What does the occurrence of Sporormiella (Preussia) spores mean in Australian fossil sequences?

Understanding the loss of the final few species of Australian megafauna is beset by a paucity of data on human arrival, well‐provenanced megafauna, human/megafauna population range and distribution (coexistence and interaction), and the range, scale and impact of environmental changes spanning the human–megafauna period. To overcome these shortcomings, the occurrence and decline of coprophilous fungal spores of Sporormiella in sediments have been used as a proxy for extinct megaherbivores. The Sporormiella evidence is presented as the key indicator of extinction timing and these reports are often from locations where there is no known archaeological record or megafauna remains. However, interpreting fungal spore occurrence is not straightforward, as demonstrated by studies investigating taphonomy, taxonomy and the types of animal dung where Sporormiella occurs. No detailed studies on these problems exist for Australia and no evidence supporting the use of Sporormiella as a valid proxy has been reported. Here we examine the occurrence of Sporormiella spores from Cuddie Springs in south‐eastern Australia. Despite a well‐preserved suite of megafauna fossils, Sporormiella occurrence is sporadic and frequencies are low. We conclude that using Sporormiella alone as an indicator for the presence of megafauna is premature for the Australian context.     

Chronological overlap between humans and megafauna in Sahul (Pleistocene Australia–New Guinea): A review of the evidence

Over 60 faunal species disappeared from the Australian continent during the Middle–Late Pleistocene. Most of these animals were large to gigantic marsupials, birds and reptiles. A terminal extinction date of 46.4 kyr has been proposed for the megafauna, with all sites containing younger fossil megafauna dismissed by some researchers because of questions over stratigraphic integrity or chronologies. The timing of the extinctions is argued to be broadly coincident with estimates of first colonization of the continent by modern humans, and explanatory extinction models involving humans have subsequently gained currency. However there is considerable evidence to suggest that in some parts of the continent, people and some species of megafauna may have co-existed well beyond 46.4 kyr. In other places, such as Tasmania and the north of the continent, there is no known record of a human–megafauna temporal overlap. A review of the available evidence indicates that only 13 species of megafauna were extant on human arrival in Australia. The archaeology of this period indicates that rather than a focus on big game hunting or ‘firestick farming’, it was characterized by regional variability in subsistence strategies consistent with the range of environmental zones. At the present time there is no substantive argument for a terminal extinction date of 46.4 kyr, the current evidence indicating that there is no specific time period that correlates to any single mass extinction event. On the basis of available evidence arguments for either human or climatic causation are entirely circumstantial and implicitly require acceptance of many unproven assumptions. Claims to have eliminated climate as a primary driver are premature and the recent focus on delivering ‘proof’ of human causation in Pleistocene faunal extinctions diverts attention from achieving a better understanding of the differential impacts of climate change and short term climatic flux in a land of environmental extremes.     

Dating megafaunal extinction on the Pleistocene Darling Downs,eastern Australia: the promise and pitfalls of dating as a test of extinction hypotheses

A key to understanding Late Pleistocene megafaunal extinction dynamics is knowledge of megafaunal ecological response(s) to long-term environmental perturbations. Strategically, that requires targeting fossil deposits that accumulated during glacial and interglacial intervals both before and after human arrival, with subsequent palaeoecological models underpinned by robust and reliable chronologies. Late Pleistocene vertebrate fossil localities from the Darling Downs, eastern Australia, provide stratigraphically-intact, abundant megafaunal sequences, which allows for testing of anthropogenic versus climate change megafauna extinction hypotheses. Each stratigraphic unit at site QML796, Kings Creek Catchment, was previously shown to have had similar sampling potential, and the basal units contain both small-sized taxa (e.g., land snails, frogs, bandicoots, rodents) and megafauna. Importantly, sequential faunal horizons show stepwise decrease in taxonomic diversity with the loss of some, but not all, megafauna in the geographically-small palaeocatchment. The purpose of this paper is to present the results of our intensive, multidisciplinary dating study of the deposits (>40 dates). Dating by means of accelerator mass spectrometry (AMS) ¹⁴C (targeting bone, freshwater molluscs, and charcoal) and thermal ionisation mass spectrometry U/Th (targeting teeth and freshwater molluscs) do not agree with each other and, in the case of AMS ¹⁴C dating, lack internal consistency. Scanning electron microscopy and rare earth element analyses demonstrate that the dated molluscs are diagenetically altered and contain aragonite cements that incorporated secondary young C, suggesting that such dates should be regarded as minimum ages. AMS ¹⁴C dated charcoals provide ages that occur out of stratigraphic order, and cluster in the upper chronological limits of the technique (～40–48 ka). Again, we suggest that such results should be regarded as suspicious and only minimum ages. Subsequent OSL and U/Th (teeth) dating provide complimentary results and demonstrate that the faunal sequences actually span ～120–83 ka, thus occurring beyond the AMS ¹⁴C dating window. Importantly, the dates suggest that the local decline in biological diversity was initiated ～75,000 years before the colonisation of humans on the continent. Collectively, the data are most parsimoniously consistent with a pre-human climate change model for local habitat change and megafauna extinction, but not with a nearly simultaneous extinction of megafauna as required by the human-induced blitzkrieg extinction hypothesis. This study demonstrates the problems inherent in dating deposits that lie near the chronological limits of the radiocarbon dating technique, and highlights the need to cross-check previously-dated archaeological and megafauna deposits within the timeframe of earliest human colonisation and latest megafaunal survival.     

