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Late Quaternary terrestrial vertebrate coprolites from New Zealand
Quaternary Science Reviews 98 (2014) 33e44
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Late Quaternary terrestrial vertebrate coprolites from New Zealand Jamie R. Wood a, *, Janet M. Wilmshurst a, b a b
Landcare Research, PO Box 69040, Lincoln, Canterbury 7640, New Zealand School of Environment, The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand
a r t i c l e i n f o
a b s t r a c t
Article history: Received 29 April 2014 Received in revised form 21 May 2014 Accepted 22 May 2014 Available online
Over the past decade, concerted efforts to find and study Late Quaternary terrestrial vertebrate coprolites in New Zealand have revealed new insights into the diets and ecologies of New Zealand's prehistoric birds. Here, we provide a broader review of the coprolites found in natural (non-archaeological) Late Quaternary deposits from New Zealand. We summarise the morphological diversity of the coprolites, and discuss the taphonomy of the sites in which they are found. Since the 1870s more than 2000 coprolites have been discovered from 30 localities, all restricted to the South Island. The distribution of coprolite localities appears to reflect the presence of geological and climatic factors that enhance the potential for coprolite preservation; coprolites require dry conditions for preservation, and have been found on the ground surface within drafting cave entrances and at shallow (<300 mm) depths beneath rock overhangs with a northerly aspect. We classify the coprolites into eleven morphotypes, each of which may represent a range of different bird and/or reptile species. A review of genetically identified specimens shows that coprolites of different bird species overlap in size and morphology, reinforcing the need for identifications to be based on ancient DNA analysis. © 2014 Elsevier Ltd. All rights reserved.
Keywords: Birds Caves Coprolites New Zealand Reptiles Rock shelters Stratigraphy Taphonomy
1. Introduction The fossil record of New Zealand's Late Quaternary terrestrial fauna is impressively complete (Worthy and Holdaway, 2002). Abundant fossil bones from caves, mires, dunes, and loess deposits have provided detailed insights into not only the identity of the faunal species that once inhabited New Zealand, but also their ecology and distribution from the last ice age until the arrival of humans in the 13th Century AD (Worthy and Holdaway, 2002). Analyses of Late Quaternary bones continue to reveal new information about the biology of New Zealand's extinct species, and technological advances (particularly in the field of ancient DNA analysis) continue to open up exciting new research possibilities (Worthy and Scofield, 2012). Evidence for New Zealand's Late Quaternary fauna is not restricted to bones. Further proxies include Maori rock art depictions (McCulloch and Trotter, 1971), preserved feathers and integument (reviewed by Rawlence et al., 2013), nests (Hartree, 1999; Wood, 2006, 2008a), eggshell (Gill, 2000, 2006; Oskam et al., 2010), footprints and trackways (reviewed by Worthy and Holdaway, 2002), sedimentary ancient DNA (Willerslev et al., 2003; Haile et al., 2007) and coprolites (Horrocks et al., 2004, *Corresponding author. Tel.: þ64 3 321 9653. E-mail address: [email protected] (J.R. Wood). http://dx.doi.org/10.1016/j.quascirev.2014.05.020 0277-3791/© 2014 Elsevier Ltd. All rights reserved.
2008; Wood et al., 2008, 2012a, 2012b, 2012c, 2013a, 2013b; Wood and Wilmshurst, 2013). Coprolites are rich sources of paleoecological information, and can reveal aspects of the paleoenvironments (Yll et al., 2006), paleodiets (Poinar et al., 1998), ecosystem function (Wood et al., 2012a) and parasites (Schmidt and Duszynski, 1992) of prehistoric extinct fauna. Late Quaternary coprolites of extinct terrestrial fauna were first discovered in New Zealand approximately 140 years ago (Cockburn-Hood, 1873), and until recently remained relatively scarce. Within the past decade, concerted efforts to identify and recover Late Quaternary coprolites from New Zealand for study have greatly expanded our understanding of this unique paleoecological resource, making it timely for a broader review. Here, we review the history of Late Quaternary coprolite (hereafter simply referred to as coprolites) discoveries in New Zealand, assess the morphological diversity of the coprolites, and discuss the taphonomy of the sites in which they occur. Many coprolites, mainly of the Polynesian dog (Canis familiaris), have also been found in association with archaeological sites throughout New Zealand (e.g. Horrocks et al., 2002, 2003) but are not reviewed here, as we focus specifically on non-archaeological specimens. 2. A brief history of coprolite discoveries in New Zealand The first coprolites discovered in New Zealand were found during excavations at Earnscleugh Cave (Fig. 1) in Central Otago
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Fig. 1. Location of sites containing natural Late Quaternary terrestrial vertebrate coprolites in New Zealand. Names of regions are shown in grey text.
during the early 1870s. Cockburn-Hood (1873) noted that “The flat ground near [the cave] had probably been a favourite camping ground, from the quantity of droppingsewhich are, no doubt, those of the large birdseswept in by the wind”. Five coprolites from Earnscleugh Cave are held by Otago Museum, and one of these has been genetically identified as having been deposited by a coastal moa (Euryapteryx curtus) (Wood et al., 2008). Around the same period, White (1875) discovered a cave near Mt. Nicholas (Fig. 1) on the western shore of Lake Wakatipu, and reported that “Excrement of a large bird was also found [in the cave] … Some of this consisted of undigested fragments of what looked like the stalk of the fern”. White (1875) also mentioned that across the lake, in a cave at Queenstown, “… a quantity of double-shafted feathers of a brown colour … appeared to be chiefly in a layer of hard-trodden excrement … Perfect droppings were also found … and a few specimens of a similar outward appearance, contained undigested vegetable fragments, some of which seemed to be branches and stalks of fern broken into short pieces of three-quarters of an inch in length”. One of the coprolites from Mt. Nicholas is held in the collections of the Museum of New Zealand Te Papa Tongarewa, however no evidence was found to indicate whether more specimens were ever collected from this site. The Mt. Nicholas cave was revisited by Wood et al. (2012c) but the sediments in the cave were found to have been heavily disturbed by rabbit (Oryctolagus cuniculus) burrowing and no further coprolites were found there. However, a new deposit of coprolites was found in a rock shelter adjacent to the cave (Wood et al., 2012c). Hamilton (1894) visited a small cave near Waikaia (Fig. 1), in northern Southland, after the desiccated leg of an upland moa
(Megalapteryx didinus) was discovered there. Hamilton reported finding many feathers and several owl pellets in the cave. Although his description (Hamilton 1894) does not mention coprolites, a small number of coprolites from this cave are held in the collections of Otago (n ¼ 3) and Canterbury (n ¼ 5) Museums. One of these coprolites was genetically identified as having been deposited by M. didinus (Wood et al., 2008). An accumulation of coprolites was discovered during the excavation of a rock shelter at Takahe Valley (Fig. 1), Fiordland, in 1949 (Duff, 1952). Plant remains from five of these coprolites were identified by Horrocks et al. (2004), and although the coprolites were inferred to have been deposited by M. didinus, this identification has only recently been confirmed by molecular analysis for one of the Takahe Valley coprolites (Supplementary Table 2 in Huynen et al., 2010). Several more discoveries of coprolites were made in the late 20th Century, during hydroelectric-scheme mitigation excavations of rock shelters in the Central Otago and North Otago regions. In 1964, coprolites were recovered from a rock shelter near Shepherd's Creek (Fig. 1), in the upper Waitaki Valley, North Otago (Trotter, 1970). Pollen analysis of one of these provided the first micropaleontological study of a coprolite from New Zealand (Trotter, 1970). In ca 1980, a sample of putative moa nesting material, preserved within a compacted earth layer believed to be moa excreta, was excavated from the Rockfall II rock shelter (Fig. 1) in Cromwell Gorge, Central Otago (Ritchie, 1982; Wood, 2008b). A sample of this material was sent to Canterbury Museum at the time for examination but almost three decades later could not be relocated by Wood (2008a). In ca 1990, a significant number of moa
Table 1 Summary of Late Quaternary terrestrial vertebrate coprolite discoveries from in New Zealand. Abbreviations: ALEX, Alexandra Museum; CM, Canterbury Museum; LCR, Landcare Research (Lincoln); MNZ, Museum of New Zealand Te Papa Tongarewa; OM, Otago Museum. Kakapo coprolites are also known from Megamania Cave system, Charleston Caves, and the Tutoko Valley, Fiordland (Horrocks et al., 2008) but no details on these deposits were available. Year
Collection
Taxa
Age
References
Comments
Earnscleugh Cave, Central Otago
c.1870
OM Av10436
Coastal moa
Late Holocene
OM holds five coprolites from this site collected during the early (1870s) excavations
Cave at Gorge Road, Queenstown, Central Otago Cave at Mt Nicholas, Wakatipu, Central Otago Cave near upper Waikaia River, Old Man Range, northern Southland Takahe Valley rock shelter A, Fiordland
c.1874
Probably moa
Late Holocene?
Wood et al. (2008), Wood and Wilmshurst (2013) White (1875)
c.1874
MNZ S.24393
Probably moa
Late Holocene?
White (1875)
MNZ holds one coprolite from this site
c.1894
CM Av9279 OM Av10436
Upland moa
Late Holocene?
Hamilton (1894), Wood et al. (2008)
Eight coprolites (five in CM; three in OM)
1949
MNZ S.25870 MNZ (Ethnology collections) CM unregistered LCR unregistered
Upland moa
<2500 years
ca 85 coprolites (ca 35 in MNZ; ca 45 in CM; five in LCR)
Dunback Cave, eastern Otago
1954
OM Av7313
Late Holocene?
