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Behaviour and ecology of the fossa, Cryptoprocta ferox (Carnivora: Viverridae) in a dry deciduous forest, western Madagascar.
Abstract
Thesis (Ph. D.)--University of Aberdeen, 1998.
... Fossa are mainly solitary [21] but have been reported to socially hunt in pairs [22]. Fossa have large home ranges, traveling up to 5-7 km per day [23][24][25]; males occupy larger territories (>20 km 2 ), which overlap with other males, and females inhabit exclusive ranges of 7-13 km 2 [26]. Despite their wide distribution in Madagascar's western and eastern forests, population densities are lower than predicted by body size [26]. ...
... Fossa have large home ranges, traveling up to 5-7 km per day [23][24][25]; males occupy larger territories (>20 km 2 ), which overlap with other males, and females inhabit exclusive ranges of 7-13 km 2 [26]. Despite their wide distribution in Madagascar's western and eastern forests, population densities are lower than predicted by body size [26]. Fossa utilize a unique mating system where females will monopolize a site (e.g., high up in a tree) and maximize the available number of mates [26,27]. ...
... Despite their wide distribution in Madagascar's western and eastern forests, population densities are lower than predicted by body size [26]. Fossa utilize a unique mating system where females will monopolize a site (e.g., high up in a tree) and maximize the available number of mates [26,27]. Transient masculinization, defined as the temporary enlargement of the spinescent clitoris supported by an os clitoridis (baubellum) of young females 1-2 years of age has been described by Hawkins [28]. ...
The fossa is a specialized Malagasy carnivore housed in ex situ facilities since the late 19th century. Moderate breeding success has occurred since the 1970s, and welfare issues (notably stereotypic pacing behaviour) are commonly documented. To understand challenges relating to fossa housing and husbandry (H&H) across global facilities and to identify areas of good practice that dovetail with available husbandry standards, a survey was distributed to ZIMS-registered zoos in 2017. Results showed that outdoor housing area and volume varied greatly across facilities, the majority of fossa expressed unnatural behaviours, with pacing behaviour the most frequently observed. All fossa received enrichment, and most had public access restricted to one or two sides of the enclosure. The majority of fossa were locked in/out as part of their daily management and forty-one percent of the fossa surveyed as breeding individuals bred at the zoo. Dense cover within an enclosure, restricted public viewing areas, a variable feeding schedule and limited view of another species from the fossa exhibit appear to reduce the risk of unnatural behavior being performed. The achievement of best practice fossa husbandry may be a challenge due to its specialized ecology, the limited wild information guiding captive care, and the range of housing dimensions and exhibit features provided by zoos that makes identification of standardized practices difficult. We recommended that holders evaluate how and when enrichment is provided and assess what they are providing for environmental complexity as well as consider how the public views their fossa.
... The unique forested ecosystems of Madagascar are threatened due to a number of anthropogenic factors including deforestation, population growth, and introduction of exotic species (Harper et al. 2007, Zinner et al. 2014, Schüßlera et al. 2018. Within many of the island's forested 35 ecosystems, including the spiny forests, eastern rainforests, and dry deciduous forests, the fosa (Cryptoprocta ferox) serves as an apex predator, consuming lemurs and exhibiting extraordinarily (Hawkins 1998, Lührs & Kappeler 2013, with males exhibiting spatial overlap with females and other males, and females occupying ranges that overlap with males but not other females (Hawkins 2003). A flexible cathemeral activity pattern has been suggested for fosas, with a slight preference for crepuscular hours (Albignac 1973, Hawkins 1998, Gerber et al. 2012). ...
... Within many of the island's forested 35 ecosystems, including the spiny forests, eastern rainforests, and dry deciduous forests, the fosa (Cryptoprocta ferox) serves as an apex predator, consuming lemurs and exhibiting extraordinarily (Hawkins 1998, Lührs & Kappeler 2013, with males exhibiting spatial overlap with females and other males, and females occupying ranges that overlap with males but not other females (Hawkins 2003). A flexible cathemeral activity pattern has been suggested for fosas, with a slight preference for crepuscular hours (Albignac 1973, Hawkins 1998, Gerber et al. 2012). 45 Fosas are currently classified as vulnerable by IUCN for a combination of reasons (Hawkins 2016). ...
... to exhibit a flexible cathemeral activity pattern with a preference for crepuscular hours, as documented in previous studies conducted elsewhere (Albignac 1973, Hawkins 1998 ...
The Vulnerable fosa Cryptoprocta ferox is the largest native carnivore in Madagascar, fulfilling a unique ecological niche in the island's remaining forests. Negative interactions with humans threaten the long-term viability of most remaining fosa populations across Madagascar. Threats to the fosa include habitat loss and persecution by humans resulting from perceived predation on domestic animals. We used GPS collars to record space use and activity patterns of five fosas in Ankarafantsika National Park, Madagascar, during the dry seasons of 2016 and 2017. The results, with up to 2,110 recorded locations per individual, indicated fosas’ home ranges and movements were not limited to the forest, and all collared individuals used networks of habitat patches and corridors to navigate deforested areas. The fosas studied in Ankarafantsika National Park had significantly larger home ranges than those reported in previous studies in other protected areas. They were rarely found within village boundaries and appeared to avoid areas of human habitation, suggesting that during the study period livestock was not a significant component of the fosas’ diet in this Park. Our results suggest that fosas have some flexibility that enables them to adapt to living near deforested and human-dominated areas by altering their space-use patterns, but they are compensating by increasing their home range size.
... Madagascar's carnivores, the Eupleridae, have been described as one of the most understudied and threatened group of carnivores worldwide [8]. The Eupleridae's largest carnivore, the fosa (Cryptoprocta ferox) weighs on average 6-7 kg [9,10] and are found across Madagascar's forested landscape. They are thought to be a keystone species due to their predation of lemurs and other species [11,12]. ...
... Fosas have reportedly been killed for straying into villages where they may be perceived to be a threat to poultry [9,18,19] and other small livestock such as lambs [20]. Recent research in villages surrounding Ranomafana National Park [21] suggests fosas are not believed to affect many poultry keepers living at distance from the forest, with less than 1% of poultry predation attributed to fosas. ...
... Recent estimates from Ranomafana suggested that < 1% of households' poultry mortality was attributable to fosas [21], supporting historical reports of anecdotally infrequent attacks [9,18]. By contrast, approximately one in six interviewees' in our study reported fosa depredation. ...
Fosas (Cryptoprocta ferox) are Madagascar’s largest carnivores, occupying much of the island’s forested landscape. This study provides the first evaluation of fosas’ conflict with humans, a problem for many small and medium sized carnivores worldwide. We examined fosas’ predation of poultry, and the subsequent retaliatory killing. Over 1750 households were interviewed across four regions, encompassing Madagascar’s major forest types (deciduous/rainforest) and protected area classifications (national park, reserve and unprotected forest). Predation by fosa was the third highest reported cause (15%) of poultry mortality, with little evidence that coops were effective in reducing predation. Predation of poultry was more prevalent in deciduous forests, and most common during the evenings of the dry season. Over half of all interviewees said they disliked fosas, with loss of poultry the most commonly stated reason. Respondents’ that had suffered poultry depredation and those with lower educational attainment were more likely to dislike fosas. Interviewees that disliked fosas and those that were wealthier were most likely to report having killed a fosa. A minimum of thirty fosas was killed in retaliation by our respondents during the year before the interviews. Given that the fosa population is in decline, and most of Madagascar’s forests are likely to be too small to support sustainable populations, these killings may be detrimental to vulnerable sub-populations. These results shed insight into the cultural perceptions and predation patterns of a medium sized carnivore, with relevance to worldwide human-wildlife conflict of often overlooked smaller carnivores. We suggest that educational programs, guard dogs, poultry disease vaccinations and robust coop construction may be effective for improving attitudes and reducing retaliatory killing.
