The Terrestrial Protected Areas of Madagascar: Their History, Description, and Biota, Volume 2: Northern and eastern Madagascar

By Steven M. Goodman

In 1989, a book written by Martin E. Nicoll and Olivier Langrand was published on the protected areas of Madagascar, which heralded in a new era of conservation for this island nation. In the subsequent three decades, there was an important increase in inventories and studies on Madagascar’s terrestrial biota. This work led to significant changes in the systematics of Malagasy plants and animals, a large percentage unique to the island, and a notable augmentation in knowledge on Malagasy biodiversity. In addition, the considerable expansion of the protected area network, reinforcement of legal tools, and the development of new management modes and tools have contributed to a modernization of the protected area network.

The purpose of The Terrestrial Protected Areas of Madagascar is to present a large-scale update of information available from 98 terrestrial protected areas, various analyses to understand general trends in the conservation of these sites, and a synthesis to assess the needs for future scientific programs. Beautifully illustrated throughout with color maps, graphs, and photos, these three volumes will be an important reference for students, researchers, protected area managers, conservationists, and visiting ecotourists. Volume two covers northern and eastern Madagascar.

• Language: English
• Publisher: Association Vahatra in Antananarivo
• Release date: Apr 15, 2021
• ISBN: 9782957099733

Book preview

1F)

PART - II

The terrestrial protected areas of Madagascar

Chapitre 17

Introduction to Part II

Steven M. Goodman & Sébastien Wohlhauser

Different chapters in Part I (Volume I) of this book give the background information needed to place Part II (Volumes II and III), the detailed descriptions of 98 terrestrial protected areas, into a clear context. In Part I, after a general introduction (Chapter 1), the different chapters include a review of the history of Madagascar’s protected areas (Chapter 2), followed by legal aspects (Chapter 3), and then a series of chapters addressing different technical aspects. The first five of these technical chapters, on geology (Chapter 4), soils (Chapter 5), meteorology (Chapter 6), vegetation (Chapter 7), and flora (Chapter 8), present information to help readers better understand the importance of these subjects in relation to the ecology of a given protected area and other aspects of biotic communities. The subsequent eight chapters in Part I (Chapters 9 à 16) provide an update on the systematics and taxonomy of the island’s terrestrial vertebrates and form the backbone to consistently present information on these groups in Part II, specifically up-to-date scientific names of organisms occurring in each protected area.

All of the sites covered in Part II are included within the Système des Aires Protégées de Madagascar (SAPM), and this aspect is discussed in Chapters 2 & 3. Apart from this protected area network, there are other conservation sites managed privately and not governed by the Code de Gestion des Aires Protégées (COAP) (Commission SAPM, 2009); these sites are not covered in this book. As private property, the non-SAPM site managers can be involved in different types of tourism and conservation activities. Important examples include Berenty in the extreme south and Tsarasaotra on the outskirts of Antananarivo.

In Part II, information is systematically presented for each of the 98 protected areas covered herein using a series of section headers. The order in which protected areas are presented follows their unique site numbers (Table 2, Figure 1), which in turn generally follows a north to south orientation for the different ecological zones of the island. A number of protected areas are divided into separate parcels and in Table 34 is presented a list of such sites and the associated logic behind whether or not the parcels are treated separately within their respective protected area account.

Table 34. Protected areas covered herein that contain multiple parcels. Information is presented on ecological similarities or differences between parcels and the reasons they are considered as a single unit or treated separately (name in bold) in the 98 site descriptions presented in Part II. One of the overriding considerations is that marine habitats, which in some cases form separate parcels for a given protected area, are not covered in this book. The mapping scheme that defines the site numbers is presented in Figure 1 and more precise details on each of the 10 blocks are given in an eastern series (1A to 1F; Figures 8 to 13), a central set (2A to 2B; Figures 14 & 15), and a western set (3A to 3B; Figures 16 & 17).

A note on authorship of different section headers

To give needed credit to individuals that took part in writing different section headers or contributed to certain texts under a given protected area and at the same time not using too much printed space to repeat names, the following system has been adopted. The individual or individuals responsible for writing all the section headers for each of the 98 sites covered herein, are cited in the introduction to those headers, which is presented in this chapter (see below), but not in the individual texts associated with each of the 98 texts. For example, Paule M. Razafimahatratra & Julie Ranivo Rakotoson were responsible for writing all of the texts for the section header legal aspects; hence, their names are cited in the introduction comments to this header (see below), but not in the individual texts for this subject for each of the 98 sites. In contrast, Sébastien Wohlhauser & Steven M. Goodman were responsible for editing the texts for the section header infrastructure, but a range of different individuals contributed to certain texts and are cited as co-authors of the respective texts under the appropriate protected area.

Different points on section headers

Official and short names (Paule M. Razafimahatratra, Julie Ranivo Rakotoson & Steven M. Goodman): The first name provided for each protected area is the official legal designation (see Table 2) presented in the governmental decree in rigor, for example, Parc National d’Ankarafantsika (site 68). As official governmental documents are written either in French or in Malagasy, the administrative languages of Madagascar, throughout the book the French names are employed for the statutes of protected areas. Verification of the official name and all associated information concerning protected areas has been carried out in close collaboration with the Direction du Système des Aires Protégées (DSAP). In numerous cases, the official name of a given protected area is not in general use and associated synonyms are presented under the next header.

The second name presented under this header is the short version or nickname (see Table 2), for example, Ankarafantsika (site 68). To save space within the text and remove a certain level of awkwardness, the short name for each protected area is used. In the English portion of the text, the French name for the type of protected area, for example parc national rather than national park, is employed.