Shepherd's Creek rock shelter, Waitaki Valley, North Otago Rockfall II rock shelter, Cromwell Gorge, Central Otago Cave near Magnesite Quarry, Cobb Valley, northwest Nelson Takahe Valley, Fiordland
1964
CM 2008.1115.31 CM 2008.1115.32
Registered as kakapo but may be moa Upland moa
Duff (1952), Horrocks et al. (2004), supp. info of Huynen et al. (2010) Wood (2009)
Late Holocene?
Trotter (1970)
Probably moa
Late Holocene?
Ritchie (1982)
Probably kakapo
Late Holocene?
A. Cooper pers comm. (2009)
ca 60 complete coprolites in CM. Three of these have been genetically identified (unpubl. data) Layer of excreta and putative nesting material. Sample sent to CM could not be relocated Kakapo feathers and moa bones found in association with the coprolites
Kakapo
Late Holocene
NZ Radiocarbon dating laboratory submission form from C. Burrows
c.1980
1980s
LCR X10/3-4
c. 1982
Rock shelter near Cairnmuir Gully, Cromwell Gorge, Central Otago
c.1990
OM Av10638 MNZ S.44597-99
Probably moa
Late Holocene?
R. Thomson pers comm. (2006)
Honeycomb Hill Cave System, Karamea, West Coast
c.1993e2010
LCR X10/7
Kakapo
Late Holocene
Sawers' rock shelter, Roxburgh Gorge, Central Otago
1994e2009
OM Av7566, Av10808 ALEX 06.49.01 LCR X09/24
Late Holocene
Euphrates Cave, Garibaldi Ridge, northwest Nelson
1994-2010
LCR X10/8
Mostly moa, but includes a range of different morphotypes that likely represent other taxa including Pacific rat Kakapo, Upland moa
Early to late Holocene
Worthy (1993), Horrocks et al. (2008), Wood et al. (2012a) Worthy (1998), Wood (2008b), Wood and Walker (2008), Wood et al. (2008) Rowe et al. (1994), Wood et al. (2012b)
Clutha River Valley, Central Otago
2001?
Late Holocene
Willerslev et al. (2003)
Upland moa
“in cave at base of limestone cliff, c.400 m NNW of E end of Lake Orbell”, “covering floor of cave and ledges to a thickness of 50 cm in places” Hundreds of coprolites collected during mitigation excavation, but since lost. Specimens in museums were donated by R. Thomson from his personal collection Hundreds of coprolites located throughout cave system. 41 in LCR
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Site
>1000 coprolites, most in ALEX. Many originate from a layer of moa nesting material. Others occur below this but are less well preserved (see Fig. 5) Kakapo coprolites discovered in side passages in 1994. Moa coprolites discovered in main entrance in 2010, ca 80 in LCR
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(continued on next page)
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Table 1 (continued ) Site
Year
Collection
Taxa
Age
References
Comments
Kawarau Gorge rock shelter A, Central Otago
2004
OM Av10637 MNZ S.44604-09
Heavy-footed moa, Pacific rat
Late Holocene
56 þ coprolites (50þ in OM; six in MNZ)
Firewood Creek rock shelter A, Central Otago Firewood Creek rock shelter B, Central Otago Roxburgh Gorge rock shelter A, Central Otago Roxburgh Gorge rock shelter B, Central Otago
2004
Probably moa
Late Holocene?
Wood (2008b), Wood et al. (2008), Wood and Wilmshurst (2013) Wood (2008b)
Probably moa
Late Holocene
Wood (2008b)
Probably moa
Late Holocene?
Wood (2008b)
One coprolite
2005
OM Av10639-41 LCR X09/22-23
Heavy-footed moa
Late Holocene
ca 40 coprolites (ca 20 in OM; 20 in LCR)
Daley's Flat, Dart River Valley, western Otago
2005e2010
2005
South Island giant moa, Heavy-footed moa, Upland moa, Little bush moa Probably moa
Roxburgh Gorge rock shelter C, Central Otago Roxburgh Gorge site D, Central Otago
OM Av10427 ALEX 06.48.01 LCR X10/12 OM Av10635
Wood (2008b), Wood et al. (2008), Wood and Wilmshurst (2013) Wood et al. (2008, 2011; 2013b)
2005e2009
LCR X09/21
Cairnmuir Gully rock shelter A, Central Otago Possum's rock shelter, Mt Nicholas, western Otago Cave on Mt. Owen, northwest Nelson
2005 2009
2004 2005
>200 coprolites (100þ in OM; seven in ALEX; 100þ in LCR).
Late Holocene?
Wood (2008b)
Red-crowned parakeet, Pacific rat
c.1500-700 years
Wood (2008b)
OM Av10642
Probably moa
Late Holocene?
Wood (2008a; 2008b)
LCR X09/3-7
South Island giant moa, Little bush moa Probably kakapo
c.1500 years
Wood et al. (2012c)
ca 40 coprolites
Late Holocene?
C. Mosen pers comm. (2014)
>100 coprolites (see Fig. 2)
200 þ coprolites associated with parakeet nesting material from rock crevice.
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2014
OM Av10637 MNZ S.44600 OM Av10644
Four coprolites
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Fig. 2. Examples of recently discovered coprolite deposits from New Zealand: A) kakapo (Strigops habroptilus) coprolites in a cave on Mt. Owen (photo courtesy of Corey Mosen); B) moa coprolites (arrowed) shallowly buried amongst plant material in a rock shelter in Roxburgh Gorge, Central Otago; C) crevice in schist outcrop, Roxburgh Gorge, Central Otago, in which cf. red-crowned parakeet (Cyanoramphus novaezelandiae) coprolites (morphotype 9) were found (camera lens cap for scale).
coprolites were discovered in another rock shelter near Cairnmuir Gully in the Cromwell Gorge (Fig. 1) by R. Thomson et al. (unpublished), and although they were collected at the time, most have since been lost (Table 1). In addition to moa coprolites, kakapo (Strigops habroptilus) coprolites have also been recorded in a number of South Island cave systems. Although likely widespread, there are few references to kakapo coprolites in scientific literature. Coprolites attributed to kakapo were collected from Dunback Cave in eastern Otago (Fig 1) in 1954 (Wood, 2009). Worthy (1997) noted the presence of kakapo coprolites in the Honeycomb Hill cave system, caves on Garibaldi Ridge, and on the cave floor in several locations through the Hodge Creek cave system, including one accumulation almost 50 cm thick. Horrocks et al. (2008) identified plant remains in putative kakapo coprolites collected from 6 different cave systems in the north and west of the South Island (Honeycomb Hill, Hodge Creek, Megamania, Charleston, Tutoko and Takahe Valley) (Fig. 1). Since 1990, coprolites have been discovered in 15 further caves and rock shelters (Figs. 1 and 2), concentrated in the Central Otago
region (Table 1). Coprolites from several of these sites have been analysed using a combination of ancient DNA (aDNA), pollen, and plant macrofossil analyses to learn about the paleoecology of the fauna represented (Wood et al., 2008, 2012a, 2012b, 2012c, 2013a, 2013b; Wood and Wilmshurst, 2013). While all specimens discovered prior to 1990 have been attributed to moa or kakapo, those discovered since 1990 include many specimens that, based on size and shape, can be attributed to taxa other than moa and kakapo. These specimens are discussed further in the following section. 3. Taxonomic representation of New Zealand coprolite assemblages 3.1. Coprolites identified using aDNA analysis The species responsible for depositing coprolites can be identified through the amplification and sequencing of short (<200 bp) species-diagnostic fragments of mitochondrial DNA. Presently, 94
Fig. 3. Sizes of genetically identified coprolites from 4 moa species, compared with measured examples of putative kakapo coprolites. The identity of the arrowed kakapo coprolite was confirmed using aDNA analysis (Wood et al., 2012a).
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Fig. 4. Morphotypes of Late Quaternary terrestrial vertebrate coprolites from New Zealand: A) morphotype 1, the shown example is a moa coprolite; B) morphotype 2, cf. moa or South Island goose (Cnemiornis calcitrans); C) morphotype 3 showing external lobing and cross section, cf. moa; D) morphotype 4, cf. moa; E) morphotype 5, kakapo (Strigops habroptilus); F) morphotype 6, cf. laughing owl (Sceloglaux albifacies), New Zealand falcon (Falco novaeseelandiae) or tuatara (Sphenodon punctatus); G) morphotype 7, cf. juvenile moa, Finsch's duck (Chenonetta finschi), kea (Nestor notabilis), parakeet (Cyanoramphus spp.) or large passerine; H) morphotype 8, cf. tuatara or large skinks (Oligosoma otagense, O. grande); I) morphotype 9, cf. red-crowned parakeet (Cyanoramphus novaezelandiae); J) morphotype 10, cf. lizard, large insect or small bird; K) morphotype 11, cf. Pacific rat (Rattus exulans). Scale bars are 10 mm.
coprolites from five moa species have been identified using aDNA analysis, representing all moa genera except Emeus (E. curtus, n ¼ 1 (Earnscleugh Cave); little bush moa Anomalopteryx didiformis, n ¼ 5 (2 Mt. Nicholas, 3 Dart River); M. didinus, n ¼ 57 (21 Dart River, 33 Euphrates Cave, 1 Takahe Valley, 1 Clutha Valley, 1 Waikaia); South Island giant moa D. robustus, n ¼ 21 (2 Mt. Nicholas, 19 Dart River); heavy-footed moa Pachyornis elephantopus, n ¼ 10 (8 Dart River, 1 Kawarau Gorge, 1 Roxburgh Gorge) (Willerslev et al., 2003; Wood et al., 2008, 2012b, 2012c, 2013b; Huynen et al., 2010). Furthermore, aDNA analysis has confirmed the identity of putative kakapo (Strigops habroptilus) coprolites from the Honeycomb Hill cave system (Wood et al., 2012a). The moa coprolites that have been identified to species exhibit variable intraspecific size and morphology. Some are distinctly lobed while others are smooth, cylindrical, or even pellet-like. There appears to be no correlation between the size of coprolites and the adult body size of the depositing moa species (Fig. 3), and therefore size is not a useful feature for discriminating between coprolites of different moa species. There is also a clear overlap between the size of moa and kakapo coprolites (Fig. 3). Kakapo coprolites often exhibit the distinct coiled spaghetti-like structure seen in modern kakapo droppings (Butler, 2006) (Fig. 4) but this is usually only present in well-preserved specimens. Ancient DNA analysis remains the only reliable method for accurately identifying the identity of the depositing species of coprolites from New Zealand.
