Naturwissenschaften (2012) 99:177–184
DOI 10.1007/s00114-012-0882-8
ORIGINAL PAPER
Dodo remains from an in situ context from Mare aux
Songes, Mauritius
Hanneke J. M. Meijer & Arike Gill &
Perry G. B. de Louw & Lars W. Van Den Hoek Ostende &
Julian P. Hume & Kenneth F. Rijsdijk
Received: 1 July 2011 / Revised: 5 January 2012 / Accepted: 11 January 2012 / Published online: 27 January 2012
# Springer-Verlag 2012
Abstract Since 2005, excavations at Mare aux Songes,
Mauritius, have revealed the presence of a very rich,
∼4,200-year-old fossil bone bed including dodo (Raphus
cucullatus) bones and bone fragments. The recently excavated dodo assemblage comprises at least 17 individuals and
is characterised by the presence of small and fragile skeletal
elements, a dominance of leg elements and an absence of
juveniles. The hydrology of the area suggests that dodos,
like many other species, were probably lured to Mare aux
Songes by the presence of freshwater during times of
drought. The most likely scenario for the origin of the fossil
deposit is that animals became trapped in the sediment in
repeated miring events, which would favour the conservation of hindlimbs. Such a scenario is fully in accordance
with the taphonomic characteristics of the bone assemblage.
Communicated by: Sven Thatje
Keywords Raphus cucullatus . Insular ecosystem . Mare aux
Songes . Mauritius . Taphonomy
Electronic supplementary material The online version of this article
(doi:10.1007/s00114-012-0882-8) contains supplementary material,
which is available to authorized users.
H. J. M. Meijer (*)
Division of Birds, National Museum of Natural History,
Smithsonian Institution,
Washington, DC 20013-70122, USA
e-mail: meijerh@si.edu
H. J. M. Meijer : A. Gill : L. W. Van Den Hoek Ostende
Department of Geology,
Netherlands Centre for Biodiversity Naturalis,
P.O. Box 9517, 2300 RA Leiden, The Netherlands
P. G. B. de Louw
Deltares, Department Soil and Groundwater,
P.O. Box 85467, 3508 AL Utrecht, The Netherlands
J. P. Hume
Bird Group, Department of Zoology,
Natural History Museum at Tring,
Akeman Street, Tring,
Herts HP23 6AP, UK
K. F. Rijsdijk
Institute of Biodiversity and Ecosystem Dynamics,
Computational Bio- and Physical Geography,
University of Amsterdam,
Sciencepark 904,
1098 XH Amsterdam, The Netherlands
Introduction
Since its demise in the seventeenth century (Hume et al.
2004), the dodo (Raphus cucullatus) has become an icon of
extinction. The small number of genuine but often contradictory and inadequate contemporary accounts and illustrations
cause our knowledge of the dodo’s morphology and ecology
to be clouded by assumptions and misinterpretations (for
instance, see the recent discussion on dodo body mass by
Angst et al. (2011a, b) and Louchart and Mourer-Chauviré
(2011)). Even though the recent history of human colonisation
and impact on Mauritius is well documented (Cheke and
Hume 2008), virtually no records of the prehuman ecosystem
exist, making it difficult to assess the magnitude of the
changes brought about by human settlement.
The first fossil remains of the dodo, together with those
of most of the now-extinct vertebrates of the island, were
discovered in 1865 by Harry Higginson and George Clark in
a richly fossiliferous horizon in the Mare aux Songes (MAS)
marsh (Fig. 1) (Clark 1866; Hume et al. 2009). Even though
large amounts of dodo material were recovered by George
Clark and subsequent excavators (Günther 1875; Carié
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Naturwissenschaften (2012) 99:177–184
suggests that accumulation of the vertebrate bones occurred
within a narrow time window between 4235 and 4100 cal.
year BP (Rijsdijk et al. 2011). However, geomorphological,
taphonomic and botanical evidence (Rijsdijk et al. 2009)
excludes a catastrophic event, such as a tsunami or volcanic
eruption, as the cause for the accumulation.
Here, we describe the dodo assemblage from recent
excavations at Mare aux Songes (2006–2008) within a
taphonomic context. We aim to explore several death
scenarios that are related to drought, including thirst, the
drinking of poor quality/toxic water, and a scenario in
which animals becoming trapped in the peat substrate
(miring). All are examined here against the taphonomic
characteristics of the dodo assemblage.
