Ampombofofo, and it was also recorded in riparian habitat. At night individuals were observed active amongst leaf litter in mango plantations edging the river. During the survey individuals were also recorded in more degraded habitat, as well as in the coastal scrub forest amongst thin leaf litter coverage on sandy soil, also in open scrubland that had once been rice padis or had been cleared for zebu grazing. Although additional specimens were not taken, morphological data for these snakes was recorded. This study also provides an important additional locality for

Amphiglossus ardouini. The genus Amphiglossus has a wide distribution and inhabits most of Madagascar’s environments. However, most species dwell in the leaf litter and are often very secretive (Glaw and Vences, 2007). As a result both the habits and distribution of A. ardouini are largely unknown. One specimen was discovered on 11 November 2006 in an area of secondary western dry forest (12°05’52”S, 049°19’48”E) as it was being consumed by a Dromicodryas quadrilineatus. Another individual was captured in a pitfall trap on 23 February 2007. The specimen was discovered early in the morning indicating nocturnal or crepuscular activity. The trap site was situated in an Acacia forest on sandy soil (12°05’57”S, 049°19’03”E). Prior to this survey this species was known from three localities only: Antsiranana, Ambilobe and Montagne des Français (Glaw and Vences, 2007).

New records: Montagne des Français

The herpetofauna of Montagne des Français [an area of herpetological importance that has been nominated as a Durban Vision Potential Site requiring some form of protection (Ministère de l’Environment, des Eaux et Forets, 2005) and granted Temporary Protected Status] has been recently surveyed intensively by a team of researchers and volunteers (D’Cruze et al., 2007). It is therefore remarkable that we discovered several species in this limestone massif during our relatively short surveys that were not previously known from this site: Reptiles include Gehyra mutilata, Leioheterodon modestus, and Paracontias hildebrandti. Amphibians include Cophylinae gen. sp., Stumpffia sp. 3 and Stumpffia sp. 4. These findings indicate that even intensive surveys, conducted by numerous researchers using different methods may often fail to obtain a complete herpetofaunal species inventory. To address this problem researchers have suggested: (1); the implementation of additional biodiversity and monitoring surveys; (2) increased collaboration with Malagasy researchers and self-funded volunteers; (3) application of molecular barcoding approaches to identify true species richness; and (4) the utilisation of approaches that speed up data availability via appropriate cyber infrastructure (D’Cruze et al., in press; Vences et al., 2008).

Relevance to patterns of biodiversity in northern Madagascar

Natural biogeographical processes are believed to be largely responsible for the current distribution of


amphibians and reptiles in Madagascar (Wilmé et al., 2006). However the disruptive anthropogenic activity that followed human invasion approximately 2000 years ago is also believed to have played a pivotal role in shaping patterns of distribution (Vallan, 2003). For example it has been suggested that prior to human invasion continuous lowland corridors of dry or transitional forest linked the lower slopes of the five major massifs of Analamera, Ankarana, Daraina, Montagne D’Ambre and Montagne des Français located in the north (Ramanamajato et al., 1999; D’Cruze et al., 2006, in press).

We suggest that prior to human invasion the forests of the Bobaomby region and Montagne des Français may have been linked via the ‘Bobaomby spine’ region (the area around Windsor Castle) (Figure 1). It is also possible that this connected area of forest may have also stretched to include Orangea. These former corridors would help to explain how these species were able to cross the distance between these sites. It is clear that to better understandIt is clear that to better understand the biogeographical patterns of the herpetofauna in the north of Madagascar more data regarding their exact distribution and phylogenetic relationships must be gathered (Vences et al., 2004; Boumans et al., 2007).

Following an extensive herpetological survey (D’Cruze et al., 2007), Montagne des Français was until recently considered as an important biological center ofimportant biological center of herpetological endemism. At least eight species were considered to be locally endemic to this calcareous massif (D’Cruze et al., 2007; Glaw and Vences, 2007).(D’Cruze et al., 2007; Glaw and Vences, 2007).. However, our results of surveys in surrounding areas have now demonstrated that many of these species are also present outside of this massif and are in fact regionally endemic to the north of Madagascar. Examples are Thamnosophis martae which has been recorded within the Forêt d’Ambre Special Reserve (D’Cruze et al., 2008) and Heteroliodon fohy and Liophidium therezienitherezieni that have been recorded from within the AmpombofofoAmpombofofo Region (Megson et al., in press; Franzen et al., in press).

This highlights both the importance and the necessity for increased herpetological survey effort in Madagascar as this type of baseline information is needed tobaseline information is needed to facilitate informed decisions for existing conservation programs and guide the course of future management strategies (Andreone et al., 2008b; Kremen et al., 2008).

