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Habitat fragmentation in Madagascar:

connecting fragmented findingsKATHERINE J. KLING1, KIMBERLY YAEGER2, AND PATRICIA C. WRIGHT2-4

1Stony Brook University, Interdepartmental Doctoral Program in Anthropological Sciences (IDPAS), Stony Brook, NY

2Stony Brook University, Department of Anthropology, Stony Brook, NY

3Stony Brook University, Institute for the Conservation of Tropical Environments, Stony Brook, NY

4Centre ValBio, Ranomafana, Madagascar

A systematic review of fragmentation studies in Madagascar, particularly

those examining lemur responses, was conducted with the following

objectives:

1. To quantify trends in the treatment of fragmentation and in the

reporting of responses of lemurs and other Malagasy taxa to

fragmentation

2. To characterize research effort across taxa, study region, and study

type

Systematic Review

― Literature search was conducted in both French and English

December 3, 2018 and April 20, 2019 to identify peer-reviewed

articles presenting research on fragmentation in Madagascar

(PRISMA guidelines for conducting literature review: [11])

― Web of Science, Google Scholar, and targeted journals searched

using search term: “Madagascar fragment*”

― From initial 5,849 records, 1,106 abstracts were reviewed manually

― Sources that did not conduct research exclusively in Madagascar

were excluded

― Further sources excluded that did not explicitly name at least one

study site as a fragment or fragmented forest in either Methods or

Results sections

Data Analyses

― Included sources split between those that statistically tested for

effects of fragmentation on study system (“Tested Articles” (TA) v.

“Untested Articles” (UA)) and for those that specifically conducted

research on lemurs (“Lemur Articles” (LA))

― Trends in included sources analyzed across following categories:- Study type, year published, geographic region, study taxon(a) and IUCN listing,

reporting of fragment metrics, stated test(s) of fragmentation, type of statistically-

tested results reported (positive, negative, not significant)

- Fragment metrics include: area of listed fragment(s), isolation distance from each

other and/or contiguous forest, # of fragments studied

- Primary classifications of tests of fragmentation on study system: by area,

isolation distance, and/or type (comparison of fragment(s) to control sites(s))

- Definition of positive, negative, and non-significant responses of study system to

fragmentation per Carretero-Pinzón [7]

― Data additionally collected regarding authorship and journals in which

studies were published (Fig. 1)

Introduction

Understanding fragmentation’s impact

― The rise in research investigating fragmentation reflects growing

presence of fragmented landscapes themselves [1]

― Biodiversity responses to fragmentation and anthropogenic habitat

change are difficult to categorize, ranging from negative [2] to positive

[2], even for the same species when in different contexts [3]

― Difficulty in defining fragmentation’s impact likely due to confounding

factors associated with fragmentation process (e.g. fragment area,

isolation distance, patch disturbance; [4])

Primates and fragmentation

― Non-human primates are especially susceptible to anthropogenic

landscape change [5]

― Growing research attention [6-7] has assessed primate response to

fragmentation: while no clear trend emerges, species richness tends

to decrease with declining fragment area [8]

Fragmentation in Madagascar

― Madagascar has an extensive and ongoing history of habitat loss and

fragmentation: est. 44% loss of forest cover from 1953-2014 [9]

― As world’s most imperiled vertebrate group [10], summative evaluation

of reported responses of the lemurs of Madagascar to fragmentation

is warranted

Study Objectives

Methods

Results

Summary and Conclusions

Works Cited

Acknowledgements

Literature search

― Search resulted in 67 articles testing impact of fragmentation in Madagascar

(TA), 42 of which focused on lemurs (LA)

― An additional 106 articles reported research within fragments in Madagascar

without explicitly testing the impact of fragmented (UA), 77 of which focused

on lemurs

Study Type, geographic regions, and taxa

― One third of LA presented genetic and biodiversity survey research each and

majority of research (85.7% of LA) concentrated in eastern Madagascar

(Fianarantsoa, Antsiranana, Toamasina, and Toliara provinces; Fig. 2)

― 453 species total were represented across sources, including 46 lemur

species, 80% of which were threatened by extinction (Fig. 3)

― 82.1% of TA included species threatened with extinction in research

― Ten introduced species (e.g. house rat, Rattus rattus, and domestic dog,

Canis familiaris) also covered by research, indicating presence in forest

fragments

Supported by a Graduate Council Fellowship (Graduate School of Stony Brook University) and an

NSF Graduate Research Fellowship (2016203971)

