A Review of Peccary-Palm Interactions and Their Ecological Ramifications across the Neotropics

Abstract

Palms (Arecaceae) are a dominant element within the neotropical plant community and because they substantially contribute to the overall and year-round fruit availability they are considered a key resource for frugivores, particularly for peccaries. Similarly, peccaries (Tayassuidae) are a dominant element within the neotropical mammal community. Their evolution of a strong mastication apparatus, unique interlocking canines, patterns of movement, and foraging ecology are viewed as adaptations to exploit hard seeds, particulary palm seeds. But how strong are the interactions between peccaries and palms, and what are the ecological ramifications? This review synthesizes over 76 papers, published between 1917 and 2004, which revealed that peccaries consumed fruits from 46 palm species, 73% of whose seeds were destroyed after ingestion. Furthermore, peccaries disperse palm seeds; eat flowers, seedlings, and roots; and trample seedlings. Thus, peccaries affect the spatiotemporal distribution and demography of palms. Local extinction of peccaries resulted in dramatic changes in the forest ecology. New conservation strategies are required to protect peccaries and prevent negative cascading effects.

Arecaceae, diversity, herbivory, Pecari tajacu, seed dispersal and predation, Tayassu pecari, trampling

Throughout the Neotropics, palms are a dominating element, with approximately 150 species in Central America and 350 species in South America (A. Henderson, in litt.). Several palm species can reach higher densities than tree species (Gentry 1988, 1990; Hartshorn and Hammel 1994; Henderson 1995, 2002; Kahn 1991; Lieberman and Lieberman 1994; Peres 1994a, 1994b; Pitman et al. 2001, 2003; Terborgh et al. 1996; Terborgh and Andresen 1998). For example, the palms Iriartea deltoidea (up to 45 mature individuals/ha) and Astrocaryum murumuru (up to 30 mature individuals/ha) are the most abundant species within the upper Amazon (Kiltie 1981a; Pitman et al. 2001; Terborgh et al. 1996). Energy-rich fruits and seeds of numerous palm species are available year-round or fruit is borne during periods (i.e., the dry season) when fruits from other trees are scarce (Bodmer and Ward, in press; Cullen et al. 2001; Galetti et al. 1999; Peres 1994a, 1994b; Terborgh 1986a). Furthermore, in the Amazon, the fruits of palms constitute between 30% and 53% of the weight of all available fruits (Bodmer and Ward, in press; Terborgh 1986a). Thus, it is not surprising that palm fruits represent a key resource for frugivores, particularly peccaries (Altrichter et al. 2001; Beck 2005; Bodmer et al. 1997; Cullen et al. 2001; Kahn 1991; Kahn and de Granville 1992; Peres 1994a, 1994b; Silman et al. 2003; Sist 1989b; Terborgh 1986a, 1986b).

Similarly, peccaries (collared peccaries, Pecari tajacu and white-lipped peccaries, Tayassu pecari) are a dominating element in the Neotropics because they form the largest foraging groups (Eisenberg and Redford 1999) and can attain the largest biomass density—over 370 kg/km²—among terrestrial mammals (Eisenberg 1980; Peres 1996; Silman et al. 2003; Terborgh 1983). Throughout their range, peccaries use fruits and seeds from over 212 plant species belonging to 53 families (Beck 2005). However, fruits and seeds from palm species are consumed more frequently (up to 24% more often) than fruits and seeds from any other plant family (Beck 2005; Wyatt and Silman 2004). Furthermore, analyses of stomach contents of peccaries indicate that palm seeds can account for over 60% of their diet (Bodmer 1990; Keuroghlian 2003; Kiltie 1981b; Kiltie and Terborgh 1983; Painter 1998). Therefore, peccaries have potentially profound effects on palms and vice versa. Throughout their range, peccaries are one of the prime targets for subsistence and commercial hunting (Bodmer 1995; Peres 1996; Sowls 1997). In addition, habitat destruction, fragmentation, and other forms of large-scale anthropogenic disturbances synergistically amplify the threats to remaining peccary populations and have resulted in population decline and local extirpation (Beck 2005; Cullen et al. 2000; Peres 1996, 2001; Sowls 1997). Thus, understanding peccary-palm interactions (Silman et al. 2003) is all the more important because of these threats, which, for instance, resulted in the almost complete extinction of white-lipped peccaries throughout Central America (Beck 2005 and citations therein), with unknown consequences for the prevalence of palms and the forest community.

Here, I first review the literature on seed predation and dispersal by collared peccaries and white-lipped peccaries across the Neotropics. I examine both negative effects, including seed predation and herbivory, and positive effects, comprising seed dispersal and trampling, of peccaries on palms. Subsequently, I review the evidence for intra- and interspecific competition between peccaries and other frugivore species for palm resources. And finally, I synthesize the findings, propose a number of ecological ramifications, highlight the ecological consequences of peccary extinction, and suggest urgently needed conservation measures.