Material and methods
The geological setting of Mare aux Songes
Fig. 1 a Google earth image of Mare aux Songes and position of subbasins I, II and III and trenches (TR). The forested parts and scrub rich
fields mark steep slopes and wet depressions. b Geomorphology of
MAS, the thickness of lines indicates the steepness classes of the
slopes. Yellow indicates a dirt road. Reprinted from Rijsdijk et al.
(2009) with permission
1930; Hoffstetter 1945, 1946a, b; Newton and Gadow 1893;
Hume et al. 2009), no contextual geological or paleoecological data were recorded at the time. Furthermore, the exact
location where these bones were found within the MAS is
still unknown, but Hume et al. (2009) suggest that the
original locality was situated farther upslope from MAS
basin I (Fig. 1). In an effort to eradicate malaria, the marsh
was in-filled with dolerite blocks and gravels in the 1940s,
after which, the site was neglected by the scientific community.
By the 1980s, the Mare aux Songes marsh was erroneously
believed to have been completely destroyed by the development of nearby Sir Seewoosagur Ramgoolam International
Airport (Cowles 1987).
The re-examination in 2005 of the MAS area provided a
unique opportunity to excavate the Mauritian vertebrate
fauna in context. Excavations at MAS revealed the presence
of a very rich fossil bed containing a diverse array of macroand micro-flora and fauna, including the dodo, giant tortoises Cylindraspis spp., and vast numbers of seeds of the
endemic Tambalacoque Sideroxylon grandiflorum and other
tree species (Rijsdijk et al. 2009). Radiocarbon dating
The marsh area (20°26′51, 2″ S; 57°41′23, 5″ E) is situated
at the south-eastern part of Mauritius within an isolated
basaltic rock basin less than 1 km from the Indian Ocean
(Fig. 1a). The basin was formed as a result of the collapse of
a lava tunnel, which resulted in three separate basins (I, II
and III), all situated at <0.5 m above mean sea level
(Fig. 1b). Basins I and III contain vertebrate fossils, whereas
basin II, filled in by a 1–2 m + accumulation of anthropogenic
dolerite boulders, did not yield any vertebrate material. Within
the basins, actual groundwater levels range from just below,
and rise during heavy rainfalls to just above, the present
surface (Rijsdijk et al. 2009). In 2005, a fossil bone bed was
discovered in basin I. Several cores and excavation trenches
(Fig. 1, Tr. 0–4) showed the bone bed to cover an area of
1.86 ha with a thickness between 0.25 and 1.5 m.
A detailed description of the sedimentology is given
by Rijsdijk et al. (2009). In summary, the rock basin is
filled with tropical clays that are overlain by eolian
coral sands (Fig. 2, unit B). On top of these carbonate
sands lies an approximately 40 cm thick discontinuous
lenticular grey lake marl (Fig. 2, unit C), which is
covered by fossil bed layer D. The top of layer D
contains small basaltic (dolerite) gravels that increase
in concentration as well as in grain size to the top of
layer D. A layer of larger, anthropogenic dolerite gravels and blocks (layer E) caps layer D and has sealed
MAS since the 1940s. At Tr. 0 and Tr. 1, the bone bed
is on average 0.5 m thick and consists almost entirely
of bones, seeds and wood (wood cross-section >1 cm)
debris. Trenches 0–1 are located at the north-western
edge of the basin. Here, bone and wood remains dominate and fine matrix material is nearly absent. Therefore, the fossil bed here is believed to represent a
Naturwissenschaften (2012) 99:177–184
179
Fig. 2 Generalised
stratigraphic profile of basin I
based on stratigraphic profiles
of three trenches within basin I
by Rijsdijk et al. (2009). Actual
thicknesses of layers A–E vary
depending upon location within
the basin. Top of profile is
located at −1 m below sea level.
Detail of layer D depicts
vertebrate remains, with
tarsometatarsus of R. cucullatus
in the middle within matrix of
floral remains and debris
shallow lake shore facies. Wind-driven currents at the lake
shore led to reworking of the fossil layer and winnowing of
the fine sediments, leading to a coarse bone-rich layer.
