What Do We Know About the Life-History Traits of Widely Hunted Tropical Mammals?

What do we know about the life-history traits of widely hunted tropical mammals?

N ATHALIE VAN V LIET and R OBERT N ASI

Abstract We synthesize information on parameters useful In ecological terms the ability of prey species to withstand for managing the hunting of two common mammal species various levels of harvest, without depletion, depends on the that are important for local people in the Neotropics and life-history traits and biological parameters of the species Africa: Cuniculus paca and Philantomba monticola, respect- (Caughley, ). Population dynamics depend on intrinsic ively. We highlight the scarcity of data available on the para- population growth, mortality and the dynamics of in–out meters needed to manage these two species sustainably. As migration based on spatial (e.g. dispersal rate, dispersal most of the studies were conducted .  years ago, we stress distance and territory size) and temporal (e.g. seasonality the need to supplement the information available using of reproduction) parameters (Novaro et al., ; Salas & methodological and technical innovations. In particular, Kim, ; van Vliet et al., ). we call for new assessments covering the possible variations A thorough understanding of the dynamics of prey po- in parameter values across the species’ distribution ranges, pulations under hunting pressure is needed for robust man- and covering various anthropogenic contexts, to test agement decision-making. In tropical forests one of the density-dependent and compensatory processes that may major impediments to sound estimates of population dy- explain the resilience of these species to hunting. namics is the paucity of available biological and ecological data for hunted species (Milner-Gulland & Akçakaya, Keywords Biological parameters, Cephalophus monticola, ; van Vliet & Nasi, ; Weinbaum et al., ; van Cuniculus paca, life-history traits, Philantomba monticola, Vliet et al., ; Mayor et al., ), and when such data sustainability assessments exist they are not available in a synthetic and comprehensive manner. Here we synthesize all the available information regard- Introduction ing parameters pertinent to the sustainable use of two of the most hunted tropical species: the lowland paca Cuniculus unting is considered to be a major threat to wildlife paca (in the Neotropics) and the blue duiker Philantomba populations in tropical forest regions (Fa et al., ; H monticola (in Africa). These are common and generalist Mallon et al., ; Benítez-Lόpez et al., ), and mammals small-sized species, with a widespread geographical range, comprise the greatest portion of hunted species (Fa et al., and are crucial for the livelihoods of many rural ; Nasi et al., ). A meta-analysis has demonstrated communities. that there have been significant declines (up to %) in mammal abundances in many regions as a result of hunting pressure (Benítez-Lόpez et al., ). The unsustainable use Study species of wild mammals in most tropical forest areas hinders the Cuniculus paca is a large nocturnal rodent that occurs in possibility that impoverished communities will be able to Mexico, Colombia, Venezuela, the Guianas, Ecuador, continue to feed on wild mammals in the future (Wilkie Peru, Bolivia, Paraguay and most of Brazil, and has been in- et al., ). troduced into Cuba and the Antilles (Patton et al., ). The After several decades of a so-called fences and fines ap- species occurs in a wide range of forest types in moist areas proach to hunting there is now a wider recognition that and is an important seed distributor, with scatter-hoarding sustainable-use approaches merit more consideration behaviour (Eisenberg & Redford ). It is categorized as (CBD, ; Mayor et al., ). Managing hunting sustain- Least Concern on the IUCN Red List, based on its wide dis- ably must address the multiple needs and desires of societies tribution, presumed large population, and occurrence in a without jeopardizing the options for future generations to number of protected areas, and because it is unlikely to be benefit from the full range of goods and services provided declining (Emmons, ). However, unsustainable hunting by hunted species (van Vliet et al., ). of the paca has been reported from several locations (e.g. Koster, ; Zapata-Ríos et al., ; Valsecchi et al., ) and has led to local depletion of the species. The NATHALIE VAN VLIET (Corresponding author) and ROBERT NASI Center for paca is sought after for its taste, nutritional value and low International Forestry Research, Bogor Barat, Indonesia   E-mail [email protected] fat content (Aguiar, ; Cordón and de Ariza, ;   Received  June . Revision requested  September . Lemire et al., ; Gálvez et al., ), and because of Accepted  September . First published online  February . these attributes is one of the most hunted and consumed