Synopsis of Peccary Ecology

Three species of peccaries (Tayassuidae) occur throughout the Neotropics. Because I did not find any publications on palm interactions with the Chacoan peccary (Catagonus wagneri), I will not consider this species here. The collared peccary can be found from Arizona to Argentina, whereas the white-lipped peccary ranges from southern Mexico to northern Argentina (Eisenberg and Redford 1999; Mayer and Wetzel 1987). Collared peccaries use a wide range of habitats, including rain forests, xeric forests, and deserts, whereas white-lipped peccaries are considered to be primarily forest-dwellers (Eisenberg and Redford 1999; Mayer and Wetzel 1987). Groups of collared peccaries range from 5 to over 50 individuals (Castellanos 1983; Judas and Henry 1999), and white-lipped peccaries can occur in groups of up to 400 individuals (Altrichter and Almeida 2002; Kiltie and Terborgh 1983). Historically, groups of over 1,000 individuals have been reported (Jardine 1836; Mayer and Wetzel 1987; Perry 1970). Individual collared peccaries can weigh up to 30 kg (Sowls 1997) and reach a biomass of up to 373 kg/km² (Eisenberg 1980) and a density of over 35 individuals/km² (Mena et al. 2000). White-lipped peccaries can weigh up to 50 kg (J. Terborgh, pers. comm.), reach a biomass of up to 219 kg/km² (Cullen et al. 2001), and attain a density of 16 individuals/km² (Eisenberg 1980). Collared peccaries have stable home ranges, which vary from 38 ha (Castellanos 1985) to 685 ha (Carrillo et al. 2002; Taber et al. 1994). White-lipped peccaries can travel up to 13 km/day and there is evidence that they are nomadic or migratory, driven by mast fruiting events of trees, and of palms in particular (Altrichter and Almeida 2002; Bodmer 1990; Kiltie and Terborgh 1983; Peres 1994a, 1994b, 1996). However, radiotracking studies, which have only been undertaken on a peninsula (Carrillo et al. 2002) and an island (Fragoso 1998a), suggest that white-lipped peccaries have stable home ranges. It remains unclear how these settings affect the movement of peccaries.

Data Acquisition and Analyses

To obtain information on peccaries and their interactions with palms, I surveyed international publications focusing on palms, peccaries, and peccary-palm interactions in several databases such as Science Citation Index, Web of Science, and Biological Abstracts. In addition, I presented a poster and solicited information at the 2003 Association for Tropical Biology and Conservation annual meeting and I had announcements published in the journal Palms (2003) and in Tropinet j (Association for Tropical Biology and Conservation supplementary publication, 2004), requesting additional information. Based on the published literature and personal communications I compiled a database on palm species whose roots, seedlings, flowers, fruits, or seeds were consumed by peccaries. In addition, I recorded the seed fate and the method employed by the author. If some data were not provided, whenever possible I contacted the author to obtain the missing information. I excluded palm species when authors could only speculate that peccaries fed on them. I excluded the data from statistical analyses when authors did not specify which peccary species consumed a given palm species.

I used chi-square goodness-of-fit tests to determine if there were differences in the frequency of seed species destroyed versus not destroyed for each peccary species. In addition, I used a 2 × 2 contingency test to quantify if there was a difference in the frequency of seeds destroyed versus not destroyed between the 2 peccary species. I performed these comparisons across the neotropical region because many palm species (i.e., Attalea butyracea, Iriartea deltoidea, and Mauritia flexuosa) or genera (i.e., Astrocaryum, Euterpe, and Socratea, which have similar properties at the species level, such as seed size or hardness of the endocarp) have large geographic distributions in which both peccary species occur sympatrically. All analyses were executed in SPSS version 11.5 (SPSS 2002).

Palm Consumption by Peccaries

I found 76 papers published between 1919 and 2004 specifying peccary-palm interactions. In total, both peccary species consumed roots, seedlings, flowers, fruits, or seeds from 46 palm species. Collared peccaries fed on 25 palm species ( Appendix I), whereas white-lipped peccaries used 37 palm species ( Appendix II). Publications where authors did not distinguish between the 2 peccary species are in  Appendix III, which lists 3 additional palm species not reported in the previous 2 appendices. Fruits and seeds of Astrocaryum, Attalea, Euterpe, and Syagrus were the 4 genera most frequently consumed by the 2 peccary species (Fig. 1).

Fig. 1

Frequency distribution of palm species with fruits consumed by collared and white-lipped peccaries across the Neotropics and the numbers of palm species whose seeds were destroyed after ingestion.

Negative Peccary-Palm Interactions

Peccaries as seed predators.—Palms use several strategies to minimize seed predation (Bodmer 1991a; Bodmer and Ward, in press; Henderson 2002). For example, Oenocarpus bataua possesses lignified fibers surrounding the seeds (Bodmer 1991a), thereby minimizing seed death through chemical digestion, whereas Phytelephas and Mauritius flexuosa have seeds with very large and extremely hard coats (Henderson 1995; Kiltie 1982), thereby preventing mastication and ingestion.

