Strength in numbers: males in a carnivore grow bigger when they associate and hunt cooperatively

Video bigger penis on carnivore



The evolution of group-living from solitary ancestors remains a hot topic in behavioral ecology ( Port et al. 2011 ; Shultz et al. 2011 ; Ebensperger et al. 2012 ; Schradin et al. 2012 ). Sociality not only yields both benefits in terms of decreased predation risk and improved resource defense but also costs in the form of enhanced feeding and mating competition (reviewed in Krause and Ruxton 2002 ). According to socio-ecological theory ( Crook and Gartlan 1966 ; Macdonald 1983 ; Terborgh and Janson 1986 ; Ims 1988 ), female sociality is largely determined by the distribution and quality of food resources, whereas male sociality is mainly a response to female distribution. Because females compete with each other over food and males compete over females ( Darwin 1871 ; Emlen and Oring 1977 ; Clutton-Brock 1989 ; Andersson 1994 ), males rarely exhibit gregarious tendencies in species where females are solitary . However, such male associations have been described, inter alia, in some carnivores with solitary females (cheetahs [ Acinonyx jubatus ]: Caro and Collins 1987 ; slender mongooses [ Galerella sanguinea ]: Rood 1989 ; Waser et al. 1994 ; kinkajous [ Potos flavus ]: Kays and Gittleman 2001 ), where cooperation among allies can outweigh the costs of competition for access to females when cooperation serves joint defense of territory and females against other males ( Axelrod and Hamilton 1981 ; Caro and Collins 1987 ; Waser et al. 1994 ; Clutton-Brock 2009 ). The fact that male sociality is a very rare phenomenon indicates, however, that the related costs are presumably high.

Precursory forms of sociality, such as facultative sociality in carnivores (e.g., felids: Caro and Collins 1986 , 1987 ; Kruuk and Parish 1987 ; herpestids: Rood 1989 ; Waser et al. 1994 ; canids: Cavallini 1996 ; mustelids: procyonids: Gompper 1996 ; Kays and Gittleman 2001 ), represent an ideal test case to identify the forces that favored the evolutionary origins of group-living. We, therefore, studied the social system of fosas ( Cryptoprocta ferox , Eupleridae), the largest Malagasy carnivore, in a seasonal environment where intense competition for food resources is expected during large parts of the year for an exclusively carnivorous species ( Hawkins and Racey 2008 ). Adult female fosas are solitary, whereas males are either solitary or associate in pairs or trios (Lührs ML, Kappeler PM, unpublished data). Male associations are stable across seasons and years, and members of the same association have been observed to hunt cooperatively ( Lührs and Dammhahn 2010 ). The fosa’s mating system is unique among mammals in that it is characterized by seasonal aggregations of males at traditional mating trees ( Hawkins and Racey 2009 ), which are temporarily occupied by up to three females (Lührs ML, Kappeler PM, unpublished data). Large numbers of males accumulate during a female’s estrous period, leading to intense pre-copulatory contest competition ( Hawkins and Racey 2009 ) and supposedly intense sperm competition due to polyandrous matings (Lührs ML, Kappeler PM, unpublished data). Copulations are prolonged and involve a copulatory tie ( Albignac 1970 ; Hawkins and Racey 2009 ). Accordingly, males achieving longer copulations may gain an advantage in sperm competition by monopolizing access to the female for the duration of the copulatory tie and accordingly increased ejaculation frequencies ( Lanier et al. 1979 ; Oglesby et al. 1981 ).

The interdependencies between male sociality and the reversed lek-like mating system in this species remain unknown. Although contest competition may favor coalition formation, small litter-size and sperm competition imply high costs of sharing access to females by unrelated males, thereby constraining sociality. In order to determine why some males in this species exhibit permanent association despite high costs of competition for food and mates, we compared solitary and associated males with respect to their morphology, relatedness, ranging and hunting behavior, diet, and mating success. Using parallel GPS-tracking and accelerometry in combination with dietary analyses based on stable isotopes and direct behavioral observation, we aimed to illuminate 1) whether males differ morphologically depending on their type of social organization, 2) whether associated males hunt cooperatively, and if so, 3) whether there is a dietary divergence between males of different types of social organization. In light of the promiscuous mating system, we were further interested in 4) whether solitary and associated males differ in mating success, and 5) how associated males compensate for the costs of mating competition. Based on these data, we will discuss why two types of male social organization coexist and how this can be related to socio-ecological theory.