... Trotzdem zeigen Studien in Kirindy und Ampijoroa, daß Graue Mausmakis Höhlen in Totholz präferieren (Schmid 1997, Radespiel 1998, Ehresmann 2000 , Ehresmann 2000, Schmelting 2000. Außerdem sind die von Weibchen genutzten Schlafplätze besser thermisch isoliert (Radespiel 1998 (Goodman et al. 1993, Hawkins 1998 (Anderson 1984, Kappeler 1998 (Barre et al. 1988, Pagès-Feuillade 1988, Fietz 1995, Peters 1999, Sarikaya 1999, während direkte Begegnungen eher selten waren (Barre et al. 1988). Bei den Männchen unterlag die Aktionsraumgröße einer starken jahreszeitlichen Dynamik im Zusammenhang mit der Reproduktion, während sie bei den Weibchen relativ konstant in ihrer Größe war , Ehresmann 2000, Schmelting 2000. ...
... Im Verlauf der Regenzeit nimmt dann das Nahrungsangebot stetig zu, um gegen Ende wieder abzunehmen , Ganzhorn 1996, Sorg & Rohner 1996, Ganzhorn et al. 1999a, Ganzhorn et al. 1999b. Als Anpassung an die Zeit der Nahrungsknappheit während der Trockenzeit wird der Trockenschlaf der Fettschwanzmakis (Cheirogaleus medius) und der saisonale Torpor weiblicher Mausmakis in Kirindy interpretiert (Petter et al. 1977, Wright & Martin 1995, Schmid & Kappeler 1998 Mausmakis sind einem extrem hohen Prädationsdruck ausgesetzt, so daß Prädation zu einem Rückgang der gefangenen Individuen beitragen könnte (Goodman et al. 1993, Hawkins 1998 (Hawkins 1998). ...
... Im Verlauf der Regenzeit nimmt dann das Nahrungsangebot stetig zu, um gegen Ende wieder abzunehmen , Ganzhorn 1996, Sorg & Rohner 1996, Ganzhorn et al. 1999a, Ganzhorn et al. 1999b. Als Anpassung an die Zeit der Nahrungsknappheit während der Trockenzeit wird der Trockenschlaf der Fettschwanzmakis (Cheirogaleus medius) und der saisonale Torpor weiblicher Mausmakis in Kirindy interpretiert (Petter et al. 1977, Wright & Martin 1995, Schmid & Kappeler 1998 Mausmakis sind einem extrem hohen Prädationsdruck ausgesetzt, so daß Prädation zu einem Rückgang der gefangenen Individuen beitragen könnte (Goodman et al. 1993, Hawkins 1998 (Hawkins 1998). ...
... External measurements were taken by two different persons from the right side of the body (if paired) using calipers or a fl exible measuring tape. Table 1 lists the morphometric measurements taken following Hawkins (1998) . The age of captured animals was estimated based on a combination of body size, tooth wear and in males, testicle size. ...
... The new fi gures might be due to the use of larger traps facilitating the capture of larger individuals in the present study. According to these measurements and an average body mass of 1500 g, G. grandidieri is Madagascar ' s third largest endemic carnivore after Cryptoprocta ferox (Hawkins 1998 ) and Fossa fossana (Kerridge et al. 2003, summarized by Goodman, Ganzhorn and Rakotondravony 2003, Goodman 2009 ). ...
... The sexual dimorphism with higher body mass of adult males than females is typical for mongoose species (Ewer 1973 ). Sexual dimorphism is pronounced in Cryptoprocta ferox (Hawkins 1998 , Hawkins andRacey 2005 ) but absent for other Malagasy carnivores as hitherto reported (Razafi manantsoa 2003, Goodman 2009 ). In most animals sexual size dimorphism indicates a promiscuous mating system (Kenagy and Trombulak 1986, Heske and Ostfeld 1990, Kappeler 1993. ...
Galidictis grandidieri (Wozencraft 1986) is a nocturnal carnivoran endemic to the threatened spiny forest ecosystem of the Mahafaly Plateau in southern Madagascar. Previous investigations estimated a total population size of about 3000 individuals restricted to an area of 440 squkm, making it one of the geographically most limited and rarest mammalian carnivoran species worldwide. Given the increasing threat through habitat destruction we compiled additional mor- phometric measurements and investigated the distribution and abundance of the species, using grid and transect cap- tures. G. grandidieri were larger than indicated by previous samples and showed signifi cant sexual dimorphism with a mean body mass of 1640 g for males and 1400 g for females (overall mean: 1500 g). The highest densities were found in the littoral forest at the western edge of the Mahafaly Plateau (six to eight individuals/squkm). From there, its abundance declined exponentially towards the east. The distribution of G. grandidieri is probably determined by decreasing water accessibility away from the cliff. We estimate a total distri- bution area of about 1500 squkm and a total population size
between 3115 and 4995 animals. Based on these estimates, G. grandidieri occurs in a much larger area than assumed so far, but reaches lower densities.
... Forest fragmentation decreases habitat quality (Harper et al. 2007), impedes gene flow (Craul et al. 2009), and even exacerbates predation events on lemurs (Irwin et al. 2009), thus studies investigating habitat fragmentation from a broad ecological standpoint are desperately needed. To date, a handful of studies have been conducted on the food habits (Hawkins and Racey 2008; Dollar et al. 2006; Goodman et al. 1997; Wright 1997; Rasoloarison et al. 1995), activity patterns (Dollar et al. 1999; Hawkins 1998; Wright 1997), and population density (Hawkins 1998) of the largest endemic carnivore, the fosa (Cryptoprocta ferox). Most of these studies come from western deciduous forest with little data collected concerning C. ferox population density, sex ratio, activity patterns, and home range within eastern rainforest habitat where the most dramatic rates of deforestation are occurring. ...
... Forest fragmentation decreases habitat quality (Harper et al. 2007), impedes gene flow (Craul et al. 2009), and even exacerbates predation events on lemurs (Irwin et al. 2009), thus studies investigating habitat fragmentation from a broad ecological standpoint are desperately needed. To date, a handful of studies have been conducted on the food habits (Hawkins and Racey 2008; Dollar et al. 2006; Goodman et al. 1997; Wright 1997; Rasoloarison et al. 1995), activity patterns (Dollar et al. 1999; Hawkins 1998; Wright 1997), and population density (Hawkins 1998) of the largest endemic carnivore, the fosa (Cryptoprocta ferox). Most of these studies come from western deciduous forest with little data collected concerning C. ferox population density, sex ratio, activity patterns, and home range within eastern rainforest habitat where the most dramatic rates of deforestation are occurring. ...
... currently classified by IUCN as Vulnerable (Hawkins and Dollar 2009), live in low population densities throughout Madagascar, with an estimated 0.26 individuals per km² in deciduous forest (Hawkins 1998) and 0.17 per km² in eastern rainforest (Gerber et al. 2010). Pilot study work in the eastern rainforests of Ranomafana National Park indicate not only low densities of this top predator within the protected area, but also a complete absence of the C. ferox from fragmented forest areas < 25 km from the park (Gerber and Karpanty, unpublished data). ...