Of a total of 22 palm species where data on seed fate were provided, collared peccaries preyed upon seeds of certain genera disproportionately more frequently on a per-species basis (χ² = 4.54, d.f = 1, P = 0.033; Fig. 1). Similarly, of 33 species consumed by white-lipped peccaries, seeds from 26 species were disproportionately more frequently destroyed (χ² = 10.8, d.f = 1, P = 0.001; Fig. 1). Thus, both peccary species are primarily seed predators, but there was no difference in the frequency of seed species killed between the 2 peccary species (χ² = 0.37, d.f. = 1, P = 0.539).

The ecological significance of peccaries as seed predators can be best illustrated by studies that quantified seed survival and seedling recruitment in the presence and absence of peccaries. For example, during the absence of peccaries, Wyatt and Silman (2004) reported a 5,340% increase of uneaten seeds of Iriartea deltoidea and a 6,000% increase of Astrocaryum murumuru seeds underneath conspecifics. Similarly, Silman et al. (2003) found that during a 12-year period in which peccaries were absent, the number of Astrocaryum seedlings increased by 70%; it was reduced by 71% after recolonization by peccaries. Furthermore, in the absence of peccaries, the distribution of Astrocaryum seedlings was uniform compared to before absence and after recolonization.

Peccaries and seedling trampling.—Peccaries trampling on palm seedlings can affect their density and spatial distribution. For example, Roldán and Simonetti (2001) demonstrated that the seedling mortality of Astrocaryum was higher in the presence than in the absence of peccaries. Higher foraging activity of peccaries in areas where Attalea maripa had a clumped distribution resulted in higher seedling mortality compared to areas with single standing individuals (Fragoso 1997, 1998b). Similar results of seedling trampling were reported in other studies (Asquith et al. 1997; Dirzo and Miranda 1990; Ticktin 2003; Wright et al. 2000; Wright and Duber 2001).

Peccaries as herbivores.—Peccaries have been reported uprooting and consuming the roots and the base of seedlings of Attalea maripa, Mauritia flexuosa (Fragoso 1999), Acrocomia aculeata, Attalea phalerata, and Euterpe edulis (A. Keuroghlian, in litt.). In Costa Rica, collared peccaries have been observed chewing on the stilt roots of Socratea exorrhiza (O. Vargas, in litt.). I observed that the root systems of numerous Socratea were so heavily damaged that 1 person alone could push over the palms. Most palm species are arborescent, so their leaves, flowers, and fruits are beyond the reach of peccaries. However, numerous palms in the genera Calyptrogyne or Geonoma are small-sized understory plants, and thus are exposed to herbivory by peccaries. Altrichter et al. (2000) reported that white-lipped peccaries frequently broke off the inflorescences of several Geonoma species and consumed their flowers ( Appendix II). The negative ecological consequences of peccary herbivory on palm roots and flowers have not yet been quantified.

Positive Peccary-Palm Interactions

Small seeds (≤ 1 cm) from numerous tree species germinated successfully after being recovered from fecal matter of peccaries (Beck 2005). However, all studies except 1 (Kiltie 1981b) found only fragments of palm seeds in stomach or fecal samples of peccaries (Altrichter et al. 2000, 2001; Barreto et al. 1997; Bodmer 1991a, 1991b; Henry 1997; Keuroghlian 2003; Martínez-Romero and Mandujano 1995; Varela and Brown 1995). Thus, peccaries appear to be poor palm seed dispersers, which is certainly related to the relatively large size of the seed of palms (diameter of 45 mm for Astrocaryum murumuru [Henderson 2002] compared to 1.5 mm for Ficus seed) and the powerful mastication apparatus of peccaries.

Peccaries frequently only chew off the fruit pulp and expectorate seeds if the endocarp is too hard to be cracked open. Examples of expectorated palm seeds include Acrocomia aculeate, Attalea butyracea (Janzen and Martin 1982), Attalea maripa (Fragoso 1997, 1998b), Attalea phalerata (A. Keuroghlian, in litt.), Attalea speciosa (Anderson 1983), Euterpe edulis (Reis 1995), Oenocarpus bataua (Bodmer 1991a), Mauritia flexuosa (Bodmer 1991a; Bodmer and Ward, in press; Fragoso 1997), and Raphia teadigera (M. Altrichter, in litt.). Some of these expectorated seeds are trampled into the ground by foraging peccaries, resulting in lower seed predation, particularly from insects (Fragoso 1997, 1998b; Mitja and Ferraz 2001; Silvius 1999; Silvius and Fragoso 2002). Thus, peccaries can enhance germination rate under parent palms, which may be 1 mechanism promoting clumped distribution (Beck and Terborgh 2002; Fragoso 1997, 1998b; Howe 1989; Hubbell 1979). Conversely, scatter-hoarding rodents such as Proechimys (Forget 1991), Myoprocta (Forget 1991; Morris 1962), and Dasyprocta (Silvius and Fragoso 2003; Smythe 1970a, 1970b) generate a leptokurtic seed shadow (dispersal declines with distance from tree—Howe and Smallwood 1982) when dispersing seeds around parent palms or trees (Forget et al. 2002; Jansen and Forget 2001).

