Madagascar Forest Conservation Programme Phase 181Science ... · entire animal groups, such as...

MGF 181 Ryan. M. Clark and Matthew Smart

Madagascar Forest Conservation Programme

Phase 181Science Report

Madagascar, Nosey Be

January–March 2018

Principal Investigator: Ryan M. Clark

Assistant Research Officer: Matthew Smart


Table of Contents

1. GENERAL INTRODUCTION ..................................................................................... 3

1.1 Madagascar................................................................................................................ 3

1.2 Measuring Biodiversity ................................................................................................... 5

2. STUDY AREA .............................................................................................................. 6

2.1 ‘Big Island’ ................................................................................................................ 6

2.2 Survey Routes ........................................................................................................... 7

3. PROJECTS .................................................................................................................. 13

3.1 Lemur Abundance ................................................................................................... 13

3.2 Herpetofauna abundance and species richness ........................................................ 15

4. Summary ..................................................................................................................... 17

REFERENCES .................................................................................................................... 19


1. GENERAL INTRODUCTION

1.1 Madagascar

“Continental Island of the first rank”—Wallace (1892, p.563).

The majority of extant flora and fauna on Madagascar are found nowhere else on earth

(Goodman & Benstead 2003; Hobbes & Dolan 2008; Phillipson et al. 2006). Many of the

island’s unique species are nested in ancient clades that host entirely native genera and families

(e.g. Callmander et al. 2011), leading some to claim that higher taxa endemism leaves

Madagascar peerless in biological uniqueness (Ganzhorn et al. 2014).

Modern estimates hold that over 90 percent of Madagascar’s endemic animals live in, or rely on

forests (Dufils 2003). Although our understanding of the historic relationship between

Madagascar’s human population and the island’s forests is mired in conflicting interpretations

and innuendo (Kline 2002), humans have undoubtedly shaped significant aspects of

Madagascar’s modern vegetative cover (Randrianja & Ellis 2009). Forest clearance for

agriculture using methods such as ‘tavy’ (slash and burn) and oxen grazing are central to

Malagasy culture, widely practiced, and thought to reduce biodiversity (Clark 2012; Gade 1996;

Marcus 2001).

The above factors, exacerbated by stochastic socioeconomic climes across Madagascar have

largely contributed to the island being classified as a biodiversity ‘hotspot’ (Myres et al. 2000)

and being considered a global conservation priority (Goodman & Benstead 2005; Mittermeier et

al. 2005).

1.1.1 Geologic Origins

Much of Madagascar’s extraordinary biological distinctness is thought to have arisen due to its

isolation from other landmasses, which has lasted at least 90 million years (Storey et al. 1995).

Compounded by the Cretaceous–Tertiary extinction event (K–T) ~65 million years ago (mya)

(Alvarez et al. 1980), geographic isolation has allowed Madagascar’s taxa to radiate into

previously impoverished niche space (Ali & Krause 2011; Fortey 1999; Renne et al. 2013).

Throughout the latter half of the 20th century, the role of dispersal in the post-Mesozoic faunal

colonisation of Madagascar was largely ignored in favour of a vicariance dominated narrative

(Yoder & Nowak 2006). The first years of the 21st century have seen this status quo reversed;

current thinking is that the majority of Madagascar’s basal fauna stocks originated from


Cenozoic colonisation events from 65 million years ago onwards (Salmonds et al. 2012; 2013).

Fossil records buttress this model by revealing that most extant Malagasy vertebrate taxa were

not present during the Cretaceous (Krause et al. 1997; 1998; 1999)—hence their arrival must

have been later. However some clades are known to have survived from Madagascar’s shared

plate tectonic history with Gondwanaland (Gaffney & Forster 2003; Noonan & Chippindale

2006).

Stochastic dispersal post-K–T has given rise to peculiar taxonomic assemblages on Madagascar

(e.g. Poux et al. 2005; Reinthal & Stiassny 1991), where a handful of speciose radiations make

up the majority of fauna (Bossuyt & Milinkovitch 2001; Nagy et al. 2003; Vences 2004) and

entire animal groups, such as large mammals, are absent. However, human driven extinctions of

certain taxa during the Holocene may compound our ability to assess ‘natural’ patterns of

Madagascan biodiversity (Crowley 2010). Nevertheless, modern biological communities on

Madagascar are widely thought to be unique in structure and composition (Ganzhorn et al.