Study system

We collected morphometric data, tissue samples, ranging and activity data, hair samples, and behavioral data from a population of 33 wild fosas (25 males, 8 females [ nf ]) trapped between 2007 and 2010 in Kirindy Forest/CNFEREF (44°39′E, 20°03′S), western Madagascar. Fosas were trapped annually during the dry season (weeks 34-48) with 10 live-traps (42 × 15 × 20 in 3 bobcat trap; Tomahawk Live Trap, Hazelhurst, WI, USA) along transects that were shifted weekly over a 9 km 2 area in the forest center. Trapped animals were briefly anesthetized, measured, and sampled for tissue, and some adults were equipped with GPS-tags in individually designed collars (150-200g, e-obs GmbH, Gruenwald, Germany) that made up less than 5% of the individuals’ body mass ( Gannon and Sikes 2007 ). After recovery from anesthesia, animals were released at the site of capture.

Solitary males were distinguished from associated males based on direct sightings, trapping, and GPS-tracking data. In contrast to solitary males, associated males were sighted in stable dyads sharing home ranges, synchronizing their activity, and showing affiliative behavior (resting and sleeping in body contact, grooming) as well as occasional coalitional support, which has never been observed among any other type of fosa dyad.


Morphometric analyses of body length, absolute body mass, scaled body mass ( Peig and Green 2009 ), canine width, and testis volume were based on 8 solitary ( ns ) and 10 associated males ( na = 10; i.e., 5 dyads), with occasional exceptions where measures were missing (see Results for precise sample sizes). Body length was measured from the base of the skull to the base of tail. If several measures of fully grown individuals were available for consecutive years, values were averaged. To control for seasonal variation in body mass and testis volume, we included only data from males trapped between weeks 38 and 43 of a given year. Canine width was preferred over canine length as a proxy for age as it is more robust and less prone to variation due to dietary differences. Individuals included into the analyses were at least 5 years old as estimated from patterns of tooth eruption and tooth wear. Although a minimum age could be calibrated based on experience in captivity (Reiter J, personal communication), older individuals could be estimated in relation to each other only or based on canine width. Because all associated males could be trapped within one trapping period, associates could be directly compared based on tooth wear. One adult male was excluded from analyses on body mass and condition because it suffered from an unknown disease at the time of trapping.

Testis volume was calculated from testis length and width using an ellipsoid formula as a proxy: V = 4/3*π*(width/2)²*length/2. We determined both absolute and relative testis volume (residual volume after correlation with body length) to account for allometric effects.


Pairwise relatedness of associated males was determined using 16 microsatellite markers applied to a population of 33 individuals (see Supplementary Table S1 ). DNA was extracted from tissue samples using the Qiagen extraction kit for tissue. We used the 16 most polymorphic (5.1 alleles per locus on average) of 26 markers developed from a captive fosa population ( Vogler et al. 2009 ) and adapted PCR protocols to local laboratory conditions (see Supplementary Table S2 ). PCR products were detected on an ABI 377 DNA Sequencer and evaluated using the software GeneMapper. PCRs were repeated one to four times per individual at homozygous loci. Relatedness coefficients were calculated with the software COANCESTRY ( Wang 2011 ), which provides the triadic likelihood estimator “TrioML” ( Wang 2007 ); a robust estimator accounting for inbreeding and genotyping errors. Male dyads of equal age estimate were tested for being littermates (paternal half- or full-siblings) using the software COLONY ( Wang 2004 ). A subset of eight males was repeated from extraction onwards to replicate our results.

Spatial data

Ranging data were obtained from three solitary and four associated males (two dyads) tracked simultaneously during the month of October. Another two associated males (a third dyad) could not be tracked simultaneously in October and were therefore evaluated based on two different years. Data on association pattern (% time spent in close proximity to the association partner and average distance to the partner) were based for this dyad on simultaneous tracking in July, whereas equivalent data were available for the other dyads in October. GPS tags recorded locations at hourly intervals with occasional loss of positions due to signal disturbance. Home ranges were determined as 100% minimum convex polygons using the Animal Movement Extension ( Hooge and Eichenlaub 2000 ) in ArcView 3.3 (Esri, Redlands, CA, USA). Daily path length was calculated as average distance covered by an individual in a day in October.