3.2. Unidentified coprolites While the focus of coprolite analyses in New Zealand has been on moa and kakapo, a diverse range of coprolite shapes and sizes have been recovered that obviously represent additional faunal species. These remain to be identified using aDNA analysis, however we provide descriptions here, and, based on comparisons with droppings of extant fauna, suggest the taxa that they most likely represent. 3.2.1. Morphotype 1 (Fig. 4A) Description: Discrete boli but with no distinctive characteristics. Can include a wide range of sizes and shapes. Probably represents poorly formed, worn or abraded coprolites (i.e. may include poorly preserved specimens that were originally morphotypes 2e11). Many examples of morphotype 1 have sediment embedded into their surface, and some (from Sawers' rock shelter, Central Otago and Dart River, West Otago) have skin impressions (see Rawlence et al., 2013), indicating they were relatively soft after deposition. Localities: Present in most coprolite assemblages. Attributed taxa: Could include coprolites from a range of different species. 3.2.2. Morphotype 2 (Fig. 4B) Description: Cyclindrical coprolites, circular in cross section, ca 20e25 mm diameter, relatively smooth surface or with a fine cauliflower-like texture. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxon: The size of the coprolites is
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indicative of a large bird. However, the absence of this morphotype in the well-characterised Dart River and Euphrates Cave assemblages suggests they may not be moa coprolites. Large waterfowl often produce relatively smooth cylindrical droppings, and therefore this morphotype may represent coprolites of the extinct South Island goose (Cnemiornis calcitrans). 3.2.3. Morphotype 3 (Fig. 4C) Description: Distinctly lobed coprolites, typically elongate or ovoid in shape, variable in size (from 25 to >90 mm long). The lobes are distinguished from coils (see morphotype 4) in that they are not continuous around the entire circumference of the coprolite but often end along a longitudinal midline. In cross section the lobes form around a central core (Fig. 3c). Localities: Most Central Otago rock shelters and Dart River, West Otago (Fig. 1). Attributed taxon: Moa. 3.2.4. Morphotype 4 (Fig. 4D) Description: Coiled coprolites, almost always spherical in shape, usually light brown to yellowish in colour. Overall size ca 23 mm in diameter, with individual coils ca 6 mm in diameter. This morphotype may represent the broken ends (or a smaller variation) of the lobed coprolites (morphotype 3). Localities: Sawers' rock shelter, Central Otago. Attributed taxon: Moa. 3.2.5. Morphotype 5 (Fig. 4E) Description: Thin tube (ca 6 mm diameter) randomly folded and coiled into an approximately spherical or ovoid shape. The resulting coprolites have a brain-like appearance, and are usually dark brown in colour. Where they occur in limestone caves they can often have a patchy white coating on the surface (Fig. 2). Localities: Cave systems on the west coast and north west region of the South Island. Attributed taxon: Kakapo. 3.2.6. Morphotype 6 (Fig. 4F) Description: Cylindrical coprolites, ca 5e20 mm in diameter, either as short straight sections or v-shaped (usually asymmetrical). Surface smooth and grey to light-brown in colour, often with white patches that may be dried urea. Often contain abundant beetle fragments visible on surface of coprolite. Microscopic examination reveals the coprolite content also includes a significant component of fine feather fragments. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: These coprolites are most likely the mutes of laughing owl (Sceloglaux albifacies), as feathers of this extinct predator were found in association with them. Laughing owl was known to feed on beetles (Worthy and Holdaway, 1996). The morphotype may also include New Zealand falcon (Falco novaeseelandiae) or tuatara (Sphenodon punctatus) coprolites, both of which were also present in the Central Otago region during the late Holocene (Worthy, 1998) and would leave similar size droppings containing beetle fragments. 3.2.7. Morphotype 7 (Fig. 4G) Description: Cylindrical coprolites, typically folded or partly coiled, sometimes straight. Blunt or slightly tapered ends. Surface smooth to slightly lumpy. The coprolites are broadly similar to morphotype 6, but are composed mostly of plant remains. Their size is variable, ranging from <3 e ca 11 mm diameter. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: Due to their variable size, these coprolites may represent a range of small terrestrial herbivore species. Larger specimens may be from juvenile moa, Finsch's duck (Chenonetta finschi) or kea (Nestor notabilis), while smaller specimens may represent parakeets (Cyanoramphus spp.) or large passerines (Passeriformes).
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3.2.8. Morphotype 8 (Fig. 4H) Description: Thin (4.5e6 mm diameter) elongate (25e30 mm long) coprolites with pointed ends. Can be straight, curved or wavy in shape (Fig. 4H) but not coiled. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: This morphotype bears similarities to reptile droppings (e.g. skink scat in Fig. 232 of Triggs, 2010). May represent tuatara or large skinks (Oligosoma otagense, O. grande). 3.2.9. Morphotype 9 (Fig. 4I) Description: Very thin and elongate coprolites forming 1e3 complete coils. Total coprolite size (ca 10 mm diameter) is 3e4 times the diameter of the coiled tube. Coils are usually neatly circular but occasionally messy and random (Fig. 4I). Colour can vary from light to very dark brown or black. Muehlenbeckia seed fragments are sometimes visible on surfaces of these coprolites. Localities: Roxburgh Gorge site D and Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxon: Probably red-crowned parakeet (Cyanoramphus novaezelandiae). Feathers attributed to this species were found in association with the coprolites at both localities, and the coprolites bear significant similarity to modern parakeet droppings (Wood, 2008b). 3.2.10. Morphotype 10 (Fig. 4J) Description: These coprolites have a similar coiled structure to morphotype 9, but are smaller (c. <6 mm diameter). Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: Possibly lizard, large insect or small bird. 3.2.11. Morphotype 11 (Fig. 4K) Description: Pellet-like coprolites, with blunt or slightly tapered ends, measuring ca 5 mm wide and 10 mm long. Shiny fragments of beetle or seed are occasionally visible on the surfaces. Localities: Roxburgh Gorge site D, Sawers' rock shelter and Kawarau Gorge rock shelter, Central Otago (Fig. 1). Attributed taxon: Probably represent early Pacific rat (Rattus exulans), based on their similarity to modern rat droppings. Unpublished radiocarbon dates for specimens of this morphotype post-date human settlement but pre-date the arrival of European rats. 4. Distribution and taphonomy of New Zealand coprolite deposits 4.1. Geological constraints In New Zealand, coprolites have been found exclusively in cave entrances and beneath rock overhangs. Therefore, it is clear that geology plays a key role in constraining the distribution of coprolite sites. The most important factors appear to be a combination of rock type and foliation/dip angle. First, the rock must either be a type that readily forms caves and overhangs (e.g. limestone or marble), or one that is cohesive or cemented enough that it can form overhangs (e.g. schist). The foliation or dip angles (with respect to the local topography) are also important, as overhangs will most frequently occur where rock layers dip into a slope. This is evident in the river gorges of Central Otago, where overhangs usually only occur on the down-dip side of creeks that cut parallel to the strike of the schist foliation. Also, most coprolites have been found in areas where the local rock types are relatively hard and resistant to weathering. Exploratory work by the authors suggests that coprolites do not appear to preserve beneath rock shelters formed from rock types that exhibit relatively rapid exfoliation (e.g. limestones in North Otago, and mudstones at Waikaremoana). The reason for this may be related to sediment moisture. We have noted that in most rock shelter sites, moisture increases with depth, and
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Fig. 5. Stratigraphic position of Late Quaternary terrestrial vertebrate coprolites in eight cave and rock shelter sites in the Central Otago region, South Island, New Zealand. Mt Nicholas Cave is based on description of White (1875), Possum's rockshelter (near Mt. Nicholas Cave) is from Wood et al. (2012c) and other sites are from Wood (2008b).