Feeding experiments on captive peccaries and field observations indicated that peccaries prefer seeds that are infested with insect larvae such as those of bruchid beetles (Fragoso 1994; Jansen 2003; Kiltie 1981a; Silvius 2002). Thus it is possible that peccaries affect or control bruchid beetle populations in a top-down fashion and thereby indirectly decrease future infestation and mortality of palm seeds. However, experimental studies are needed to test this notion.

Intra- and Interspecific Competition

Both peccary species consumed fruits and seeds of the same 23 palm species, and their overlap in palm species use was 59% (Appendices I and II). When considering only those palm species where both peccary species destroyed the seeds, overlap was still 52% (12 out of 23). The decline in overlap of fruits and seeds consumed versus seeds destroyed might indicate some resource partitioning. White-lipped peccaries are able to crack the stony endocarp of several palm species including Attalea and Socratea, which is rarely possible for collared peccaries (Appendices I and II). For instance, to crack open seeds of Mauritia flexuosa or Iriartea deltoidea, peccaries have to overcome a crushing resistance of over 350 kg. Using a lever model of the jaw, Kiltie (1982) estimated that white-lipped peccaries have a bite force 1.3 times stronger than that of collared peccaries (Kiltie 1981c). Sicuro et al. (2002) reported parallel findings based on different craniological comparisons. Kiltie and Terborgh (1983) suggested that the different bite forces of these 2 peccary species and the hardness of palm endocarps was 1 mechanism to maintain a diet niche divergence of sympatric peccary species (see also Sicuro et al. 2002). Numerous studies provide empirical evidence for resource partitioning based on endocarp hardness (Barreto et al. 1997; Bodmer 1989; Fragoso 1998b, 1999; Olmos 1993; Painter 1998), whereas Fragoso (1999) indicated that differences in scale perception might also lead to resource partitioning.

It is well known that many tropical frugivores other than peccaries consume the fruit pulp and prey upon palm seeds. For example, spiny rats (Proechimys), paca (Agouti paca), agouti (Dasyprocta), brocket deer (Mazama), and tapir (Tapirus) consume fruits and seeds of palms (Beck 2002, 2005; Beck and Terborgh 2002; Beck-King et al. 1999; Bodmer 1991b; Bodmer and Ward, in press; Galetti et al. 2001; Gayot et al. 2004; Henderson 1995; Painter 1998; Silvius and Fragoso 2002; Smythe 1978, 1986; Smythe et al. 1985; Wright 2003; Zona and Henderson 1989). Therefore, peccaries may compete exploitatively with other terrestrial frugivores for palm fruits and seeds. One mechanism through which rodents, including Proechimys, Myoprocta (Adler and Kestell 1998; Forget 1990, 1991; Morris 1962; Smythe 1978), and Dasyprocta (Smythe 1970a, 1970b, 1978) might minimize competition is by scatter-or larder-hoarding seeds. Those seeds are frequently cached near landmarks such as fallen logs, possibly to facilitate their recovery (Forget 1990, 1991; Jansen 2003; Kiltie 1981a; Kiltie and Terborgh 1983). Interestingly, there is empirical evidence that peccaries preferentially forage near objects (Kiltie 1981a) for those hidden seeds (Fragoso 1999; Kiltie 1981a; Kiltie and Terborgh 1983; Smythe 1986, 1989). Thus, peccaries might also compete with other frugivorous in a kleptoparasitic fashion, a yet underappreciated mechanism.

Conclusions

An earlier literature review of animal-palm interactions on a worldwide scale located 3 papers indicating that peccaries consumed fruits and seeds from 5 palm species (Zona and Henderson 1989). This review located 76 papers that revealed that peccaries consumed resources from 46 palm species across the Neotropics (Appendices I—III). Both peccary species destroyed seeds of over 73% of species that they interacted with (Fig. 1), highlighting the ecological role of peccaries as palm seed predators (Fig. 2). Several authors have linked the evolution of the strong mastication apparatus and the unique interlocking canines (which prevents dislocation of the lower jaw when cracking seeds) of peccaries as adaptations to crack open hard seeds, in particular those of palms, which are among the hardest (Bodmer 1991b; Kiltie 1981a, 1981c, 1982; Kiltie and Terborgh 1983; Sicuro et al. 2002; Silman et al. 2003; Terborgh 1986a). Furthermore, the movement and foraging ecology of white-lipped peccaries is primarily driven by mast fruiting events of several palm species, including Astrocaryum murumuru, Astrocaryum tucuma, Attalea maripa, Lepidocaryum, Mauritia flexuosa, Orbygynia phalerata, and Raphia taedigera (Beck and Terborgh 2002; Fragoso 1994, 1998a, 1998b, 1999; Janzen and Wilson 1983; Kiltie 1982; Kiltie and Terborgh 1983; Peres 1994a, 1994b, 1996).