Acceleration data were obtained from three-axes accelerometers embedded in the GPS-RF-collars (e-obs GmbH, Gruenwald, Germany), which sampled at 3-min intervals at 3 Hz with a resolution of 54 bytes. Absolute differences of acceleration measures were averaged for each sequence and categorized into classes of acceleration of ≥1 (class I), ≥2 (II), ≥4 (III), and ≥8g (IV). These categories were chosen as equivalent to the following activities (see Supplementary Material for details on calibration): head movements during resting—walking (class I), walking-slow running (II), running (III), and jumping-falling (IV). We then tested whether associated males synchronize their activities across all four of these acceleration classes, which for classes III and IV would imply joint fast running and jumping, that is, hunting or fighting. Matrices of pair-wise comparison of the two partners of an association counting occurrences of each partner in each class were determined for hours in which they were tracked in close proximity (≤100 m) and when separated (>100 m). We tested for deviations from random expectation probability matrices using a chi-squared test.

Stable isotope analyses

For dietary analyses, stable carbon and nitrogen isotope ratios were measured in hair samples of 6 associated males (3 dyads), 10 solitary males, 8 females, and 7 common mammalian prey species ( Hawkins and Racey 2008 ) using mass spectrometer analyses at the KOSI (Centre for Stable Isotope Research & Analysis, Göttingen, Germany). We assessed whether individual spread (Euclidean distances to the group centroid [CD]) in the δ 13 C-δ 15 N bi-plot differed between associated and solitary males by comparing mean group distances to null distributions generated by residual permutation procedure ( Turner et al. 2010 ). Using nearest-neighbor analyses ( Krebs 1999 ), we assessed whether associated males represent only a part of the trophic diversity of the population. Using Bayesian isotope mixing models ( Parnell et al. 2010 ), we estimated proportions of prey species in the diet of associated and solitary males.

Behavioral observations

Observational data were obtained by continuous behavioral sampling of mating activity of seven females (one female remaining unidentified) between 2007 and 2010. Six females were observed continuously over the course of their mating activity (530 observation hours) and one female was observed continuously for the last three nights of mating (30 observation hours), whereby length and number of copulations indicated that these included peak mating activity. Marked individuals could be identified by individual earmarks, tail shaving patterns, or GPS collars. Unmarked fosas could be distinguished by scars, natural earmarks, body size, and fur color. In order to assess differences in male mating success, we first investigated individual mean copulation duration based a subset of 207 matings where the social category of the mating male was known ( ns = 9, na = 7). Copulation duration was measured as the time between intromission and external emergence of the penis. We assumed mean copulation duration a proxy of a male’s contest competitive abilities because 70% of all matings were terminated by other males disturbing the mating couple (Lührs ML, Kappeler PM, unpublished data). Because a male’s likelihood of successful fertilization presumably increases with the total time, it manages to monopolize the female ( Altmann et al. 1996 ), that is, its total mating time per female and year, we further tested for differences in total monopolization duration for seven solitary and five associated males mating with five females for which continuous observational data were available. Two of the seven females in our sample mated in two different years of the study period and were annually regarded as independent samples because of inter-annual variability in male communities and the high importance of male contest competition in the system.

We used linear mixed models (LMMs) with female and male identity as random factors and male social organization (solitary, associated) as fixed factor for normally distributed response variables. For the analysis of single copulation duration, the activity period of the female (whether the copulation was observed when the female’s daily mating activity peaked or not) and the absolute number of heavier males present (in comparison to the mating male) were added as fixed factors to the model to account for the female’s receptivity and the potential for male-male competition. Copulation durations were log-transformed to normal distribution. For the analysis of total monopolization duration, a male’s total number of matings and the interaction of the latter with the type of social organization were added as fixed factors to the model to illuminate possible differences in the number of copulations needed for each group to achieve a certain mating time. Monopolization durations were z-transformed to account for differential monopolization potential of different females and female-years. This standardization was performed on the basis of the complete continuous raw data set, that is, including monopolization durations of all males present.

Model selection was based on AICs and normality of residual distribution. If not otherwise stated, all statistical calculations were performed in R 2.14.1 ( R Development Core Team 2011 ). P values for LMMs were calculated from 1000 Monte Carlo simulations. The accepted significance level was P < 0.05.