Megafauna demography and late Quaternary climatic change in Australia: A predisposition to extinction

Arguments about the extinction of Australia's megafauna have largely rested on anthropogenic factors consequent upon the arrival of humans there, and have lacked any appreciation of the possibilities of climate/environmental changes taking place during the late Quaternary. Moreover, the status of the megafauna at the extinction and in the period leading up to it has largely been ignored. This article assesses the species that existed during the late Quaternary, their continental dispersal, the likely impact of negative climate change during that time and the effect this had on their demography and variety. These factors are discussed together with a synthesis of present data regarding Australia's mega 2004: fauna demography and which species may have reached the extinction threshold. One interpretation of the data suggests a mid–late Quaternary process of demographic fragmentation, disjunction and fluctuation, a restricted continental distribution among a diminishing group and a limited and reducing species variety due to climate and environmental change. It is argued that increasing continental aridity during the mid–late Quaternary was a forcing mechanism behind species distribution, changes to that distribution and population reduction through episodic but widespread drought and vegetation change. This resulted in alteration of the biogeographic status of the megafauna, with increasing stress on and reduction of the population as a whole. In particular, it changed population composition and reduced species variety and overall population size by the beginning of the last glaciation, such that at the time of human entry the population had reached a precarious stage vulnerable to any level of subsequent anthropogenic activity with the arrival of humans in Australia.     

Revisiting the late Pleistocene mammal extinction record at Tight Entrance Cave, southwestern Australia

Tight Entrance Cave (TEC) in southwestern Australia provides a Pleistocene sequence documenting the extinction of 14 large mammal species. This record has been interpreted as indicating that extinctions did not occur during or before the penultimate glacial maximum (PGM) and that humans played a primary role in the extinctions. However, it remains possible that the majority of extinct megafauna persisted no later than the PGM. The TEC extinctions correspond with vegetation change, a cooling/drying trend, increased biomass burning, and increasingly unstable small mammal communities. The initiation of these trends predates human arrival on the continent and implies environmentally mediated extinctions.     

The timing of Late Pleistocene mammalian extinctions in North America

More than 375 ¹⁴C dates from 150 fossil sites in North America have been analyzed to evaluate the question of extinction of Late Pleistocene megafauna. When critically evaluated, no ¹⁴C ages for any extinct Pleistocene genera are younger than 10,000 yr B.P.     

Last glacial megafaunal death assemblage and early human occupation at Lake Menindee, southeastern Australia

The Tedford subfossil locality at Lake Menindee preserves a diverse assemblage of marsupials, monotremes and placental rodents. Of the 38 mammal taxa recorded at the site, almost a third are of extinct megafauna. Some of the bones are articulated or semi-articulated and include almost complete skeletons, indicating that aeolian sediments rapidly buried the animals following death. New optical ages show the site dates to the early part of the last glacial (55,700 ± 1300 yr weighted mean age). This is close to the 51,200-39,800 yr Australia-wide extinction age for megafauna suggested by Roberts et al. [2001, Science 292:1888-1892], but like all previous researchers, we cannot conclusively determine whether humans were implicated in the deaths of the animals. Although an intrusive hearth at the site dating to 45,100 ± 1400 yr ago is the oldest evidence of human occupation of the Darling River, no artifacts were identified in situ within the sub-fossil-bearing unit. Non-anthropogenic causes, such as natural senescence or ecosystem stress due to climatic aridity, probably explain the mortality of the faunal assemblage at Lake Menindee.     