IUCN category (Paule M. Razafimahatratra & Julie Ranivo Rakotoson): The statutes of Malagasy protected areas are defined by Article 10 of the Code de Gestion des Aires Protégées (COAP) (Law No. 2005-005 of 22 January 2015). There are six different legal categories under the code and six equivalent categories based on the International Union for Conservation of Nature (IUCN) and following the Système des Aires Protégées de Madagascar (Commission SAPM, 2009); these different categories are presented in Table 35.

Table 35. The protected areas legal status types and associated IUCN categories.

In addition to the IUCN categories, certain protected areas may include other international statutes, such as UNESCO World Heritage sites, Biosphere Reserves, Ramsar sites, Key Biodiversity Areas (KBA), Alliance for Zero Extinction (AZE), Important Bird Areas (IBA), and Important Plant Areas (IPA) – see Chapter 3 for further details on these different international statutes.

Legal aspects (Paule M. Razafimahatratra & Julie Ranivo Rakotoson): This section traces the date of creation, subsequent modifications associated with protected area type or delimitation, and the precise citations to different governmental legal documents; these different aspects are discussed in detail in Chapter 3. The following entries are presented under this section header: 1) current manager, which is the organization at least up to mid-2017 was responsible for the management of the site; 2) former management, which is the organization that was previously responsible for the management of the protected area – in cases that different managers have been involved, the organization names are presented in chronological order; 3) creation, which is the date associated with the first legal action to put in place a given protected area; this includes temporary status for those with the former statute nouvelle aire protégée or NAP; 4) status change, which is related to a modification of the type of protected area, such as from réserve spéciale to parc national, and governed by Article 29 of the Code de Gestion des Aires Protégées (COAP) (Law No. 2005-005 of 22 January 2015), which allows for different types of statute changes (Articles 30 & 31). Certain sites covered in this book and before their inclusion in the modern protected area system had different management or protection statutes, such as within the Domaine Forestier National (DFN), specifically forêt classée, réserve forestière, périmètres de reboisement et de restauration, station forestière, station d’essais sylvicoles, etc. (Direction des Eaux et Forêts et de la Conservation des Sols, 1969); 5) last status change, which is the most up-to-date status of the site; and 6) current surface, which is the surface area of the site according to the governmental decree in force.

All of the different legal aspects presented in this section are derived from a considerable number of unpublished and published documents. Within the official government decrees and articles, a number of different acronyms are presented associated with the organizations and different codes used in these documents. Rather than giving the full spellings of these acronyms in the text, they are defined in Table 36.

Table 36. Definitions of acronyms used in official governmental decrees and articles.

Total surface area and elevational zonation (Jean Clarck N. Rabenandrasana & Steven M. Goodman): In certain cases, important differences were found in the calculated total surface area of different circulating shapefiles for a given protected area; this problem is discussed in detail in Chapter 3 (Size of protected areas). Under this section header, the calculated total surface area based on the 2017 shapefiles provided by Direction du Système des Aires Protégées (DSAP) are used. In certain cases, these values differ from those presented in different portions of this book (e.g. Tables 2 & 20 and in Part II for each protected area under the entry legal aspects) and these differences are often associated with technical aspects associated with topography or particular topologies and the resolution of computing sources. In most cases, the variation does not exceed 1% of the calculated surface area of a site (Table 21).

Calculations on the elevational zonation of a site were made based on downloadable images from the Shuttle Radar Topography Mission (SRTM, see https://lta.cr.usgs.gov/srtm/mission_summary) and associated with the National Imaging, Mapping and Cartography Agency (NIMA) and the National Aeronautics and Space Administration (NASA). One of the objectives of this project was to produce elevation topographic data for 80% of the earth using a technique known as interferometry radar. In the case of the SRTM 1 Arc-Second Global system, which was used herein for these calculations, the elevation data offers high resolution of 1 arc second (30 m). Using WGS 84-UTM 38S projections, elevations were classified using ARCGIS software at intervals of 250 m, starting with a minimum of 0 m (sea-level) to a maximum of greater than 2000 m. Elevation classes were superimposed on the respective protected area shapefile, and the surface area were calculated for each elevational zone.

Under this section header, only the elevational zones found in a given protected area are presented. For example, Ambohitantely (site 71) falls within the elevational range from 1001 to 1750 m and, hence, no figures are presented for lowland or high mountain zones. An additional example includes Tampolo (site 30), which is strictly lowland and no figures are presented for areas above the 0-250 m zone. The elevational zones of each protected area are also represented on a map associated with the site. The global representation of the different elevational bands across the 98 protected area sites covered herein is presented in Table 37.

Table 37. Total representation within the 98 protected areas included herein of nine different elevation bands.

Synonyms (Steven M. Goodman, Paule M. Razafimahatratra & Jeannie Raharimampionona): These include different names for the 98 protected areas covered in this book and used in the published literature, historical maps, and unpublished documents. In general, the original language the name was presented is given and translation into the other language of this bilingual book are not given. When the number of synonyms for a given site exceeds four, these are presented in a separate table.

Site synonyms have different origins. In some cases, they are derived from the transformation of the legal statute, for example, from réserve naturelle intégrale to parc national. In other cases, they include former designations of site types (e.g. forêt classée, réserve forestière, station forestière, etc.) within the limits of the protected area recognized today. Some variants exist based on differences in pronunciation between dialects of Malagasy and the manner of transcription (e.g., Tsimanampetsotsa as compared to Tsimanampesotse). Different historical and modern names exist in the literature for forest blocks that today fall within a delimited protected area; an effort has been made to include these names among the site synonyms (e.g., Forêt de Périnet or Forêt d’Andasibe for the Parc National d’Analamazaotra [site 42] or Didy or Forêt de Didy for the Réserve de Resources Naturelles du Corridor Ankehiny-Zahamena [site 34]). Names have also been included associated with different types of conservation zones outside of the International Union for the Conservation of Nature and Natural Resources (IUCN) protected area designations (see Chapter 3, International designations for protected sites), but occurring at least in part within a recognized protected area (e.g., Ramsar, World Heritage site, etc.). To help readers negotiate these names and to provide quick cross-referencing, there is a place-name index at the end of the book.