Trenches 2 and 4 are located in the centre of basin I, where
Fig. 3 Proportions of cranial,
wing, leg and axial skeletal
elements within the MAS dodo
assemblage (a), and in a nonbias setting (b), and for trenches
0 and 1 (c) and trenches 2
and 4 (d)
the fossil layer is thicker and the bones are dispersed in a
chestnut brown organic matrix (gyttja). The bones are estimated to constitute 10–30% of the total weight of the fossil
layer. The gyttja consists of fine organic debris indicating
180
accumulation under low-energy conditions that prevailed in
the deeper centre of the lake. These contrasting lake shore and
centre depositional environments may have affected the taphonomy of the vertebrate assemblage.
Hydrological and geological data indicate that MAS
formed ca. 5000 cal. year BP ago as a shallow freshwater
lake in response to eustatic sea level rise (Rijsdijk et al.
2009). Radiocarbon dating of nine vertebrate bones yielded
a weighted average 14C age of 3850± 15 14C year BP,
indicating that fossil deposition occurred over a very short
period of time between 4235 and 4100 cal. year BP (Rijsdijk
et al. 2009, 2011). The age of the MAS assemblage coincides with a period of regional aridification (Gasse 2000;
Marchant and Hooghiemstra 2004; Rijsdijk et al. 2011). On
volcanic islands, stagnant freshwater is rare due to the great
permeability of the basaltic rocks, which cause rainwater to
infiltrate easily into the soil. On Mauritius, however,
groundwater flowed via aquifers towards the dry coastal
regions where it emerged at depressions, such as MAS,
forming freshwater lakes (Rijsdijk et al. 2011). Because
MAS provided a rare source of freshwater within the dry
coastal lowlands, it attracted a diverse fauna, especially
during the dry season.
Naturwissenschaften (2012) 99:177–184
bone fragments from the excavations in basin 1 from 2006
to 2008 (n0235) (see ESM). Bones handpicked during the
initial discovery in 2005 (n014) have been excluded in
order to reduce taphonomic bias relating to size sorting.
From 2009, bones were excavated in an in situ situation,
which differs from the excavation techniques employed
previously. For the taphonomic analyses, the following
parameters were scored: side of the bone (dexter or sinister),
completeness (in case of bone fragment, proximal, middle or
distal part), colour [six categories, ranging from 0.5 (light
ochre) to 3 (dark brown)], bone weathering class following
Behrensmeyer (1978), degree of erosion [eight classes ranging
from 1 (no erosion) to 8 (severe erosion)], degree of fusion and
the presence of pathologies and gnawing, cutting and/or burning marks. We assessed skeletal part survivorship as expressed
as the percentages of anterior elements (%A) relative to the
total number of (A) and posterior (P) elements following
Ericson (1987), where A 0 ulna + carpometacarpus +
humerus and P 0 femur + tibiotarsus + tarsometatarsus.
The ratio of anterior and posterior limb elements provides
evidence for taphonomic processes that affect paleoecological signals and values that deviate significantly from
50% indicate taphonomic overprinting.
Excavation methods
Results
All excavations and subsequent analyses were carried out
with permission of the Mauritius National Heritage Fund
and Omnicane. As a result of the coincidental discovery of
the fossil bed in 2005, bones were handpicked from material
that was brought above surface with a mechanical digger’s
scoop. In 2006, trenches were dug at basins 1 (Fig. 1, Tr. 1–2)
and 3 (Fig. 1, Tr. 3) using a mechanical shovel scoop to
penetrate the upper layer of gravel and boulders and expose
the fossil layer. As a result of the rapid infilling of groundwater, it was not possible to excavate under dry conditions.
Initially, a bulk sampling method was employed in which
the relatively undisturbed sedimentary structures within the
centre of the shovel scoop were used for sedimentological and
stratigraphic analyses. However, association, articulation and
orientation of skeletal material were not assessed systematically, as the in situ vertebrate material was disturbed during
scooping and removal. The scooped bulk samples were sieved
with a nested sieve system using 15, 10 and 4 mm meshes to
retrieve macrofossil remains. Sieve residue subsamples were
dried and microfossils were handpicked.