Of the unique wildlife in Madagascar carnivores are important as they may exert significant pressure on ecosystem structure; however, a paucity of information currently exists for these endemic species. The Masoala-Makira landscape, Madagascar’s largest protected area complex, has the highest levels of biodiversity and safeguards 6 of Madagascar’s 8 carnivore species. We provide the results of a pilot study using camera trapping methods to investigate carnivore population state variables across the Makira Protected Area. From September 2008 through November 2009 we conducted camera trapping surveys across three study sites (Anjanaharibe, Soavera, and Vinanibe), having varying degrees of fragmentation, human encroachment, poaching, and invasive species. We found that carnivore trapping rates varied significantly across the three study sites. More specifically, sites having increases in fragmentation, human and invasive trapping rates, and poaching rates (i.e. Vinanibe) had the lowest carnivore trapping rates, including the lowest occupancy estimates for Cryptoprocta ferox and Fossa fossana. Trapping rates for all six carnivore species were negatively correlated with human and invasive species trapping rates, though only F. fossana was found to be significant. MMDM density estimates for F. fossana at the Anjanaharibe site (2.71 ± 0.39) were analogous to recent estimates from populations at Ranomafana National Park. In this report we provide the first assessment of how fragmentation, human encroachment, invasive species, and poaching influence Madagascar’s carnivore populations. The results of this study highlight the growing threat to Madagascar’s biodiversity resulting from the on-going trends in habitat fragmentation and poaching, particularly in eastern rainforest habitat.
... The fossa Cryptoprocta ferox is an Endangered (IUCN, 2003), solitary viverrid with both arboreal and terrestrial habits, found in undisturbed forests throughout Madagascar, where it is endemic (Hawkins, 2003). It is the island's largest terrestrial animal species, with a mean body mass of 7.4 kg (n = 17, SD = 1.2) in males and 6.1 kg (n = 11, SD = 0.6) in females (Hawkins, 1998). The IUCN Mustelid and Viverrid Action Plan recommended field research to assess the population size of the fossa within Madagascar's conservation areas (Schreiber et al., 1989). ...
... The IUCN Mustelid and Viverrid Action Plan recommended field research to assess the population size of the fossa within Madagascar's conservation areas (Schreiber et al., 1989). Its size and highly carnivorous diet (Albignac, 1973;Rasolonandrasana, 1994;Rasoloarison et al., 1995;Hawkins, 1998) suggest that fossa population densities are likely to be lower than those of any other terrestrial species in Madagascar. These populations may therefore be the most sensitive to habitat fragmentation and to the size of protected areas. ...
... Fossas were tracked for periods ranging from 6 weeks to 13 months between 1995 and 1996. Fixes were taken a minimum of 8 hours apart, evenly spread across day and night because fossa activity was unaffected by time of day (Hawkins, 1998). As a result of the distribution of the roads in Kirindy and the transmitter signal range, it was possible to radio-track fossas reliably within a total area of 56 km2. ...
The widespread geographical distributions of mammalian carnivores such as the Carnivora and the Dasyuridae have often been erroneously equated with abundance. Their low densities and high demands on habitat area can render mammalian carnivores especially vulnerable to habitat fragmentation and destruction. The fossa Cryptoprocta
ferox (Viverridae) is a mammalian carnivore threatened by the rapid loss of Madagascar's forests, to which it is endemic. A 3-year mark-recapture study, comprising four censuses, generated an estimate of fossa population density at 0.18 adults km−2, or 0.26 individuals km−2. This was supported by a similar estimate from home range data. The fossa is thought to be unusually common in the study area, yet the estimated density was lower than that predicted for a typical tropical carnivorous mammal of the body mass of a fossa. Ecologists are frequently under pressure to provide estimates of local and global population numbers of their study species; we discuss the numerous factors that limit our ability to do this on the basis of a single population estimate. Nonetheless, our findings are sufficient to suggest that none of Madagascar's 46 protected areas supports a viable population of fossas, indicating a demand for corridors and enlarged reserves to ensure this species' long-term survival. Loss of the top predator can have a knock-on effect on an ecosystem. The findings indicate that, to maintain intact tropical forest ecosystems, it may be essential to consider the requirements of their often little-known mammalian carnivores. These requirements could be far greater than previously assumed.
... Soderquist & Ealey, 1994). This large (head-body length c. 70 cm) euplerid (Hawkins, 1998), endemic to the forests of Madagascar, was reported to exhibit an unusual, relatively easily observed mating behaviour. Reports from captive (Vosseler, 1929) and wild observations (Albignac, 1970;C. ...
... Clutton-Brock's (1989) scheme predicts male fossas to exhibit facultative monogamy/polygyny, or a 'staying' strategy, on the basis of the defensibility of female ranges by males. While defensibility of range is more or less impossible to measure, a recent study (Hawkins, 1998) found the ranges of female fossas to be exclusive, and therefore presumably defensible by males. As further evidence for this, males appear capable of territoriality, scent-marking frequently throughout the year in captivity (Albignac, 1973;Klock-enk¨amper, 1988), and occupy ranges more than twice the size of those of females (Hawkins, 1998). ...
... While defensibility of range is more or less impossible to measure, a recent study (Hawkins, 1998) found the ranges of female fossas to be exclusive, and therefore presumably defensible by males. As further evidence for this, males appear capable of territoriality, scent-marking frequently throughout the year in captivity (Albignac, 1973;Klock-enk¨amper, 1988), and occupy ranges more than twice the size of those of females (Hawkins, 1998). Sandell's (1989) argument, on the other hand, predicts fossa males to exhibit 'roaming', because fossa population density relative to body size (Hawkins & Raccy, 2005) falls well below the regression line for solitary carnivores (Sandell, 1989) as well as for tropical carnivorous mammals (Peters, 1983). ...
The mating strategies of male mammals have long been treated as broadly predictable on the basis of just two factors: the dispersion of females and the benefit of paternal care to male reproductive success. Female strategies and finer scale variations in mating systems remain poorly understood. In the fossa Cryptoprocta ferox, we had the rare opportunity to examine the mating system of a wild solitary carnivore directly, and identified features not classified or predicted by mating system theory. Males competed for mating opportunities at a traditional site monopolized by a female, high in a tree. The female mated with multiple males, repeatedly mated with some individuals and appeared to express mate choice. We observed three females thus, one replacing another on the site after each was seen to mate with four to five males over a period of 1–6 days. Copulations were prolonged (up to 3 h 8 min), involving a weak copulatory tie, and males appeared to guard females briefly after mating. Fossas are at low population density and do not use a den regularly; we suggest that both these factors impede individuals from locating a mate. We hypothesize that the observed mating system reduces this problem for both sexes, and increases the number of mates available to a female while ensuring a low risk of sexual harassment.
... The modern native Carnivora fauna of Madagascar is composed of eight species, the largest of which is Cryptoprocta ferox Bennett, 1833. This animal shows sexual dimorphism in body mass with adult males reaching between 6.2-8.6 kg and adult females between 5.5-6.7 kg; average body mass across adults is 6.8 kg (Hawkins 1998(Hawkins , 2003. C. ferox occurs in a variety of natural and human modified habitats. ...
... On the basis of comparisons he made, it appears that a maximum of three skeletons of recent C. ferox were available to him. Given the range of sexual dimorphism in this species (Hawkins 1998) and the possibility of geographic variation associated with the island's dramatic bioclimatic zonation (Ljungquist 1930), this number of specimens was insufficient to capture the possible range of variation in this species. Subsequently certain authors have considered C. spelea to be a synonym of C. ferox (Savage 1978;Köhncke & Leonhardt 1986). ...
... In a few cases individuals classified as sub-adults fell within the measurement range of adults, but in order to maintain a rigid separation of the age classes it was decided not to include these individuals in the analysis -this had the consequence of reducing sample sizes. Hawkins (1998) has already established that adult male C. ferox is larger on average than adult females based on a series of external morphological measurements. For most of her linear measurements males tended to be less than 10% larger than females. ...