Electron spin resonance dating of the fossil deposits in the Naracoorte Caves,South Australia

The caves near Naracoorte, South Australia, contain one of the richest and most diverse fossil faunal assemblages on the Australian continent. Three sites were selected for electron spin resonance (ESR) dating because clastic, fossiliferous sediments were sandwiched between speleothem layers. This allows independent age control by highly precise thermal ionization mass‐spectrometry (TIMS) U‐series dating. We find that all ESR results agree within the constraints given by the U‐series dates, and allow further refinement of the age of the fauna analysed, indicating that most of the fauna in the large Victoria Cave Fossil Chamber is twice as old as reported previously. Our dating results, spanning from 280 to 500 ka for the Fossil Chamber, Victoria Cave, to about 125 ka for the Grant Hall, Victoria Cave, and 170 to 280 ka for the Fossil Chamber, Cathedral Cave, indicate little change, if any, in the megafaunal assemblage from the early Middle to the early Late Pleistocene. This changed dramatically after the last interglacial, when a large proportion of the megafauna suddenly disappeared. Copyright © 2001 John Wiley & Sons, Ltd.     

Arctic Siberia: refuge of the Mammoth fauna in the Holocene

Global climate change at the end of Pleistocene led to extinction in the huge territories of Northern Eurasia of the typical representatives of the Mammoth fauna: mammoth, woolly rhinoceros, wild horse, bison, musk-ox, and cave lion. Undoubtedly the Mammoth fauna underwent pressure from Upper Paleolithic humans, whose hunting activity could also have played a role in decreasing the number of mammoths and other representatives of megafauna. Formerly it was supposed that the megafauna of the “Mammoth complex” had become extinct by the beginning of the Holocene. Nevertheless the latest data indicate that extinction of the Mammoth fauna was significantly delayed in the north of Eastern Siberia. In the 1990s some radiocarbon dates established that mammoths existed in the Holocene on Wrangel Island—from 7700 until 3700 yBP. Radiocarbon data show that wild horses inhabited the north of Eastern Siberia 4600–2000 yBP. Muskoxen lived here about 3000 yBP. Some bison remains from Eastern Siberia belong to the Holocene. The following circumstances could promote the survival of representatives of Mammoth fauna. Cool and dry climate in this region promotes the maintenance of steppe associations—the habitats of those mammals. Late Paleolithic and Mesolithic settlements are not found in the Arctic zone of Eastern Siberia from Taimyr Peninsula to the lower Yana River; they are very rare in basins of the Indigirka and Kolyma Rivers. The small number of Stone Age hunting tribes in the northern part of Eastern Siberia was probably another factor that contributed to the survival of some Mammoth fauna representatives.     

New U/Th ages for Pleistocene megafauna deposits of southeastern Queensland,Australia

Arguments over the extinction of Pleistocene megafauna have become particularly polarised in recent years. Causes for the extinctions are widely debated with climate change, human hunting and/or habitat modification, or a combination of those factors, being the dominant hypotheses. However, a lack of a spatially constrained chronology for many megafauna renders most hypotheses difficult to test. Here, we present several new U/Th dates for a series of previously undated, megafauna-bearing localities from southeastern Queensland, Australia. The sites were previously used to argue for or against various megafauna extinction hypotheses, and are the type localities for two now-extinct Pleistocene marsupials (including the giant koala, Phascolarctos stirtoni). The new dating allows the deposits to be placed in a spatially- and temporally constrained context relevant to the understanding of Australian megafaunal extinctions. The results indicate that The Joint (Texas Caves) megafaunal assemblage is middle Pleistocene or older (>292 ky); the Cement Mills (Gore) megafaunal assemblage is late Pleistocene or older (>53 ky); and the Russenden Cave Bone Chamber (Texas Caves) megafaunal assemblage is late Pleistocene (～55 ky). Importantly, the new results broadly show that the sites date prior to the hypothesised megafaunal extinction ‘window’ (i.e., ～30–50 ky), and therefore, cannot be used to argue exclusively for or against human/climate change extinction models, without first exploring their palaeoecological significance on wider temporal and spatial scales.     