During the 2006–2008 field campaigns of the Dodo Research
Programme, a total of 235 dodo bones and bone fragments
were retrieved from the MAS (see ESM). The assemblage
comprises cranial bone elements, including mandibula, nasal
bar, palatine, premaxillary, pterygoid and jugal bones, sternal
elements, vertebrae, humeri, scapulae, radii, carpometacarpi,
pelves, femora, fibulae, tibiotarsi, tarsometatarsi and phalanges (Figs. 3a, 4 and 5). Only 26 fragments could not be
assigned to a bone element. All bones are fully ossified and
no bones of juvenile dodos were found. The retrieval of
numerous small elements such as phalanges confirms that
the dominance of larger elements in previously collected
museum dodo material was due to sampling bias (Hume
2005). The total number of identifiable specimens is 209.
The minimum number of individuals (MNI) is based on the
most common element, which was the left tibiotarsus (n017).
Note that the 36 pelvic fragments (ESM) may not represent 36
individuals, as we cannot rule out that some may have come
from the same individual pelvis.
Taphonomic analyses
Taphonomical characteristics
All bone material from the excavations at Mare aux Songes
is stored at Omnicane, Mauritius, and all dodo bones have
been numbered and categorised. We have focused our taphonomic analyses on a dataset that is limited to bone and
In situ articulation and association could not be assessed
systematically within the scoop samples, but the presence of
at least three partly associated pelvic bone constellations (i.e.
bone clusters belonging to one individual) suggests that partial
Naturwissenschaften (2012) 99:177–184
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Fig. 4 Selected dodo upper
body remains from Mare aux
Songes: a sternum (AG 94); b
cervical vertebra (AG 83); c left
scapula (AG 57); d left humerus
(AG 112); e left radius (AG 16);
f left carpometacarpus (AG 62).
Scale bar notes 5 cm
in situ association occurs (MAS-AG 171-173, 184-189 and
183-196-472). Throughout the dodo assemblage, bone preservation is good. The majority of the MAS dodo bones
(>70%) shows a medium to dark brown colouration (colour
classes 2.0–3.0), with lighter colours being less common.
There are no signs of subaerial weathering: the surface of
the bones is very smooth and small bone structures are still
present. The dodo bones were therefore all assigned to
weathering stage 0 as defined by Behrensmeyer (1978),
i.e. no signs of cracking and flaking. The majority of the
bones show small, non-parallel scratches (Fig. 6) on the
surface. They range in length from several millimetres to
10 mm in length and are generally similar in colour or
darker than the bone. No gnawing, cutting or burning marks
were found on the bones. Most of the bones show no to
medium erosion (68%). Heavy and severe erosion is less
common. Erosion is most evident on the proximal and distal
articular surfaces of the bones due to the thinness of the
cortical bone compared to the shaft. Increasing stages of
erosion resulted in holes in the bones and sometimes
even the removal of the articular surface. It was not clear if a
fragmented bone resulted from friction (accumulated erosion)
or actual breakage (quick break caused by the pressure of a
large object on top of the bone). However, the bones show
recent fractures that are thought to have been caused by the
dumping of rocks and the sampling with a mechanical digger
scoop. These ‘new’ fractures are generally oblique to the bone
long axis, relatively sharp and generally lighter in colour than
the undamaged bone surface.
Bias towards hind limb elements
The MAS dodo assemblage is dominated by leg elements,
constituting 76% (n0158) (Fig. 3a). Elements from the axial
skeleton (including the sternum) are, after the leg elements,
far less common (14%; n029). Cranial bones constitute
only 6% (n013) of the assemblage, whereas wing elements
are scarcest with only 4% (n09). The number of leg elements in MAS is almost twice as many as might be expected
if there were no preservational bias (Fig. 3b), whereas wing
elements and bones from the axial column are notably
under-represented. When skeletal part survivorship is
assessed, the %A in the MAS dodo assemblage is 5%
suggesting a strong degree of taphonomic overprinting.
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trenches (Fig. 3c–d) shows a dominance of leg elements in
both Tr. 0 and 1 (n026, leg elements 67%, %A00) and Tr. 2
and 4 (n0209, leg elements 76%, %A05). Dominance of hind
limb elements was also true of the original nineteenth
century collections in which the most common element
recovered was the tarsometatarsus (MNI 0300) (Hume
2005; Rijsdijk et al. 2009).