Fig. 2

Summary of peccary-palm interactions and their ecological ramifications. On the top are predictions of the ecological consequences of seed dispersal, predation, and herbivory. Below are predictions based on the different foraging strategies of peccaries.

The strength of the revolutionary forces that have been operating between peccaries and palms can best be assessed by studies that use a comparative approach. As mentioned above, Silman et al. (2003) and Wyatt and Silman (2004) found that in areas where peccaries were locally extinct, seed survival increased over 5,000%, also resulting in dramatic changes in the spatial distribution of seedlings underneath parent palms compared to areas with peccaries. Numerous other studies found similar effects for peccaries or other frugivores (Asquith et al. 1997; Dirzo and Miranda 1990, 1991; Terborgh 1988, 1992; Terborgh et al. 1997, 1999, 2001; Ticktin 2003; Wright et al. 2000; Wright and Duber 2001; Wright 2003).

Because peccaries affect seed survival, seedling recruitment, and distribution underneath and in close proximity to parent palms (Fig. 2), peccaries may be a crucial force for the Janzen-Connell effects (Connell 1971; Janzen 1970). In brief, the Janzen-Connell hypothesis predicts that seed predators prevent competitive exclusion by affecting distance- or density-dependent seed survival. Thereby seed predators influence the recruitment, distribution, and species richness of plants. Thus extinction events of peccaries appear to offset the Janzen-Connell mechanisms, as demonstrated by Wyatt (2002), and thereby not only affect the spatiotemporal distribution and demography of palms, but also the community composition and diversity of tropical ecosystems (see also Asquith et al. 1997; Dirzo and Miranda 1990, 1991; Silman et al. 2003; Terborgh 1988, 1992; Terborgh et al. 1997, 1999, 2001; Ticktin 2003; Wright 2003; Wright et al. 2000; Wright and Duber 2001).

In Figure 2, I summarized peccary-palm interactions and their ecological ramifications, and derived a new hypothesis which should be tested in the future. Results from these studies will not only elucidate peccary-palm interactions in more detail, but may also provide new conservation implications.

Conservation Implications

Because overhunting, habitat destruction, and fragmentation have led to a continuous decline and local extinctions of peccary populations throughout the Neotropics (Beck 2005 and citations therein), several authors consider peccaries one of the most endangered neotropical species (Ceballos and Navarro 1991; Sowls 1997; Taber 1991), which may explain why both species are listed in  Appendix II of the Convention for International Trade in Endangered Species (CITES 2003—http://www.cites.org). However, additional threats must also be considered. In 1853, Alfred R. Wallace described in detail the usefulness and extensive harvesting of palms by humans across the Amazon. More recently, widespread and destructive harvesting of palm fruits (i.e., practices such as cutting down female Mauritia palms to gain access to their seeds) and palm hearts pose an additional threat to many frugivores that depend heavily on palm food resources, as outlined here (Bodmer et al. 1999; Bodmer and Ward, in press; Galetti and Fernandez 1998).

Because of their large group sizes, foraging ecology, and area requirements, white-lipped peccaries are much more susceptible to human threats than are collared peccaries (Cullen et al. 2000; Peres 1996, 2001; Sowls 1997). In fact, they are almost completely extinct in Central America. Therefore, white-lipped peccaries should be listed on  Appendix I of the Convention for International Trade in Endangered Species. Furthermore, the issues of overhunting, habitat destruction, and destructive palm harvesting must become international conservation priorities.

Acknowledgments

I thank M. Altrichter, R. Bodmer, S. Brewer, F. Feer, M. Galetti, A. Keuroghlian, S. Mori, F. Olmos, C. Peres, M. Silman, J. Terborgh, J. Vandermeer, O. Vargas, and S. Zona for providing field observations or unpublished manuscripts. Thanks to F. Reid for permission to reproduce her drawing. M. Silman and 3 anonymous reviewers provided valuable comments.

Appendix I

Herbivory, frugivory, and seed fate for palm fruits eaten by collared peccaries (Pecari tajacu) across the Neotropics. Abbreviations defined in footnotes. Henderson et al. (1995) are followed for nomenclature, using names indicated in footnotes ^g–^m in place of names used in references for palm species.