Grandidier (1902) described a large form of Cryptoprocta ferox Bennett, 1833, a genus of Carnivora endemic to Madagascar, from subfossil material dating from the presumed Holocene that he referred to the variety spelea. Subsequently there has been varying opinion on the validity of this taxon. In this paper 159 subfossil and 32 modern osteological specimens of Cryptoprocta are examined and analyzed to determine if indeed two separate forms can be morphologically distinguished within the sample. On the basis of these tests we conclude that C. spelea Grandidier, 1902 is a valid species that existed on the island in the recent geological past. It was presumably a formidable predator that may well have feed on a variety of large lemur species that are now extinct.
... Masculinization in 2 female museum specimens of the fossa (Cryptoprocta ferox) was described early in the last century [17,18]. The fossa is a large, solitary civet endemic to Madagascar, with a head-body length of approximately 70 cm [19]. To confirm these reports, we examined genital morphology in individuals of both sexes and a range of ages from a wild population and a captive group. ...
... There is, as yet, no reported evidence in the sparse literature that increased aggression confers higher fitness in female fossas. Adult females appear to have home ranges that are more exclusive than those of adult males [19], suggesting that females may be more territorial, but this territoriality does not appear to be extreme. We found no other information on fossa aggression levels, other than from workers at Duisburg Zoo (A. ...
... If androgen levels are not raised, the alternative explanation is that there are direct benefits of the masculinization. However, the benefits so far proposed (participation in the greeting ceremony, female dominance) apply to females in the context of a socially structured group, whereas the fossa is solitary [19]. Furthermore, the benefit that East et al. [13] proposed for female hyaenas, that the resulting impossibility of rape leads to beneficially submissive behavior by males, could not be expected in the fossa. ...
In at least 9 mammalian species, females are masculinized throughout life, but the benefits of this remain unclear despite decades of thorough study, in particular of the spotted hyaena (Crocuta crocuta) in which the phenomenon has been associated with a high fitness cost. Through examination of wild and captive fossas (Cryptoprocta ferox, Viverridae), androgen assays, and DNA typing for confirmation of gender, we made the first discovery of transient masculinization of a female mammal. Juvenile female fossas exhibited an enlarged, spinescent clitoris supported by an os clitoridis and a pigmented secretion on the underpart fur that in adults was confined to males. These features appeared to diminish with age. The majority of adult females lacked them, and os clitoridis length was inversely related to head-body length. No evidence was found to link this masculinization to elevated female androgen levels. Circulating concentrations of testosterone and androstenedione, but not dihydrotestosterone, were significantly lower in females than in males. No significant differences in testosterone, androstenedione, or dihydrotestosterone levels were found between juvenile (masculinized) and adult (nonmasculinized) females. There are several possible physiological mechanisms for this masculinization. None of the hypotheses so far proposed to explain the evolutionary basis of female masculinization in mammals are applicable to our findings. We present 2 new hypotheses for testing and development.
... Covering 64 km 2 , Andranomena Special Reserve (ASR) is situated approximately 30 km north of Morondava. Two species of euplerids persist, the fosa and the endangered Bokiboky Mungotictis decemlineata (Hawkins, 1998;Razafimanantsoa, 2003). The camera-trap grid in ASR covered 35.45 km 2 of mostly contiguous forest, although sections of ASR were experiencing high levels of illegal logging during surveys. ...
... Of particular concern are free-ranging, mostly feral cats, a formidable worldwide invasive predator (Medina et al., 2011;Nogales et al., 2013) and a recognized predator of lemurs (Brockman et al., 2008) and other endemic species (Merson, 2018). At ANP, feral cats have been recorded to weigh 2.9-5.5 kg (Brockman et al., 2008), significantly larger than local domestic cats and less than the average adult 6-7.4 kg female and male fosa (Hawkins, 1998;Dollar, 2006). Presumably, fosas' larger size would mostly prohibit interference competition, with male adult fosas (weighing up to and over 11 kg) potentially capable of consuming smaller cats. ...
Western Madagascar's deciduous forest is an important, threatened habitat to little-known Eupleridae, such as the fosa Cryptoprocta ferox. Using camera-trap grids established in two important deciduous forests, Ankarafantsika National Park and Andranomena Special Reserve, we explore the activity patterns of endemic and exotic carnivores and evaluate how exotics may be influencing fosa temporal activity across these two protected areas. Three methodologies, kernel density, circular and wave analysis, were used to evaluate the activity pattern overlap between exotic free-ranging cats Felis sp. and dogs Canis familiaris, exotic small Indian civets Viverricula indica and the endemic fosa. Our results showed fosas to be cathemeral, peaking in activity in the evening and at dawn. Cats were nocturnal, whilst also highly active at dusk. Civets did not display a dominant period of activity, but were relatively more active in the night, and at dusk. In contrast, dogs were diurnal and also active at dawn, in alignment with human activity. Fosa activity significantly overlapped with the nocturnal cat and civet, whilst being distinct from the diurnal dog and humans. Our results suggest that there is potentially a competitive impact of exotic free-ranging dogs and cats when present across fosas’ dry forest habitat. This highlights the need for further examination into the impact of Madagascar's exotic carnivores, and the experimental trial to investigate the impact of their removal (or reduced abundance), on Madagascar's wildlife.
... These crepuscular peaks in activity could result from an adaptive response to fossa predation that could ultimately explain the evolution of cathemeral activity patterns. Although cathemeral, recent evidence suggests that the fossa exhibits a peak in activity during the middle of the nocturnal phase [Hawkins, 1998] . As a consequence , risk of predation is also likely to peak at this time for nocturnal species. ...
... It is certainly true that cathemeral lemurs (E. fulvus) are underrepresented in the diet of fossa in relation to their availability at Kirindy, with most nocturnal genera (Microcebus, Lepilemur, Phaner and Mirza) as well as the diurnal sifaka (Propithecus verreauxi) overrepresented in relation to their abundance [Hawkins, 1998]. This suggests that an ancestral shift from a nocturnal bimodal activity pattern to a crepuscular bimodal pattern with peaks centred on sunrise and sunset may have been an adaptive response to temporal variation in the risk of predation posed by the fossa. ...
As a behavioural strategy, cathemerality is thought to confer considerable advantages by allowing animals to extend activity flexibly into either the diurnal or nocturnal phase in response to the prevailing ecological conditions. Factors such as temperature, access to food sources and minimising the risk of predation are all thought to be important in promoting cathemerality, although previous studies have produced inconsistent results. This paper adopts a different approach by first asking whether an obligate diurnal species, the chacma baboon (Papio hamadryas ursinus), exhibits seasonal variation in behavioural flexibility in response to annual cycles of day length. While short day lengths are an important constraint on the activity of the baboons at De Hoop Nature Reserve, South Africa, long summer days permit considerable flexibility in thermoregulatory response, diet selection and patterns of habitat choice. Given that baboons adapt flexibly in response to a relaxation of time constraints, the question thus arises as to why diurnal and nocturnal primates do not adopt cathemeral activity patterns when time is constrained? For baboons, the costs of predation appear to prohibit exploitation of the nocturnal phase and it is likely that such constraints are true of most primates. It thus follows that Madagascar's predatory environment must in some way permit or select for a cathemeral lifestyle. The importance of the predation by fossa (Cryptoprocta ferox) on the evolution of cathemerality is discussed.
