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Trade and habitat change virtually eliminate the Grey Parrot Psittacus erithacus from Ghana

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  • University of Environment and Sustainable Development

Abstract and Figures

The heavily traded Grey Parrot Psittacus erithacus is believed to have undergone rapid population decline, yet there are almost no quantitative data on abundance changes over time from anywhere within its huge range. We reviewed the species' historical abundance across Ghana, undertook targeted searches during 3- to 5-day visits to 42 100-km(2) cells across the country's forest zone, repeated counts at 22 parrot roosts first performed two decades ago and gauged around 900 people's perceptions of the decline and its causes. In over 150days of fieldwork, just 32 groups (maximum group size=12) were recorded in 10 cells. Encounter rates averaged 0.15 individuals per hour of targeted search, around 15 times lower than those recorded in the early 1990s. No active roosts were found, and only 18 individuals were recorded in three roost areas that each harboured 700-1200 birds two decades ago. Interviewees stressed the importance of very tall trees of commercially important species such as Terminalia superba and Ceiba pentandra for nesting and roosting, and believed that the felling of large trees on farmland (42% of responses) and trapping for trade (37%) were the two main causes of decline. Ghana has lost 90-99% of its Grey Parrots since 1992, a time when the population had presumably already been seriously reduced by two decades of extremely heavy trade. There is no evidence that, away from one or two localities, declines are less severe anywhere else within the West African range of P.erithacus, or across the entire range of the recently split Timneh Parrot Psittacus timneh.
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Trade and habitat change virtually eliminate the Grey
Parrot Psittacus erithacus from Ghana
NATHANIEL N. D. ANNORBAH,
1
* NIGEL J. COLLAR
2
& STUART J. MARSDEN
1
1
School of Science and the Environment, Manchester Metropolitan University, Chester Street,
Manchester M1 5GD, UK
2
BirdLife International, Girton Road, Cambridge CB3 0NA, UK
The heavily traded Grey Parrot Psittacus erithacus is believed to have undergone rapid
population decline, yet there are almost no quantitative data on abundance changes over
time from anywhere within its huge range. We reviewed the specieshistorical
abundance across Ghana, undertook targeted searches during 3- to 5-day visits to 42
100-km
2
cells across the countrys forest zone, repeated counts at 22 parrot roosts rst
performed two decades ago and gauged around 900 peoples perceptions of the decline
and its causes. In over 150 days of eldwork, just 32 groups (maximum group size =12)
were recorded in 10 cells. Encounter rates averaged 0.15 individuals per hour of targeted
search, around 15 times lower than those recorded in the early 1990s. No active roosts
were found, and only 18 individuals were recorded in three roost areas that each har-
boured 7001200 birds two decades ago. Interviewees stressed the importance of very
tall trees of commercially important species such as Terminalia superba and Ceiba pentan-
dra for nesting and roosting, and believed that the felling of large trees on farmland
(42% of responses) and trapping for trade (37%) were the two main causes of decline.
Ghana has lost 9099% of its Grey Parrots since 1992, a time when the population had
presumably already been seriously reduced by two decades of extremely heavy trade.
There is no evidence that, away from one or two localities, declines are less severe any-
where else within the West African range of P. erithacus, or across the entire range of
the recently split Timneh Parrot Psittacus timneh.
Keywords: CITES, encounter rate, parrot trade, population decline, roosts, West Africa.
Efforts to assess long-term changes of global popu-
lations of birds are often hampered by a lack of
quantitative data on historical abundance. Baseline
historical data against which contemporary repeat
surveys can be directly compared are powerful
tools in quantifying the extent and degree of decli-
nes (Prakash et al. 2003, Alderman et al. 2011)
but the existence of such data is rare. More usu-
ally, concern for the status of a species triggers
dedicated surveys, which have to be compared
with anecdotal information about historical abun-
dance (e.g. Cuthbert et al. 2009). An exacerbating
issue with very wide-ranging species is that abun-
dance and population trajectories may be uneven
across their extent of occurrence, making it dif-
cult to characterize declines accurately and assign
causes (Senyatso et al. 2013).