Palm species	Growth forma	Part usedb	Seed fatec	Sited	Methode	Habitatf	Reference
Acrocomia aculeatag	tall	fr, se,	surv, not det	Pan	observ, fecal	Rain	Enders 1930, 1935; A. Keuroghlian, in litt.
Acrocomia vinifera	med	sl, ro fr, se	killed	CR	observ	Rain	Baker 1983
Astrocaryum aculeatum	med	fr	surv	Bra	observ	Rain	Silvius 1999
Astrocaryum chambira	med	fr	not det	Per	observ	Rain	Lleellish et al. 2003
Astrocaryum jauari	med	fr, se	killed	Ecu	observ	Rain	Pedersen and Balslev 1990
Astrocaryum mexicanum	med	fr, se	killed	Bel, Mex	observ, feeding	Rain	Brewer 2001; Martínez-Ramos 1997; Sánchez-Cordero and Martinez-Gallardo 1998
Astrocaryum murumuru	med	fr, se	killed	Per, Bol	observ, stomach	Rain	Cintra 1997; Kiltie 1981a, 1981b, 1981c, 1982; Kiltie and Terborgh 1983; Roldán and Simonetti 2001
Astrocaryum paramaca	med	fr, se	killed	FG	feeding	—	F. Feer, in litt.
Astrocaryum	med	fr, se	killed, not det	Per, Bra	observ, feeding	Rain	Schaller 1983, Terborgh and Kiltie 1976
Astrocaryum standleyanum	med	fr, se	killed	Pan	observ	Rain	Smythe 1970a, 1970b, 1978, 1989
Attalea butyraceah	med	fr, se	surv, killed,	Pan, Mex,	observ	Rain,	Coe and Due 1980; Enders 1930, 1935;
			not det	CR		Dry	Janzen and Martin 1982
Attalea maripai	med	fr, sl, ro	surv	Bra	observ	Rain	Fragoso 1997, 1999; Silvius 1999
Attalea phalerata	med	fr, se,	surv, killed	Bol	observ	Rain	A. Keuroghlian, in litt.; Quiroga-Castro and
		sl, ro					Roldán 2001
Attalea speciosaj	med	fr	surv	Bra	observ	Dry	Anderson 1983
Attalea sp.k	med	fr, se	surv, not det	Bra, Per,	observ, feeding,	Rain	Duke 1967; Kiltie 1981a, 1981b, 1982;
				Hon	stomach		Schaller 1983
Bactris maraja	small	se	killed	Bra	observ	Rain	Fragoso 1999
Copernicia tectorum	tall	fr	not det	Ven	observ	Gal	Robinson and Eisenberg 1985
Euterpe edulis	tall	fr, se, sl	surv, killed	Bra, Per	observ	Rain	Galetti et al. 1999; Keuroghlian 2003;
							Lleellish et al. 2003; Reis 1995
Euterpe sp.	med	fr	not det	Per	stomach	Rain	Bodmer 1990
Iriartea deltoideal	tall	fr, se	surv, killed	Per	observ, stomach	Rain	Cintra 1998; Kiltie 1981a, 1981b, 1982;
							O. Vargas, in litt.
Jessenia	tall	fr, se	killed	Per	observ, feeding,	Rain	Kiltie 1981a, 1981b, 1981c, 1982;
					stomach		Kiltie and Terborgh 1983; Terborgh and Kiltie 1976
Mauritia flexuosa	tall	fr, se,	surv, killed	Ecu, Bra,	observ, feeding	Rain	Bodmer 1991a; Bodmer and Ward, in press;
		sl, ro		FG, Per			F. Feer, in litt.; Fragoso 1999;
							Kiltie 1981b, 1982; Lleellish et al. 2003;
							Peres 1994a; Urrego 1987
Oenocarpus batauam	tall	fr	surv, not det	Per	observ, feeding	Rain	Bodmer 1991a; Lleellish et al. 2003
Oenocarpus	tall	fr	not det	Hon	observ	Rain	Duke 1967
Phytelephas microcarpa	med	fr	surv	Per	observ, feeding	Rain	Kiltie 1982
Raphia taedigera	med	fr, se	killed	CR	observ	Rain	Janzen and Wilson 1983
Socratea exorrhiza	tall	fr, se, ro	killed, not det	CR, Per	observ	Rain	Lleellish et al. 2003; O. Vargas, in litt.
Syagrus oleracea	tall	fr	surv	Bra	observ, fecal	Rain	Keuroghlian 2003
Syagrus romanzoffiana	tall	fr, se	surv, killed	Bra	observ, fecal, feeding	Rain	Keuroghlian 2003; Olmos et al. 1999
Welfia georgii	tall	fr, se	killed	CR	observ	Rain	Vandermeer et al. 1979; O. Vargas, in litt.

a

Growth form of palms: tall = reaching upper canopy, med = reaching medium canopy, small = reaching subcanopy, underst = understory plant.

b

Part used by peccaries: flow = flower, fr = fruit pulp, ro = root, se = seed, sl = seedlings.

c

Seed fate: killed = seed killed, surv = seed survived digestion and potentially dispersed, not det = not determined.