Infrastructure (Sébastien Wohlhauser & Steven M. Goodman): This section header presents information on infrastructure provided by site managers or obtained from various reports (e.g., MNP 2005, 2014) and with a cut-off date of late November 2017. The details include management infrastructure (main administrative office and sector operational offices), conservation and restoration devices (control posts, barriers, watchtowers, firewalls, and tree nurseries), tourist reception facilities (circuits or trails, points, offices and reception kiosks, interpretation centers, bungalows, lodging, camping sites, viewpoints, and belvederes), and research and monitoring facilities (stations, associated structures, camps, and permanent plots or transects for research and monitoring). In most cases, this section has been written or verified in collaboration with the site manager or partner organization.

Access (Steven M. Goodman & Sébastien Wohlhauser): The information given on access to the different protected areas is not meant to be comprehensive, and it is suggested that tourists or researchers consult recent editions of Madagascar guidebooks that contain details that may be more precise or up-to-date. For information on accommodations, information is generally restricted to those found within a given protected area, which are based on the editors’ experience or gleaned from different published books (Nicoll & Langrand, 1989; Bradt, 2011, Petit Futé, 2016) and websites (http://www.travelmadagascar.org/ and http://www.parcs-madagascar.com/).

At the time this text was written (late 2017), the number of cities and towns served by Air Madagascar flights was notably reduced as compared to the recent past, rendering certain details in guide books on domestic air connections no longer valid. Hence, it is recommended that tourists check on the availability of national flights with travel agencies, the Air Madagascar website or their principal office in Analakely, Antananarivo (Tananarive).

Tourist tickets to enter protected areas under the direction of Madagascar National Parks can be purchased in Antananarivo at their main office in Ambatobe (near the Lycée Français); at the kiosk of the Regional Tourist Office of Analamanga in Antananarivo (Antaninarenina); or in most cases from the reception office at the entrance of a given protected area, with the exception of those off the beaten-track and seldom visited by tourists. It is important to note that all sites with the statute réserve intégrale naturelle are closed to tourists. Information on local protected areas is available in most Office Régional du Tourisme located in the various regional administrative centers or former provincial capitals.

For visitors traveling on their own, it is recommended to obtain the standard FTM (Foibe Tao-tsaritan’i Madagasikara) road map (Carte Routière) from their main office in Antananarivo (Ambanidia, Route Circulaire) or from tourist shops around the country. In our texts, this map has generally been used in describing the road networks. The numeration system for principal national roads (Route Nationale in French = RN), tends to be consistent on maps and in guidebooks. However, this is not the case for secondary or tertiary roads, generally with different acronyms (e.g. RIP = Route d’intérêt provincial, RNS = Route nationale secondaire or RNT = Route nationale temporaire). Hence, off the main national roads, it is best to request information or directions, rather than relying on road numbers.

Changes in weather, specifically rain, can modify rapidly road conditions. This is most important during the passage of cyclones or tropical depressions. In general, the rainy season in the eastern and central portions of the island is from early December through April and in the west, southwest, and extreme south from January through March. However, different events, such as early or late tropical storms, can occur outside these seasonal periods.

We have employed a systematic mapping scheme for the 98 protected areas presented herein. The island-wide base map that defines the system is presented in Figure 1 and on the inside front cover, which shows the placement of the individual protected areas, each with its unique site number (Table 2). Within this mapping scheme, Madagascar is divided into 10 different blocks following a north to south system and keyed to Figure 1; an eastern series, including the extreme north (blocks 1A to 1F; Figures 8 to 13), a central set (2A to 2B; Figures 14 & 15), and a western set (3A to 3B; Figures 16 & 17). Within Part II, each of the 98 protected areas has an individual map presenting different details, which include localities mentioned in the associated text within or outside the site, principal access routes, and elevational zonation. Part II is divided into two books, with Volume II covering the eastern and central series (sites 1 to 62) and Volume III the western series (sites 63 to 98).

Today, Madagascar is divided into 22 different official regions, which represent after the central government, the primary administrative divisions for the governance of this island nation. This is compared to the former system of six provinces, which was suspended and replaced by the region system in early October 2009. For a map of the placement of the 98 protected areas covered herein based on the older province system see Figure 18a and on the current administrative region system see Figure 18b.

Visitation rates (Madagascar National Parks & Steven M. Goodman): The figures for the number of visitors buying entry tickets to a protected area are derived from statistics provided by Madagascar National Parks (MNP) for most sites under their management; a review of these data is presented in Chapter 1 (Visitation of protected areas). This information provides insights into patterns of ecotourism. Visitation data are presented for three different periods (1992-2001, 2002-2011, and 2012-2016), and with three different descriptive statistics: total number for period, average number for period, and extremes (ranges) per period (minimum and maximum values). In the case of notably low or out-lying minimum values, the year has been added to indicate if this may have been associated with political problems on the island, which include, for example, the periods of late 2001-2002 and 2009. At some sites, data for certain years are not available. Visitation rates are not consistently available for protected areas falling outside the Madagascar National Park network and no information for these sites is presented herein.

Geology (Brooke Crowley & Janine Sparks): This section includes detailed descriptions of the geology of each site following aspects and technical terms presented in Chapter 4. The geological classification employed is based on the recent U.S. Geological Survey map of Madagascar and published by Roig et al. (2012). In Figure 105, the 98 sites are presented on the simplified geological base map. In certain cases, for protected areas with complex underlying geology, individual maps are given within the site texts. Finally, in the geology texts, geographic place name terms presented by Roig et al. (2012) have been employed, rather than the standard spellings used elsewhere in the book (e.g. Tolanaro as compared to Tolagnaro).