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41
Fig. 6. Rose diagram of aspect for 11 rock shelters from New Zealand that contained late Holocene coprolites. The outer ring shows the passage of the sun across the sky over a year.
the implications of this for coprolite preservation are discussed further below. 4.2. Preservation conditions All coprolites from New Zealand caves and rock shelters have been preserved through desiccation, and therefore aridity is a key factor determining whether or not coprolites will preserve within a particular site. The aridity of a site is usually (but not always) influenced by regional aridity. There are some examples of coprolites that have been found in dry rock shelters and caves within relatively high rainfall zones (e.g. Takahe Valley, and the northwest Nelson and west coast regions of the South Island), but most are concentrated in the semi-arid zone of Central Otago (Fig. 1). Preservation can also vary within sites. In most Central Otago rock shelters coprolites are preserved only at shallow depths, typically <300 mm (Fig. 5). With increasing depth the sediments generally become damper and the condition of coprolites deteriorates accordingly. Drafting in caves can also create suitable microaridity situations in situations where the regional climate is moist. An example of this is Euphrates Cave, where moa coprolites were found over a discrete area on a rocky slope within a zone of cold breeze. Above this slope was a ledge, which was out of the breeze and several degrees warmer, and although it was likely to have been a prime moa roosting site, no coprolites were found there (Wood et al., 2012b). The aspect of a cave entrance or rock shelter also appears to be an important factor in constraining where coprolites will be preserved. All eleven rock shelters with preserved coprolites for which aspect data was available had aspects within the northwestnortheast quadrant, being sheltered from the prevailing weather while receiving maximum exposure to the sun (Fig. 6). 4.3. Ages of New Zealand coprolites The oldest moa coprolites from New Zealand are three specimens from the subalpine Euphrates Cave with calibrated age ranges
(95.4% confidence) of 5290e4,968, 7266e7022 and 7315e7165 years BP. All other radiocarbon dated coprolites from New Zealand are of late Holocene (
Fig. 7. Temporal distribution of calibrated ages for radiocarbon dated bird coprolites from New Zealand (Horrocks et al., 2004, 2008; Wood et al., 2011, 2012a; 2012b, 2012c; 2013b; unpubl. data). Note that dates for kakapo (Strigops habroptilus) coprolites presented by Horrocks et al. (2008) (arrowed) are for amalgamated samples from different sites rather than individual coprolites and may therefore represent a broader temporal span than the calibrated age would indicate. a) Calibrated age probability distributions for individual coprolites; b) Histogram summarising age distribution of coprolites in 500 year bins.
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used more for roosting rather than nesting, as eggshell fragments were rarer in such sites. Using radiocarbon dating, mitochondrial DNA haplotypes of the depositing birds, and pollen assemblages, Wood et al. (2012) showed that Euphrates Cave deposit represented a continuous (albeit punctuated) accumulation, reflecting periods of deposition by individual birds separated by hundreds or thousands of years. This may reflect a lower population density of moa in subalpine areas, compared with the Dart River site which appears to contain coprolites from many individual birds in a relatively short time period (Wood et al., 2013b). 5. Conclusions Over 2000 Late Quaternary coprolites have been discovered in New Zealand over the past 140 years. We recognise eleven different morphotypes within these coprolites, which likely represent a range of bird, reptile, and early introduced mammal species. Analysis of these coprolites will continue to provide detailed insights into the prehistoric ecosystems of New Zealand. Although a number of coprolite deposits are now known from New Zealand (Table 1), it is clear that they are dependent on specific environmental conditions, and are a significantly rarer palaeoecological resource than bone deposits. Therefore, care needs to be taken to ensure that they are preserved for future analyses. Part of this includes the need for awareness of the likely sites in which coprolite deposits exist, and an understanding that they are often shallowly buried and could be mistaken for postEuropean mammal dung. Also, in some sites, they may be difficult to distinguish from the surrounding sediment and can be very fragile, requiring them to be excavated carefully using dental probes and fine brushes; such coprolites could easily be destroyed by trowel scraping. There are few published data on the taphonomy of Late Quaternary coprolites, and so our observations on these factors in relation to the distribution and occurrence of coprolites in New Zealand (e.g. geological constraints, site aspect, stratigraphic position of coprolite horizons) may be of particular interest to international researchers, and help inform where coprolites are most likely to be encountered on other landmasses. Acknowledgements We would like to thank the large number of people and research institutions who have assisted with the discovery of, and analysis of, coprolites from New Zealand's Late Quaternary fauna. In particular Daphne Lee, Geoff Rogers, Roydon Thomson, Jeff Sawers, the New Zealand Department of Conservation, Otago Museum, and the Australian Centre for Ancient DNA at the University of Adelaide. P. Scofield and A. Tennyson provided information of specimens in Canterbury Museum and Te Papa. Corey Mosen provided the photograph of kakapo coprolites used in Fig. 2. Various aspects of this work have been funded by Otago University, Landcare Research, National Geographic and Royal Society of New Zealand Marsden Fund. References Butler, D.J., 2006. The habitat, food and feeding ecology of kakapo in Fiordland: a synopsis from the unpublished MSc thesis of Richard Gray. Notornis 53, 55e79. Cockburn-Hood, T.H., 1873. Letter read to Wellington Philosophical Society 16 January, 1874. Trans. Proc. N. Z. Inst. 6, 387e388. Duff, R.S., 1952. Recent maori occupation of Notornis Valley, Te Anau. J. Polyn. Soc. 61, 90e119. Gill, B.J., 2000. Morphometrics of moa eggshell fragments (Aves: Dinornithiformes) from Late Holocene dune-sands of the Karikari Peninsula, New Zealand. J. Roy. Soc. N.Z. 30, 131e145. Gill, B.J., 2006. A catalogue of moa eggs (Aves: Dinornithiformes). Rec. Auckl. Mus. 43, 55e80.
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Haile, J., Holdaway, R., Oliver, K., Bunce, M., Gilbert, M.T.P., Nielsen, R., Munch, K., Ho, S.Y.W., Shapiro, B., Willerslev, E., 2007. Ancient DNA chronology within sediment deposits: are paleobiological reconstructions possible and is DNA leaching a factor? Mol. Biol. Evol. 24, 982e989. Hamilton, A., 1894. On the feathers of a small species of Moa (Megalapteryx) found in a cave at the head of the Waikaia River, with a notice of a moa-hunters camping place on the Old Man Range. Trans. Proc. N.Z. Inst. 27, 232e238. Hartree, W.H., 1999. A preliminary report on the nesting habits of moas in the East Coast of the North Island. Notornis 46, 457e460. Horrocks, M., Jones, M.D., Beever, R.E., Sutton, D.G., 2002. Analysis of plant microfossils in prehistoric coprolites from Harataonga Bay, Great Barrier island, New Zealand. J. Roy. Soc. N. Z. 32, 617e628. Horrocks, M., Irwin, G.J., McGlone, M.S., Nichol, S.L., Williams, L.J., 2003. Pollen, phytoliths and diatoms in prehistoric coprolites from Kohika, Bay of Plenty, New Zealand. J. Archaeol. Sci. 30, 13e20. Horrocks, M., D'Costa, D., Wallace, R., Gardner, R., Kondo, R., 2004. Plant remains in coprolites: diet of a subalpine moa (Dinornithiformes) from southern New Zealand. Emu 104, 149e156. Horrocks, M., Salter, J., Braggins, J., Nichol, S., Moorhouse, R., Elliot, G., 2008. Plant microfossil analysis of coprolites of the critically endangered kakapo (Strigops habroptilus) parrot from New Zealand. Rev. Palaeobot. Palynol. 149, 229e245. Huynen, L., Gill, B.J., Millar, C.D., Lambert, D.M., 2010. Ancient DNA reveals extreme egg morphology and nesting behaviour in New Zealand‘s extinct moa. Proc. Natl. Acad. Sci. U. S. A. 107, 16201e16206. McCulloch, B.A., Trotter, M.M., 1971. Prehistoric Rock Art of New Zealand. AH & AW Reed, Wellington. Oskam, C.L., Haile, J., McLay, E., Rigby, P., Allentoft, M.E., Olsen, M.E., Bengtsson, C., Miller, G.H., Schwenninger, J., Jacomb, C., Walter, R., Baynes, A., Dotch, J., ParkerPearson, M., Gilbert, M.T.P., Holdaway, R.N., Willerslev, E., Bunce, M., 2010. Fossil avian eggshell preserves ancient DNA. Proc. Roy. Soc. B 277, 1991e2000. Poinar, H.N., Hofreiter, M., Spaulding, W.G., Martin, P.S., Stankiewicz, B.A., Bland, H., Evershed, R.P., Possnert, G., Paabo, S., 1998. Molecular coproscopy: dung and diet of the extinct ground sloth Nothrotheriops shastensis. Science 218, 402e406. Rawlence, N.J., Wood, J.R., Scofield, R.P., Fraser, C., Tennyson, A.J.D., 2013. Soft-tissue specimens from pre-European extinct birds of New Zealand. J. Roy. Soc. N. Z. 43, 154e181. Ritchie, N.A., 1982. The prehistoric role of the Cromwell gorge, New Zealand. N. Z. J. Archaeol. 4, 21e43. Rowe, P., Millar, I., Worthy, T., 1994. Exploration on garibaldi Ridge e euphrates cave, Kahurangi National Park. N. Z. Speleol. Bull. 9, 271e290. Schmidt, G.D., Duszynski, D.W., 1992. Parasites of the extinct Shasta ground sloth, Nothrotheriops shastensis, in Rampart Cave, Arizona. J. Parasitol. 78, 811e816. Triggs, B., 2010. Tracks, Scats and Other Traces: a Fieldguide to Australian Mammals. Oxford University Press, Melbourne, 340 pp. Trotter, M.M., 1970. Archaeological investigations in the Aviemore area, South Island. Rec. Canterb. Mus. 8, 439e453. White, T., 1875. Notes on moa caves, etc., in the wakatipu District. Trans. Proc. N. Z. Inst. 8, 97e102. Willerslev, E., Hansen, A.J., Binladen, J., Brand, T.B., Gilbert, M.T.P., Shapiro, B., Bunce, M., Wiuf, C., Gilichinsky, D.A., Cooper, A., 2003. Diverse plant and animal genetic records from Holocene and Pleistocene sediments. Science 300, 791e795. Wood, J.R., 2006. Subfossil kakapo (Strigops habroptilus) remains from near Gibraltar Rock, Cromwell Gorge, Central Otago, New Zealand. Notornis 53, 191e193. Wood, J.R., 2008a. Moa (Aves: Dinornithiformes) nesting material from the semiarid interior of South Island, New Zealand. J. Roy. Soc. N. Z. 38, 115e129. Wood, J.R., 2008b. Pre-settlement Paleoecology of Central Otago's Semi-arid Lowlands, with Emphasis on the Pre-settlement Role of Avian Herbivory in South Island Dryland Ecosystems, New Zealand. Unpublished PhD Thesis. Univerisity of Otago, Dunedin. Wood, J.R., 2009. Two late quaternary avifaunal assemblages from the dunback region, eastern otago, south island, New Zealand. Notornis 56, 154e157. Wood, J.R., Walker, S., 2008. Macrofossil evidence for pre-settlement vegetation of Central Otago's basin floors and gorges. N. Z. J. Bot. 46, 239e255. Wood, J.R., Wilmshurst, J.M., 2013. Pollen analysis of coprolites reveals dietary details of heavy-footed moa (Pachyornis elephantopus) and coastal moa (Euryapteryx curtus) from Central Otago. N. Z. J. Ecol. 37, 151e155. Wood, J.R., Rawlence, N.J., Rogers, G.M., Worthy, T.H., Austin, J.J., Cooper, A., 2008. Coprolites reveal the diet and ecology of New Zealand's extinct megaherbivore moas (Aves: Dinornithiformes). Quat. Sci. Rev. 27, 2593e2602. Wood, J.R., Wilmshurst, J.M., Rawlence, N.J., 2011. Radiocarbon-dated faunal remains correlate very large rock avalanche deposit with prehistoric Alpine fault rupture. N. Z. J. Geol. Geophys. 54, 431e434. Wood, J.R., Wilmshurst, J.M., Holzapfel, A.S., Worthy, T.H., Cooper, A., 2012a. A lost link between a flightless parrot and a parasitic plant and the potential role of coprolites in conservation paleoecology. Cons. Biol. 26, 1091e1099. Wood, J.R., Wilmshurst, J.M., Wagstaff, S.J., Worthy, T.H., Rawlence, N.J., Cooper, A., 2012b. High-resolution coproecology: using coprolites to reconstruct the habits and habitats of New Zealand's extinct upland moa (Megalapteryx didinus). PLoS ONE 7, e40025. Wood, J.R., Wilmshurst, J.M., Worthy, T.H., Cooper, A., 2012c. First coprolite evidence for the diet of little bush moa (Anomalopteryx didiformis); an extinct forest ratite from New Zealand. N. Z. J. Ecol. 36, 164e170.