d

Site: Arg = Argentina, Bel = Belize, Bol = Bolivia, Bra = Brazil, Col = Colombia, CR = Costa Rica, Ecu = Ecuador, FG = French Guiana, Hon = Honduras, Mex = Mexico, an = Panama, Per = Peru, Sur = Suriname, Ven = Venezuela.

e

Methods: fecal = fecal analysis, feeding = feeding experiment (because experiments were performed on captive animals no habitat is indicated), observ = observation, omach = stomach contents analysis.

f

Habitat: Cer = cerrado, Dry = tropical dry forest, Gal = gallery forest, Humid = humid tropical forest, Rain = tropical rain forest, not det = not determined.

g

Acrocomia aculeata = Acrocomia sclerocarpa.

h

Attalea butyracea = Attalea gymphococca, Scheelea liebermanii, Scheelea rostrata, Scheelea zonensis, or Orbignya martiana.

i

Attalea maripa = Attalea regia or Maximiiiana maripa.

j

Attalea speciosa = Orbignya martiana.

k

Attalea sp. — Scheeleae.

l

Iriartea deltoidea = Iriartea ventricosa.

m

Oenocarpus bataua — Jessenia bataua.

Appendix II

Herbivory, frugivory, and seed fate for palm fruits eaten by white-lipped peccaries (Tayassu pecari) across the Neotropics. Abbreviations and other details as for  Appendix I.

Palm species	Growth form	Part used	Seed fate	Site	Method	Habitat	Reference
Acrocomia aculeatag	tall	fir, sl, ro	surv	Bra	observ	Rain	A. Keuroghlian in litt.
Astrocaryum aculeatum	med	fir, se	killed	Bol, Bra	observ	Rain	Painter 1998; Silvius 1999
Astrocaryum chambira	med	fr	not det	Per	observ	Rain	Lleellish et al. 2003
Astrocaryum jauari	med	fir, se	killed	Ecu	observ	Rain	Pedersen and Baislev 1990
Astrocaryum mexicanum	med	fir, se	killed	Bel	observ	Rain	Brewer 2001
Astrocaryum murumuru	med	fr, se	killed	Per	observ, feeding,	Rain	Cintra 1997, 1998; Cintra and Horna 1997;
					stomach		Kiltie 1981a, 1981b, 1981c, 1982;
							Kiltie and Terborgh 1983; Wyatt 2002;
							Wyatt and Silman 2004
Astrocaryum paramaca	med	fr, se	killed	FG	feeding	—	F. Feer, in litt.
Astrocaryum sciophilum	med	fr, se	killed	FG	observ	Rain	Sist 1989a
Astrocaryum	med	fr, se	killed,	Bra, CR,	observ, fecal,	Humid,	Altrichter et al. 1999, 2000; Duke 1967;
			not det	Hon, Per	feeding	Rain	D. Moskovits, in litt.; Schaller 1983;
							Terborgh and Kiltie 1976
Astrocaryum standleyanum	med	fr, se	killed	Pan	observ	Rain	Smythe 1970a, 1970b, 1989
Astrocaryum tucuma	med	fr, se	not det	Bra	observ	Rain	Peres 1996
Astrocaryum vulgare	med	fr, se	not det	Sur	observ	Rain	Husson 1978; Penard et al. in
							Husson 1978
Attalea butyraceah	med	fr, se	killed	CR, Per	observ, fecal	Humid,	Altrichter et al. 2000; Wyatt 2002;
						Rain	Wyatt and Silman 2004
Attalea maripai	med	fr, se,	surv, killed,	Bra, Sur	observ	Rain	Husson 1978; Fragoso 1998a, 1998b,
		sl, ro	not det				1999; Penard et al. in Husson 1978; Silvius 1999, 2002
Attalea phalerata	med	fr, se, ro	surv, killed	Bol	observ	Rain	A. Keuroghlian, in litt.;
							Quiroga-Castro and Roldán 2001
Attalea speciosaj	med	fr	surv	Bra	observ	Dry	Anderson 1983
Attalea sp.k	med	fr, se	surv, not det	Bra, Per	observ, feeding	Rain	Kiltie 1981a, 1982; Schaller 1983
Bactris glaucescens	small	fr, se	killed	Bra	observ	Rain	A. Keuroghlian, in litt.
Bactris sp.	small	se	killed	CR	observ, fecal	Humid	Altrichter et al. 1999, 2000
Copernicia alba	tall	fir, se	killed	Bra	observ	Rain	A. Keuroghlian, in litt.
Copernicia tectorum	tall	fr	surv	Ven	fecal	Dry	Barreto et al. 1997
Crysophila guagara	tall	se	killed	CR	observ, fecal	Humid	Altrichter et al. 1999, 2000
Dictyocaryum lamarckianum	med	fir, se	killed	Pan, Per	observ	Rain	S. Mori, in litt., M. Silman, in litt.
Elaeis guineensis	tall	fr	surv	FG	feeding	—	F. Feer, in litt.
Euterpe edulis	tall	fr, se, sl	surv, killed	Bra, Per	observ, fecal	Rain	Galetti et al. 1999; Keuroghlian 2003;
							Lleellish et al. 2003; Reis 1995
Euterpe oleracea	tall	fr, se	killed	Bra	observ	Rain	Smith 1976
Euterpe precatoria	tall	fr, se	killed	Bra	observ	Rain	Silvius 1999
Geonoma cuneata	underst	flow	—	CR	observ	Humid	Altrichter et al. 2000
Geonoma	underst	flow	—	CR	observ	Humid	Altrichter et al. 2000
Iriartea deltoideal	tall	fr, se	killed	CR, Per	observ, fecal,	Humid,	Altrichter et al. 1999, 2000; Cintra 1998;
					feeding,	Rain	Kiltie 1981a, 1981b, 1981c, 1982;
					stomach		Kiltie and Terborgh 1983; Wyatt 2002;
							Wyatt and Silman 2004
Iriartea	tall	fr, se	killed,	Bra, Per	observ, feeding,	Rain	Bodmer 1990; Smith 1976; Terborgh and Kiltie 1976
			not det		stomach
Jessenia	med	fr, se	killed	Per	observ, stomach,	Rain	Terborgh and Kiltie 1976; Kiltie 1981b, 1981c, 1982; Kiltie and Terborgh 1983
					feeding
Lepidocatyum	med	fr	not det	Bra	observ	Rain	Peres 1996
Mauritia flexuosa	tall	fr, se,	surv, killed	Bra, Ecu,	observ, feeding	Rain	Bodmer 1991a; Bodmer and Ward, in press; F. Feer, in litt.;
		sl, ro		FG, Per
							Fragoso 1998a, 1998b,
							1999; Kiltie 1981b, 1981c, 1982;
							Kiltie and Terborgh 1983; Lleellish et al. 2003; Silvius 1999; Urrego 1987
Oenocarpus batauam	tall	fr	surv, not det	Hon, Per	observ, feeding	Rain	Bodmer 1991a; Duke 1967;
							Lleellish et al. 2003
Phytelephas microcarpa	med	fr	surv	Per	observ, feeding	Rain	Kiltie 1982
Raphia taedigera	med	fr, se	killed, not det	CR	observ	Rain	Janzen and Wilson 1983; Carrillo 2000
Socratea durissima	tall	fr, se	killed	Per	observ	Rain	Kiltie 1981c; Kiltie and Terborgh 1983
Socratea exorrhiza	tall	fr, se	surv, killed	Bra, Bol,	observ, feeding,	Rain	F. Feer, in litt.; Kiltie 1981b,
				FG, Per	stomach		1981c, 1982; Lleellish et al. 2003;
							Painter 1998; Silvius 1999
Socratea sp.	tall	fr, se	killed	Per	feeding	Rain	Terborgh and Kiltie 1976
Syagrus oleracea	tall	fr	surv	Bra	observ, fecal	Rain	Keuroghlian 2003
Syagrus romanzoffiana	tall	fr, se	surv, killed,	Bra	observ, fecal, feeding	Rain	Cullen et al. 2001; Keuroghlian 2003;
			not det				F. Olmos, in litt.
Syagrus sancona	tall	fr, se	killed	Bol	observ	Rain	Painter 1998