The Pleistocene/Holocene transition in Peru and its effects upon human use of the landscape

This paper reviews the state of our knowledge regarding the effects of the Pleistocene/Holocene transition on the kinds and range of human adaptations in what is now Peru. Following a discussion of geological, paleoclimatic, and archaeological data, the paper focuses upon four aspects of environmental change with specific regard to how these would have affected human adaptations: environmental changes on the Pacific littoral, the extinction of Pleistocene faunal and floral species and their replacement with modern faunas, the adaptive radiation of these modern flora and fauna, especially into the Andean highlands, and characteristics of newly available high elevation environments that would have affected the process by which foraging peoples moved into them. Although Pleistocene fauna have been discovered in Peru, none of these finds have been made in the context of indisputable human activity, and therefore, the effect of their extinction on early foraging peoples is unknown. The earliest acceptable archaeological sites in Peru date around 11–12,000 years ago, and are found on the coastal lowlands. The highlands were not occupied until after 11,000 years ago. While high elevation environments were attractive after 11,000 years ago, they were only slowly occupied by humans due to the constraints of the combined effects of hypoxia and cold.     

Late Pleistocene biota from Pubenza,Colombia; turtles,mammals, birds,invertebrates and plant remains

Pubenza is a remarkable palaeontological site of Colombia, and a place that could hold some of the potentially oldest evidence of humans in northern South America. Previous palaeontological research at this site has mainly focused on the megafauna. Here we describe and establish the systematic palaeontology for the small fauna that inhabited this ancient lacustrine ecosystem, including the first report of birds, tortoises and vipers for the Late Pleistocene in Colombia. Furthermore, exceptionally well preserved fossilised wasp nests are morphologically and elementally characterised, which is the first report of an ichnofossil of this kind in northern South America. In addition, new material of kinosternid turtles, armadillos and rodents is also described. Our results reveal that the Bogotá River Basin, where Pubenza is located, was a rich ecosystem during the Late Pleistocene and a region of great interest for future articulated palaeontological and archaeological studies.     

Geochronology,sediment provenance,and fossil emplacement at Sumidouro Cave,a classic late Pleistocene/early Holocene Paleoanthropological site in eastern Brazil

Peter Wilhelm Lund's (1845a) heavily debated suggestion of a contemporaneity between Paleo‐Indians and extinct Pleistocene fauna at Sumidouro Cave was re‐examined through detailed sedimentological and geochronological analyses of sediment and both human and faunal remains. Sources of the cave's sediment include both entrances as well as ceiling fissures. Non‐human fossils, on the other hand, were probably carried by floodwater through the once more‐spacious swallet entrance. Seasonal flooding reworked and mixed these two highly asynchronous assemblages. U‐series and radiocarbon ages indicate that there are at least two distinct episodes of sediment input in the cave, at ˜240,000 yr B.P. and ˜8000 yr B.P. Human remains represent a later emplacement event, probably at ˜8400 cal yr B.P. Although the human remains are of considerable age, the cave's complex stratigraphy, flooding dynamics, and extensive removal of the cave's filling during earlier excavations do not allow the determination of an unequivocal co‐existence between Paleo‐Indians and extinct megafauna at the site. © 2005 Wiley Periodicals, Inc.     

Late Pleistocene sedimentology,taphonomy and megafauna extinction on the Darling Downs,southeastern Queensland

The Kings Creek catchment, southeastern Queensland, contains a variety of Pleistocene – Holocene depositional settings. Fluvial depositional accumulation processes in the catchment reflect both high-energy channel and low-energy episodic overbank deposition. The lithofacies and depositional environments of locality QML796 were examined in detail to aid interpretation of taphonomic accumulation patterns of large and small taxa in the deposit. The basal fossiliferous unit was deposited in a meandering channel and passes upward into overbank deposits that include ephemeral interfluve channels and splays. The most striking taphonomic observations on vertebrates at the locality include: (i) low representation of post-cranial elements; (ii) high degree of bone breakage; (iii) variable abrasion with most identifiable bone elements having a low to moderate degree of abrasion; (iv) low rates of bone weathering; (v) a low degree of carnivore bone modification; and (vi) a low degree of articulated or associated specimens. Collectively, these data suggest that the material was transported into the deposit from the surrounding proximal floodplain and that the assemblages reflect substantial hydraulic sorting. However, despite that, sequential faunal horizons show a stepwise decrease in taxonomic diversity that cannot be explained by sampling or taphonomic bias. The decreasing diversity includes loss of some, but not all, megafauna and is consistent with a progressive local loss of megafauna in the catchment over an extended interval of time. Data are consistent with a climate change model for megafauna extinction but not with nearly simultaneous extinction of megafauna as required by the human-induced blitzkrieg extinction hypothesis.     