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Wood, J.R., Wilmshurst, J.M., Rawlence, N.J., Bonner, K.I., Worthy, T.H., Kinsella, J.M., Cooper, A., 2013a. A megafauna's microfauna: gastrointestinal parasites of New Zealand extinct moa (Aves: Dinornithiformes). PLoS ONE 8, e57315. Wood, J.R., Wilmshurst, J.M., Richardson, S.J., Rawlence, N.J., Wagstaff, S.J., Worthy, T.H., Cooper, A., 2013b. Resolving lost herbivore community structure using coprolites of four sympatric moa species (Aves: Dinornithiformes). Proc. Natl. Acad. Sci. U. S. A. 110, 16910e16915. Worthy, T.H., 1993. Fossils of Honeycomb Hill. Museum of New Zealand Te Papa Tongarewa. Wellington. 56 pp. Worthy, T.H., 1997. Fossil deposits in the Hodges creek cave system, on the northern foothills of Mt Arthur, Nelson, south island, New Zealand. Notornis 44, 111e124. Worthy, T.H., 1998. Quaternary fossil faunas of otago, south island, New Zealand. J. Royal Soc. N. Z. 28, 421e521.
Worthy, T.H., Holdaway, R.N., 1996. Quaternary fossil faunas, overlapping taphonomies, and palaeofaunal reconstruction in North Canterbury, South Island, New Zealand. J. Roy. Soc. N. Z. 26, 275e361. Worthy, T.H., Holdaway, R.N., 2002. The Lost World of the Moa. Canterbury University Press, Christchurch. 718 pp.. Worthy, T.H., Scofield, R.P., 2012. 21st Century advances in knowledge of the biology of moa (Aves: Dinornithiformes), diagnoses revised, and a new morphological analysis. N. Z. J. Zool. 39, 87e153. Yll, R., Carrion, J.S., Marra, A.C., Bonfiglio, L., 2006. Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy). Palaeogeog. Palaeoclim. Palaeoecol. 237, 32e39.
Contents lists available at ScienceDirect
Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev
Late Quaternary terrestrial vertebrate coprolites from New Zealand Jamie R. Wood a, *, Janet M. Wilmshurst a, b a b
Landcare Research, PO Box 69040, Lincoln, Canterbury 7640, New Zealand School of Environment, The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand
a r t i c l e i n f o
a b s t r a c t
Article history: Received 29 April 2014 Received in revised form 21 May 2014 Accepted 22 May 2014 Available online
Over the past decade, concerted efforts to find and study Late Quaternary terrestrial vertebrate coprolites in New Zealand have revealed new insights into the diets and ecologies of New Zealand's prehistoric birds. Here, we provide a broader review of the coprolites found in natural (non-archaeological) Late Quaternary deposits from New Zealand. We summarise the morphological diversity of the coprolites, and discuss the taphonomy of the sites in which they are found. Since the 1870s more than 2000 coprolites have been discovered from 30 localities, all restricted to the South Island. The distribution of coprolite localities appears to reflect the presence of geological and climatic factors that enhance the potential for coprolite preservation; coprolites require dry conditions for preservation, and have been found on the ground surface within drafting cave entrances and at shallow (<300 mm) depths beneath rock overhangs with a northerly aspect. We classify the coprolites into eleven morphotypes, each of which may represent a range of different bird and/or reptile species. A review of genetically identified specimens shows that coprolites of different bird species overlap in size and morphology, reinforcing the need for identifications to be based on ancient DNA analysis. © 2014 Elsevier Ltd. All rights reserved.
Keywords: Birds Caves Coprolites New Zealand Reptiles Rock shelters Stratigraphy Taphonomy
1. Introduction The fossil record of New Zealand's Late Quaternary terrestrial fauna is impressively complete (Worthy and Holdaway, 2002). Abundant fossil bones from caves, mires, dunes, and loess deposits have provided detailed insights into not only the identity of the faunal species that once inhabited New Zealand, but also their ecology and distribution from the last ice age until the arrival of humans in the 13th Century AD (Worthy and Holdaway, 2002). Analyses of Late Quaternary bones continue to reveal new information about the biology of New Zealand's extinct species, and technological advances (particularly in the field of ancient DNA analysis) continue to open up exciting new research possibilities (Worthy and Scofield, 2012). Evidence for New Zealand's Late Quaternary fauna is not restricted to bones. Further proxies include Maori rock art depictions (McCulloch and Trotter, 1971), preserved feathers and integument (reviewed by Rawlence et al., 2013), nests (Hartree, 1999; Wood, 2006, 2008a), eggshell (Gill, 2000, 2006; Oskam et al., 2010), footprints and trackways (reviewed by Worthy and Holdaway, 2002), sedimentary ancient DNA (Willerslev et al., 2003; Haile et al., 2007) and coprolites (Horrocks et al., 2004, *Corresponding author. Tel.: þ64 3 321 9653. E-mail address: [email protected] (J.R. Wood). http://dx.doi.org/10.1016/j.quascirev.2014.05.020 0277-3791/© 2014 Elsevier Ltd. All rights reserved.
2008; Wood et al., 2008, 2012a, 2012b, 2012c, 2013a, 2013b; Wood and Wilmshurst, 2013). Coprolites are rich sources of paleoecological information, and can reveal aspects of the paleoenvironments (Yll et al., 2006), paleodiets (Poinar et al., 1998), ecosystem function (Wood et al., 2012a) and parasites (Schmidt and Duszynski, 1992) of prehistoric extinct fauna. Late Quaternary coprolites of extinct terrestrial fauna were first discovered in New Zealand approximately 140 years ago (Cockburn-Hood, 1873), and until recently remained relatively scarce. Within the past decade, concerted efforts to identify and recover Late Quaternary coprolites from New Zealand for study have greatly expanded our understanding of this unique paleoecological resource, making it timely for a broader review. Here, we review the history of Late Quaternary coprolite (hereafter simply referred to as coprolites) discoveries in New Zealand, assess the morphological diversity of the coprolites, and discuss the taphonomy of the sites in which they occur. Many coprolites, mainly of the Polynesian dog (Canis familiaris), have also been found in association with archaeological sites throughout New Zealand (e.g. Horrocks et al., 2002, 2003) but are not reviewed here, as we focus specifically on non-archaeological specimens. 2. A brief history of coprolite discoveries in New Zealand The first coprolites discovered in New Zealand were found during excavations at Earnscleugh Cave (Fig. 1) in Central Otago
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Fig. 1. Location of sites containing natural Late Quaternary terrestrial vertebrate coprolites in New Zealand. Names of regions are shown in grey text.