In total, we found 10 out of 22 adult males trapped to be associated in either dyads or triads (four dyads and one triad with a third individual remaining untrapped). Genetic analyses and age estimation based on tooth wear revealed that four out of six dyads in the population were composed of littermates (TrioML, r ≥ 0.40; see Supplementary Table S1 ). We could not detect potential littermates among solitary males in the population, but one litter of paternal half-siblings appeared to span two different associations (see Supplementary Table S1 ).

Simultaneous GPS tracking of associated males indicated a highly variable degree of association, ranging from low cohesion (21% of time tracked spent in close proximity of ≤50 m) to near full-time association (91% of time tracked in close proximity, Table 1 ). All individual ranges of associated males were about twice as large as those of solitary males (median minimum convex polygons: 61.7 vs. 33.0 km², na = 6, ns = 3; Table 1 ). However, daily travel distances within ranges were similar for associated and solitary males (median 6.5 vs. 5.3 km; Table 1 ), despite higher energy demands for male dyads. As this indicates that associated males manage to compensate for the costs of food competition, we explored morphological consequences in both types of males.

Morphometric analyses revealed that associated males were considerably larger and heavier than solitary males (mean body length 66.1±3.3cm vs. 59.1±2.2cm; Mann-Whitney U test, Z = 3.18, P = 0.001, na = 9, ns = 8; mean body mass 9.6±0.8kg vs. 7.3±0.7kg, Z = −3.12, P = 0.002, na = 9, ns = 6; Figure 1 ). Because both classes of males neither differed in body condition (scaled body mass Mi [ Peig and Green 2009 ], Mann-Whitney U test, Z = −0.24, P = 0.810, na = 9, ns = 8) nor in age (operationalized by canine width; Z = 0.00, na = 10, ns = 9, P = 1.000), there was a pronounced morphological difference between males of different type of social organization with correspondent consequences for the degree of sexual dimorphism in fosas. Although solitary males did not differ from females in body mass (Mann-Whitney U test, Z = 0.58, ns = 8, nf = 8, P = 0.564) and body size (body length: Z = 0.68, P = 0.495; Figure 1 ), associated males exhibited pronounced sexual dimorphism with females (body mass: Z = −2.65, P = 0.006, na = 9, nf = 8; body length: Z = −3.03, P = 0.002; Figure 1 ). In addition, associated males had higher absolute testis volume than solitary males (Mann-Whitney U test, Z = 2.65, na = 9, ns = 8, P = 0.008), whereas relative testis volume did not differ between the two groups ( Z = −0.47, P = 0.641). Given these results, we further explored whether the physical advantage of associated males could be mediated by benefits of cooperative hunting.

In order to quantify cooperative hunting, we determined the degree of synchrony between four classes of body acceleration among pairs of associated males and found that individuals of all three dyads synchronized their activity across all four classes when in close proximity (<100 m; χ2 -tests: P < 0.01 for all simultaneous occurrences across all classes), but not when they were spatially separated ( Table 2 ). If synchronized bouts of extreme acceleration (classes III and IV) were indeed related to cooperative hunting, we predicted associated males to be more similar in their diet than solitary males and to have higher proportions of more profitable prey (i.e., large lemurs) in their diet.

Comparison of time-integrated information based on stable carbon and nitrogen isotope analyses revealed that 1) associated males were more similar in their diet than solitary males and covered only a subset of the population’s isotopic space, reflected by a smaller mean Euclidean distance of individuals to the groups’ CDs in the δ 13 C-δ 15 N bi-plot (residual permutation procedure, | CDassociated males – CD solitary males | = 1.21, P = 0.001) and a clumped pattern of associated males in the δ 13 C-δ 15 N bi-plot (nearest neighbor analysis, index of aggregation R = 0.47, Z = −2.50, P = 0.012), whereas the distribution of solitary males did not deviate from a random pattern ( R = 0.73, Z = −1.66, P = 0.10; Figure 2 ). 2) Associated males had higher proportions of the locally largest lemur (≈ 3kg Propithecus verreauxi ) in their diet, whereas the most important prey species for solitary males were medium-sized lemurs and rodents (Bayesian isotope mixing models; Figure 3 ).