Figure 105. Map of the 98 protected areas covered in Part II overlaid on the simplified U.S. Geological Survey (Roig et al., 2012).

Soils (Amanda H. Armstrong & Megan E. McGroddy): This section includes descriptions of the soils of each site following the classification and technical terms presented in Chapter 5, specifically Table 22. The soil classification employed is based on the World Reference Base classification (IUSS Working Group WRB, 2015).

Climate (Marie Louise Rakotondrafara, Luc Y. A. Randriamarolaza, Herilalaina Rasolonjatovo, Christian L. Rakotoarimalala & Fidison S. Razanakiniana): An overview of meteorological patterns is presented for each site following parameters defined in Chapter 6. More specifically, these include descriptions of weather patterns for the period from 1985 to 2014 – rainfall periodicity, shifts in rainfall patterns over time, and different data concerning annual mean, maximum, and minimum temperatures. Insights into recent climatic events or trends related to climate change are also given.

The only aspect that does not follow the parameters outlined in Chapter 6 is annual precipitation for each site. These values are derived from Climate Engine (https://clim-engine-development.appspot.com/), which calculates average precipitation at a pixel resolution of 4800 m x 4800 m; analyses are restricted to the period from 1981 to 2017. The reported average rainfall figures use downscaled satellite data combined with ground station data and values are area averaged based on the shapefile for each protected area. The descriptions of local climatic regimes for each protected area employ Cornet’s (1974) bioclimatic scheme, which is based on calculations of hydric deficit (precipitation minus potential evapotranspiration), mean minimum temperature in the coldest month, and length of the dry season.

Vegetation (Laurent Gautier, Sebastien Wohlhauser, Jacquis Tahinarivony & Patrick Ranirison, with assistance of Yannie Andriamiarantsoa for collecting and organizing information): This entry includes a description of main vegetation types for each site, forest cover, and special habitats. When appropriate, details are given on aspects of altitudinal variation when they relate to changes in vegetation. Given the heterogeneous level of available information for the different protected areas, these texts are based on (in order of priority): published scientific articles and monographs, if available, specific to the area, unpublished reports, university memoirs and theses, and the personal experience of the authors.

In several cases, there was little or no information for a given site and in these cases, satellite imagery accessible through Google Earth has been used, adding valuable high resolution information to what can be deduced from vegetation maps of the island (Humbert & Cours Darne, 1965; Faramalala et al., 1995; Moat & Smith, 2007). For aspects of vegetation seasonality, the Land Cover Map of the European Space Agency Climate Change Initiative (http://maps.elie.ucl.ac.be/CCI/viewer/index.php) has been used. In some cases, fire occurrence and dynamics provide an important window to interpret satellite imagery and the origin of data is explained in another section header (see Fire, below).

For each protected area, the vegetation text usually follows three separate entries, depending on available information. The overview section includes general information on the level of knowledge (descriptive studies, quantitative studies, and vegetation maps), as well as information on climate, soils, and physiographic aspects that influence primary vegetation. The natural vegetation section describes the natural and primary vegetation types following the classification given in Chapter 7, which is summarized in Table 23, as well as specific habitats, distribution, seasonality, structure, main species, biological types, and a brief eco-geographic description of vegetation types. The entry modified vegetation is associated with human transformation of vegetation, and includes some insight into the processes involved. In several cases, a concluding text highlights main trends in vegetation dynamics, particularly threatened vegetation types or habitats, as well as relevant directions for further research. For different reasons, for certain sites the entries concerning natural and modified vegetation are combined.

The composition or specific richness are not presented here, but are treated in the Flora section (see below). Taxa mentioned within the vegetation section header are those that are critical for structure and ecologically determinant or discriminating taxa. Given different problems in the published literature associated with plant identification, including the recent description of many new species, cited taxa are often only presented at the generic or familial level. Although not consistent with other portions of this book, to reduce the length of the vegetation texts, when presenting families, the expression family has not been included.

Flora (Peter B. Phillipson, Porter P. Lowry II, Lalao Andriamahefarivo, Marina Rabarimanarivo & Fano Rajaonary)

Methodology and synthesis

Endemism within the flora of Madagascar at the levels of species, genus, and family is among the highest in the world (see Chapter 8 for further details). However, with few exceptions, little or no information is available on the relative levels of plant endemism of different protected areas, as well as for those species that are rare or with restricted distributions. In this section header, profiles are presented for each site, providing details on botanical collecting history; a snap-shot of the extent of current knowledge and notes on the flora and its affinities; and lists of noteworthy species, including site endemics, range-restricted species, threatened species, and the representatives of endemic plant families. The primary source used to compile this information was a GIS analysis of the available collection data for Madagascar and occurrence data from the Madagascar Catalogue (2018), which was supplemented by details in reports and scientific publications, and knowledge of the authors, as well as other botanists for certain sites and plant groups. Although not consistent with other portions of this book, to reduce the length of the flora texts, when presenting families, the expression family has not been included.

Collection data for each site

Data extracted from the Madagascar Catalogue (2018) provided the basis for the flora summaries presented in table format for the majority of the 98 sites. This dataset contains information on an estimated 60% of all vascular plant collections made on Madagascar, and represents the most comprehensive source available.

Plant specimens are generally accompanied by data on their geographic origin, although in the early decades of botanical exploration, this information was generally limited and sometimes with no more than Madagascar or Central Madagascar. During the 20th century, botanists gradually started to provide more detailed information, often using place names that can be associated with precise localities and sometimes recording geographic coordinates obtained from maps. However, geographic coordinates have only been recorded systematically since the advent of hand-held GPS units in the mid-1990s. When possible, coordinates for older collections have been estimated post-facto using the geographic information provide on specimen labels in conjunction with itineraries of collecting expeditions, maps, gazetteers, and online resources such as Google Earth.