during the early 1870s. Cockburn-Hood (1873) noted that “The flat ground near [the cave] had probably been a favourite camping ground, from the quantity of droppingsewhich are, no doubt, those of the large birdseswept in by the wind”. Five coprolites from Earnscleugh Cave are held by Otago Museum, and one of these has been genetically identified as having been deposited by a coastal moa (Euryapteryx curtus) (Wood et al., 2008). Around the same period, White (1875) discovered a cave near Mt. Nicholas (Fig. 1) on the western shore of Lake Wakatipu, and reported that “Excrement of a large bird was also found [in the cave] … Some of this consisted of undigested fragments of what looked like the stalk of the fern”. White (1875) also mentioned that across the lake, in a cave at Queenstown, “… a quantity of double-shafted feathers of a brown colour … appeared to be chiefly in a layer of hard-trodden excrement … Perfect droppings were also found … and a few specimens of a similar outward appearance, contained undigested vegetable fragments, some of which seemed to be branches and stalks of fern broken into short pieces of three-quarters of an inch in length”. One of the coprolites from Mt. Nicholas is held in the collections of the Museum of New Zealand Te Papa Tongarewa, however no evidence was found to indicate whether more specimens were ever collected from this site. The Mt. Nicholas cave was revisited by Wood et al. (2012c) but the sediments in the cave were found to have been heavily disturbed by rabbit (Oryctolagus cuniculus) burrowing and no further coprolites were found there. However, a new deposit of coprolites was found in a rock shelter adjacent to the cave (Wood et al., 2012c). Hamilton (1894) visited a small cave near Waikaia (Fig. 1), in northern Southland, after the desiccated leg of an upland moa
(Megalapteryx didinus) was discovered there. Hamilton reported finding many feathers and several owl pellets in the cave. Although his description (Hamilton 1894) does not mention coprolites, a small number of coprolites from this cave are held in the collections of Otago (n ¼ 3) and Canterbury (n ¼ 5) Museums. One of these coprolites was genetically identified as having been deposited by M. didinus (Wood et al., 2008). An accumulation of coprolites was discovered during the excavation of a rock shelter at Takahe Valley (Fig. 1), Fiordland, in 1949 (Duff, 1952). Plant remains from five of these coprolites were identified by Horrocks et al. (2004), and although the coprolites were inferred to have been deposited by M. didinus, this identification has only recently been confirmed by molecular analysis for one of the Takahe Valley coprolites (Supplementary Table 2 in Huynen et al., 2010). Several more discoveries of coprolites were made in the late 20th Century, during hydroelectric-scheme mitigation excavations of rock shelters in the Central Otago and North Otago regions. In 1964, coprolites were recovered from a rock shelter near Shepherd's Creek (Fig. 1), in the upper Waitaki Valley, North Otago (Trotter, 1970). Pollen analysis of one of these provided the first micropaleontological study of a coprolite from New Zealand (Trotter, 1970). In ca 1980, a sample of putative moa nesting material, preserved within a compacted earth layer believed to be moa excreta, was excavated from the Rockfall II rock shelter (Fig. 1) in Cromwell Gorge, Central Otago (Ritchie, 1982; Wood, 2008b). A sample of this material was sent to Canterbury Museum at the time for examination but almost three decades later could not be relocated by Wood (2008a). In ca 1990, a significant number of moa
Table 1 Summary of Late Quaternary terrestrial vertebrate coprolite discoveries from in New Zealand. Abbreviations: ALEX, Alexandra Museum; CM, Canterbury Museum; LCR, Landcare Research (Lincoln); MNZ, Museum of New Zealand Te Papa Tongarewa; OM, Otago Museum. Kakapo coprolites are also known from Megamania Cave system, Charleston Caves, and the Tutoko Valley, Fiordland (Horrocks et al., 2008) but no details on these deposits were available. Year
Collection
Taxa
Age
References
Comments
Earnscleugh Cave, Central Otago
c.1870
OM Av10436
Coastal moa
Late Holocene
OM holds five coprolites from this site collected during the early (1870s) excavations
Cave at Gorge Road, Queenstown, Central Otago Cave at Mt Nicholas, Wakatipu, Central Otago Cave near upper Waikaia River, Old Man Range, northern Southland Takahe Valley rock shelter A, Fiordland
c.1874
Probably moa
Late Holocene?
Wood et al. (2008), Wood and Wilmshurst (2013) White (1875)
c.1874
MNZ S.24393
Probably moa
Late Holocene?
White (1875)
MNZ holds one coprolite from this site
c.1894
CM Av9279 OM Av10436
Upland moa
Late Holocene?
Hamilton (1894), Wood et al. (2008)
Eight coprolites (five in CM; three in OM)
1949
MNZ S.25870 MNZ (Ethnology collections) CM unregistered LCR unregistered
Upland moa
<2500 years
ca 85 coprolites (ca 35 in MNZ; ca 45 in CM; five in LCR)
Dunback Cave, eastern Otago
1954
OM Av7313
Late Holocene?
Shepherd's Creek rock shelter, Waitaki Valley, North Otago Rockfall II rock shelter, Cromwell Gorge, Central Otago Cave near Magnesite Quarry, Cobb Valley, northwest Nelson Takahe Valley, Fiordland
1964
CM 2008.1115.31 CM 2008.1115.32
Registered as kakapo but may be moa Upland moa
Duff (1952), Horrocks et al. (2004), supp. info of Huynen et al. (2010) Wood (2009)
Late Holocene?
Trotter (1970)
Probably moa
Late Holocene?
Ritchie (1982)
Probably kakapo
Late Holocene?
A. Cooper pers comm. (2009)
ca 60 complete coprolites in CM. Three of these have been genetically identified (unpubl. data) Layer of excreta and putative nesting material. Sample sent to CM could not be relocated Kakapo feathers and moa bones found in association with the coprolites
Kakapo
Late Holocene
NZ Radiocarbon dating laboratory submission form from C. Burrows
c.1980
1980s
LCR X10/3-4
c. 1982
Rock shelter near Cairnmuir Gully, Cromwell Gorge, Central Otago
c.1990
OM Av10638 MNZ S.44597-99
Probably moa
Late Holocene?
R. Thomson pers comm. (2006)
Honeycomb Hill Cave System, Karamea, West Coast
c.1993e2010
LCR X10/7
Kakapo
Late Holocene
Sawers' rock shelter, Roxburgh Gorge, Central Otago
1994e2009
OM Av7566, Av10808 ALEX 06.49.01 LCR X09/24
Late Holocene
Euphrates Cave, Garibaldi Ridge, northwest Nelson
1994-2010
LCR X10/8
Mostly moa, but includes a range of different morphotypes that likely represent other taxa including Pacific rat Kakapo, Upland moa
Early to late Holocene
Worthy (1993), Horrocks et al. (2008), Wood et al. (2012a) Worthy (1998), Wood (2008b), Wood and Walker (2008), Wood et al. (2008) Rowe et al. (1994), Wood et al. (2012b)
Clutha River Valley, Central Otago
2001?
Late Holocene
Willerslev et al. (2003)
Upland moa
“in cave at base of limestone cliff, c.400 m NNW of E end of Lake Orbell”, “covering floor of cave and ledges to a thickness of 50 cm in places” Hundreds of coprolites collected during mitigation excavation, but since lost. Specimens in museums were donated by R. Thomson from his personal collection Hundreds of coprolites located throughout cave system. 41 in LCR
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Site
>1000 coprolites, most in ALEX. Many originate from a layer of moa nesting material. Others occur below this but are less well preserved (see Fig. 5) Kakapo coprolites discovered in side passages in 1994. Moa coprolites discovered in main entrance in 2010, ca 80 in LCR
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(continued on next page)
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Table 1 (continued ) Site
Year
Collection
Taxa
Age
References
Comments
Kawarau Gorge rock shelter A, Central Otago
2004
OM Av10637 MNZ S.44604-09
Heavy-footed moa, Pacific rat
Late Holocene
56 þ coprolites (50þ in OM; six in MNZ)
Firewood Creek rock shelter A, Central Otago Firewood Creek rock shelter B, Central Otago Roxburgh Gorge rock shelter A, Central Otago Roxburgh Gorge rock shelter B, Central Otago
2004
Probably moa
Late Holocene?
Wood (2008b), Wood et al. (2008), Wood and Wilmshurst (2013) Wood (2008b)
Probably moa
Late Holocene
Wood (2008b)
Probably moa
Late Holocene?
Wood (2008b)
One coprolite
2005
OM Av10639-41 LCR X09/22-23
Heavy-footed moa
Late Holocene
ca 40 coprolites (ca 20 in OM; 20 in LCR)
Daley's Flat, Dart River Valley, western Otago
2005e2010
2005
South Island giant moa, Heavy-footed moa, Upland moa, Little bush moa Probably moa
Roxburgh Gorge rock shelter C, Central Otago Roxburgh Gorge site D, Central Otago
OM Av10427 ALEX 06.48.01 LCR X10/12 OM Av10635
Wood (2008b), Wood et al. (2008), Wood and Wilmshurst (2013) Wood et al. (2008, 2011; 2013b)
2005e2009
LCR X09/21
Cairnmuir Gully rock shelter A, Central Otago Possum's rock shelter, Mt Nicholas, western Otago Cave on Mt. Owen, northwest Nelson
2005 2009
2004 2005
>200 coprolites (100þ in OM; seven in ALEX; 100þ in LCR).
Late Holocene?
Wood (2008b)
Red-crowned parakeet, Pacific rat
c.1500-700 years
Wood (2008b)
OM Av10642
Probably moa
Late Holocene?
Wood (2008a; 2008b)
LCR X09/3-7
South Island giant moa, Little bush moa Probably kakapo
c.1500 years
Wood et al. (2012c)
ca 40 coprolites
Late Holocene?
C. Mosen pers comm. (2014)
>100 coprolites (see Fig. 2)
200 þ coprolites associated with parakeet nesting material from rock crevice.