2014; Horvath et al. 2008; Reddy et al. 2012; Salmonds et al. 2012; 2013).

1.1.2 Modern Biogeography

Often cited as a pseudo-continent (de Wit 2003), Madagascar is the world’s fourth largest island

(587,000 km2), lying ~400km east of mainland Africa in the Indian Ocean. Madagascar’s

topography is dominated by an eastern massif running north-south, dividing the island into two

slopes: a steep eastern escarpment and a gentler western slope (27% and 73% of land cover

respectively (Ganzhorn et al. 2014)). This topography determines large aspects of moisture

deposition across Madagascar. Humid prevailing winds rolling in off the Indian Ocean are

forced upward by eastern mountains and unload precipitation on the east coast and central

highlands; western Madagascar is left in a seasonal rain shadow and experiences pseudo-

monsoon conditions (Jury 2003). What results is a fine-grain diversity of microclimes: discrete

abiotic pockets of different weather conditions (Dewar & Richard 2007; Rakoto-Joseph et al.

2009). These heterogeneous moisture patterns shape much of Madagascar’s local landscape

diversity. Although ~65% of the island’s land cover is savannah (Moat & Smith 2007), habitat

mosaics of scrub, wetland, and forests typify most regions (Mayaux 2000; 2004). The

centrepieces of these landscapes are scattered woodland blocks; forest archipelagos that support

the majority of Madagascar’s species (Dufils 2003).


The unique structure, composition, and history of Madagascar’s biological communities leaves

them as exquisite examples of radiation in isolation, and bolster calls for conservation of these

ecosystems.

1.2 Measuring Biodiversity

More biodiverse communities are considered to be ecologically healthier; generally having

greater stability, productivity, and resistance to disturbance (Levins 2013; Purvis & Hector

2000). Thus, measuring biodiversity can provide insights into the condition of biological

communities, track changes over space and time, and inform practical management and

conservation (Casey 1999; Gotteli & Chao 2013).

Most often biodiversity is measured in two ways: richness and evenness. Richness, often

synonymous with species richness, is recorded by a total tally of species in an area, ecosystem,

or landscape. While this method hints at biodiversity levels, the structure and composition of

wild populations is not eluded to. Evenness, describes the proportions of individuals in an area:

the more equal groups are the greater the evenness of the site. Both measures are required to

give a relatively robust estimate of biodiversity, and indeed there are many more detailed

methods to quantify more specific aspects of biodiversity.

Estimating biodiversity provides a snapshot in biological time but regular and consistent repeat

measures can provide a window onto the trends and trajectories of species in an area with

reference to external factors, such as human driven reordering of biological communities. Ergo,

at the beginning of this, our project’s new five year research plan, we build surveys around a

framework of measuring the abundance and proportions of various taxa in our research area

over time; various estimates that denote richness and evenness will be made at least once per

dry and wet season for core animal clades (lemurs, reptiles and amphibians). In exploring finer-

grained aspects of ecological dynamics we hope to expand our repertoire of surveys to include

habitat descriptions, more complete inventories of terrestrial invertebrates, and

ecomorphological investigations of certain reptiles. Further, we aim to measure different aspects

of human disturbance in our study area and to pair this data with that of our ecological

investigations. It is hoped that elucidating these relationships, and others, will not only develop

a more complete understanding of these ecosystems but also spread awareness of these forests

and support future conservation management in the area if required.


2. STUDY AREA

2.1 ‘Big Island’

The Madagascar Frontier project (MGF) is currently located on Nosy Be, Madagascar’s largest

offshore island (25, 000 ha), Nosy Be lies ~8km from the mainland and is itself part of an

offshore archipelago of several islands and many more small islets (Fig. 2.1).

Within Madagascar Nosy Be is a relatively developed area. The island’s thriving tourist

industry has fuelled the development of several large holiday resort areas that provide a large

proportion of the islands 75,000 inhabitants a sustainable income. Ecotourism on Nosy Be takes

the form of guided tours of the islands forests or seas; these are very popular as Nosy Be has a

large area of pristine woodland and has several coral reefs in its coastal waters.