Conservation Implications

Socio-economic factors such as rapid population growth, poor education and other particular aspects of Malagasy culture are partly responsible for obliging local residents to employ harmful agricultural methods

and other activities that lead to the exploitation of natural resources (Durbin et al., 2003) which can be a serious threat to amphibians and reptiles (Glaw and Vences, 2007). As a result of its close proximity to the administrative capital of the Antsiranana province (ca. 30 km) and neighbouring communes such as the villages of Ampombofofo and Ramena, the unique herpetofauna found in our study sites located in the extreme north Madagascar are vulnerable to these anthropogenic pressures. The major threats to the integrity of the herpetofauna of these areas are: (1) agricultural clearance for maize and rice cultivation; (2) charcoal production; (3) timber production; and (4) zebu grazing (during which sites are either selectively logged or cleared of all trees) (D’Cruze et al., 2007). All of these threats have resulted in the degradation or clearance of large areas of forest in all three of our study sites. Continued forest clearance will lead to the eventual fragmentation of the remaining areas of forest (with serious consequences for the herpetofauna; e.g. Vallan, 2000; Vallan, 2003). This will be followed by local extirpation, and possibly extinctions, that will in turn place increased pressure on the existing system of protected areas in the north of Madagascar.

Due to its remote location Ampombofofo remains far less affected by the rapid population growth compared to Montagne des Français or Orangea. The main pressure on the forest is anthropogenic but may be linked to climate change. After meetings with members of the local village it was explained that they had noticed a significant drop in rainfall and in the height of the river water within the last 15 years particularly. It is believed in the village that the reduction in rainfall is punishment from the Ancestors for the younger members of the village disobeying the local taboos (Fadis). It was explained that this lack of available water meant that once successful padi fields were simply too dry to yield an adequate crop. This lack of appropriate rice padis has forced the village to find alternate ways to generate income, e.g. charcoal production and the clearing of land to encourage breeding of Zebu, which are sold in Antsiranana to purchase rice. It was also explained that since charcoal production and the clearing of land for zebu grazing has become more frequent, ‘soil is disappearing’.

Future action

The reptile and amphibian rich fauna found in the north of Madagascar is already recognized as a highly speciose region of the biodiversity hotspot that is Madagascar (D’Cruze et al., 2008). To safeguard this diversity we


suggest that a detailed conservation management plan should be developed for the extreme north of Madagascar. As part of this plan we strongly recommend that Montagne des Français should be granted full-protected status so that it is able to function as a biological refuge for the threatened species that increasingly seem to be endemic to the extreme north of the island. In addition increased conservation management action is required to protect the biological diversity found within the existing protected areas found in the north of the country (D’Cruze et al., 2008). Furthermore habitat restoration activity should be encouraged to improve connectivity with Orangea and the Bobaomby Region so as to address habitat fragmentation. In order to implement this in a sustainable and holistic manner we make the following additional conservation recommendations:

• Further assessment and monitoring of natural resource use activities. In particular, clearance of forest needs to be restricted and the remaining areas must be carefully monitored (e.g. by remote sensing using satellite data). \

• Development and implementation of a large-scale, regionally connected, more sustainable community-focused management system, which includes payments for ecosystem services, including biodiversity conservation and carbon sequestration.

• Development efforts, to promote sustainable agriculture practices and to improve human conditions. Village-based education programmes aimed at demonstrating alternative energy systems and crops, which are compatible with local traditions.

• Raising awareness about environmental problems. Village-based programmes targeting all socioeconomic groups.

Acknowledgements. We would like to thank Sally Eaton, Amanda Martin, Edmond Randriamalala, Robert Devas, Adam Mushi, Stuart Brooker, The Village of Ampombofofo and all of the Frontier staff and volunteers in 2006-2007. Furthermore, Parfait Bora, Hildegard Enting, Angelika Knoll, and Zoltan Nagy assisted during fieldwork in 2007 and 2008. The work was carried out in collaboration with the Département de Biologie Animale, Université d’Antananarivo, and the Association Nationale pour la Gestion des Aires Protégées (ANGAP). We are grateful to the Malagasy authorities, in particular the Ministère de l’Environnement et des Eaux et Forêts, for research and export permits.

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Appendix 1. Voucher specimens from Ampombofofo, Orangea and Montagne des Français.

Ampombofofo

Amphibia

Mantellidae

Aglyptodactylus securifer ZSM 1637/2008 ZSM 1636/2008, Blommersia cf. wittei UADBA 1937/2008, Boophis tephraeomystax ZSM 1645/2008, Laliostma labrosum UADBA 1955/2008, Mantella viridis FGZC 1948.

Microhylidae

Stumpffia sp.1 ZSM 1661/2008 ZSM 1663/2008, Stumpffia sp. 2 UADBA 1959

Reptilia

Chamaeleonidae

Brookesia stumpffi ZSM 1507/2008.