For the present study, the compiled dataset comprises 220,073 collection records of vascular plants extracted in March 2018 from the Madagascar Catalogue (2018), of which 211,260 were analyzed using Geographic Information System (GIS) technology to determine occurrence within the 98 protected areas (the other 8813 records lacked coordinate data). A total of 91,170 specimens (ca. 40%) were placed within the official Direction du Système des Aires Protégées (DSAP) shapefiles defining these sites. Three protected areas are without any documented collection records: Sahamalaza-Iles Radama (site 15), Ampanangandehibe-Behasina (site 37), and Ambondrobe (site 84), whereas many others possess thousands of records, including Loky Manambato (site 6) having the highest number with over 6200 records. The number of documented collections per km² for each site was calculated as a measure of collecting intensity, taking into account only the terrestrial portion of sites that also include marine areas, notably Nosy Hara (site 1) and Nosy Mangabe (site 25), for which the terrestrial portion is less than 50% of the total surface area. Four of the 98 sites have over 300 collections per km², including Nosy Mangabe, Analalava (site 33), Analamazaotra (site 42), and Mandena (site 59), while 39 sites have less than one documented collection per km².

Many of the collections from a given site have been identified to the species level (or infraspecific taxon when appropriate), whereas others only to genus or family, and a small portion are unidentified. This information was used to assess the level of identification effort for each site, both in the field and subsequently in the herbarium. Some of the relatively well-collected sites have more than 80% of the collections fully identified, including Zahamena (site 32), Torotorofotsy (site 41), Manjakatompo Ankaratra (site 74), Itremo (site 78), Beanka (site 81), and Bezà-Mahafaly (site 96). By contrast, the following well-collected sites have only about 50% of their specimens identified to species: Makira (site 23), Agnakatrika (site 54), Tsitongambarika (site 56), and Alandraza Analavelo (site 92). While some of these comparatively low values result from relatively less effort having been made to identify collections, they also likely reflect the presence of a large number of undescribed (and therefore unidentifiable) species.

Compilation and analysis of species lists for each site

Based on the compiled dataset, an initial species list was produced for each site, which was then supplemented by information from the following sources:

1) Recorded occurrence in protected areas from the Madagascar Catalogue (2018). Since the launch of the Madagascar Catalogue in 2005 (Phillipson et al., 2006), data on occurrence in protected areas has been captured for each species. These data were obtained from herbarium specimens and from information cited in recent taxonomic revisions. This was initially limited to the sites included in Madagascar’s protected area network prior to 2015, but was then expanded to include newly designated protected areas, as well as designated extensions of the older protected areas. This information inevitably evolves as new collections are incorporated and new taxonomic work is completed. The cut-off date for the dataset used herein is March 2018.

2) Additional data on species occurrence in protected areas obtained from published sources dealing with specific sites, as well as unpublished reports and university memoires and theses.

3) Specialist knowledge provided by botanists with expertise on the flora of certain sites and regions.

4) Detailed data on the five currently recognized endemic plant families on Madagascar obtained from Ramananjanahary et al. (2010).

The Madagascar Catalogue (2018) systematically indicates global distribution range and rarity for all plant species documented on the island. Using the species list compiled for each site, data were extracted for each recorded species with respect to these criteria and analyzed to provide information on the total number of documented species along with totals for those that are: 1) native to Madagascar, 2) endemic to Madagascar, 3) introduced and naturalized on Madagascar, 4) known only from the site (either from multiple collections or from just the type collection), or 5) known from two to five other sites. When relevant, the percentage of the native flora for each of these categories was calculated for each protected area and presented in table form.

Compilation of noteworthy information on the flora of each protected area

In addition to providing quantitative data for each protected area, which indicate the levels of endemism and uniqueness of its flora, notes on general characteristics and interesting or remarkable floristic features of each site are also given. Rather than listing the known plant species for a site, in many cases, these would be notably long, this information is available through the Madagascar Catalogue (2018) following the provided link. The inclusion of details on selected families or genera have been chosen to illustrate the exceptional floristic interest of a given site.

Three protected areas have more than 100 species that are currently considered site endemics: Tsaratanàna (site 13), Marojejy (site 19), and Masoala (site 24). Moreover, among the 98 sites, 66 are currently known to have at least one locally endemic species. Eight sites harbor more than 100 range-restricted species known to occur at the site and at no more than four other protected areas: Loky Manambato, COMATSA Nord (site 11), Tsaratanàna, Manongarivo (site 14), Marojejy, Masoala, Analamazaotra, and Andohahela (site 57). Risk of extinction assessments based on the IUCN categories and criteria (IUCN, 2012) and published on the IUCN Red List (2018) have been presented for selected species.

With respect to Madagascar’s five endemic plant families, 71 of the 98 protected areas treated herein have at least one of these families represented and seven sites have at least one member of all five families. Four protected areas have at least 19 of the 97 species belonging to the endemic families: Makira, Masoala, Zahamena, and Betampona (site 35).

Fauna (Steven M. Goodman, Sébastien Wohlhauser & Achille P. Raselimanana): Aspects of the land vertebrate fauna (excluding fish) of each protected area are summarized in this section header and mostly derived from the associated table presented for each site. Species richness and local levels of endemicity are emphasized. The term endemic might cause some confusion. A large percentage of the Malagasy land vertebrate fauna is unique to the island and, hence, endemic. More specifically, local endemic species are the focus of such texts, that is to say those that based on current information only occur at a given site (micro-endemic) or local regional endemic species; put in another manner, those with limited geographical distribution and unique to a relatively confined area (regional micro-endemic). When listing local endemic species in the text, only those that have been formally named are cited; hence, candidate species are excluded (see Chapter 1, The concept of candidate species for further details). Comments are also provided in some site texts on the local native carnivoran (family Eupleridae) communities, with the assumption that when a protected area contains the full complement of species known from a given region, this is a sign of relatively intact forest ecosystems; such cases are almost exclusively associated with the largest protected areas.