... In two cases, around RNP, owners reported that their own dogs killed fossa (a cat-like mammal endemic and unique to Madagascar), Madagascar's largest carnivore, and a top predator with no known natural enemies. Previous studies of human attitudes toward fossa indicate that many people view them negatively, because they are perceived threats to poultry (Albignac 1973;Hawkins 1998;Jones et al. 2008) and small livestock, such as lambs (Gardner and Davies 2014), due to fady that they resemble dogs (Gardner and Davies 2014), and consume the remains of villagers' ancestors (Jones et al. 2008;Ruud 1960). Across four sites (comprising deciduous forest and rainforests) in Madagascar, respondents had killed fossas, sometimes by training dogs to kill them to retaliate for the consumption of their poultry (Merson et al. 2019b). ...
Domestic dogs (Canis lupus familiaris) are estimated to be one of the most globally abundant invasive carnivores that threaten wildlife. Madagascar is home to large populations of free-roaming dogs and is a highly diverse and anthropogenically threatened environment, making it one of the world's top conservation priorities. Comparatively little is known about human-dog relationships in developing countries such as Madagascar. We surveyed non-dog owners and dog owners visiting free mobile veterinary clinics in their communities around Ranomafana National Park (RNP) and Andasibe-Mantadia National Park (AMNP) to understand human-dog relationships, gain insight on free-roaming dog behavior, and to assess the feasibility of humane population control measures. Amongst dog owners, the vast majority of respondents reported owning their dog for protection and a significant number had dogs for companionship. Our results indicate that free-roaming (owned, unconfined) dogs may be an underappreciated threat to endemic wildlife in the National Parks of Madagascar, as nearly half of dog owners reported that their dog killed at least one wild animal a month. Most dog owners in surveyed communities approve of spay/neuter/vaccine programs and state that they would use them if freely available, indicating that veterinary intervention can be an important tool in humanely controlling free-roaming dog populations in these regions.
... Estimates from the ASSR were comparable to Masoala-Makira (ψ = 0.68; Farris et al. 2015b), Betampona (ψ = 0.95; Rasambainarivo et al. 2017) and Ranomafana (ψ = 0.93; Gerber et al. 2012). The lower occupancy in the ASSR may be related to factors such as their large home range, as the species is known to traverse large areas of intact forests (Gerber et al. 2012) and is not limited to just forested areas (Hawkins 1998, Hawkins & Racey 2005. Canis familiaris has been shown to negatively influence the temporal and spatial patterns of C. ferox at multiple sites in Madagascar (Farris et al. 2016, 2017, 2020, Merson 2017, and the lower occupancy and TS estimates at ASSR may be the result of the widespread distribution and activity of C. familiaris across the surveyed area. ...
ABSTRACT: Madagascar is an island nation renowned for its biodiversity and species endemism, yet it is still largely understudied, despite intense anthropogenic threats, including forest loss and edge effects. Anjanaharibe-Sud Special Reserve is a recently expanded rainforest protected area that is lacking detailed surveys and assessments of the native carnivore community of the endemic family Eupleridae. To identify which terrestrial carnivores occupy the reserve and what anthropogenic disturbances and factors best explain their occurrence patterns, we deployed 35 motion-activated cameras to detect native and introduced carnivores. From November 2018 to February 2019, we collected 2918 unique capture events (all species) and confirmed the presence of 5 euplerids: Galidia elegans, Galidictis fasciata, Eupleres goudotii, Fossa fossana and Cryptoprocta ferox. These results extend the known range of E. goudotii and G. fasciata. In the reserve, F. fossana and G. elegans are the most common and widespread native carnivores, while E. goudotii was the rarest. We highlight the negative impact of edge effects on G. fasciata and F. fossana and the threat posed by the free-ranging non-native carnivore, C. familiaris. This study represents the first detailed survey and occurrence estimates of the carnivore community of this protected area, allowing comparison with other protected areas in Madagascar. Our empirical findings show that anthropogenic disturbance negatively impacts carnivore existence within Anjanaharibe-Sud Special Reserve and suggest important management recommendations for protecting the carnivore community and the co-occurring wildlife living within Anjanaharibe-Sud Special Reserve.
... For analysis in the program FragStats (McGarigal et al. 2012), we placed a 500 m buffer around individual camera stations (based on estimated home range of native carnivores using ranging data and body size from camera traps) and clipped the classified imagery for each of the resulting seven camera grid buffers (each providing an approximately 10-15 km 2 area). We calculated the following landscape metrics from FragStats: (1) number of habitat patches: total number of rainforest, degraded forest, and matrix patches (based on habitat classifications from satellite imagery) within the buffer, where a patch is an area of habitat type separated from similar habitat by C 50 m, (2) largest patch index: the percentage of total buffered area comprised by the largest rainforest patch, (3) landscape shape index (LSI) or the standardized measure of total edge adjusted for the size of the buffered area (McGarigal et al. 2012), (4) percent rainforest within the buffered area, (5) percent matrix or non-forest, cultivated area within the buffered area, (6) total rainforest core area: the sum of the core areas (accounting for edge of depth of 500 m) of each rainforest patch within the buffer, and (7) (Gerber et al. 2010;Gerber 2011;Gerber et al. 2012b;Goodman 2012;Hawkins 1998;Hawkins and Racey 2005) explored how landscape and habitat variables influence native carnivore spatial distribution. However, we still lack an understanding of exactly which habitat and landscape variables best explain space use for each individual carnivore species (both native and invasive), including how anthropogenic changes to these landscapes affect native carnivore spatial distribution. ...
Studies of elusive carnivores often rely on passive sampling when investigating either spatial or temporal interactions. However, inference on behavioral mechanisms are usually lacking. We present an analysis that combines previously published spatial co-occurrence estimates and temporal kernel density estimates to explore spatiotemporal interspecific interactions. We do so by deriving a spatiotemporal value (STV) that is a relative measure of potential interaction in both niche dimensions, across a gradient of degradation, for rainforest carnivore pairs in Madagascar. We also use a conceptual framework to provide insight into the potential behavioral mechanisms of habitat selection. Of the six native and three invasive carnivores, we estimate the spatiotemporal interactions for twelve pairings, which range from no spatial/temporal relationship (n = 5) to spatiotemporal aggregation or segregation (n = 7). We visualized these spatiotemporal interactions along a fragmentation gradient and demonstrate that these interactions are not static, as STV overlap increases with increasing anthropogenic disturbance. Of the three invasive carnivores (free-ranging dogs Canis familiaris, cats Felis species, and small Indian civets Viverricula indica) the latter had the highest number of spatial occurrence (n = 4) and spatiotemporal overlap (n = 4) relationships with native carnivores. Our results highlight the potential for increasing direct and indirect interactions between native and invasive species as forest degradation and invasive predators increase. Our approach allows us to better understand adaptive behaviors, plasticity in temporal activity, community assemblage, and to develop targeted conservation strategies to manage ecological communities in rapidly changing ecosystems.
... For analysis in the program FragStats (McGarigal et al. 2012), we placed a 500 m buffer around individual camera stations (based on estimated home range of native carnivores using ranging data and body size from camera traps) and clipped the classified imagery for each of the resulting seven camera grid buffers (each providing an approximately 10-15 km 2 area). We calculated the following landscape metrics from FragStats: (1) number of habitat patches: total number of rainforest, degraded forest, and matrix patches (based on habitat classifications from satellite imagery) within the buffer, where a patch is an area of habitat type separated from similar habitat by C 50 m, (2) largest patch index: the percentage of total buffered area comprised by the largest rainforest patch, (3) landscape shape index (LSI) or the standardized measure of total edge adjusted for the size of the buffered area (McGarigal et al. 2012), (4) percent rainforest within the buffered area, (5) percent matrix or non-forest, cultivated area within the buffered area, (6) total rainforest core area: the sum of the core areas (accounting for edge of depth of 500 m) of each rainforest patch within the buffer, and (7) (Gerber et al. 2010;Gerber 2011;Gerber et al. 2012b;Goodman 2012;Hawkins 1998;Hawkins and Racey 2005) explored how landscape and habitat variables influence native carnivore spatial distribution. However, we still lack an understanding of exactly which habitat and landscape variables best explain space use for each individual carnivore species (both native and invasive), including how anthropogenic changes to these landscapes affect native carnivore spatial distribution. ...