The MGF study area is located at the southeast tip of Nosy Be, on a peninsula bordered by

Lokobe National Park. The small adjacent village of Ambalahonko has a population of ~100

Figure 2.1: Map showing Nosy Be, its surrounding islands, and MGF’s camp.


people that are largely a subsistence community. No road access exists for this area; hence,

small local boats are the only method of reaching our study area.

2.1.2 Sambirano

Named after a mainland watershed, the Sambirano region is a semi-distinct biogeographic

domain of which Nosy Be occupies the northern extreme locality. This region is a transitional

zone between the dry deciduous forests of western Madagascar and the humid eastern

rainforests. As such, characteristics’ of both habitats are found on Nosy Be. This transitional

arrangement defines much of the ecology in our research area.

2.2 Survey Routes

The beginning of phase 181 marked the beginning of MGF’s new five year research plan. This

plan will refocus the research methodology and aims of the project. As such a variety of new

projects, with new study methods will be implemented from this period onwards; new forest

survey routes are a major component of this enterprise.

MGF’s previous forest survey routes were separated into ‘lemur’, ‘bird’, and ‘herp’ routes and

were different (arbitrary) lengths. These methods were defenestrated at the beginning of phase

181 by the creation of nine new survey routes that are each 400 m in length and are suitable for

both lemur and herpetofauna surveys. Bird surveys will begin in phase 182 and will be based on

a random point count methodology; not ‘transects’.

As the main focus of our new five year plan will be focused on the long-term biodiversity

monitoring of lemuriforms and herpetofauna, these new routes will take centre stage in MGF’s

core study effort in phases to come. These new survey routes follow MGF’s old method of

categorising habitat along routes as ‘primary’, ‘secondary’, or ‘degraded’ forests based on past

histories of the routes, current use by human populations, and observations of ‘forest health’ by

trained research staff. Survey routes run along pre-existing forest trails to align with MGF’s ‘no

chop’ policy—thereby avoiding unnecessary deforestation and/or habitat degradation. Routes

were chosen to run as straight as possible; this (a). reduces variation in visual overlap between

observers, i.e. over/under-surveying on bends (b). simplifies routes so as to flatten the learning

curve for new researchers attempting to memorise the complex network of forest trails in our

study area.


Next we describe each new terrestrial forest survey route:

2.2.1 Degraded 1 (D1)

D1 begins at the rear of Ambalahonko (the village where our camp is based). This path is well

sheltered by the forest canopy (~5-15 m high) and later opens up into a rocky gorge-type habitat

in its final 100 m. Vegetation bordering this route while initially thick and comprised mostly of

relatively wide trees, wanes in its later section and gives way to large shrubs and more arid

biological communities. D1 ends adjacent to a small patch of agricultural land. The diameter

and condition of this path varies greatly along its length: there are many patches of thick mud,

small streams, and wide openings dispersed throughout. The prevalence of open areas of

habitat, frequent use by human populations, and proximity to agricultural land justifies this

survey route as being classified ‘degraded’.


D2 begins at the end of D1: adjacent to a small agricultural patch. From here the route follows a

medium incline over rocky terrain for ~50 m (vegetation only borders one side of the path here).

The remaining majority of the route runs a straight forest path where the canopy height rarely

exceeds 8 m. On one side this band of flora is no more than 10 m thick and overlooks a large

human altered landscape comprising of rice paddies and areas for oxen grazing (Fig. 2.4?). A

sharp 45o bend is made before the final ~30 m of the survey route; the habitat characteristics do

not change following this. Muddy areas and rocky outcrops are common along this route. The

relatively low canopy density, frequency of open areas and gaps in vegetation, and nearby

human use of land warrant that this route be classified ‘degraded’.

Figure 2.2: D1 path.



This route is the ‘healthiest’ of the degraded routes. D3 runs a straight path through forest that

backs on to several areas of farmland then curves through a flooded area and ends in an ylang-

ylang plantation. The condition of this path is poor, mud ridden patches and streams are very

common along all of this path—likely due to heavy human use of the path and surrounding area.

