identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038BEA16FFAB7808FCF2FC99FCBFFBAB.text	038BEA16FFAB7808FCF2FC99FCBFFBAB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeinae Latreille 1802	<div><p>Scarabaeinae: Onthophagini</p> <p>Onthophagus (Figs. 35–36) is a large genus found throughout the world and consists of approximately 2,000 species (Tarasov and Kabakov 2010). The two species taken in this study (Table 1) were found only in the dung traps, and neither was abundant, seeming instead to prefer more open habitats as in my other pitfall study in a secondary forest remnant on the grounds of INPA in Manaus (unpublished data).</p> </div>	https://treatment.plazi.org/id/038BEA16FFAB7808FCF2FC99FCBFFBAB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
038BEA16FFAB7808FCE4FBC6FCF4F9D2.text	038BEA16FFAB7808FCE4FBC6FCF4F9D2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeinae Latreille 1802	<div><p>Scarabaeinae: Oniticellini</p> <p>The genus Eurysternus Dalman (Fig. 37) was recently reclassified in the Oniticellini and contains 53 species that are found only in the Neotropics (Génier 2009). Several species of Eurysternus are abundant in the Amazon Basin where most seem to be coprophages. The nidification behavior of only 10 species are known, and this includes complex nesting behavior. Five species of Eurysternus were collected at Reserva Ducke, primarily in the dung traps but occasionally in the carrion traps (Table 1). Eurysternus atrosericus Génier (Amazon basin of Brazil, Ecuador, Guyana, Venezuela) was the most abundant species collected. It was found throughout the year and represented 92% of all Eurysternus taken in the dung traps.</p> </div>	https://treatment.plazi.org/id/038BEA16FFAB7808FCE4FBC6FCF4F9D2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
038BEA16FFAB780FFCE7F9CEFDDAFDDE.text	038BEA16FFAB780FFCE7F9CEFDDAFDDE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeinae Latreille 1802	<div><p>Scarabaeinae: Canthonini</p> <p>Canthon (Figs. 38–39) consists of about 150 species found from Canada to Argentina (Gill 2002). Three species of Canthon were taken at Reserva Ducke, with C. triangularis being the most abundant (Table 1). Canthon triangularis (Fig. 39, frontispiece) (Amazonian parts of Brazil, Suriname, French Guiana, Colombia, Bolivia) is diurnal, often seen resting on leaves, and is a dung roller; it represented 85% of all Canthon collected in the dung traps and 15% taken in the carrion traps.</p> <p>Canthonella Chapin is a small genus of 16 species that occur in the West Indies and South America (Ratcliffe and Smith 1999). Matthews (1965) observed them opportunistically feeding on bird, snail, and mammal dung, and the five specimens of Canthonella silphoides (Harold) taken in this study were collected in both dung and carrion traps (Table 1).</p> <p>Cryptocanthon Balthasar (Fig. 40) is another small genus with 35 species occurring from Mexico to Brazil (Cook 2002). They inhabit the litter of lowland rainforests and montane cloud forests. Cryptocanthon peckorum Howden (Brazil, Colombia, Bolivia; Cook 2002) was col- lected in both dung (89%) and carrion traps (11%) at Reserva Ducke (Table 1).</p> <p>Deltochilum Eschscholtz (Figs. 41–43) is a New World genus with more than 80 species (Gill 2002). Most species feed on carrion, but all five species taken at Reserva Ducke were also collected in the dung traps. Interestingly, the few specimens of Deltochilum orbiculare Lansberge were collected only in the dung traps. Deltochilum diringshofeni (Brazil) (61% on carrion) and D. septemstriatum (French Guiana, Brazil) (78% on carrion) were the two most abundant species of Deltochilum at Reserva Ducke (Table 1).</p> <p>Scybalocanthon Martínez has approximately 19 species that occur in Central and South America (Silva 2011). Molano Rendón and Medina Uribe (2010) reported Scybalocanthon species are usually diurnal, and that they are known to feed on vertebrate dung preferentially but also on carrion and fungi. Scybalocanthon pygidialis (Schmidt) (Brazil, Colombia, Suriname) was collected from both dung (92%) and carrion traps (8%) at Reserva Ducke (Table 1).</p> </div>	https://treatment.plazi.org/id/038BEA16FFAB780FFCE7F9CEFDDAFDDE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