Studies of elusive carnivores often rely on passive sampling when investigating either spatial or temporal interactions. However, inference on behavioral mechanisms are usually lacking. We present an analysis that combines previously published spatial co-occurrence estimates and temporal kernel density estimates to explore spatiotemporal interspecific interactions. We do so by deriving a spatiotemporal value (STV) that is a relative measure of potential interaction in both niche dimensions, across a gradient of degradation, for rainforest carnivore pairs in Madagascar. We also use a conceptual framework to provide insight into the potential behavioral mechanisms of habitat selection. Of the six native and three invasive carnivores, we estimate the spatiotemporal interactions for twelve pairings, which range from no spatial/temporal relationship (n = 5) to spatiotemporal aggregation or segregation (n = 7). We visualized these spatiotemporal interactions along a fragmentation gradient and demonstrate that these interactions are not static, as STV overlap increases with increasing anthropogenic disturbance. Of the three invasive carnivores (free-ranging dogs Canis familiaris, cats Felis species, and small Indian civets Viverricula indica) the latter had the highest number of spatial occurrence (n = 4) and spatiotemporal overlap (n = 4) relationships with native carnivores. Our results highlight the potential for increasing direct and indirect interactions between native and invasive species as forest degradation and invasive predators increase. Our approach allows us to better understand adaptive behaviors, plasticity in temporal activity, community assemblage, and to develop targeted conservation strategies to manage ecological communities in rapidly changing ecosystems.
... In one species (Northern river otter, Lontra canadensis), the baubellum did not begin development until 2 years after birth ( Lauhachinda, 1978), in contrast to the male baculum which was present at birth and continued to grow throughout the animal's life ( Friley, 1949;Stephenson, 1977). In two species (Walrus, Odobenus rosmarus; Fossa, Cryptoprocta fossa), baubellum size decreased with age, opposite the developmental patterns of the baculum ( Fay, 1955Fay, , 1982Hawkins, 1998;Hawkins et al., 2002). Across 13 species, 12 had a higher baubellum CV versus baculum CV, seven of which were significantly higher by the asymptotic test and six of which were significantly higher by the modified signed-likelihood ratio test ( Table 1). ...
Understanding the evolutionary forces that influence sexual dimorphism is a fundamental goal in biology. Here, we focus on one particularly extreme example of sexual dimorphism. Many mammal species possess a bone in their penis called a baculum. The female equivalent of this bone is called the baubellum and occurs in the clitoris, which is developmentally homologous to the male penis. To understand the potential linkage between these two structures, we scored baculum/baubellum presence/absence across 163 species and analyzed their distribution in a phylogenetic framework. The majority of species (N = 134) shared the same state in males and females (both baculum and baubellum present or absent). However, the baubellum has experienced significantly more transitions, and more recent transitions, so that the remaining 29 species have a baculum but not a well-developed baubellum. Even in species where both bones are present, the baubellum shows more ontogenetic variability and harbors more morphological variation than the baculum. Our study demonstrates that the baculum and baubellum are generally correlated across mammals, but that the baubellum is more evolutionarily and developmentally labile than the baculum. The accumulation of more evolutionary transitions, especially losses in the baubellum, as well as noisier developmental patterns, suggests that the baubellum may be nonfunctional, and lost over time.
... The present study seeks to inform this debate by assessing the predicted niche overlap between this novel, toxic potential prey and native, endemic carnivores, a group that is likely to be directly affected by this incipient invasion. (Rasoloarison et al. 1995;Goodman et al. 1997Goodman et al. , 2003cGoodman et al. , 2012Wright et al. 1997;Hawkins 1998;Britt et al. 2001;Hawkins and Racey 2005;Garbutt 2007); Falanouc- (Albignac et al. 1984;Dollar 1999;Garbutt 2007;Goodman 2012); Spotted fanaloka- (Albignac 1972;Albignac et al. 1984;Goodman and Pidgeon 1999;Goodman 2003cGoodman , 2012Kerridge et al. 2003;Garbutt 2007); Ring-tail vontsira- (Albignac 1972;Dunham 1998;Goodman and Pidgeon 1999;Britt and Virkaitis 2003;Goodman 2003aGoodman , 2012Garbutt 2007); Broad-stripe vontsira- (Goodman 2003b(Goodman , 2012; Brown-tail vontsira- (Britt and Virkaitis 2003;Goodman 2012) b Sources for carnivore habitat preference include: (Gerber et al. 2012c;Goodman 2012;Farris et al. 2014) c IUCN (2016) d Carnivore weights taken from: (Goodman and Pidgeon 1999;Goodman 2003aGoodman , b, c, 2012Kerridge et al. 2003;Garbutt 2007;Lührs and Kappeler 2013) First, we predicted the degree of niche overlap between environmental niches for D. melanostictus and the carnivores using a multivariate niche modeling approach. We expected this relationship to be modulated by different factors, such as abundance or microhabitat differences (e.g., site-specific temperature or forest cover). ...
Biological invasions can represent important threats to endemic species,
including those within the invaders’ food webs. The Asian common toad (Duttaphrynus
melanostictus) was introduced to Madagascar in 2011. This introduction presents a
potentially dangerous prey item to a relatively naı¨ve, highly diverse endemic carnivore
fauna. Using a multivariate niche modeling approach (background test), we assessed the
predicted niche overlap between D. melanostictus and six endemic carnivores in eastern
Madagascar. The overlap between this potential prey and predators was assessed on four
environmental niche axes: temperature, precipitation, vegetation cover and elevation. Our results showed a mixture of niche overlap and divergence between D. melanostictus and
the six carnivores for environmental axes tested. There was significant overlap with five of
the carnivores on temperature and NDVI axes. On the precipitation axis, there was significant
overlap between D. melanostictus with two species. Our results suggested that
wide-ranging, locally rare carnivores may overlap extensively with D. melanostictus. The
six carnivores that inhabit the eastern rainforest of Madagascar will likely share multiple,
niche axes with this novel potential prey item. Species that eat the non-native common toad
and are susceptible to its toxins are at conservation risk because their populations may not
be robust enough to adapt quickly to this threat. We advocate closely monitoring these
emerging interactions and suggest a preemptive conservation strategy for carnivores
potentially at risk.
... e (Albignac, 1972). f (Hawkins, 1998). g (Dollar, 1999). ...
... e (Albignac, 1972). f (Hawkins, 1998). g (Dollar, 1999). ...