Despite this the forest bordering D3 is seemingly in good condition; canopy cover is relatively

dense, and canopy height relatively high along the majority of this route. Heavy human traffic

along this path and several human-altered landscapes being in the vicinity allows us to classify

this route as ‘degraded’.

2.2.4 Secondary 1 (S1)

S1 begins in close proximity to D2; as this route bends left in its final quarter S1 begins right of

this turning. S1 follows a straight path that runs slightly down hill and ends close to a coastline.

The habitat is all healthy forest excluding a ~50 m section at its terminus that is bordered by a

Figure 2.3: agricultural land adjacent to D2.

Figure 2.4: deforestation on D3.


human plantation. The forest that envelops S1 is ~10–15 m in height and canopy density rarely

falls below 20% cover. While a small area of this route is used as agricultural land, and the path

itself is used not infrequently by the local population, the relative health of the forest and

thickness of the vegetation band that borders this route indicate that it should be classified

‘secondary’.


S2 is by far in the best condition of our secondary routes. S2 begins on the edge of a forest

stream and runs up a semi-steep incline into montane tropical forest. Vegetation is very thick on

both sides of this path. Canopy cover is constantly >60%; canopy height rarely falls below 5 m.

The factor leading us to classify this route as ‘secondary’ and not ‘primary’ is heavy use by

humans. This route is used by locals to ascend to an adjacent mountain top where there is a

regional phone mast and a small compound; not only is this path heavily used by people but also

heavy equipment is frequently transported up the mountain along S2. It is reasonable therefore

to infer that disturbance along this path may be relatively high. Further, canopy height along S2

is slightly below what would be expected from a primary forest.

Figure 2.5: (A). Start of S1 path (B). End of S1 path; farmland runs along one side of

this section.

A B



Of our secondary routes S3 is in the worst condition. This survey route runs adjacent to several

plantations and in fact bisects a section of agricultural land at its midway point. Beginning at the

foot of a rice paddy S3 travels a short incline in its first 50 m then steadily declines as the route

continues. Despite this vicinity to human-altered landscapes the majority of S3 is covered by

healthy secondary forest; canopy height is for the most part above 10 m. However canopy cover

is not as dense as our other secondary routes. The width and condition of this path vary greatly.

Muddy and flooded sections of this route are frequent. The structure of the habitat along S3

allow us to term this route ‘secondary’.

2.2.7 Primary 1 (P1)

Figure 2.6: S2 path.

Figure 2.7: (A). water-logged section of S3 (B). beginning of S3.

A B


P1 runs adjacent to the border of Lokobe National Park (LNP), a protected area of montane and

coastal tropical rainforest. This habitat is considered to be relatively pristine. As the habitat of

LNP runs continuously across the border adjacent to our research area, it is reasonable to infer

that the environment along our survey route is similar, if not identical. P1 begins close to the

coast and steeply ascends into thick rainforest. Punctuated by a few <50 m ‘flat’ sections in the

first half of P1, towards the second, incline generally decreases until a level plateau is reached at

the end of the route that is adjacent to a waterfall (Fig.). The canopy height along P1 is very

high, often stretching beyond 20–25 m. Canopy density is also relatively high, mostly being

>70% cover except above water bodies. The vegetation on either side of the path is very thick

but does thin out on some steep valley sections towards the last 100 m of P1. High and dense

canopy are key indicators of ‘primary’ habitat.


P2 is very close and very similar in habitat type to P1. Thus, characteristics of vegetative

structure are analogous to P1. P2 runs through several montane streams and thus is composed of

Figure 2.8: (A). beginning of P1, adjacent to the coast (B). level section of P1 (C). waterfall

at the end of P1.

C

B A


numerous small valleys. Other than this the route is relatively flat. The condition of the path

varies and frequently changes with stormy conditions—likely due to flooding and receding of

the streams that punctuate its length. This route has been a challenge to track on occasion.

Similar high and dense canopy to that seen on P1 allows us to label P2 a ‘primary’ route.


A site for P3 has been selected however this route has not been measured or tracked; this work

will be completed at the beginning of phase 182.

2.2.10 Summary

The new survey route described here are a core framework of the projects to emerge in 2018.