038BEA16FFAC780FFF2BFDF2FC24FB26.text	038BEA16FFAC780FFF2BFDF2FC24FB26.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeinae Latreille 1802	<div><p>Scarabaeinae: Dichotomiini</p> <p>Aphengium Harold is a small South American genus with only three species. Only two specimens of Aphengium cupreum Shipp were collected at Reserva Ducke, one each at dung and carrion (Table 1). Virtually nothing is known of these rare beetles.</p> <p>Ateuchus (Fig. 44) contains about 92 species distributed from the USA to Argentina (Gill 2002; Larsen 2012a). This large genus is in great need of revision, and identification for many of the species is difficult. Ten species were taken at Reserva Ducke, but only four could be reliably identified (Table 1). All but three species came exclusively to dung. Ateuchus murrayi (Amazon basin of Brazil, Colombia, Ecuador) was collected in large numbers in dung (96%), but some individuals also were collected in the carrion traps (4%).</p> <p>Canthidium is comprised of approximately 170 species found from southern Arizona to South America (Gill 2002). Like Ateuchus, this genus is in great need of revision, and it is currently difficult to identify many species with any reliability. Fourteen species were collected at Reserva Ducke, mostly in the dung traps; Canthidium sp. #2 was the exception with nearly five times more individuals collected in the carrion traps (Table 1). Canthidium bicolor Boucomont (Fig. 45) (Brazil, Colombia, Peru, Bolivia) and Canthidium sp. near gigas Balthasar (French Guiana, Brazil) are coprophages (this study), whereas Canthidium gerstaeckeri Harold (Fig. 46) (Brazil, Peru, Colombia, Venezuela, Guyana, Bolivia) feeds primarily on dung but also less frequently on carrion, dead invertebrate animals, fruit, and fungi (Larsen 2012b).</p> <p>Dichotomius is a large genus of 150–200 species found from the USA to southern South America (Gill 2002).The genus is in great need of revisionary studies, and many species probably remain to be described. Most of the species are large, nocturnal coprophages. Four species were collected at Reserva Ducke, and the vast majority were taken throughout the year in the dung traps (Table 1). Dichotomius boreus (Olivier) (Fig. 47) (Brazil, Suriname, French Guiana, Colombia, Ecuador) (99% on dung) and Dichotomius sp. near mamillatus (Felsche) (Fig. 48) (Amazon basin of Brazil, Colombia, Ecuador, Peru, Bolivia) (98% on dung) were the two most abundant species in the study.</p> <p>Ontherus Erichson has 58 species that occur from Mexico to Argentina (Génier 1996). Only one species, Ontherus carinifrons Lüderwaldt (Brazil, one record for Peru; Génier 1996), was found at Reserva Ducke (Table 1), exclusively on dung.</p> <p>Trichillum Harold has, depending on the authority one consults, 50–100 species (e.g., Vaz-de-Mello 2003) distributed in the Neotropics. Two species were collected at Reserva Ducke. Trichillum boucomonti Saylor (Brazil, Colombia, Bolivia, Paraguay) was found almost exclusively in the carrion traps (99%), and Trichillum hirsutum Boucomont (Brazil, Colombia, Bolivia, Paraguay) was divided between the dung and carrion traps (62% on dung versus 38% on carrion) (Table 1).</p> <p>Uroxys is comprised of about 60 species that are found from Mexico to Argentina. Three species were collected at Reserva Ducke, and two of these were represented by singletons taken in dung traps (Table 1). One of these, Uroxys besti Ratcliffe (Amazonian Brazil), is known to inhabit the fur of sloths (Ratcliffe 1980) and may not normally come to dung in pitfall traps. Uroxys pygmaeus (Amazonian Brazil) (Fig. 49), on the other hand, was taken in huge numbers throughout the year (7,721 specimens), primarily on dung (98% versus 2% on carrion).</p> </div>	https://treatment.plazi.org/id/038BEA16FFAC780FFF2BFDF2FC24FB26	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
038BEA16FFAC780DFCF4FB5AFF3CFEB3.text	038BEA16FFAC780DFCF4FB5AFF3CFEB3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Scarabaeinae Latreille 1802	<div><p>Scarabaeinae: Phanaeini</p> <p>Coprophanaeus has 38 species distributed from the southern USA to Argentina (Edmonds and Zídek 2010). Three species were collected at Reserva Ducke, all fairly abundantly (Table 1) with C. jasius exceeding a thousand specimens. Coprophanaeus jasius (Brazil, Guyana, French Guiana, Suriname, Argentina, Bolivia, Ecuador, Peru, Venezuela, Colombia) is a nocturnal burrower and feeds primarily on carrion but also on dung (86% on carrion versus 14% on dung). It was active throughout the year.</p> <p>Coprophanaeus lancifer (frontispiece, Fig. 50) (Brazil, Venezuela, French Guiana, Guyana, Suriname, Bolivia, Peru, probably Colombia and Ecuador) is the largest species (30–50 mm) in the study area and one of the most conspicuous with its bright metallic blue coloration. It is crepuscular and cruises just above ground while foraging. It is primarily necrophagous, but it has also been observed feeding on dung (64% on carrion versus 36% on dung in this study). It was active throughout the year and had a peak collecting event in mid-January. Vulinec et al. (2009) also observed it feeding on rotting bananas and occasionally attracted to lights at night. Edmonds and Zídek (2010) characterized this species as the best choice as an indicator species of Amazonia.</p> <p>Oxysternon (Fig. 51) is comprised of 11 mostly brightly colored species that are found in the Neotropics (Edmonds and Zídek 2004). Two species were collected exclusively in the dung traps at Reserva Ducke (Table 1), and neither was abundant.</p> <p>Phanaeus MacLeay has 54 primarily metalliccolored species occurring from the USA to Argentina (Edmonds and Zídek 2012). Only Phanaeus chalcomelas (Perty) (Fig. 52) (Brazil, Bolivia, Colombia, Ecuador, French Guiana, Peru, Suriname) was collected at Reserva Ducke (Table 1) and only in the dung traps. This species is diurnal.</p></div> 	https://treatment.plazi.org/id/038BEA16FFAC780DFCF4FB5AFF3CFEB3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