Gekkonidae

Blaesodactylus boivini ZSM 1519/2008, Ebenavia inunguis ZSM 1535/2008, Geckolepis cf. maculata ZSM 1521/2008, Hemidactylus frenatus UADBA 1929/2008, Lygodactylus heterurus ZSM 1543/2008 ZSM 1544/2008 ZSM 1545/2008, Paroedura cf. karstophila ZSM 1531/2008, Paroedura stumpffi UADBA 1936/2008, Uroplatus sp. nov. aff. henkeli ZSM 1924/2008.

Scincidae

Amphiglossus ardouini ZSM 1555/2008, Cryptoscincus minimus ZSM 1583/2008, Madascincus cf. intermedius 1 ZSM 1564/2008, Madascincus cf. intermedius 2 ZSM 1570/2008, Trachylepis cf. tavaratra ZSM 1552/2008 ZSM 1553/2008.


Heteroliodon fohy ZSM 1624/2008, Langaha madagascariensis ZSM 1722/2008, Liophidium therezieni ZSM 1612/2008, Liophidium torquatum ZSM 1616/2008, Pseudoxyrhopus cf. quinquelineatus ZSM 1609/2008, Stenophis granuliceps ZSM 1604/2008, Stenophis inopinae UADBA 1720/2008, Stenophis variabilis ZSM 1606/2008 ZSM 1607/2008.

Typhlopidae

Typhlops microcephalus ZSM 1588/2008, Typhlops sp. 2 ZSM 1592/2008.

Montagne des Français

Amphibia

Mantellidae

Mantella viridis FGZC 1761

Microhylidae

Stumpffia sp.2 ZSM 1671/2008, Stumpffia sp.3 ZSM 1672/2008, Stumpffia sp.4 ZSM 1667/2008, Cophylinae gen. sp. ZSM 1674/2008

Ranidae sensu lato

Ptychadena mascareniensis ZSM 2066/2007

Reptilia

Chamaeleonidae

Brookesia ebenaui ZSM 2110/2007, Brookesia sp. nov. ZSM 1505/2008 ZSM 2146/2007 ZSM 2147/2007 ZSM 2148/2007 ZSM 2149/2007

Gekkonidae

Geckolepis sp. ZSM 1523/2008 ZSM 2112/2007 Gehyra mutilata ZSM 1536/2008, Lygodactylus heterurus ZSM 1546/2008 Lygodactylus sp. ZSM 1547/2008, Paroedura cf. karstophila ZSM 2106/2007 ZSM 2107/2007, Paroedura stumpffi ZSM 1525/2008, Uroplatus sp. nov. aff. henkeli ZSM 2111/2007.

Scincidae

Amphiglossus ardouini ZSM 1554/2008, Amphiglossus sp. nov. 1 ZSM 1556/2008, Madascincus cf. intermedius 2 ZSM 1573/2008 ZSM 1574/2008, Madascincus cf. polleni ZSM 1559/2008 ZSM 1560/2008 ZSM 1561/2008, Paracontias hildebrandti ZSM 1578/2008,


Alluaudina bellyi ZSM 1627/2008, Ithycyphus miniatus ZSM 1625/2008, Langaha madagascariensis ZSM 1621/2008 ZSM 2133/2007, Liophidium therezieni ZSM 2053/2007, Liophidium torquatum ZSM 1618/2008, Madagascarophis colubrinus ZSM 1620/2008, Madagascarophis sp. nov. ZSM 2130/2007, Mimophis mahfalensis ZSM 2101/2007, Pseudoxyrhopus cf. quinquelineatus ZSM 1610/2008, Thamnosophis martae ZSM 1595/2008 ZSM 1596/2008.

Typhlopidae

Typhlops microcephalus ZSM 1587/2008.


Orangea

Reptilia

Chamaeleonidae

Furcifer petteri ZSM 2136/2007 ZSM 2137/2007 ZSM 2141/2007 ZSM 2142/2007.

Gekkonidae

Geckolepis sp. ZSM 2138/2007, Hemidactylus mercatorius ZSM 2140/2007, Paroedura stumpffi ZSM 2139/2007, Uroplatus sp. nov. aff. henkeli ZSM 2134/2007 ZSM 2192/2007.

Scincidae

Cryptoblepharus boutonii cognatus ZSM 2077/2007, Cryptoscincus minimus ZSM 1585/2008 ZSM 1586/2008 ZSM 1584/2008 ZSM 2248/2007 ZSM 2249/2007, Madascincus cf. intermedius 1 ZSM 1565/2008 ZSM 1566/2008, Paracontias rothschildi ZSM 1580/2008 ZSM 1581/2008 ZSM 1582/2008, Paracontias sp. nov. ZSM 2256/2007, Trachylepis elegans ZSM 1548/2008 ZSM 2127/2007.


Liophidium therezieni ZSM 1613/2008, Liophidium torquatum ZSM 2129/2007,

Typhlodiae

Xenotyphlops grandidieri ZSM 1594/2008 ZSM 2075/2007 ZSM 2125/2007 ZSM 2193/2007.

Accepted by Angelica Crottini; Managing Editor: Zoltan T. Nagy


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