Once every route has been properly marked out and mapped a complete map of these trails will

be made and included in phase 182’s report. In the meantime MGF will look to explore new

locations within our study area that could serve as additional routes; eventually a roster of 4–5

survey routes per habitat type should be maintained and walked regularly by the MGF team.

3. PROJECTS

3.1 Lemur Abundance

Lemuriforms are old world prosimians endemic to Madagascar. Placing their date of divergence

from the other primates is still the subject of robust scientific debate; some studies argue this is

around 60 million years ago (Yoder & Nowak 2006; Horvath et al. 2008), while others have

suggest dates tens of millions of years later (Godinot 2006; Sussman 2003). Nevertheless, most

Figure 2.9: (A). & (B). Forest streams bisecting P2.


authors on the subject agree that this date was after Madagascar split from India. Hence, lemurs

are thought to have colonised the island transoceanicaly sometime later, most likely by rafting

on vegetation (Tattersall 2006).

Once on mainland Madagascar, lemurs underwent several million years of adaptive innovation

and radiated into diverse niche space provided by the island’s fine-grained habitat diversity and

stochastic climate. Modern lemurs are near exclusively arboreal and have a broad repertoire of

ecological roles within Madagascar’s forests. From tiny mouse lemurs that are the world’s

smallest primates, to the largest living lemur, the Indri, these animals occupy diverse and

intricate positions in their respective trophic webs and are linchpins in many of their woodland

ecosystems. A good example of this is the role of lemuriforms as seed dispersers (Dew and

Wright 2006).

Three lemur species are found on Nosy be: the black lemur (Eulemur macaco macaco), Hawk’s

sportive lemur (Lepilemur tymerlachsonorum), and Clair’s mouse lemur (Microcebus

mamiratra). These species have different niches and life histories, and may use their habitat in

different ways. For instance, both L. tymerlachsonorum and M. mamiratra are nocturnal and are

found in relatively small groups (often solitary); this is not true for diurnal, group-living E.

macaco macaco. Thus, impacts from external stimuli such as human disturbance or

environmental catastrophe may be different among these species. We aim to address this

question.

Aims:

Determine if lemuriform abundance varies in our research area as a function of habitat

type.

Determine if lemuriform abundance varies in our research area as a function of human

disturbance level.

3.1.1 Methods

During phase 181 lemur surveys were walked along several 400 m pre-existing forest routes

that run through the forest in our study area. Teams of trained research staff walked these routes

at a regular pace and visually scanned the path and surrounding area for lemurs. Observers left

at least 2 m between each other. When an individual was spotted, the survey leader—a member

of research staff—identified the species and confirmed the group size and measurements. A tape

measure was used to measure the distance from path. Height of individuals was estimated by


eye as most lemurs were well above a height that could be recorded using a tape measure. Both

day and night surveys were conducted; electric light was used to assist observations during

night routes.

3.1.2 Outcome

Due to restriction in time and resources (heavily influenced by several large storms that hit our

research area over this period) the total number of surveys completed during phase 181 was

insufficient to draw any conclusions from. The data from this phase will be carried over to the

next and included in the interpretation of phase 182’s research.

3.2 Herpetofauna abundance and species richness

Madagascar’s trend of hosting peculiar assemblages owing to stochastic transoceanic dispersal

following separation from other landmasses and biological impoverishment after a catastrophic

environmental event ~65 million years ago also hold true for its reptiles and amphibians—for

instance, Madagascar is the global hub of chameleon diversity yet hosts no salamanders (Glaw

& Vences 2007).

All wild amphibians on Madagascar are frogs; these species are near exclusively restricted to

woodland habitat, not being able to withstand long periods in direct sunlight and needing humid

conditions and access to waterbodies for reproduction. As such, amphibian diversity is heavily

concentrated in the islands eastern rainforests. The heterogeneity of these habitats has helped

catalyse micro endemism of frogs along Madagascar’s eastern coast. These amphibians fill a

diverse range of niches in their habitats, from leaf litter dwelling Stumpfia, to Boophids capable

of bounding between high tree tops, Madagascar’s frogs can be very locally abundant and

occupy important positions in their trophic webs.