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2014
OM Av10637 MNZ S.44600 OM Av10644
Four coprolites
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Fig. 2. Examples of recently discovered coprolite deposits from New Zealand: A) kakapo (Strigops habroptilus) coprolites in a cave on Mt. Owen (photo courtesy of Corey Mosen); B) moa coprolites (arrowed) shallowly buried amongst plant material in a rock shelter in Roxburgh Gorge, Central Otago; C) crevice in schist outcrop, Roxburgh Gorge, Central Otago, in which cf. red-crowned parakeet (Cyanoramphus novaezelandiae) coprolites (morphotype 9) were found (camera lens cap for scale).
coprolites were discovered in another rock shelter near Cairnmuir Gully in the Cromwell Gorge (Fig. 1) by R. Thomson et al. (unpublished), and although they were collected at the time, most have since been lost (Table 1). In addition to moa coprolites, kakapo (Strigops habroptilus) coprolites have also been recorded in a number of South Island cave systems. Although likely widespread, there are few references to kakapo coprolites in scientific literature. Coprolites attributed to kakapo were collected from Dunback Cave in eastern Otago (Fig 1) in 1954 (Wood, 2009). Worthy (1997) noted the presence of kakapo coprolites in the Honeycomb Hill cave system, caves on Garibaldi Ridge, and on the cave floor in several locations through the Hodge Creek cave system, including one accumulation almost 50 cm thick. Horrocks et al. (2008) identified plant remains in putative kakapo coprolites collected from 6 different cave systems in the north and west of the South Island (Honeycomb Hill, Hodge Creek, Megamania, Charleston, Tutoko and Takahe Valley) (Fig. 1). Since 1990, coprolites have been discovered in 15 further caves and rock shelters (Figs. 1 and 2), concentrated in the Central Otago
region (Table 1). Coprolites from several of these sites have been analysed using a combination of ancient DNA (aDNA), pollen, and plant macrofossil analyses to learn about the paleoecology of the fauna represented (Wood et al., 2008, 2012a, 2012b, 2012c, 2013a, 2013b; Wood and Wilmshurst, 2013). While all specimens discovered prior to 1990 have been attributed to moa or kakapo, those discovered since 1990 include many specimens that, based on size and shape, can be attributed to taxa other than moa and kakapo. These specimens are discussed further in the following section. 3. Taxonomic representation of New Zealand coprolite assemblages 3.1. Coprolites identified using aDNA analysis The species responsible for depositing coprolites can be identified through the amplification and sequencing of short (<200 bp) species-diagnostic fragments of mitochondrial DNA. Presently, 94
Fig. 3. Sizes of genetically identified coprolites from 4 moa species, compared with measured examples of putative kakapo coprolites. The identity of the arrowed kakapo coprolite was confirmed using aDNA analysis (Wood et al., 2012a).
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Fig. 4. Morphotypes of Late Quaternary terrestrial vertebrate coprolites from New Zealand: A) morphotype 1, the shown example is a moa coprolite; B) morphotype 2, cf. moa or South Island goose (Cnemiornis calcitrans); C) morphotype 3 showing external lobing and cross section, cf. moa; D) morphotype 4, cf. moa; E) morphotype 5, kakapo (Strigops habroptilus); F) morphotype 6, cf. laughing owl (Sceloglaux albifacies), New Zealand falcon (Falco novaeseelandiae) or tuatara (Sphenodon punctatus); G) morphotype 7, cf. juvenile moa, Finsch's duck (Chenonetta finschi), kea (Nestor notabilis), parakeet (Cyanoramphus spp.) or large passerine; H) morphotype 8, cf. tuatara or large skinks (Oligosoma otagense, O. grande); I) morphotype 9, cf. red-crowned parakeet (Cyanoramphus novaezelandiae); J) morphotype 10, cf. lizard, large insect or small bird; K) morphotype 11, cf. Pacific rat (Rattus exulans). Scale bars are 10 mm.
coprolites from five moa species have been identified using aDNA analysis, representing all moa genera except Emeus (E. curtus, n ¼ 1 (Earnscleugh Cave); little bush moa Anomalopteryx didiformis, n ¼ 5 (2 Mt. Nicholas, 3 Dart River); M. didinus, n ¼ 57 (21 Dart River, 33 Euphrates Cave, 1 Takahe Valley, 1 Clutha Valley, 1 Waikaia); South Island giant moa D. robustus, n ¼ 21 (2 Mt. Nicholas, 19 Dart River); heavy-footed moa Pachyornis elephantopus, n ¼ 10 (8 Dart River, 1 Kawarau Gorge, 1 Roxburgh Gorge) (Willerslev et al., 2003; Wood et al., 2008, 2012b, 2012c, 2013b; Huynen et al., 2010). Furthermore, aDNA analysis has confirmed the identity of putative kakapo (Strigops habroptilus) coprolites from the Honeycomb Hill cave system (Wood et al., 2012a). The moa coprolites that have been identified to species exhibit variable intraspecific size and morphology. Some are distinctly lobed while others are smooth, cylindrical, or even pellet-like. There appears to be no correlation between the size of coprolites and the adult body size of the depositing moa species (Fig. 3), and therefore size is not a useful feature for discriminating between coprolites of different moa species. There is also a clear overlap between the size of moa and kakapo coprolites (Fig. 3). Kakapo coprolites often exhibit the distinct coiled spaghetti-like structure seen in modern kakapo droppings (Butler, 2006) (Fig. 4) but this is usually only present in well-preserved specimens. Ancient DNA analysis remains the only reliable method for accurately identifying the identity of the depositing species of coprolites from New Zealand.
3.2. Unidentified coprolites While the focus of coprolite analyses in New Zealand has been on moa and kakapo, a diverse range of coprolite shapes and sizes have been recovered that obviously represent additional faunal species. These remain to be identified using aDNA analysis, however we provide descriptions here, and, based on comparisons with droppings of extant fauna, suggest the taxa that they most likely represent. 3.2.1. Morphotype 1 (Fig. 4A) Description: Discrete boli but with no distinctive characteristics. Can include a wide range of sizes and shapes. Probably represents poorly formed, worn or abraded coprolites (i.e. may include poorly preserved specimens that were originally morphotypes 2e11). Many examples of morphotype 1 have sediment embedded into their surface, and some (from Sawers' rock shelter, Central Otago and Dart River, West Otago) have skin impressions (see Rawlence et al., 2013), indicating they were relatively soft after deposition. Localities: Present in most coprolite assemblages. Attributed taxa: Could include coprolites from a range of different species. 3.2.2. Morphotype 2 (Fig. 4B) Description: Cyclindrical coprolites, circular in cross section, ca 20e25 mm diameter, relatively smooth surface or with a fine cauliflower-like texture. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxon: The size of the coprolites is
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indicative of a large bird. However, the absence of this morphotype in the well-characterised Dart River and Euphrates Cave assemblages suggests they may not be moa coprolites. Large waterfowl often produce relatively smooth cylindrical droppings, and therefore this morphotype may represent coprolites of the extinct South Island goose (Cnemiornis calcitrans). 3.2.3. Morphotype 3 (Fig. 4C) Description: Distinctly lobed coprolites, typically elongate or ovoid in shape, variable in size (from 25 to >90 mm long). The lobes are distinguished from coils (see morphotype 4) in that they are not continuous around the entire circumference of the coprolite but often end along a longitudinal midline. In cross section the lobes form around a central core (Fig. 3c). Localities: Most Central Otago rock shelters and Dart River, West Otago (Fig. 1). Attributed taxon: Moa. 3.2.4. Morphotype 4 (Fig. 4D) Description: Coiled coprolites, almost always spherical in shape, usually light brown to yellowish in colour. Overall size ca 23 mm in diameter, with individual coils ca 6 mm in diameter. This morphotype may represent the broken ends (or a smaller variation) of the lobed coprolites (morphotype 3). Localities: Sawers' rock shelter, Central Otago. Attributed taxon: Moa. 3.2.5. Morphotype 5 (Fig. 4E) Description: Thin tube (ca 6 mm diameter) randomly folded and coiled into an approximately spherical or ovoid shape. The resulting coprolites have a brain-like appearance, and are usually dark brown in colour. Where they occur in limestone caves they can often have a patchy white coating on the surface (Fig. 2). Localities: Cave systems on the west coast and north west region of the South Island. Attributed taxon: Kakapo. 3.2.6. Morphotype 6 (Fig. 4F) Description: Cylindrical coprolites, ca 5e20 mm in diameter, either as short straight sections or v-shaped (usually asymmetrical). Surface smooth and grey to light-brown in colour, often with white patches that may be dried urea. Often contain abundant beetle fragments visible on surface of coprolite. Microscopic examination reveals the coprolite content also includes a significant component of fine feather fragments. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: These coprolites are most likely the mutes of laughing owl (Sceloglaux albifacies), as feathers of this extinct predator were found in association with them. Laughing owl was known to feed on beetles (Worthy and Holdaway, 1996). The morphotype may also include New Zealand falcon (Falco novaeseelandiae) or tuatara (Sphenodon punctatus) coprolites, both of which were also present in the Central Otago region during the late Holocene (Worthy, 1998) and would leave similar size droppings containing beetle fragments. 3.2.7. Morphotype 7 (Fig. 4G) Description: Cylindrical coprolites, typically folded or partly coiled, sometimes straight. Blunt or slightly tapered ends. Surface smooth to slightly lumpy. The coprolites are broadly similar to morphotype 6, but are composed mostly of plant remains. Their size is variable, ranging from <3 e ca 11 mm diameter. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: Due to their variable size, these coprolites may represent a range of small terrestrial herbivore species. Larger specimens may be from juvenile moa, Finsch's duck (Chenonetta finschi) or kea (Nestor notabilis), while smaller specimens may represent parakeets (Cyanoramphus spp.) or large passerines (Passeriformes).