Discussion
The recently excavated MAS assemblage
and existing collections
Fig. 5 Selected dodo pelvic girdle remains from Mare aux Songes: a
pelvic fragment with the acetabular region (AG 128-130); b right
femur (AG 125); c right tibiotarsus (AG 100); d right tarsometatarsus
(AG 480); e phalanges (AG 39-40). Scale bar notes 5 cm
A difference in location, and thus depositional setting,
between Tr. 0 and 1 (located near the north-western edge of
the basin) and Tr. 2 and 4 (located more towards the middle)
may account for an over-representation of leg elements in one
of the two sites. However, a breakdown of the assemblage into
Fig. 6 Dodo left femur with
polydirectional scratches (inset)
interpreted as derived from
bioturbation such as giant
tortoise trampling and/or the
anthropogenic deposition of
boulders
The retrieval of small and fragile skeletal elements and the
colouration of the recently excavated MAS bone material is
in contrast to the material collected in the nineteenth and
early twentieth century (Hume et al. 2009). The variation in
colour observed in dodo bones in museum collections, ranging from dark brown to almost white, is greater than in the
recently excavated assemblage. Also, despite the fact that
Carié’s collection included small specimens (i.e. Angst and
Buffetaut 2010), museum collections appear to be biased
towards large elements. These differences confirm the
observations by Hume et al. (2009) that the dodo bones
excavated in the nineteenth and early twentieth century are
derived from another locality outside basin 1 at the eastern
margin of the basin complex (Hume et al. 2009). There is no
clear indication that sediments were fine sieved during these
excavations, as Clark (1866) even described that his workers
located fossils in the marshy sediments by using their bare
feet. These practices lead to the assumption that there was a
taphonomic bias towards larger elements. The recent
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excavation of small elements at MAS confirms that the overrepresentation of larger elements in previously collected fossil
material was at least partly due to sampling bias (Hume 2005).
Dodo death scenarios
Before exploring possible hypotheses for the deposit’s formation during the megadrought event, it is important to note that
there is very little evidence for post-depositional transport,
indicating that the birds died at, or close to, the marsh. Partial
observation was only observed incidentally. This is thought to
result from in situ disturbances, such as the wallowing of giant
tortoises (Hume 2005). Also, due to the insular nature of the
fauna, predators as accumulating agents would have contributed little to dodo mortality. Hydrological evidence (Rijsdijk
et al. 2009, 2011) indicates that MAS was a rare source of
freshwater in a dry, lowland coastal setting which, certainly in
times of a megadrought (4235–4100 cal. year BP, Rijsdijk et
al. 2011), would have attracted a range of animals.
The simplest death scenario would entail animals dying
of starvation and/or thirst, as animals are tied to the water
and cannot range far enough to forage. In such a scenario,
the bones of the deceased animals would have been exposed
at the surface, resulting in weathering of the bone surface
(Behrensmeyer 1978; Behrensmeyer et al. 2003). This is in
contrast to the good preservation of the bones, indicating that
exposure of the bones was limited and burial occurred rapidly
after death. A drought scenario alone also does not account for
the over-representation of leg bones, as the ratio of anterior
and posterior elements (%A and %P) should have been more
or less equal. Furthermore, the absence of juveniles, which as a
vulnerable subset of the population would certainly have been
severely affected, does not support drought-induced death.
Another possibility to explain such a dense assemblage
of animals is that in times of drought, animals that gathered
at MAS resorted to drinking low quality or even toxic water.
As MAS is connected to the sea, brackish groundwater may
have flowed into the lake during droughts when lake level
dropped below ocean level due to high evaporation rates.
Alternatively, toxic cyanobacteria may bloom in standing
freshwater lakes during dry, warm periods. There is at least
one fossil locality (Neumark Nord, Germany) at which toxic
cyanophycean blooms are linked to mass death of vertebrates (Braun and Pfeiffer 2002). So far, there is no evidence
to support the presence of toxic algae in MAS. Also, the
Neumark Nord vertebrate assemblage was characterised by
the presence of complete and articulated skeletons, which
have not been found at MAS.
An alternative scenario for the formation of the fossil
deposit is that the MAS functioned as a mire in which
(heavy) animals became trapped. Mire is an ecological term
broadly synonymous with peatland (Johnson and Gerbeaux
2004), and the term ‘miring’ refers to the trapping of
183
animals in such sites. Miring can lead to well-preserved,
associated remains as the sediments would protect the submerged lower parts of the animal’s body from subaerial
exposure and weathering. After the animal’s death, the
exposed parts of the carcass may have attracted scavengers,
and any remaining exposed parts of the animal on the
surface (vertebral column and cranial elements) would have
been prone to erosion and scavenging. A miring scenario
explains the overrepresentation of hind limb elements and
the preservation of small phalanges, and repeated miring
events at MAS, over perhaps long periods of time, may
have led to the final accumulation.