Oryx, 2019, 53(4), 670–676 © 2018 Fauna & Flora International doi:10.1017/S0030605317001545 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.229, on 24 Sep 2021 at 22:18:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605317001545 Life history of hunted tropical mammals 671

species in Latin America (Koster, ; Read et al., ;El (1) Only studies that contained information on the follow- Bizri et al., ; Quiceno-Mesa et al, ; van Vliet et al., ing biological and demographic parameters useful for ; Gómez et al., ; Vanegas et al., ). prey population management (Sutherland, ): (i) Variables used in one-off intrinsic population growth Philantomba monticola is an abundant ungulate, widely dis- models (Robinson & Redford, ): age at first and tributed in central, eastern and southern Africa, from the last reproduction, gestation period, litter size, interval Cross River in Nigeria to south-west South Sudan and south- between births, sex ratio at birth (female/male), lon- wards to central Angola, and Zambia, Malawi, eastern gevity, mortality rate. (ii) Variables used in spatially Zimbabwe, parts of central Mozambique, and on the islands explicit demographic models (Novaro et al., ; of Pemba, Zanzibar and Mafia (East, ; Wilson, ; Hart Salas & Kim, ; van Vliet et al., ): mean terri- &Kingdon,). It thrives in a wide range of forested and tory size; dispersal distance, dispersal rate, dispersal wooded habitats, including primary and secondary forests, age. (iii) Variables influencing temporal variations: gallery forests, dry forest patches, coastal scrub farmland, re- seasonality in reproduction, dispersal and mortality. generating forest and degraded forest patches, even near (2) Only studies providing primary data. We selected human settlements (Hart & Kingdon, ). The species is studies only if they provided primary information on among the most hunted throughout its range and is an im- the biological and demographic characteristics de- portant source of meat and income for rural people (de scribed above. If no primary data were provided but Merode et al., ; Kümpel, ; van Vliet et al., ; the primary source was cited (e.g. in the AnAge data- van Vliet & Nasi, ). It withstands hunting pressure better base (Tacutu et al., ) or in Weigl, ), then the than most of the larger duikers (van Vliet et al., ; Mockrin, primary source was searched and, if available, screened ) and is categorized as Least Concern on the IUCN Red based on the same primary and secondary criteria de- List (IUCN SSC Antelope Specialist Group, ). scribed above. For each of the studies that passed our filters ( for P. Methods monticola and  for C. paca) we extracted the information on each of the  variables (Tables  and ), and re- During March–April  we carried out a bibliographic corded whether the information came from wild or search using seven databases: ISI Web of Science, Science captive populations, and the size of the sample studied, Direct, EBSCO, Scielo, Redalyc, Scopus and Google if available. Scholar. The search used a combination of words in English, Spanish, Portuguese and French: (‘species scientific name’ OR ‘common name’) AND (reproducti* OR dispers* Results OR behaviour OR mortality OR gestati* OR offspring OR longevity OR ‘litter size’ OR ‘biological parameters’ Philantomba monticola A total of  studies published OR ‘life-history traits’). For the blue duiker we used the during – were found to have generated primary scientific names Cephalophus monticola and Philantomba data on biological and demographic parameters for P. monticola, as both are commonly used in the literature. monticola, of which eight were based on individuals in We screened the references using titles and abstracts captivity, five on wild individuals from South Africa, two according to the following primary inclusion criteria: on wild individuals in Gabon and one on wild individuals in Republic of Congo. Most of the information for each (1) Only studies for which we were able to source the full variable was gleaned from – studies, mostly from text. We searched for the full online text or the PDF, individuals in captivity or from a limited number of wild contacting the authors if necessary. We were able to individuals (–). No information was found on age at source full documents from as far back as . last reproduction, mortality rate, seasonality of mortality (2) Only studies with scientific merit. To ensure the scien- or seasonality of dispersal. The results are synthesized in tific quality of the information reported, we selected Table . only peer-reviewed documents such as scientific journal articles, published records of zoological data pro- Cuniculus paca A total of  studies published during – duced by zoos, book chapters or theses for Master’s  were found to have generated primary data on the or PhD degrees. biological and demographic parameters of C. paca,ofwhich (3) Only focused on the species of interest to this study.  were based on individuals in captivity and five on wild We selected only papers that provided information individuals (in Peru, Colombia, French Guiana, Costa Rica). on C. paca or P. monticola. Reproduction variables were derived from sample sizes of – We screened the resulting references based on the full  for individuals in the wild and – for individuals in text and used the following secondary criteria: captivity. For longevity, a sample of  individuals was used.