In the closing sentences of a given site text, suggestions are sometimes given on future work. These recommendations are from several perspectives: filling in missing information for a given site; utilizing long-term faunal research to advance on a variety of questions; and, based on sites with considerable threats of human pressures as measured by forest loss (more than 20% loss since 1996), the need for inventories to obtain what is almost certain to be archival information before a forest block is largely lost. Comments are provided for certain sites on aspects related to biogeography or more specifically zoogeography, long-term research, and different proposed actions for management and scientific advancements. As the synthesis of information associated with putting together the tables of the known vertebrates of a site are in certain cases complicated and not the same for each group covered, the next section presents these details.

Vertebrate lists

For each protected area, lists are presented in table form for the different terrestrial vertebrate groups and following the taxonomy presented in Part I, specifically amphibians (Chapter 9), reptiles (Chapter 10), birds (Chapter 11), tenrecs (Chapter 12), native rodents (Chapter 13), bats (Chapter 14), native carnivorans (Chapter 15), and lemurs (Chapter 16). These lists include species documented from a given site, rather than those inferred to occur based on their known distribution. In most cases, these lists were completed or verified in collaboration with the site manager or partner organizations. To help the reader negotiate scientific names previously used in the published literature and no longer in use, a table has been included in Part I in the respective chapter for each vertebrate group listing these synonyms.

As is evident from many of the tables of land vertebrates for the 98 sites treated herein, considerable work remains to resolve taxonomic questions, and this situation is most evident for groups such as amphibians, reptiles, and lemurs. Decisions associated with the recognition of different species in the lists based on published and unpublished literature, particularly for the three groups mentioned above, was often difficult without voucher specimens or photographs. Hence, in uncertain cases, this was done in a conservative manner and these lists should be considered as a starting point for future refinement of the taxa occurring at a given site.

For several protected areas, the commencement of research and the collection of specimens date from more than 100 years ago. At certain sites, over the intervening years, forest cover has been reduced drastically, and different biotic changes followed, including local extirpations. As it is difficult to draw an artificial line between these different periods and data sources, both historical and modern information was used construct the lists. Some clear problems can be cited associated with this system: a species that was historically recorded from a protected area, but for which there is no modern record (e.g. Eutriorchis astur at Analamazaotra [site 18]); a taxon that has subsequently gone extinct (e.g. Tachybaptus rufolavatus at Lac Alaotra [site 31]); a species has been locally extirpated (e.g. Aythya innotata at Lac Alaotra [site 31]; or a taxon has been locally extirpated and subsequently reintroduced (e.g. Propithecus diadema at Analamazaotra [site 18]).

Codes – For each species entry within a fauna list table, the following coding system was used: an asterisk (*) before the species name designates a Malagasy endemic; scientific names in bold are those that are strictly endemic to the protected area; underlined scientific names are unique or relatively unique to the site; and + before a species name indicate taxa with an IUCN statute of at least Vulnerable or higher (see Chapter 1, Conservation status). It is important to keep in mind that the IUCN statute for many recently described species have yet to be assessed. In the case of lemurs, when an asterisk (*) appears after a species name this designates a taxon that is tentatively accepted herein and in need of further data to verify its validity (see Chapter 16, Lemur species and their recognition).

Compilers and sources of data – For each list, the information compilers are noted, which include individuals and organizations, and are those that contributed data, including in many cases unpublished records. When multiple compilers are given, the order reflects relative levels of input. In contrast, contributor names are presented alphabetically. For most species of amphibians, reptiles, non-spiny tenrecs, native rodents, and bats, the lists are derived from specimen data or at least captured animals identified in the hand. In contrast, for birds and diurnal lemurs, most records are based on direct observation in nature.

Amphibians and reptiles – Given the many taxonomic uncertainties surrounding Madagascar’s amphibians, especially at the species level, elaborating lists for each protected areas is a complex task, and at this stage, many of the tables are not free of errors. Information relied on herpetological surveys, available for the majority of Madagascar’s protected areas through the remarkable and intensive work of numerous researchers and students during the last 25 years, and complemented with historical records summarized in Glaw & Vences (2007), as well as the unpublished observations of Frank Glaw, Achille Raselimanana, and Miguel Vences. Hence, these lists are based on verified records and not on probable occurrences extrapolated from models or distributional maps. However, a certain level of subjectivity has been introduced, as explained below.

The published and unpublished inventory work cited as original sources to create the species lists of amphibians and reptiles are generally based on the accepted taxonomy at the time of the site inventory. Due to changing taxonomy and presence of large numbers of morphologically similar species, particularly among many amphibians from moist evergreen forest sites, critical evaluation was required. Thus, the lists were adjusted based on the following criteria:

1) If a given record was considered doubtful, and could not be assigned with some confidence to another, morphologically similar species known to occur in the respective geographic region, the record was not included in the respective protected area list.

2) If a record was considered doubtful, but occurrence of the species in the respective region was not unlikely, the scientific name was placed in parentheses, which in turn indicates it requires confirmation.

3) If a recorded species was unknown to occur in the respective geographic region and its occurrence was unlikely, but another, morphologically similar species is known to occur, the species name was modified accordingly. In such cases, the species name was placed in parentheses, which in turn indicates it requires confirmation.

4) If species-level identification of a record seemed unacceptable, but genus-level identification was deemed reliable, and if this was the only record for a certain genus at the site, the species was added to the respective list as sp. (one species recorded) or spp. (more than one species of that genus recorded). A good example is the amphibian genus Stumpffia (family Microhylidae), whose species-level identification in the past was almost impossible, but which in most cases can easily be recognized as belonging to this genus. Following this example, information was included for a protected area where one or several species of Stumpffia had been recorded, although species-level identification requires further study.