A miring scenario has also been proposed for Pleistocene
and Holocene assemblages of the large, flightless moa
(Family Dinornithidae) from New Zealand (Holdaway and
Worthy 1997; Worthy 1998; Wood et al. 2008). Particularly,
the sites of Pyramid Valley (Holdaway and Worthy 1997)
and Te Aute (Worthy 2000) are similar to Mare aux Songes
in many respects. Both Pyramyid Valley and Te Aute are
swamp deposits, Holocene of age, dominated by large,
flightless birds, characterised by an over-representation of
leg elements and dense accumulations of mostly disarticulated bones (Holdaway and Worthy 1997; Worthy 2000).
For the Pyramid Valley, it is hypothesized that the mire operated seasonally and that mostly heavy, adult females were
trapped in summer when water levels were low (Holdaway
and Worthy 1997; Turvey and Holdaway 2005). A seasonally
active mire at MAS could explain the absence of juvenile
dodos in the MAS assemblage. Alternatively, the absence of
juvenile dodos may also be indicative of a very low reproductive rate and rapid growth to adulthood, as appears to be the
case in the endemic Hawaiian geese, the Moa-nalos (S. Olson,
personal communication), or that the breeding grounds were
elsewhere. However, it should be noted that the population
structure of excavated samples does not necessarily reflect that
of the surrounding population, as stratigraphic and taphonomic variation can preferentially affect the preservation of groups
within populations (Turvey and Holdaway 2005).
Although the time window for MAS is much shorter than
for the NZ sites, a miring scenario explains both the excellent preservation of the bones and the over-representation of
posterior elements (%P). Ideally, miring would also lead to
finding leg bones articulated and in vertical position. As yet,
however, there is no evidence for this at Mare aux Songes.
The presence of small polydirectional scratches on the bones
(Fig. 6) indicates that there was significant bioturbation during
and after deposition of the accumulation (Behrensmeyer et al.
1989). Animals becoming trapped and fighting to get out, as
well as giant tortoises scavenging on the carcasses would have
caused significant bioturbation at the time of deposition
(Hume 2005). The anthropogenic deposition of boulders in
the 1940s also might have caused disturbance of the sediments
post-deposition.
184
Conclusions
Taphonomic characteristics of the Mare aux Songes dodo
assemblages suggest that the animal remains are found close
to the place they died. The hydrology of the area indicates
that the presence of freshwater during times of drought drew
the animals to the site. Of the various scenarios presented
here that might explain the origin of the Mare aux Songes
fossil bed, only a miring scenario agrees with the taphonomic evidence thus far. Miring would have specifically
affected heavy and flightless animals such as adult dodos
and giant tortoises, and repeated miring events at MAS led
to the final accumulation.
Acknowledgements We are very grateful to C. Foo Kune, former
CEO of Mon Trésor Mon Désert Sugar Estate (now Omnicane), and
his staff for their hospitality and invaluable support in the field. The
excavation would not have been possible without the technical supervision by R. Floore. We thank A. Janoo (National Heritage Fund), C.
Baider, F.B.V. Florens, A. Grihault, J. de Vos and J. Brinkkemper for
their support in the field and valuable discussions regarding the extinct
and extant Mauritian ecosystem. I. Prins provided invaluable logistic
support. Fieldwork and research was funded by Omnicane, TNO—the
Geological Survey of the Netherland, the Treub Foundation for
Research in the Tropics, World Wildlife Fund—The Netherlands,
Deltares, Mauritius Museums Council, Hollandia Archaeology,
Taylor Smith Group (Mauritius), Air Mauritius, Mauritius Sugar Industry
Research Institute and the Netherlands Centre for Biodiversity Naturalis.
The funders had no role in study design, data collection and analysis,
decision to publish or preparation of the manuscript. We thank S. Olson
for insightful comments on an earlier version of the manuscript, and A.
Cheke for help on correct citation details. Four anonymous reviewers
provided their constructive input.
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