038BEA16FFAE7832FE88FC8CFB6EFB21.text	038BEA16FFAE7832FE88FC8CFB6EFB21.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trogidae MacLeay 1819	<div><p>Trogidae</p> <p>Trogid adults typically feed on carrion in the late stages of decay, although they are occasionally encountered on fresher remains. Only two species of Trogidae, Omorgus suberosus (F.) and Omorgus batesi (Harold), were known from Brazil’ s Amazon Basin until Polynoncus juglans (Ratcliffe) was described (Ratcliffe 1978) based upon 11 specimens captured during this study (Table 1). Most trogids prefer drier habitats than those of Amazonia and, as necrophages, are typically outcompeted by ants and vultures in tropical rainforests, hence their rarity.</p> <p>CONCLUSIONS</p> <p>In this paper, I discuss trapping methodology and contrast the results from traps using two different solutions for preservation. I also review the species of scarabaeoids that were taken and relate their abundance and taxonomic diversity to local climatic factors. Concurrent with this study, I also conducted a similar trapping and inventory program for 43 weeks in a small remnant patch of secondary forest on the INPA campus 28 km from the Reserva Ducke site (unpublished data). As expected, fewer numbers (5,901 specimens) and fewer species (34) were collected on the INPA campus due to the lack of many dung producing mammals (agoutis were the largest), fewer number of traps (six as opposed to 20), and absence of a reservoir population of replacement scarabaeines. Eight species were collected on the INPA campus that were not taken in the Ducke study site, again reflecting the difference in biotopes as it affects species composition.</p> <p>The guild of scarabaeoids in tropical rainforests relying on dung is more diverse in species and larger in numbers that those relying on carrion. The Reserva Ducke study supports that hypothesis with 100% of the species and 74% of the numbers collected in dung traps and 52% of the species and 26% of the numbers taken in carrion traps. The comparisons of picric acid and chloral hydrate solutions resulted in virtually no significant differences in species diversity or numbers of individuals, and so the preferred solution is chloral hydrate because it is more readily available and safer to use.</p> <p>Dung and carrion in tropical rainforests are generally more limited in space and time because the biomass of mostly mammals producing the dung and carrion is less than in, for example, the tropical savannas of Africa (Cambefort and Walter 1991; Davis and Sutton 1997). Competitive success depends on the ability of a beetle to rapidly locate dung or carrion. Some beetles have evolved strategies that reduce competition. The most obvious is removal and burial of dung and carrion. Davis and Sutton (1997) observed that while most species are generalist coprophages, some species evolved to feed on fruit, fungi, carrion, and plant detritus or specialized on omnivore or herbivore dung to enhance coexistence of potential competitors with the result of increased species richness. Others developed arboreal lifestyles that may have evolved as a response to competition with the ground fauna for a common resource that is transitory and limited (Ratcliffe 1980; Howden and Young 1981; Davis et al. 1997; Jacobs et al. 2008). Peck and Forsyth (1982) and Feer and Pincebourde (2005) concluded that most species use a spectrum of dung types, and it is the fine partitioning of different activity periods, foraging behaviors, body sizes, dung removal methods, feeding behaviors, and microclimate that contribute to so many species co-existing in the same habitat.</p> <p>Subsequent to this study in the late 1970s, many large mammals were extirpated from the study area, thus reducing the dung resources needed for scarabaeine diversity. The total number of species (66 in the traps, others observed but not trapped) at the Reserva Ducke study site in the 1970s is today probably reduced due to the reduction in the populations of the dung producers. Removal of large-bodied mammals reduces dung resource availability that will then differentially affect dung beetles using either the “perch and wait” strategy observed in small-bodied species or the foraging on-the-wing strategy for locating large dung pats common to larger species of dung beetles (Peck and Howden 1984; Gill 1991). Perching species, primarily diurnal and smaller, usually