TABLE 1 Primary data available on basic biological and ecological parameters (reproduction, mortality, dispersal and seasonality) for the blue duiker Philantomba monticola. No data were . https://doi.org/10.1017/S0030605317001545 available on age at last reproduction, mortality rate, seasonality of mortality and seasonality of dispersal. Blank cells indicate data were not provided in the source document.

Interval . IPaddress: Age at first Gestation between Mean Dispersal Dispersal reproduction period Litter births Sex Longevity territory age distance Dispersal Seasonality of Country Captivity n (months) (days) size (days) ratio (years) size (ha) (months) (km) rate (%) reproduction Source

170.106.35.229 Yes Year round von Ketelhodt (1977) Congo No 2 1.5 25 Mockrin (2010) Gabon No 9 2.5–412–18 0.27 69 Dubost (1980) Gabon No . 1500 204 1 Year round Dubost & Feer (1992)

, on Germany Yes 205 Dittrich (1972)

24 Sep2021 at22:18:15 Germany Yes 1 12.9 265.5 Year round; von Ketelhodt (1977) peak during –

Oryx Aug. Dec. Germany Yes 16 6–17 207 265 21 Year round Boehner et al. (1984) 09 34,670 53(4), 2019, , South No Year round; Sclater & Thomas Africa peak during (1900) Sep.–Oct. , subjectto theCambridgeCore termsofuse,available at South Yes 12 Year round Brand (1963)

– Africa 676 South No 12 Grzimek (1990) ©

08Fua&FoaItrainldoi:10.1017/S0030605317001545 International Flora & Fauna 2018 Africa South No 10 1 0.75 18 0.15 Bowland (1990) Africa South No 8 0.695 Bowland and Perrin Africa (1995) South No 0.88 Lawes et al. (2000) Africa UK Yes 16 2.9 Jarvis & Morris (1961) USA Yes 6.5 Crandall (1965) Zimbabwe Yes 6 1 Bowman & Plowman (2002) Life history of hunted tropical mammals 673

) For seasonality in reproduction, data were derived from – 1977 samples of individuals. We found no information on

) dispersal (age, rate, distance, seasonality). The results are 

) synthesized in Table . ) 1982 ) ) 1984 1999 2003 2007 ) ) No data were available Discussion 1989 1981

) ) We summarize the available published data on the main

1993 1993 variables needed to assess productivity and management Merritt ( Smythe & Brown De La Guanti( et al. Smythe & Brown De La( Guanti Oliveira et al. ( Oliveira et al. ( Matamoros-Hidalgo ( Matamoros-Hidalgo & Pashov-Nicheva ( for the sustainable use of two common and non-threatened Cuniculus paca. species used as a source of food in African and Neotropical Sep. Sep. May – – forests. Researchers and managers may refer to the synthetic – tables produced, in which we cite the primary sources of data, with information they provide on sample size, geographical origin of the assessment and whether the data ori- during Mar. in Sep. during Mar. during Apr. Seasonality of reproduction Source ginated from captive or wild individuals. In general terms, our review suggests that both species 3 Year round; peak – 3.4reach Beck-King et al. ( maturity after c.  year following birth, reproduce – 1.5 Mean territory size (ha) twice per year all year round, for the whole duration of their mature life (c.  years) and give birth to one offspring per year. In addition, the available data suggest that P. mon-