The Grey Parrot Psittacus erithacus (until
recently considered conspecic with the Timneh
Parrot Psittacus timneh) is widely distributed, with
a range of nearly 3 million km
2
from C^
ote dIvoire
and Ghana in West Africa, through Nigeria and
Cameroon and the Congo forests, to Uganda and
western Kenya (del Hoyo & Collar 2014). Over
recent decades, it has been among the most traded
of all birds listed under CITES (Convention on
International Trade in Endangered Species of Wild
Fauna and Flora). CITES-reported imports of Grey
Parrots from all range countries totalled 847 525
between 1980 and 2014 (http://trade.cites.org/).
However, owing to unreported domestic trade,
*Corresponding author.
Email: n.annorbah@mmu.ac.uk
© 2015 British OrnithologistsUnion
Ibis (2016), 158,8291
mortality between capture and sale, and consider-
able unreported international trade, a more realis-
tic gure for the numbers taken from the wild
between 1982 and 2001 alone could be more than
1 million birds (BirdLife International 2015).
The impact of this trade on wild populations is
easy to imagine but difcult to quantify. A recent
review lists multiple reports of severe declines
across much of the Grey Parrot range, but these
are based on anecdotal evidence and/or question-
able or non-comparable methods (Martin et al.
2014). Added to this, recent extensive surveys of
Grey Parrot, using distance sampling (e.g. Buck-
land et al. 2008), found highly variable local abun-
dances very low densities at sites in Liberia,
Cote dIvoire and Democratic Republic of the
Congo (DRC), but remarkably healthy densities in
parts of Cameroon and on the island of Pr
ıncipe in
the Gulf of Guinea (Marsden et al. 2015). An
accurate rate of overall decline has therefore
proved elusive, but given the enormous levels of
offtake for the trade acting concurrently with habi-
tat loss, a decline of 3049% in three generations
(47 years) is currently assumed, qualifying the spe-
cies as Vulnerableaccording to IUCN Red List
criteria (BirdLife International 2015).
An unpublished but technically meticulous
CITES study which reported Grey Parrot encoun-
ter rates and examined parrot abundance at 22
roosts in the early 1990s (D
andliker 1992) pro-
vides a unique opportunity to measure the degree
of decline in Ghana over the past 2025 years.
Using this baseline, the objectives of our study
were fourfold. First, we compared the current dis-
tribution and abundance of the species across
Ghana with historical data from various sources.
Second, we repeated the series of the 1990sroost
surveys undertaken by D
andliker (1992). Third,
we used structured interviews to record knowledge
and perceptions of the decline in Grey Parrots and
its possible causes. Fourth, we examined historical
and current trade and other factors likely to have
contributed to the decline in the species.
METHODS
Historical and current status of the Grey
Parrot in Ghana
We rst reviewed information on the specieshis-
torical status from published and unpublished
reports of previous surveys, involving some of the
earliest scientic information on the species from
the last century, through surveys conducted in the
1970s, to the most recent work on the species in
the 1990s. These included encounter rates for the
species calculated by D
andliker (1992), both from
his surveys and from those from studies between
1976 and 1978 conducted by the Ghana Wildlife
Division (GWD) in and around the village of
Achiase (Aykease) in the Central Region. D
and-
liker himself surveyed parrots at fairly randomly
chosenpoints across his study area during around
30 mornings and evenings. He counted mostly y-
ing parrots, which included both movements of
birds to roosts and ights of local breeding birds.
His surveys were conducted during 24 days in
Western and Central regions, 5 days in the Volta
region, 10 days in the Northern region, and 6 days
in Ashanti and Brong-Ahafo regions.
To assess current distribution and abundance,
50 000 km
2
of the 75 000 km
2
of the historical
range of the species within Ghana (areas in the
west and southwest that were most likely still to
hold the species) were partitioned into a grid of
100-km
2
cells. A sample of 42 cells was selected,
not randomly but to maximize the likelihood of
encountering Grey Parrot populations (Fig. 1).
This was itself based on historical records of the
species and the presence of substantial forest
cover. Each cell was visited for periods of 3
5 days, between April 2012 and June 2014, with
parrot surveys conducted on each day between
05:45 and 11:00 h. Surveys were conducted by
walking along hunter or farmer footpaths and
along drivable pathways connecting villages or
towns. Survey routes traversed a variety of habitats
including small-scale agroecosystems, large-scale oil
palm plantations, forest reserves and settlements.