Appendix III

Herbivory, frugivory, and seed fate for palm fruits eaten by both peccary species (Pecari tajacu and/or Tayassu pecari) across the Neotropics— In these references authors did not specify which peccary species was involved. Abbreviations and other details as for  Appendix I.

Palm species	Grow form	Part used	Seed fate	Site	Method	Habitat	Reference
Acrocomia mexicana	med	fr, se	killed	Mex	observ	Dry	Gaumer 1917
Astrocaryum murumuru	med	fr, se	killed	Per	observ	Rain	Beck and Terborgh 2002
Astrocaryum sciophilum	med	fr, se	killed	FG	observ	Rain	Sist 1989a
Astrocaryum sp.	med	fr, se	not det	Per	observ	Rain	Terborgh 1986a
Attalea butyraceag	med	fr, se	surv, killed	CR	observ	Dry	Janzen and Martin 1982
Butia leiospatha	med	fr, se	surv	Bra	observ	Cer	Gottsberger and Silberbauer-
							Gottsberger 1983
Iriartea deltoideah	tall	fr, se	killed	Per	observ	Rain	Terborgh 1986a
Jessenia polycarpa	med	fr, se	surv	Col	observ	Rain	West 1957
Syagrus romanzoffiana	tall	fr, se	surv	Arg	observ	Humid	Varela and Brown 1995

Author notes

Present address: Department of Biological Sciences, Towson University, 8000 York Road, Towson, MD 21252, USA

Associate Editor was Gerardo Ceballos.