To summarize, while the amphibian and reptile species lists presented herein are an important step forward in the understanding of the biodiversity in Madagascar’s protected areas, users of these lists need to keep in mind that they contain numerous unresolved taxonomic questions. In the case of amphibians, the often short lists of species in western protected areas with dry forest formations are probably rather complete and taxonomically less controversial. The data compilation presented herein highlights that many gaps exist in current understanding of the species-rich amphibian fauna of protected areas in the east, warranting more field and taxonomic work.

Birds – We have extracted some records for the bird species lists from the ebird site (http://ebird.org/content/ebird/), organized by The Cornell Lab of Ornithology and National Audubon Society, and the individuals that submitted the information have been cited. Doubtful records and problematic localities are not included from this website. However, the vast majority of the data used to compose the species lists come from numerous published or unpublished site inventories conducted by researchers, students, and ecotourists. Synanthropic species, specifically fowl, and pigeons (Columba livia, family Columbidae) living in a feral state, have not been included.

Domestic mammals – We do not include in the lists any domestic mammals, specifically ungulates (cattle, goats, and sheep); in some cases, these animals find pasture in protected areas or live in a feral state. In addition, the African bush pig (Potamochoerus larvatus, family Suidae) has not been included.

Lemurs – The taxonomy of nocturnal primate genera is largely based on molecular genetics (see Chapter 16 for further details). For populations of these animals that have yet to be genotyped, it was preferred not to infer the local species and, hence, designations such as "Microcebus sp." are used.

References – A number of references for different vertebrate group have been used to formulate the lists for different protected areas, and these are not presented at the end of each site text: amphibians (Glaw & Vences, 2007), reptiles (Glaw & Vences, 2007), birds (Raherilalao & Goodman, 2011; Goodman & Raherilalao, 2013; Safford & Hawkins, 2013), tenrecs (Soarimalala & Goodman, 2011; Goodman et al., 2013), endemic rodents (Soarimalala & Goodman, 2011; Goodman et al., 2013), bats (Goodman, 2011; Goodman & Ramasindrazana, 2013); carnivorans (Goodman 2012, 2013), and lemurs (Ganzhorn et al., 1996/1997; Mittermeier et al., 2010, 2013; Schwitzer et al., 2013). For numerous sites that were included in Nicoll & Langrand (1989), the lists they provided form the basis of those presented herein. In most cases, taxonomic references to different scientific paper used to create the different species lists are not presented in the bibliography at the end of the site text, but rather those to faunal inventories.

Cultural aspects (Steven M. Goodman & Sébastien Wohlhauser): The different aspects presented under this header are largely based on the replies of the site managers to a questionnaire they were sent on the importance of a given protected area and its immediate surroundings to local people; these texts have been condensed and edited. The responses differed considerably in detail and subject matter. In many cases, the provided information is associated with historical aspects, points related to the daily lives of local ethnic groups, and, more broadly, Malagasy beliefs, customs, and narrations. For certain sites, no response was received from the site manager organizations, and this is indicated. When appropriate, words or expressions in Malagasy have been included. In some cases, additional information has been added, particularly when this fills in details not mentioned by the site managers.

Anthropogenic pressures (Steven M. Goodman & Sébastien Wohlhauser): This section header, which focuses primarily on terrestrial habitats, relies mostly on information supplied by the protected area managers associated with a questionnaire. In most cases, these texts have been condensed and edited, in part assessing different causal levels (Salafsky et al., 2008). For certain sites, no response was received from the site managers, and this aspect is indicated under the section header. For some sites, additional details have been added from the published literature or unpublished reports, specifically when this adds important insights into aspects cited by the site managers. Reference to coastal zones and marine aspects are rarely made, with the exception of mangrove exploitation. The names of different vegetation types follow the classification presented in Chapter 7.

In portions of Madagascar, traditional taboos (fady in Malagasy) are quickly eroding away. In some cases, certain portions of local society, particularly the impoverished younger population associated with deficits in land access and education, do not follow the classical cultural dictates of right and wrong, and, most importantly for this section header the exploitation of natural resources. This change in cultural dictates has different ramifications for threats to local ecosystems and anticipated tendencies of conservation programs. The importance of this change in certain regions or with particular ethnic groups is that 10 to 20 years ago different types of anthropogenic pressures were minor, but today are major challenges to management of natural resources. These vicissitudes are related with broad patterns of change in Malagasy society and socio-economic structure, which include population demographics and lack of access to education via a decline in state actions and institutions. At a finer level, in certain circumstances, the non-recognition of local taboos is also related to the immigration of people with different cultural norms into areas of other ethnic groups. For example, the displacement of a considerable number of Antandroy from the extreme south into the region of the Sakalava Menabe in the central west, specifically in and around the Menabe Antimena (site 85) protected area.

Forest loss (Jean Clarck N. Rabenandrasana, Sébastien Wohlhauser & Steven M. Goodman)

Overview

In this section, details are presented on an investigation of forest loss in protected areas, which focus on two periods: 1996 to 2006 and 2006 to 2016. These periods were chosen in order to cover different timelines than previously published studies on forest loss, often based on different analytical techniques than used herein (e.g. Green & Sussman, 1990; Nelson & Horning, 1993; Horning, 2000; Dufils, 2003; Harper et al., 2007; Allnutt et al., 2008; Grinand et al., 2013, Zinner et al., 2014). Further, most of these previous studies have had a regional or island-wide focus and not on specific protected areas. Subsequent to these publications, there has been considerable expansion of the protected areas network and changes in anthropogenic pressures at certain sites; this is a dynamic process. Hence, the intent of this section header is to provide insights into aspects of forest loss in the 98 protected areas covered herein.