Survey routes were walked by NNDA in the com-
pany of a local guide, and all individual Grey
Parrots heard or seen along the route, perched or
in ight, were recorded together with direction of
ight and GPS coordinates at the point of detec-
tion. Furthermore, long watches were conducted
from vantage points between 16:00 and 18:00 h
on some days, timed to correspond with the peri-
ods of afternoon ight towards roosts (Wirming-
haus et al. 2001, Amuno et al. 2007).
Roost counts in 199192 and 2014
D
andliker (1992) travelled around Ghana in
December 1991 and January 1992 identifying 60
© 2015 British OrnithologistsUnion
Grey Parrot declines 83
roosts or roost areas, using the local knowledge of
parrot trappers and ofcials from the Ghana Wild-
life Division (see Supporting Information
Appendix S1 for locations). He and his assistants
visited 22 roost areas and counted parrots (on
either one evening or an evening and one morn-
ing) at 15 of these. NNDA visited 42 of the 60
roost areas between 2012 and 2014, including all
22 visited by D
andliker. Roost areas were sought,
based on D
andlikers descriptions, for periods of
12 days. Former trappers or other knowledgeable
local people would guide NNDA to the known
roost or former roost site and/or potential alterna-
tive areas. If local people did not know any
remaining roosts, bird surveys were conducted for
two consecutive days, focusing on potentially suit-
able habitats for parrot roosts, including swampy
habitats (known often to be favoured by Psittacus:
Forshaw & Cooper 1989).
Public perception and knowledge of
Grey Parrots
Structured interviews were conducted in all the
study cells and roost areas were surveyed to obtain
information about local knowledge of Grey Parrots
and to investigate aspects of the parrot trade. The
mean number of interviewees per cell was
17 10 sd (range 345). On arrival at a village
in a study cell, the chief or village leader and his/
her elders were informed about the project.
Aeld assistant subsequently walked around the
village and interviewed any residents willing to
respond to a set of questions from a prepared
questionnaire.
A total of 906 people were interviewed within
squares and during roost searches, although not all
interviewees answered every question. Intervie-
wees were asked when they had last observed
Grey Parrots, whether they thought the species
had declined, and the species of trees in which
they knew Grey Parrots to nest, roost and feed,
and the height of trees used (>30 m =Very tall;
2030 m =Tall;<20 m =Not tall). Questions
and possible responses are given in Supporting
Information Appendix S2. The great majority
(97%) of respondents were male, 87% were aged
30 years or more and 52% were over 40 years old.
Most (92%) had lived in their communities for
over 10 years, and 85% were farmers. Just 0.2%
Figure 1. Map of southern Ghana showing positions of 42 sampled cells, parrot records, roost sites visited and protected areas.
Note that surveys and roost searches were focused on the historical range of the species in the southwest of the country.
© 2015 British OrnithologistsUnion
84 N. N. D. Annorbah, N. J. Collar & S. J. Marsden
were professionalhunters, although 1% stated
hunting as a secondary occupation.
Historical and current trade in Ghanaian
Grey Parrots
We explored the history of the Grey Parrot
trade in Ghana using relevant literature (Ussher
1874, Lowe 1937, Grimes 1987, D
andliker
1992, Dowsett-Lemaire & Dowsett 2014), and
by interrogating the CITES trade database (ac-
cessed on 16 November 2014 with a search for
reported live exports of the species from
Ghana). NNDA conducted detailed interviews
with 29 bird traders in eight major urban centres
in southern Ghana: Kumasi Central Market in
the Ashanti Region; Kantamanto, Malata and
Madina Markets in the Greater Accra Region;
Assin Fosu, Twifo Praso and Kotokoraba Cape-
Coast Markets in the Central Region; and Tako-
radi Market Circle in the Western Region (see
Supporting Information Appendix S3 for GPS
coordinates). Grey Parrots on sale in these mar-
kets or in villages during eld visits were
counted and vendors interviewed. Interviews
were held with active, or previously active, Grey
Parrot trappers throughout the study area.