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3.2.8. Morphotype 8 (Fig. 4H) Description: Thin (4.5e6 mm diameter) elongate (25e30 mm long) coprolites with pointed ends. Can be straight, curved or wavy in shape (Fig. 4H) but not coiled. Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: This morphotype bears similarities to reptile droppings (e.g. skink scat in Fig. 232 of Triggs, 2010). May represent tuatara or large skinks (Oligosoma otagense, O. grande). 3.2.9. Morphotype 9 (Fig. 4I) Description: Very thin and elongate coprolites forming 1e3 complete coils. Total coprolite size (ca 10 mm diameter) is 3e4 times the diameter of the coiled tube. Coils are usually neatly circular but occasionally messy and random (Fig. 4I). Colour can vary from light to very dark brown or black. Muehlenbeckia seed fragments are sometimes visible on surfaces of these coprolites. Localities: Roxburgh Gorge site D and Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxon: Probably red-crowned parakeet (Cyanoramphus novaezelandiae). Feathers attributed to this species were found in association with the coprolites at both localities, and the coprolites bear significant similarity to modern parakeet droppings (Wood, 2008b). 3.2.10. Morphotype 10 (Fig. 4J) Description: These coprolites have a similar coiled structure to morphotype 9, but are smaller (c. <6 mm diameter). Localities: Sawers' rock shelter, Central Otago (Fig. 1). Attributed taxa: Possibly lizard, large insect or small bird. 3.2.11. Morphotype 11 (Fig. 4K) Description: Pellet-like coprolites, with blunt or slightly tapered ends, measuring ca 5 mm wide and 10 mm long. Shiny fragments of beetle or seed are occasionally visible on the surfaces. Localities: Roxburgh Gorge site D, Sawers' rock shelter and Kawarau Gorge rock shelter, Central Otago (Fig. 1). Attributed taxon: Probably represent early Pacific rat (Rattus exulans), based on their similarity to modern rat droppings. Unpublished radiocarbon dates for specimens of this morphotype post-date human settlement but pre-date the arrival of European rats. 4. Distribution and taphonomy of New Zealand coprolite deposits 4.1. Geological constraints In New Zealand, coprolites have been found exclusively in cave entrances and beneath rock overhangs. Therefore, it is clear that geology plays a key role in constraining the distribution of coprolite sites. The most important factors appear to be a combination of rock type and foliation/dip angle. First, the rock must either be a type that readily forms caves and overhangs (e.g. limestone or marble), or one that is cohesive or cemented enough that it can form overhangs (e.g. schist). The foliation or dip angles (with respect to the local topography) are also important, as overhangs will most frequently occur where rock layers dip into a slope. This is evident in the river gorges of Central Otago, where overhangs usually only occur on the down-dip side of creeks that cut parallel to the strike of the schist foliation. Also, most coprolites have been found in areas where the local rock types are relatively hard and resistant to weathering. Exploratory work by the authors suggests that coprolites do not appear to preserve beneath rock shelters formed from rock types that exhibit relatively rapid exfoliation (e.g. limestones in North Otago, and mudstones at Waikaremoana). The reason for this may be related to sediment moisture. We have noted that in most rock shelter sites, moisture increases with depth, and
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Fig. 5. Stratigraphic position of Late Quaternary terrestrial vertebrate coprolites in eight cave and rock shelter sites in the Central Otago region, South Island, New Zealand. Mt Nicholas Cave is based on description of White (1875), Possum's rockshelter (near Mt. Nicholas Cave) is from Wood et al. (2012c) and other sites are from Wood (2008b).
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Fig. 6. Rose diagram of aspect for 11 rock shelters from New Zealand that contained late Holocene coprolites. The outer ring shows the passage of the sun across the sky over a year.
the implications of this for coprolite preservation are discussed further below. 4.2. Preservation conditions All coprolites from New Zealand caves and rock shelters have been preserved through desiccation, and therefore aridity is a key factor determining whether or not coprolites will preserve within a particular site. The aridity of a site is usually (but not always) influenced by regional aridity. There are some examples of coprolites that have been found in dry rock shelters and caves within relatively high rainfall zones (e.g. Takahe Valley, and the northwest Nelson and west coast regions of the South Island), but most are concentrated in the semi-arid zone of Central Otago (Fig. 1). Preservation can also vary within sites. In most Central Otago rock shelters coprolites are preserved only at shallow depths, typically <300 mm (Fig. 5). With increasing depth the sediments generally become damper and the condition of coprolites deteriorates accordingly. Drafting in caves can also create suitable microaridity situations in situations where the regional climate is moist. An example of this is Euphrates Cave, where moa coprolites were found over a discrete area on a rocky slope within a zone of cold breeze. Above this slope was a ledge, which was out of the breeze and several degrees warmer, and although it was likely to have been a prime moa roosting site, no coprolites were found there (Wood et al., 2012b). The aspect of a cave entrance or rock shelter also appears to be an important factor in constraining where coprolites will be preserved. All eleven rock shelters with preserved coprolites for which aspect data was available had aspects within the northwestnortheast quadrant, being sheltered from the prevailing weather while receiving maximum exposure to the sun (Fig. 6). 4.3. Ages of New Zealand coprolites The oldest moa coprolites from New Zealand are three specimens from the subalpine Euphrates Cave with calibrated age ranges
(95.4% confidence) of 5290e4,968, 7266e7022 and 7315e7165 years BP. All other radiocarbon dated coprolites from New Zealand are of late Holocene (
Fig. 7. Temporal distribution of calibrated ages for radiocarbon dated bird coprolites from New Zealand (Horrocks et al., 2004, 2008; Wood et al., 2011, 2012a; 2012b, 2012c; 2013b; unpubl. data). Note that dates for kakapo (Strigops habroptilus) coprolites presented by Horrocks et al. (2008) (arrowed) are for amalgamated samples from different sites rather than individual coprolites and may therefore represent a broader temporal span than the calibrated age would indicate. a) Calibrated age probability distributions for individual coprolites; b) Histogram summarising age distribution of coprolites in 500 year bins.
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used more for roosting rather than nesting, as eggshell fragments were rarer in such sites. Using radiocarbon dating, mitochondrial DNA haplotypes of the depositing birds, and pollen assemblages, Wood et al. (2012) showed that Euphrates Cave deposit represented a continuous (albeit punctuated) accumulation, reflecting periods of deposition by individual birds separated by hundreds or thousands of years. This may reflect a lower population density of moa in subalpine areas, compared with the Dart River site which appears to contain coprolites from many individual birds in a relatively short time period (Wood et al., 2013b). 5. Conclusions Over 2000 Late Quaternary coprolites have been discovered in New Zealand over the past 140 years. We recognise eleven different morphotypes within these coprolites, which likely represent a range of bird, reptile, and early introduced mammal species. Analysis of these coprolites will continue to provide detailed insights into the prehistoric ecosystems of New Zealand. Although a number of coprolite deposits are now known from New Zealand (Table 1), it is clear that they are dependent on specific environmental conditions, and are a significantly rarer palaeoecological resource than bone deposits. Therefore, care needs to be taken to ensure that they are preserved for future analyses. Part of this includes the need for awareness of the likely sites in which coprolite deposits exist, and an understanding that they are often shallowly buried and could be mistaken for postEuropean mammal dung. Also, in some sites, they may be difficult to distinguish from the surrounding sediment and can be very fragile, requiring them to be excavated carefully using dental probes and fine brushes; such coprolites could easily be destroyed by trowel scraping. There are few published data on the taphonomy of Late Quaternary coprolites, and so our observations on these factors in relation to the distribution and occurrence of coprolites in New Zealand (e.g. geological constraints, site aspect, stratigraphic position of coprolite horizons) may be of particular interest to international researchers, and help inform where coprolites are most likely to be encountered on other landmasses. Acknowledgements We would like to thank the large number of people and research institutions who have assisted with the discovery of, and analysis of, coprolites from New Zealand's Late Quaternary fauna. In particular Daphne Lee, Geoff Rogers, Roydon Thomson, Jeff Sawers, the New Zealand Department of Conservation, Otago Museum, and the Australian Centre for Ancient DNA at the University of Adelaide. P. Scofield and A. Tennyson provided information of specimens in Canterbury Museum and Te Papa. Corey Mosen provided the photograph of kakapo coprolites used in Fig. 2. Various aspects of this work have been funded by Otago University, Landcare Research, National Geographic and Royal Society of New Zealand Marsden Fund. References Butler, D.J., 2006. The habitat, food and feeding ecology of kakapo in Fiordland: a synopsis from the unpublished MSc thesis of Richard Gray. Notornis 53, 55e79. Cockburn-Hood, T.H., 1873. Letter read to Wellington Philosophical Society 16 January, 1874. Trans. Proc. N. Z. Inst. 6, 387e388. Duff, R.S., 1952. Recent maori occupation of Notornis Valley, Te Anau. J. Polyn. Soc. 61, 90e119. Gill, B.J., 2000. Morphometrics of moa eggshell fragments (Aves: Dinornithiformes) from Late Holocene dune-sands of the Karikari Peninsula, New Zealand. J. Roy. Soc. N.Z. 30, 131e145. Gill, B.J., 2006. A catalogue of moa eggs (Aves: Dinornithiformes). Rec. Auckl. Mus. 43, 55e80.
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Worthy, T.H., Holdaway, R.N., 1996. Quaternary fossil faunas, overlapping taphonomies, and palaeofaunal reconstruction in North Canterbury, South Island, New Zealand. J. Roy. Soc. N. Z. 26, 275e361. Worthy, T.H., Holdaway, R.N., 2002. The Lost World of the Moa. Canterbury University Press, Christchurch. 718 pp.. Worthy, T.H., Scofield, R.P., 2012. 21st Century advances in knowledge of the biology of moa (Aves: Dinornithiformes), diagnoses revised, and a new morphological analysis. N. Z. J. Zool. 39, 87e153. Yll, R., Carrion, J.S., Marra, A.C., Bonfiglio, L., 2006. Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy). Palaeogeog. Palaeoclim. Palaeoecol. 237, 32e39.