feed on pelleted droppings typical of rodents and small ungulates, and so these species may persist after the larger mammals have been removed by hunting. Conversely, larger species that are primarily nocturnal (Cambefort and Walter 1991; Feer and Pincebourde 2005) may disappear because the larger dung pats upon which they rely will no longer be available (Halffter et al. 1992; Nichols et al. 2009). Today, only the smaller species of mammals remain at Reserva Ducke (Vulinec 2000), and the genera of large-bodied beetles most likely to be deleteriously affected would be Dichotomius, Phanaeus, Coprophanaeus, and Oxysternon. Nichols et al. (2009) concluded that this kind of restructuring of dung beetle communities has profound implications for the maintenance of several ecosystem functions, including reduction in nutrient recycling and reduction in burial of excreted seeds with concomitant change in plant associations and patterns of regrowth.</p> <p>Chazdon et al. (2009) noted that degraded old-growth forests (affected by road building, selective logging, recurrent fires, fragmentation, and hunting) and secondary regrowth forests comprise roughly half of the world’ s remaining tropical forests. Quintero and Roslin (2005) suggested that the regrowth of fragments of secondary tropical vegetation in Amazonia a mere decade after clear cutting of forest supported dung beetle assemblages similar to those in formerly continuous forest. They did not mention what mammals remained in these fragments, but the dung beetles captured in their study were all smallbodied species with the exception of C. jasius. While they demonstrated that secondary forest growth can restore some conditions favorable for certain species of dung beetles, I believe the original fauna of large-bodied dung beetles cannot be restored due to the absence of larger mammals and the dung resources they provide. In another study in Amazonia, Gardner et al. (2008) concluded from their data that tropical secondary and plantation forests were of low value for restoring dung beetle populations. In view of the detrimental consequences for maintenance of dung beetlemediated ecosystem services by these impoverished dung beetle communities, they urged that conservation strategies prioritize the protection of remaining areas of primary forest rather than relying on secondary forests. I concur.</p> <p>Some colleagues have expressed concerns to me of “trapping out” the beetles during such a long-term, intensive survey. I believe the species composition and abundance data clearly show that this did not happen, even though collecting was done for slightly more than a year. However, this could be a concern in forest fragments where the reservoir of dung-producing mammals and additional species or individuals of scarabs is limited by lack of suitable habitat or adequate food resources. In Reserva Ducke at the time of this study, the collecting grid represented only a tiny area of the forest, and so the reservoir of dung- and carrionfeeding scarabs was substantial and always available to replace the 19,000 specimens collected.</p> <p>Goodland and Irwin (1975), Myers (1984), Klein (1989), Whitmore (1997), Fearnside (1990, 2009), and many others have observed that fragmentation of Amazonian rainforests, indeed all tropical forests, is continuing unabated at an alarming rate. The cascading consequences of selective extinction in remaining forest fragments results in a depauperate fauna and flora and a concomitant decrease in attendant ecosystem services. Unlike some of those disappearing dung beetles, humankind is in deep shit if this continues to happen. I am reminded of a sentiment from the World Wildlife Fund:</p> <p>All that lives beneath Earth’ s fragile canopy is, in some elemental fashion, related.</p> <p>Is born, moves, feeds, reproduces, dies.</p> <p>Tiger and turtle dove; each tiny flower and homely frog; the running child, father to the man and, in ways as yet unknown, brother to the salamander.</p> <p>If mankind continues to allow whole species to perish, when does their peril also become ours?</p> <p>Humans need to step up and assume enhanced stewardship for the biota with which we share the planet... if it is not already too late. As H. G. Wells said long ago before there was a biodiversity crisis, “Human history more and more becomes a race between education and catastrophe”. And the answer to the WWF soliloquy above is NOW.</p> </div>	https://treatment.plazi.org/id/038BEA16FFAE7832FE88FC8CFB6EFB21	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Ratcliffe, Brett C.	Ratcliffe, Brett C. (2013): The Dung- and Carrion-Feeding Scarabs (Coleoptera: Scarabaeoidea) of an Amazonian Blackwater Rainforest: Results of a Continuous, 56 - Week, Baited-PitfallTrap Study. The Coleopterists Bulletin 67 (4): 481-520, DOI: 10.1649/0010-065x-67.4.481, URL: http://dx.doi.org/10.1649/0010-065x-67.4.481