Reptiles on Madagascar are generally less restricted to the islands forests, their scaled dermis

allows them to thrive in areas of intense ultraviolet exposure and to store water so as to go much

longer between needing hydration. Therefore, Madagascar’s reptiles can occupy areas of scrub

and savannah—the most common habitat type on the island. These reptiles include a number of

charismatic and well-known species, such as the chameleons; Madagascar holds ~50% of the

globes chameleon species.


Nosy Be hosts ~60 reptile species; these range from tiny leaf litter skinks to the largest predator

present on the island and Madagascar as a whole: the Nile crocodile. Within our study area the

majority of reptile species are lizards; geckos, chameleons and skinks are by far the most

common species seen on surveys.

Aims

Determine if herpetile abundance and species diversity varies in our research area as a

function of habitat type

Determine if herpetile abundance and species diversity varies in our research area as a

function of human disturbance level.

3.2.1 Methods

During phase 181 herpetofauna surveys were walked along several 400 m pre-existing forest

routes that run through the forest in our study area. Teams of trained research staff walked these

routes at a regular pace and visually scanned the path and surrounding area for reptiles and

amphibians. Observers left at least 2 m between each other. When an individual was spotted the

survey leader—a member of research staff—identified the species. A tape measure was used to

measure the distance from path and height of individual. Both day and night surveys were

conducted; electric light was used to assist observations during night routes.

3.2.2 Results

In total ten herpetofauna surveys were walked (four night; six day). Four surveys were done in

degraded habitat, three in secondary, and three in primary. These surveys observed 103

individuals; 43 in degraded, 22 in secondary, and 38 in primary. However as degraded routes

were walked once more than other habitats, total encounter tallies cannot be used to draw

conclusions about the biological communities in our research area. On average 10.75

individuals were observed on each degraded route, 7.3 in secondary, and 12.7 in primary. On

average 6.5 species were observed on each degraded survey, 5.3 in secondary, and 6.3 in

primary.

Unfortunately due to a relatively poor sample size (only having one or zero surveys walked for

some routes), it would be misleading to analyse these results with more detailed methods,


looking at species richness for example. The results from phase 181 will be joined with 182 and

looked at as one data set.

Some species were found exclusively in certain habitat types. Phelsuma laticauda was only

found on primary surveys, while Langaha madagascariensis was only found on degraded

routes. As these results were only taken from several specific surveys that span a relatively short

period of time, it is difficult to make any lasting statements about whether these species will

turn up in other habitat types on later surveys. Specialisation to specific habitat types will be

revisited in later reports.

3.2.3 Discussion

While our data set was too small to elude to biodiversity values as discussed in section 1.2, we

can see that both the number of individuals and species observed on average on secondary

routes is lower than that of degraded or primary routes. This could be due to gaps in surveying

but there are many primary and degraded specialist species that may not be able to persist in

secondary habitat. Next phase we must look to expand our surveying effort and gather more

data to compare assemblages across habitat types.

4. Summary

The major achievement of phase 181 was the construction of nine new terrestrial survey routes

to be used for herpetile and lemuriform surveys; with the intent of further incorporating lemur

behaviour and invertebrate projects. This will underlie MGF’s core biodiversity monitoring

research that aims to explore the changes in our study area’s biological community with

reference to human influences over time. While is a long-term aim, the infrastructure used here

can allow smaller, more hypotheses based projects to take place; this is our aim in the coming

phases.

Completion of terrestrial survey route tracking/mapping is an immediate priority at the

beginning of phase 182, with the aim of producing a comprehensive map of all our routes. We

also aim to explore new routes for each habitat type.

Lemur behaviour surveys will be introduced during phase 182. The central aim of this

investigation will be to determine if black lemur behaviour towards human observers differs in

its composition and/or frequency along a human disturbance gradient. This project will run in


tandem with an attempt to measure disturbance on our routes. This project is expected to be

completed during phase 182.

Introducing butterfly and dragonfly surveying is also a priority for the next phase. This will take

the form of point counts where random GPS points are chosen and teams survey the

surrounding area for target species; water way surveys where forest streams into the forest are

followed for an extended period to scan for target species; and opportunistic surveying when

teams are out in our study area with different objectives.

With these research plans in place our understanding of ecological workings in our research area

will be greater than ever before and provide a platform for continued elucidating of biological

dynamics on Nosy Be.


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