Based on the numerical and physical differences between solitary and associated males, we further predicted an advantage for associated males over solitary males in contest competition over access to females. We found higher individual copulation durations for associated males compared with solitary individuals (LMM, t = 2.01, P = 0.045, na = 7, ns = 9, nf = 7, 207 matings; Table 3a ), with a general increase in copulation length during a female’s peak activity ( t = 2.20, P = 0.029) but no effect for the number of heavier males present on the tree ( t = 0.78, P = 0.435). Within dyads, dominant individuals mated on average longer than their subordinate associates (median 81.3±27.5min vs. 49.4±31.1min, na = 6), whereas the latter achieved mating durations comparable to solitary males (median 34.3±66.4min, ns = 6).

Summing up copulations to total monopolization duration revealed that associated males monopolized females for longer than solitary males (LMM, t = 2.47, P = 0.019, na = 5, ns = 7, nf = 5, 178 matings; Table 3b , Figure 4 ). Because both male categories did not differ in the total number of copulations (Mann-Whitney U test, Z = −0.11, na = 5, ns = 7, P = 0.913), associated males tended to need fewer copulations to achieve comparable mating durations (LMM, t = −1.95, P = 0.059, Table 3b ). Furthermore, associated males interrupted matings of solitary males more often than vice versa (Mann-Whitney U test, Z = 2.01, na = 7, ns = 3, P = 0.045).


We showed that two male phenotypes can be distinguished in fosas, which differ in their type of social organization. Although solitary males reach similar size as adult females, associated males grow considerably larger and exhibit sexual size dimorphism with females. The present analyses indicate that this somatic advantage of associated males is linked to cooperative hunting, allowing them to take down larger-bodied and more agile prey. These benefits of cooperative hunting may compensate for the costs of sociality related to feeding competition. As a result of their physical superiority, associated males also gain reproductive advantages, if mating duration in fosas is positively correlated with fertilization success. Hence, their somatic advantage is likely to translate directly into reproductive benefits.

Depending on their type of social organization, male fosas may physically develop differentially with a pronounced advantage for associated males with respect to body mass and size. As this difference is not explained by age, it is most likely due to dietary divergence because associated males differ remarkably in their diet from solitary individuals of either sex. A combination of a higher proportion of larger-bodied prey in their diet and no difference in searching time (as reflected in daily travel distances) indicated higher foraging efficiency of associated males, which is most likely achieved by cooperative hunting. This hunting technique may allow associated males to hunt large lemurs at higher success rates than solitary males. In cheetahs, another hypercarnivorous specialized hunter, cooperative hunting has also been shown to allow associated males to hunt larger prey ( Caro 1994 ). Yet, an increase in prey intake rate per capita could not be shown for this species and thus male association in cheetahs has been attributed to benefits of cooperative territory defense rather than benefits of cooperative hunting ( Caro 1989 ).

Because there was no difference in condition between solitary and associated fosa males, both strategies most likely differ in their energy allocation to growth. As a result, two male phenotypes exist; solitary males, which do not exceed adult females in size and associated males that are on average 38% heavier and 13% larger than females. In meerkats, growth rates have been shown to be individually variable over even longer life trajectories with significant environmental influence ( English et al. 2012 ). In fosas, variability in growth patterns may be similarly related to highly seasonal environmental conditions in Madagascar, which has been linked to various behavioral, physiological, and morphological adaptations in other endemic taxa ( Wright 1999;,Dewar and Richard 2007 ). The fact that fosa females remain solitary indicates that the potential energetic costs of feeding competition for communally raising offspring are too high. Similar constraints seem to apply to males. Despite obvious benefits of sociality for each member of a male dyad, the costs of food sharing cannot be outweighed by any larger association, as indicated by the observation that each of the two male triads in this study comprised one individual that was in very poor condition (data not shown). Sociality in fosas may thus function as a means to overcome energy limitation. We, therefore, propose that fosas represent an example of a species, where energetic and nutritional factors have a tangible impact on male sociality.

Patterns of male sociality in fosas can contribute to a better understanding of the evolution of carnivore sociality and cooperation in general as the trade-off between the costs and benefits of group-living seems to offer very limited opportunities for sociality in this species. Intrasexual competition for reproductive opportunities is intense in this highly promiscuous mating system, raising the question of how associated males share limited mating opportunities. In four out of six associations, which were all composed of likely litter-mates, inclusive fitness benefits may compensate for the costs of mating competition ( Hamilton 1964 ). The only unrelated adult dyad in this study achieved generally high mean mating durations compared with the population average (92min for the dominant individual and 69min for the subordinate) and balanced mean monopolization durations per female (216min for the dominant individual and 207min for the subordinate), indicating both efficient monopolization of access to females as well as equitable share of mating between the two associates. Unrelated associates may, therefore, enjoy higher fertilization probabilities than solitary individuals due to longer monopolization times.