Bearing up well? Understanding the past,present and future of Australia's koalas

The modern Koala Phascolarctos cinereus is the last surviving member of a once diverse family Phascolarctidae (Marsupialia, Phascolarctomorphia). Nine genera and at least 16 species of koala are known. Late Oligocene sediments of central Australia record the oldest fossils and highest species diversity. Five species are known from the early to middle Miocene rainforest assemblages of the Riversleigh World Heritage Area, Queensland. With the onset of dryer conditions after the middle Miocene climatic optimum (~ 16 Ma), rainforest habitats contracted resulting in the apparent extinction of three koala lineages (Litokoala, Nimiokoala, Priscakoala). Phascolarctos first appears in the fossil record during the Pliocene and the modern species around 350 ka. Despite a dramatic decline in taxonomic diversity to a single extant species, the fossil record indicates that at most only three koala species coexisted in any given faunal assemblage throughout their 24 million year history. Within these assemblages, the vast majority of extinct koalas are extremely rare (some known from only a single specimen) which may reflect a general rarity within their palaeohabitats compared with the modern species which is represented by an estimated 400,000 individuals spread over most of eastern mainland Australia. Be that as it may, P. cinereus, although once geographically more widespread, occurring for example in Western Australia in the Pleistocene, underwent significant range contractions and localized population extinctions during the stressful climatic conditions of the late Pleistocene and more recently through human-induced habitat destruction. Combined with threats of disease, reduced genetic diversity and climate change, the survival of this iconic Australian marsupial is arguably a cause for concern.     

The North of Eastern Siberia: Refuge of Mammoth Fauna in the Holocene

The global climate changings at the end of Pleistocene led to extinction of the typical representatives of Mammoth fauna–mammoth, woolly rhinoceros, wild horse, bison, muskox, cave lion, etc.–on the huge territories of Northern Eurasia. Undoubtedly the Mammoth fauna underwent pressure from the Upper Paleolithic Man, whose hunting activity also could play the role in decreasing the number of mammoths and other representatives of megafauna (large mammals). Archaeological data testify that the typical representatives of Mammoth fauna were the Man's hunting objects only till the end of the Pleistocene. Their bone remains are not usually found on the settlements of Mesolithic Man. Formerly it was supposed that the megafauna of ‘Mammoth complex’ was extinct by the beginning of Holocene. Nevertheless the latest data testify that the global extinction of the Mammoth fauna was sufficiently delayed in the north of Eastern Siberia. In the 1990s some radiocarbon data testified that the mammoths on the Wrangel Island existed for a long time during the Holocene from 8000 till 3700 y. BP. The present radiocarbon data show that wild horses inhabited the north of Eastern Siberia (the lower stream of the Enissey river, the Novosibirskie Islands, the East Siberian sea-shore) 3000–2000 y. BP. Musk-oxen lived on the Taimyr Peninsula and the Lena River delta about 3000 y. BP. Some bison remains from Eastern Siberia belong to the Holocene. The following circumstances could promote the process of preservation of the Mammoth fauna representatives. The cool and dry climate of this region promotes the maintenance of steppe associations – habitats of those mammals. The Late Paleolithic and Mesolithic settlements are not found in the Arctic zone of Eastern Siberia from the Taimyr Peninsula to a lower stream of the Yana River; they are very rare in the basins of the Indigirka and Kolyma Rivers. So, the small number of the Stone Age hunting tribes on the North of Eastern Siberia was another factor in the long-term preservation of some Mammoth fauna representatives.     