12 2 ticola has the capacity to disperse and occupy available – –

Longevity (years) empty areas through a small-scale source sink process (Mockrin, ). This behavioural characteristic probably contributes to the resilience of the species to hunting (van Vliet et al., ). No information on dispersal is available

Sex ratio (F/M) for C. paca. The resilience to hunting observed for this species may be linked to its generalist behaviour and capacity to exist in high population densities (– individuals 251 1 Year round; peak

–  per km ; Eisenberg & Redford, ). Interval between births (days) This review highlights the paucity of data available for the parameters needed to manage these species sustainably. 210 2 Year round Contrera Perez & Hernandez ( – – There is little information available on parameters that influ- Litter size ence demographic patterns, such as reproduction, dispersal, home range characteristics, mortality, longevity and seasonal variations. Most of the available data are from captive indi- 173 150 1 310 139 1 117 1 Year round; peak – – – – – viduals, and most, particularly for P. monticola, are from

Gestation period (days) studies conducted during the s and s or earlier. The lack of robust data on life-history traits hinders efforts to propose sustainable management practices for these two species, which are both important nutritional as- 23 1 137

– sets for forest people. Nevertheless, scientists and decision

Age at first reproduction (months) makers continue to make decisions based on erroneous estimates of maximum sustainable yields. Without a significant investment in estimations of life-history traits under varying contexts, quota setting efforts are prone to failure as they will be based on best guesses rather than on sound scientific evidence. We call for new assessments covering the possible variations in parameter values across the distribution range of these two species, and covering various anthropogenic con- 2 Primary data available on basic biological and ecological parameters (reproduction, mortality, dispersal and seasonality) for the lowland paca texts; for example, to address hypotheses on forms of ABLE Costa Rica Yes 49 114 Year round; peak Costa RicaPanama No Panama YesUSA Yes 40 Yes 40 2 20 138 117 Brazil Yes 5 215 Brazil Yes 9 135 T Country Captivity n Colombia Yes 115 1 ColombiaCosta Rica No Yes 49 12 12 118 114 1 12.5 Year round Collett ( on dispersal (age, rate, distance, seasonality). Blank cells indicate data were not provided in the source document. density-dependent mortality and reproduction, and

compensatory vs additive mortality effects in tropical har- (Rodentia: Agoutidae) in Costa Rica: a study using alternative vested species (Weinbaum et al., ). Life-history trait methods. Biotropica, , –. data are also needed for many other species, particularly BENÍTEZ-LÓPEZ, A., ALKEMADE, R., SCHIPPER, A.M., INGRAM, D.J., VERWEIJ, P.A., EIKELBOOM, J.A.J. & HUIJBREGTS, M.A.J. () those of conservation concern, as the use of mean values The impact of hunting on tropical mammal and bird populations. from a few studies is insufficient to assess intra-specific var- Science, , –. iations and adaptations of different populations in relation BOEHNER, J., VOLGER,K.&HENDRICHS,H.() Breeding dates of to environmental and anthropogenic gradients. blue duikers (Cephalophus monticola). Zeitschrift fuer  – Methodological and technical innovations already Saeugetierkunde, , . BOWLAND, A.E. () The ecology and conservation of the blue duiker developed could help produce new assessments; for and red duiker in Natal. PhD thesis. University of KwaZulu Natal, example, participatory sampling involving hunters (e.g. to Durban, South Africa. collect reproductive organs or monitor pregnant females) BOWLAND, A.E. & PERRIN, M.R. () Temporal and spatial patterns is an underestimated and under-used approach that could in blue duikers Philatomba monticola and red duikers Cephalophus natalensis. Journal of Zoology, (), –. be an efficient means of gathering information about repro-    BOWMAN,V.&PLOWMAN,A.( ) Captive duiker management at duction patterns (Mayor et al., ; van Vliet et al., ). the duiker and mini‐antelope breeding and research institute ę Technologies such as camera traps (J drzejewski et al., (Dambari), Bulawayo, Zimbabwe. 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