Photo Credits: Nick Garbutt: P. diadema, D. madagascariensis, P. simus; R. A. Mittermeier: I. indri;

Paul Bratescu: E. collaris; Andy Rouse: M. murinus. K. Kling: Fragmented landscape

Figure 2. Map of Madagascar with relative percentage of number of lemur studies (LA) per type

displayed per region. Madagascar and the study sample are divided over each of its six

administrative provinces: 1) Antananarivo Province, 2) Antsiranana Province, 3) Fianarantsoa

Province, 4) Mahajanga Province, 5) Toamasina Province, 6) Toliara Province. Total number of LA

per study type are as follows: Behavior, n = 8; Biodiversity Surveys, n = 14; Genetics: n = 14;

Health: n = 6; Modelling: n = 5; and Other, n = 4. N = 42 LA studies total.

Figure 3. Most-commonly studied lemur species across articles, A) diademed sifaka (Propithecus

diadema; n = 8 studies), B) the collared brown lemur (Eulemur collaris; n = 6 studies), and C) from

top to bottom all n = 4 studies: the grey mouse lemur (Microcebus murinus), greater bamboo lemur

(Prolemur simus), aye-aye (Daubentonia madagascariensis), and indri (Indri indri). Photo credits in

Acknowledgements.

― Fragmentation research in Madagascar covers a wealth of taxa, with

individual studies predominantly focusing on lemurs

― Variable reporting of responses by lemurs to fragmentation results in

difficulty in generalizing its effects

- Wide variety of fragment metrics employed across examined

studies, an unclear distinction between ‘fragment’ and ‘control’

forests, and breadth of tests employed indicates challenge of

working across non-standardized definitions of fragmentation [4]

― The majority of studies report negative responses of Malagasy

biodiversity to fragmentation, yet few study fragmentation per se― Sensu Fahrig [13], fragmentation per se involves testing the impacts of

fragmentation itself, independent of patch size and isolation

― Two studies in this sample examined fragmentation per se via modelling

techniques to determine the relative impact of factors such as patch size, removal,

and isolation [14-15]

― This study emphasizes importance of continued examination of

impact of fragmentation on biodiversity, while encouraging such

research to employ explicit definitions of terms related to

fragmentation and to use comparable testing methods

Kling

Figure 1. All key words that appeared two or more times across TA (n = 59 studies). Key

words are sized according to the number of times they appeared (most frequent:

“fragment*,” 38 times). Note that the stems of words were analyzed to match derivations

of the same concept (e.g. “connect*” could be grouped with “connectivity” or

“connection”).

Defining fragments and fragmentation

― TA conducted research over an average of 8 ± 8.8 fragments each; 5 studies

did not indicate number of fragments studied

― Isolation distances reported (n = 11 studies) ranged from 15-219,000 m

between fragments and/or nearest continuous forest

― Fragment area ranged from 0.16-11,200 ha (five orders of magnitude

difference; median: 57.1 ha; n = 35 studies)

― Control area ranged from 297-104,000 ha (median: 4,143 ha; n = 9 studies)

and did not differ significantly from fragment area (t(12.1) = -2.0, p = 0.07;

Fig. 4)

Tests and impact of fragmentation

― Impact of fragment size most commonly tested (49.3% TA), followed by

studies comparing fragments to one or more control sites (32.8%; note: some

studies applied more than one category of test)

― 23.9% each of TA tested fragmentation by distance and other means (e.g.

consideration of whether land cover types were isolated or connected,

simulated removal of fragments, spatial transformation of landscape across

time)

― For LA, 68.1% of studies reported one or more significantly negative response

of taxa to fragmentation (n = 32 LA; Fig. 5) while only one reported a positive

impact [12]

Figure 4. Area (in hectares) of

study land units by author-

defined type (fragment vs.

control) for total articles (TA)

testing fragmentation by type.

Dark lines indicate means.

Boxes represent standard

error and whiskers standard

deviation. Note all land units,

even replicates used across

studies, are included to

represent what is tested and

reported for overall research

on fragmentation in

Madagascar. N = 187

fragments; 13 control sites. N

= 8 studies (TA).

Figure 3. Number of

lemur-specific sources

(LA) that list the presence

of one or more positive,

not significant, or negative

result as an effect of

fragmentation on study

variables. Sources that

indicated ‘Mixed

responses’ had results of

more than one type

(positive, not significant,

negative). Study that

included ‘Only this

response,’ reported only

responses in one

category. Mixed

responses, n = 17; Only

this response, n = 25.

A

B

C

Figure 6. Fragmented

landscape in southeastern

Madagascar.