RESULTS
Historical and current status of the Grey
Parrot in Ghana
Historically, the Grey Parrot occupied much of
the forest zone of Ghana, occurring over approxi-
mately 75 000 km
2
of forest, including the whole
of the Western and Central Regions of the coun-
try, nearly the whole of Ashanti Region as well as
the semi-deciduous forest areas of southern Brong-
Ahafo Region, the western part of the Eastern
Region, and parts of the Greater Accra Region
(Bannerman 1931, Bouet 1961). The species is
also reported to have been present in riparian for-
ests of the savanna zone of the Northern Region,
but there are no data indicating that it was present
in seemingly suitable habitat in the Volta Region
of (easternmost) Ghana (D
andliker 1992). Despite
the sense in the 1940s that the species might have
been declining in some areas owing to trade, large
ocks of 5001000 parrots could still be observed
(Grimes 1987). Four decades later (1988), large
ocks of 20003000 continued to occur in parts of
the speciesrange in Ghana (D
andliker 1992).
Data from studies between 1976 and 1978 con-
ducted by the GWD in and around the village of
Achiase (Aykease) in the Central Region, and anal-
ysed by D
andliker (1992), produced an average
Grey Parrot encounter rate of 1.51.9 birds per
hour (mean of 52 counts). Fifteen years after the
GWD studies, D
andliker (1992) recorded Grey
Parrot encounter rates of 1.11.3 birds per hour
near the Achiase area. From 30 morning/evening
counts across his whole study area, encounter rates
averaged 1.92.4 per hour.
By 1992, around 60 Grey Parrot roosts (per-
haps not all active at one time) had been identi-
ed within Ghana. Individual roosting areas in
Ghana appear to be used over several years
(D
andliker 1992), as they are elsewhere (e.g.
Cameroon: Fotso 1998), although the precise
location of the roost within the area might shift.
It might also be expected that numbers of Grey
Parrots using roosts might change seasonally or
even daily, as in other parrots (Amuno et al.
2007). Roosts were distributed throughout the
forest zone, with the distance of any given roost
to the three nearest roosts averaging 27 km; alto-
gether, 130210 roosts were thought to exist
within the forest zone (D
andliker 1992). Roost
density was estimated at 0.180.27 per 100 km
2
and, allowing for varying abundances in different
forest types, the national population was extrap-
olated to be 30 00080 000 Grey Parrots (D
and-
liker 1992). In a multi-species forest bird study
between June 1995 and August 1996, Grey
Parrots were found in more than 20of the 28
forest reserves studied, but only 87 groups were
recorded during the entire survey period (Nti-
amoa-Baidu et al. 2000).
In our survey, a total of 32 groups, comprising
103 individual Grey Parrots, was recorded in 10
(24%) of the 42 squares surveyed (Fig. 1). Only
one of these groups was actually recorded perched.
The amount of time spent surveying in each
square ranged between 12 and 21 h of eld effort.
Mean encounter rate across all sites was
0.047 0.14 (sd) groups per hour (0.15 0.51
individuals per hour). Half of all birds encountered
(51 of 103) were recorded in one square, and
exclusion of this square reduces mean encounter
rates to 0.026 0.059 (groups) and 0.077 0.18
(individuals). Mean group size was 3.2 (modal
group size, occurring 13 times, =2; maximum
group size =12).
© 2015 British OrnithologistsUnion
Grey Parrot declines 85
Roost counts in 1992 and 2014
Of the 60 roosts and roost areas identied by
D
andliker (1992), NNDA visited 42 (70%;
Appendix S1). No active roosts were found,
although Grey Parrots were seen within a few kilo-
metres of ve former roosts. The largest number
of birds (51) recorded during 4 days of surveys
within a single square was counted in and around
the village of Douahohorodo, a roost identied by
D
andliker (1992) but one which he failed to
locate in his surveys. Only 11 and seven Grey Par-
rots, respectively, were counted near two of D
and-
likers (1992) three biggest roosts of 7001200
birds each. No parrots were found at the third
roost area, where the vegetation is currently domi-
nated by unshaded Cocoa Theobroma cacao planta-
tions. A proper assessment of changes in
vegetation at roosts between 1992 and 2014 was
impossible, but D
andlikers original descriptions of
habitat type tended to match fairly well with that
found in 2014 (Appendix S1). However, habitat
changes are likely to have occurred, including
increased prevalence of oil palm, drainage of and
planting on rafa swamp, and the cutting of indi-
vidual large trees which were the focus of some
roosts in 1992 (Appendix S1).