Madagascar's Eupleridae carnivores are perhaps the least studied and most threatened family of Carnivora. Investigating potential direct and indirect competition among these native species and sympatric exotic carnivores is necessary to better direct conservation actions. From 2008 to 2013, we photographically surveyed a diverse rainforest landscape, comparing six native and three exotic carni-vores' activity patterns throughout the diel cycle. We used hierarchical Bayesian Poisson analysis to describe the activity patterns of Madagascar's carnivore community , assessed effects of season and site on temporal activity patterns, and estimated coefficients of overlap between carnivore pairings to assess effects of body size and ecological niche on temporal overlap among native and exotic carnivores. We observed changes in temporal activity patterns across seasons particularly during the austral summer (hot–dry season) for four native and two exotic carnivores, including evidence of fossa Cryptoprocta ferox altering their temporal activity during their mating season (hot–dry season). We found evidence of high overlap between natives and exotics indicating the potential for increased interactions and competition. The greatest overlap in temporal activity occurred between both ring-tail Galidia elegans and brown-tail vontsira Salanoia concolor and exotic dogs Canis familiaris. Cr. ferox, falanouc Eupleres goudotii and spotted fanaloka Fossa fossana also overlapped in activity with the nocturnal, exotic Indian civet Viverricula indica. Cr. ferox avoided humans and Ca. familiaris across all seasons. Unexpectedly, carnivore body size and ecological niche were not important predictors of temporal overlap. Previous research has shown these native and exotic carnivores overlap spatially and these new findings of temporal overlap among native and exotic carnivores add urgency to the need to manage exotic carnivores across Madagascar.
... Despite some pioneering field studies (e.g. Albignac, 1976;Hawkins, 1998;Razafimanantsoa, 2003), the size and composition of social units has not been determined for the majority of the Eupleridae. Similarly, limited life-history data from the Eupleridae indicate small litter sizes (i.e. in most species only one young) and low reproductive rates. ...
The diversity of extant carnivores provides valuable opportunities for comparative research to illuminate general patterns of mammalian social evolution. Recent field studies on mongooses (Herpestidae), in particular, have generated detailed behavioural and demographic data allowing tests of assumptions and predictions of theories of social evolution. The first studies of the social systems of their closest relatives, the Malagasy Eupleridae, also have been initiated. The literature on mongooses was last reviewed over 25 years ago. In this review, we summarise the current state of knowledge on the social organisation, mating systems and social structure (especially competition and cooperation) of the two mongoose families. Our second aim is to evaluate the contributions of these studies to a better understanding of mammalian social evolution in general. Based on published reports or anecdotal information, we can classify 16 of the 34 species of Herpestidae as solitary and nine as group-living; there are insufficient data available for the remainder. There is a strong phylogenetic signal of sociality with permanent complex groups being limited to the genera Crossarchus, Helogale, Liberiictis, Mungos, and Suricata. Our review also indicates that studies of solitary and social mongooses have been conducted within different theoretical frameworks: whereas solitary species and transitions to gregariousness have been mainly investigated in relation to ecological determinants, the study of social patterns of highly social mongooses has instead been based on reproductive skew theory. In some group-living species, group size and composition were found to determine reproductive competition and cooperative breeding through group augmentation. Infanticide risk and inbreeding avoidance connect social organisation and social structure with reproductive tactics and life histories, but their specific impact on mongoose sociality is still difficult to evaluate. However, the level of reproductive skew in social mongooses is not only determined by the costs and benefits of suppressing each other's breeding attempts, but also influenced by resource abundance. Thus, dispersal, as a consequence of eviction, is also linked to the costs of co-breeding in the context of food competition. By linking these facts, we show that the socio-ecological model and reproductive skew theory share some determinants of social patterns. We also conclude that due to their long bio-geographical isolation and divergent selection pressures, future studies of the social systems of the Eupleridae will be of great value for the elucidation of general patterns in carnivore social evolution.
... The fossa is the only Malagasy predator capable of hunting adult lemurs of any species (Table 1). In fact, lemurs can comprise in excess of 50 % of its diet (Hawkins, 1998). Amongst lemurs, the fossa is the most common predator of Propithecus species, although the smaller bodied dry forest sifakas are occasionally killed by raptors and snakes as well (Table 1). ...
... Short-winged birds, such as most owls, are generally more capable of rapid changes of direction (see eg. Norberg, 1985), whereas long-winged species may only be capable of taking lemurs from the very top of the canopy. Cryptoprocta, the fossa, is a large-bodied but short-legged viverrid, den-living but competent arboreally and capable of leaping (see eg. Wright et al, 1997; Hawkins, 1998;. Dollar 1999; Dollar et al. this volume; Patel 2005). ...
Predator pressure is normally very difficult to assess, and most reports tend to be anecdotal. However, it has been estimated
that an annual predation rate of 25% may apply to Microcebus populations (Goodman et al., 1993). Such a rate, albeit for a particularly small prosimian, implies strong selective pressure in favor of adaptations that
reduce predation, and it seems reasonable to assess adaptations with predation in mind. Predator avoidance by vigilance is
usually seen as an attribute of social foragers (see, e.g., Terborgh & Janson, 1986), to which category many of the Lemuridae, and arguably some Indriidae and Lepilemuridae, belong. However, the small body
size and nocturnality of those prosimians described as “solitary foragers” are often regarded as facilitating alternative
predator avoidance strategy, crypsis (e.g., Clutton-Brock & Harvey, 1977; Stanford, 2002).
... The vast majority of animals considered here are classified as strict natural forest inhabitants, yet there are conflicting interpretations of the level of disturbance species can tolerate. Amongst carnivorans, certain taxa with large home ranges, such as Cryptoprocta ferox, occur in disturbed forests and move across anthropogenic formations (Hawkins, 1998;). The specialized shrew-tenrec Microgale gracilis has been considered restricted to undisturbed montane forest, but was found at the ecotone between native tree and pine plantations (Goodman et al., 1996). ...
The emergence of the genus Homo is widely linked to the colonization of 'new' highly seasonal savannah habitats. However, until recently, our understanding of the possible impact of seasonality on this shift has been limited because we have little general knowledge of how seasonality affects the lives of primates. This 2005 book documents the extent of seasonality in food abundance in tropical woody vegetation, and then presents systematic analyses of the impact of seasonality in food supply on the behavioural ecology of non-human primates. Syntheses in this volume then produce broad generalizations concerning the impact of seasonality on behavioural ecology and reproduction in both human and non-human primates, and apply these insights to primate and human evolution. Written for graduate students and researchers in biological anthropology and behavioural ecology, this is an absorbing account of how seasonality may have affected an important episode in our own evolution.
Anthropogenic habitat alteration and invasive species are threatening carnivores globally. Understanding the impact of these factors is critical for creating localized, effective conservation programmes. Madagascar's Eupleridae have been described as the least studied and most threatened group of carnivores. We investigated the effects of habitat degradation and the presence of people and exotic species on the modelled occupancy of the endemic fosa Cryptoprocta ferox , conducting camera-trap surveys in two western deciduous forests, Ankarafantsika National Park and Andranomena Special Reserve. Our results indicated no clear patterns between habitat degradation and fosa occupancy but a strong negative association between cats Felis sp. and fosas. Cat occupancy was negatively associated with birds and positively associated with contiguous forest and narrow trails. In contrast, dog Canis lupus familiaris occupancy was best predicted by wide trails, degraded forest and exotic civets. Our results suggest fosas are capable of traversing degraded landscapes and, in the short term, are resilient to contiguous forest disturbance. However, high occupancy of cats and dogs in the landscape leads to resource competition through prey exploitation and interference, increasing the risk of transmission of potentially fatal diseases. Management strategies for exotic carnivores should be considered, to reduce the widespread predation of endemic species and the transmission of disease.