Some terminology needs to be clarified for this section to avoid confusion, and certain terms follow the definitions of Salafsky et al. (2008). By forest loss (also referred to as ecosystem conversion), it is specifically meant the process of complete ecosystem conversion associated with relatively broad scale activities, such as clear cutting, mining, or agricultural expansion. In contrast, forest degradation (also referred to as ecosystem degradation), indicates partial ecosystem conversion related to small scale activities, such as selective logging, localized mineral extraction, etc. These terms are related to the scale of different measures of land cover change. In the context of the investigation discussed in detail below, the broad scale analysis was designed to identify change in land cover and focuses specifically on forest loss, while forest degradation is generally at too small of scale to be measured by the techniques employed. For example, in order for any change in forest statute to be discernable, it has to be notably larger than one pixel, which is 30 x 30 m. Hence, for example, small scale mineral extraction under forest cover might not be detected, as compared to a zone the size of a soccer field and largely devoid of canopy cover.

A number of points are important associated with the technical aspects of this section header. The first is that the analysis was conducted at the level of Madagascar, which provides important insights into broad scale patterns, but is distinctly less precise at a fine scale and may also lead to unclear, and sometimes incorrect, land cover classifications. This is particularly the case when available cloud-free satellite images were taken at different foliation stages and lead to false interpretation, when using automated classification system: dry vs. evergreen forest, secondary vs. primary forest, natural forest vs. tree plantation (Pinus at Manjakatompo Ankaratra, site 78), secondary forest vs. non-forest, and sometimes even forest vs. non-forest. This is also the case in mixed open and closed vegetation areas (sclerophyllous woodland vs. semi-woody vegetation, sites 77 & 78) and in transitional areas, where evergreen and dry forests are imbricated and vegetation shows a notably seasonal pattern (north: sites 1 to 8, 10, 12, 14, 15, 17, 18, and 21; south: sites 57 and 62; central west: site 72). For some protected areas, there was no element of ground-truthing, which would have been helpful to correct the resulting classification. Specifically, it was difficult to identify semi-deciduous evergreen forests, which were sometimes classified as dry forests or moist evergreen forests.

Technical aspects

The description of the technical aspects presented below is simplified. For more details on the approach used, see the analysis undertaken by a consortium of historical deforestation for the years 2005-2010-2013, which was validated during different workshops (Grinand et al., 2013). Herein, LANDSAT 5, 7, and 8 images were used because of their resolution and the availability of scenes for different periods. They are freely downloadable from https://earthexplorer.usgs.gov/. The image selection associated with the chosen windows (1996, 2006, and 2016) and criteria (10% maximum of cloud cover, images without striping) led to the downloading of 105 images (35 images per year). During the pre-processing phase, the images were validated based on their geometric and spectral quality and underwent some geometric and atmospheric corrections. In some cases, in the absence of quality images for the relevant periods, mosaics of anterior or posterior images were built to complete the coverage. To improve the image classification, the SRTM Digital Elevation Model (DEM) (https://lta.cr.usgs.gov/srtm/mission_summary) was used to extract three layers (altitude, slope, and roughness) and to improve the indicators created from the LANDSAT primary channels.

The classification approach for the images is based on the RandomForest algorithm, coded by Brieman & Cukter (2005) scripted in the free software R developed by Liaw & Wiener (2002). The image classification was supervised using the vegetation map of the Royal Botanic Gardens (Moat & Smith, 2007), from which 11 classes were selected, including five forest classes (humid forest, dry forest, spiny forest, littoral forest, and mangrove) and six non-forest classes (fallow lands, wooded savannas, and croplands; grasslands and savannas; cloud; water; shadows; and rocks, sand, and bare rock). On the final maps, classes were simplified to highlight forest loss rather than the other land-use changes. Only the changes related to forest cover were considered within the classes forest (F) and non-forest (N), which resulted in the following three temporal scenes: FFF = forest in 1996, 2006, and 2016; FFN = forest in 1996 and 2006, and non-forest in 2016; and FNN = forest in 1996, non-forest in 2006 and 2016.

The post-processing phase involved some image smoothing and the suppression of random pixels using a filter of 3 x 3 pixels, as well as the removal of forest units of a surface area below 1 ha. The results were controlled and validated through an accuracy assessment, based on 1/3 of the test parcels, aiming at a minimum overall accuracy of 75%. A final improvement phase included verification on Google Earth and feedback from field and analysis experts, managers, biologists, or geomatics specialists.

The results led to the development of the deforestation maps and the tables of forest and deforested areas for each scene (1996, 2006, 2016), as well as the calculation of deforestation rate between each period (between each scene). The deforestation rate for a period is the ratio between the loss in hectares during the period and the initial forest surface in hectares. The annual deforestation rate is calculated by dividing this inter-period rate by 10 (10 years between each scene). These calculations were performed for each protected area using the official boundaries provided by the Système des Aires Protégées de Madagascar (SAPM).

Interpretation of results

In the site texts associated with this section header, short interpretations of the results are presented and focus on forest formations. In the associated tables, the reported figures are rounded to the nearest hectare, rather than using decimals. One repeated pattern is the degradation of natural forest as a direct result of timber and charcoal exploitation or from fires to rejuvenate pasture, which enter into the forest understory (Figure 106). This type of pressure opens the cycle of incursion of invasive species, followed by an increase in flammable biomass, erosion, and landslides; under these conditions, wildfires can then more readily enter into degraded habitats and result in deforestation.

Figure 106. Litter fire gradually extending into the undergrowth destroying the roots in the northwestern portion of COMATSA North (site 11) at about 2170 m altitude. This type of habitat destruction has not been widely recognized and might explain wide scale and recent degradation of montane areas where human populations are notably reduced. (Photo by Sébastien Wohlhauser.)

In the text for each protected area, figures are given on the