Taxonomic analysis of the Quaternary archaeofauna found at The Lapa do Santo site,Lagoa Santa region,Brazil

The Lapa do Santo archaeological site, located in the Lagoa Santa region, Brazil, represents an important hunter-gatherer occupation dated from the Early and Middle Holocene. Prior studies of archaeofauna dating to this period are few and most of them only provide basic faunal identification with limited information on taphonomic processes. The main goal of this study is to identify the archaeofauna, record the taphonomic processes, and make inferences about its natural (interpreted as the death of the animal in the rockshelter area due to natural causes or due to predation) or anthropic origins. No extinct species (including megamastofauna) were identified. Taxonomic analysis indicated that most faunal remains consist of taxa that may be of mixed origin (natural or anthropic), such as microvertebrates and carnivorous mammals. The Cervidae are the main vertebrate family found in the studied material and these are likely the result of hunting. The Cervidae Ozotoceros support the presence of a fauna that is typical of savannah environments, although the assemblage contains mostly generalist fauna that can thrive in a variety of environments. Very few remains showed clear evidence for taphonomic processes related to human interaction, such as burnt bones.     

Palaeotrophic reconstruction and climatic forcing of mega‐Lake Eyre in late Quaternary Central Australia: a review

Webb, S. 2009: Palaeotrophic reconstruction and climatic forcing of mega‐Lake Eyre in late Quaternary Central Australia: a review. Boreas, 10.1111/j.1502‐3885.2009.00120.x. ISSN 0300‐9483. Extreme Quaternary climatic variation in Australia brought radical environmental changes to various parts of the continent. In this article, I discuss these changes in terms of mega‐lake development in Central Australia, and in particular the southern Lake Eyre Basin (SLEB). The formation of these features, together with the fossil record of the region, throws light on the palaeoclimatic and palaeobiological relationships of megafauna and other animal groups, and the trophic development required to support them. Australian continental drying during the late Quaternary has been noted by many workers, but this process was punctuated by strong pluvial episodes of decreasing strength from MIS 5e. Mega‐lake development during MIS 5 resulted from unusual monsoonal and evaporative patterns at that time. However, the climatic forcing behind mega‐lake formation and the rate of lake growth is not well understood, although species composition in SLEB aquatic fossil fauna assemblages attests to the size and development of these lakes and indicates their long‐term persistence. The degree of trophic development and the maintenance of broad, well‐bedded aquatic and terrestrial ecological frameworks and biotic variety support that conclusion. The fossil record contributes to our understanding of mega‐lake and palaeoriverine trophic complexity, the speed and duration of lake‐fill and the intensity and persistence of the supporting intracontinental moisture balance. Although other more remote mega‐lakes formed in Central Australia, they were not populated by complex trophic systems or megafauna populations. This discrepancy between the various geographic areas sheds light on the biogeography and population distribution of megafauna, thus helping form a better picture of the reasons behind the final extinction of relict populations of this group in MIS 4.     

内蒙古赤峰东村更新世早期哺乳动物群

内蒙古赤峰西北地区的东村、东梁一带更新世早期河-湖相沉积物相当发育,为一套白色砂与棕红色亚粘土互层,其中含有丰富的哺乳动物化石,计11科18属21种。动物群性质与泥河湾(狭义)的相当。这是我国最东和最北的一个三趾马与真马共生的化石地点。     

New Quaternary Mammalian Faunas and Cave Deposits in the Zhoukoudian Area,Beijing

From 1985 to 1987, four new localities with abundant fossil mammals were dis-covered by Cao, Tian and others in the Zhoukoudian (Choukoutien) area, Beijing. They are theEast, West, Shangdian and Donglingzi caves. The East Cave fauna consists of 28 speices ofmammals and its age is middle Early Pleistocene. The East Cave assemblage shows that a tem-perature-falling event took place at around 1.20 Ma B.P. at Zhoukoudian. Sixteen species ofmammals were collected from the West Cave, which are mainly forms of late Early Pleistoceneage. The West Cave fauna represents a transitional fauna from the East Cave fauna (dry-cold)to the fauna (warm) at locality 9. The Shangdian Cave fauna is composed of four forms, beingMiddle Pleistocene in age. The Donglingzi Cave fauna contains 21 Late Pleistocene forms. Inthe cave two fossil horizons may be distinguished. The age of the lower horizon is early LatePleistocene, which is equivalent to that of the New Cave fauna; while the fauna of the upper ho-rizon may be correlated with the Upper Cave fauna.