Madagascar is renowned for its biodiversity, but also for forest loss, fragmentation, and degradation, making it a global conservation priority. With few studies dedicated to Madagascar‟s carnivores, little is known about their ecology. My objectives were to 1) compare density estimation techniques applicable to enumerating rare and/or elusive carnivores, 2) investigate Malagasy carnivore distributions, abundance and density, and occupancy/use across four sites that vary in forest disturbance, and 3) explore temporal activity patterns of rainforest carnivores. I found the spatially-explicit-capture-recapture models were empirically superior, as they are flexible and account for spatial variation in detection probability and area estimation. I found both endemic and exotic carnivore composition varied among four rainforest sites: Primary, Selectively-logged, Fragments <2.5 km and Fragments >15 km from contiguous primaryrainforest. All endemic carnivores were present in the Primary and Selectively-logged rainforest, while endemic carnivore species richness decreased and exotic carnivore species richness increased in the fragmented forests. Malagasy civet (Fossa fossana) density ± SE was significantly less in the Selectively-logged compared to the Primary rainforest (1.38 ± 0.22, 3.19 ± 0.55 civets/km2, respectively); they were absent from both fragmented forests. Fossa (Cryptoprocta ferox) density ± SE was not different between the Primary and Selectively-logged rainforests (0.12 ± 0.05, 0.09 ± 0.04 adults/km2 , respectively); a single animal was detected in
the Fragments <2.5 km, while none were detected in the Fragments >15 km. Malagasy carnivores had varied temporal activity overlap (5.8-88.8%). C. ferox preferred crepuscular activity, but overall exhibited a cathemeral activity pattern.
The carnivores of Madagascar are likely the least studied of the world’s carnivores, thus little is known about threats to their persistence. I provide the first long-term assessment of Madagascar’s rainforest carnivore community, including: 1) how multiple forms of habitat degradation (i.e., fragmentation, exotic carnivores, human encroachment, and hunting) affect native and exotic carnivore occupancy; 2) how native and exotic carnivore temporal activity overlap and how body size and niche explain these patterns; 3) how native and exotic carnivores spatially co-occur across the landscape and which variables explain these relationships; and 4) how native and exotic carnivores and humans co-occur with lemurs across Madagascar’s largest protected landscape: the Masoala-Makira landscape. From 2008 to 2013 I photographically sampled carnivores and conducted line-transect surveys of lemurs at seven study sites with varying degrees of degradation and human encroachment, including repeat surveys of two sites. As degradation increased, exotic carnivores showed increases in activity and occupancy while endemic carnivore, small mammal, and lemur occupancy and/or activity decreased. Wild/feral cats (Felis sp.) and dogs (Canis familiaris) had higher occupancy (0.37 ± SE 0.08 and 0.61 ± SE 0.07, respectively) than half of the endemic carnivore species across the landscape. Additionally, exotic carnivores had both direct and indirect negative effects on native carnivore occupancy. For example, spotted fanaloka (Fossa fossana) occupancy (0.70 ± SE 0.07) was negatively impacted by both wild/feral cat (beta = -2.65) and Indian civets
iii
(beta = -1.20). My results revealed intense pressure from hunting (ex. n = 31 fosa Cryptoprocta ferox consumed per year from 2005-2011 across four villages), including evidence that hunters target intact forest where native carnivore and lemur occupancy and/or activity are highest. I found evidence of high temporal overlap between native and exotic carnivores (ex. temporal overlap between brown-tail vontsira Salanoia concolor and dogs is 0.88), including fosa (Cryptoprocta ferox) avoiding dogs and humans across all seasons. However, I found no evidence of body size or correlates of ecological niche explaining temporal overlap among carnivores. Estimates of spatial co-occurrence among native and exotic carnivores in rainforest habitat revealed strong evidence that native and exotic carnivores occur together less often than expected and that exotic carnivores may be replacing native carnivores in forests close to human settlements. For example, falanouc show a strong increase in occupancy when dogs are absent (0.69 ± SE 0.11) compared to when they are present (0.23 ± SE 0.05). Finally, the two-species interaction occupancy models for carnivores and lemurs, revealed a higher number of interactions among species across contiguous forest where carnivore and lemur occupancy were highest. These various anthropogenic pressures and their effects on carnivore and lemur populations, particularly increases in exotic carnivores and hunting, have wide-ranging, global implications and demand effective management plans to target the influx of exotic carnivores and unsustainable hunting affecting carnivore and primate populations across Madagascar and worldwide.
In hypercarnivorous species, females have large spatial requirements to meet their nutritional needs, and food competition among females is intense. As a result, females are typically solitary and territorial, and solitary males compete for access to dispersed females. Yet, largely anecdotal reports indicate that facultative male sociality may be more common in solitary carnivores than previously thought. We studied spatial interactions among fossas (Cryptoprocta ferox), Madagascar's largest carnivore, using simultaneous GPS tracking of 13 adult individuals to determine patterns of sex-specific spatial distribution and sociality. Male home ranges were larger than those of females, male home ranges overlapped more with those of other males than those of females with other females. Whereas some males were solitary, a subset of adult males was found to have very high home range overlap, high rates of co-location within <50 m, low minimum inter-individual distances, and significantly positive “dynamic interaction”. These associated dyads sometimes, but not always, were close relatives. The fact that solitary and associated males coexist in this population raises interesting questions concerning constraints and flexibility of social tolerance. This study yielded preliminary indications that female distribution appears to be primarily structured by resource competition, whereas male sociality seems to depend on demographic chance events, yet unknown proximate determinants of social tolerance, and it is associated with somatic and reproductive advantages. Male associations among carnivores are therefore more widespread and appear to be based on a wider range of factors than previously thought.
Temporal partitioning of activity among sympatric species can be an important mechanism for species coexistence. Further, if exotic and native species overlap temporally, there is potential for direct competition and antagonism, which may lead to native species extirpation. We 1st assessed if ecologically similar native carnivores of Madagascar demonstrated activity pattern overlap and then explored whether overlap in activity might lead to negative impacts of exotic carnivores on native carnivores. We used photographic sampling to quantify the temporal activity patterns of carnivores at 4 study sites. The activity of the 2 smaller-bodied native species, Galidia elegans and Galidictis fasciata, overlapped minimally; these 2 carnivores share a similar generalist diet, which may drive their divergent temporal activity. In contrast, the medium-sized native species, Fossa fossana and Eupleres goudotii, were both highly nocturnal; these 2 species appear segregated in their diets. The largest native carnivore, Cryptoprocta ferox, selectively used crepuscular hours, but overall was cathemeral; it was notably absent or basically so at sites where dogs were most abundant and active throughout the diel cycle. We found G. elegans to shift from preferred activity periods in the presence of dogs and the exotic Viverricula indica. Our results suggest that the presence and activity of exotic carnivores can negatively impact native carnivores in fragmented rain forests.
The puma-like Cryptoprocta ferox is the largest living Carnivora on Madagascar (Goodman et al., 2003). Cryptoprocta has been a taxonomic enigma until recently (cf. Veron & Catzeflis, 1993; Veron, 1995), showing numerous convergent morphological characters with members of the Felidae. Some of these attributes, such as semi-retractable
claws used in both climbing and hunting, contributed to the long-running uncertainty as to the phylogenetic relationships
of this animal. Recent molecular studies indicate that Cryptoprocta is part of a radiation of Carnivora endemic to Madagascar, which unites all of the native species on the island into a single
clade (Yoder et al., 2003), now recognized as the endemic family Eupleridae (Wozencraft, in press). On the basis of molecular data this radiation of Carnivora is slightly younger than that of lemurs, but the two groups
have co-existed on Madagascar for something on the order of 20 million years (Yoder et al., 2003). Until the Holocene a second member of Cryptoprocta occurred on the island that was notably larger than the living species (Goodman et al., 2004).
Keywords:Cryptoprocta ferox;Madagascar;microsatellites;PCR;Viverridae
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