Given somatic and potentially reproductive benefits of male sociality in fosas, the persistence of solitary males raises the question whether these males are solitary because of a lack of opportunity (e.g., because they were the only male of their litter), or whether they may represent an alternative strategy maintained by frequency-dependent selection ( Gross 1996 ). Because we did not detect male littermates for any solitary male in our sample, they are likely constrained in their opportunities to associate with a familiar male. Interestingly, our genetic analyses revealed that male CF20 of the only unrelated adult association was closely related to the dyad CF27-CF31 but not to its associate (see Supplementary Table S1 ). CF20 is a confirmed ( P = 1.000) half-sibling of CF27 even though it could not be statistically confirmed as a half-sibling of CF31 ( P = 0.047). If CF20 is a third litter-mate of CF27 and CF31, early development in an all-male litter may have facilitated social tolerance toward other males in this case. A putative brother of its partner CF21 could not be found in the population, however, and the triadic relationship could not be resolved. The importance of familiarity and social tolerance for male sociality, therefore, remains speculative.

On the other hand, our results also indicate that solitary males remain small but do not suffer from reduced body condition, and that they seem to be highly diverse in their diet and hunting strategies (as indicated by a wide spread of solitary individuals in stable isotope space; Figure 2 ). Their small body size may allow solitary males to extract small lemurs from tree-holes or rodents from their burrows more effectively. When it comes to mating, however, solitary males face a significant disadvantage in contest competition. In some mammals, physically inferior males have been shown to invest more in sperm competition by producing more sperm ( Parker 1990 ; Stockley and Purvis 1993 ). In fosas, however, the two types of males appear to invest equally in relative testes size. Based on the currently available evidence, we conclude that the demographic constraint as a result of a lack of male littermates may, therefore, explain the existence of solitary males.


In summary, we described a rare phenomenon of male sociality with functional significance for growth and sexual dimorphism in a carnivore. Most likely, physical superiority of associated males over solitary individuals of either sex is determined by higher foraging efficiency due to cooperative hunting. Sociality is most common among littermates, however, as food limitation generally promotes a solitary lifestyle in this strictly carnivorous species, and social tolerance among unrelated males is likely constrained. Based on the link of facilitated access to food by cooperative hunting and subsequent mating benefits due to physical superiority, we suggest that food resources can also be important ultimate determinants of male sociality. Furthermore, this example indicates that cooperative hunting can precede group-living rather than being a consequence or a by-product of the latter ( Packer and Ruttan 1988 ) and can act as an evolutionary force promoting it ( Creel and Creel 1995 ).


Supplementary material can be found at Supplementary Data .

This work was supported by the Deutsche Forschungsgemeinschaft (DFG KA 1082/17-1), the Fossa Fund of Zoo Duisburg AG, and the German Primate Center GmbH (DPZ). We thank Rémy Ampataka, Tianasoa Andrianjanahary, Nielsen Rabarijaona, and Jean-Pierre Tolojanahary for field assistance; Léonard Razafimanantsoa, Rodin Rasoloarison, and Heike Klensang for administrative support; Reinhard Langel from the Centre for Stable Isotope Research & Analysis (KOSI) for technical help in the laboratory; Christina Oberdieck for support in the genetic laboratory; Cornelia Kraus for statistical advice; Elise Huchard for veterinary assistance and helpful discussions; Franz Kümmeth from e-obs GmbH for technical support and two anonymous reviewers for constructive comments on the manuscript. We thank Daniel Rakotondravony from the Département de Biologie Animale de l’Université d’Antananarivo, the Commission Tripartite CAFF, and CNFEREF Morondava for their authorization and support of this study. All research protocols were approved by the appropriate Animal Use and Care committees of Germany (Bundesministerium für Naturschutz, BfN) and Madagascar (Ministère de l’Environnement et des Eaux et Forêts, MINEEF). This study is in compliance with animal care regulations and applicable national laws of Germany and Madagascar. The authors declare no competing financial interests.


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