Public perception and knowledge of
Grey Parrots
Of 866 respondents, 27% stated that they had last
observed Grey Parrots within the previous year,
41% 15 years previously, 22% 610 years previ-
ously and 9% over 10 years previously. In only ve
(12%) of the 42 squares surveyed was there no
report of a parrot sighting in the previous year.
However, 94% of 846 respondents believed there
to have been a decline in Grey Parrot abundance
in the last 5 years, and 99% over the past 10
20 years. All 26 bird traders interviewed in major
market centres in the south of Ghana thought
there had been a major population decline in Grey
Parrots, and that the current supply of birds was
negligible.
Respondents stated that the most important
tree species for nesting and roosting are Terminalia
superba and Ceiba pentandra, with Elaeis guineensis
the most important for feeding (Table 1). A strong
preference for very tall trees for nesting (93% of
respondents) and roosting (95%) was reported, but
there was no stated preference for tree height for
feeding. Of 889 respondents, 70% said nest trees
were located outside rather than within forest
reserves. While we acknowledge potential upward
bias in recording rates outside over those inside
the forest, this at least indicates that Grey Parrots
in Ghana nest outside reserves, as they do else-
where (Twanza & Pomeroy 2011, Irumba 2013).
Respondents attributed the decline in Grey Parrots
to trapping for trade, felling of large trees on farms
or both (Table 2). Hunting for food was very sel-
dom cited as a reason for declines, and nor was
mechanized logging, although the latter was
deemed more important when coupled with loss
of farm trees.
Historical and current trade in Ghanaian
Grey Parrots
Large numbers of Grey Parrots were already
traded in Accra and the Cape Coast in the 1870s
(Ussher 1874, Dowsett-Lemaire & Dowsett 2014).
In his account of expeditions to the Gold Coast in
the 1930s, Lowe (1937) expressed hopes for the
recovery of Grey Parrot populations in the country
owing to the closure of the live bird trade in Eng-
land, indicative of potentially appreciable exports
to the latter country. The large numbersof
Table 1. Main tree species used by Grey Parrots for nesting,
roosting and feeding. The relative importance of each species
is indicated by the number/percentage of times it was men-
tioned by the 906 respondents.
Species Nesting Roosting Feeding
Terminalia
superba (Ofram)
433 (34.6%) 468 (36.5%) 37 (3.0%)
Ceiba pentandra
(Onyina)
236 (18.9%) 232 (18.1%) 46 (3.7%)
Milicia
(Chlorophora)
excelsa (Odum)
221 (17.7%) 204 (15.9%) 113 (9.1%)
Celtis mildbraedii
(Celtis zenkeri)
(Essah)
61 (4.8%) 57 (4.5%) 172 (13.8%)
Elaeis guineensis
(Oil Palm)
3 (0.2%) 58 (4.5%) 497 (40.0%)
Raphia spp.
(Rafa)
2 (0.2%) 41 (3.2%) 185 (14.9%)
Other species
(Other)
295 (23.6%) 222 (17.3%) 193 (15.5%)
Tree heights
Very tall
(>30 m)
743 (92.8%) 732 (95.2%) 216 (26.4%)
Tall (2030 m) 45 (5.6%) 22 (2.9%) 291 (35.6%)
Not tall (<20 m) 13 (1.6%) 15 (2.0%) 311 (38.0%)
© 2015 British OrnithologistsUnion
86 N. N. D. Annorbah, N. J. Collar & S. J. Marsden
parrots traded historically from Ghana remain,
however, unquantied (Ussher 1874, Lowe 1937).
A trade with neighbouring countries in Grey Par-
rot body parts may have begun in the mid-1970s
(D
andliker 1992). In the early 1970s, Ghanaian
Muslims on the Hajj pilgrimage to Mecca began
exporting Grey Parrots to Saudi Arabia for resale
(D
andliker 1992). This novel and lucrative trade
went largely unscrutinized by Ghanaian ofcial-
dom and involved large numbers of parrots; at its
peak, up to 20 000 birds annually were being
moved without documentation by air, both as
cargo and as hand luggage. Trade via this route
ourished until Saudi Arabia placed an embargo
on parrot imports in 1986 (D
andliker 1992).
Meanwhile, several thousand Grey Parrots were
also being exported annually through Ghanas
neighbouring countries such as C^
ote dIvoire,
Togo, Mali and Benin, as far as Senegal, for re-
export to Europe and the USA (D
andliker 1992).
Commercial trade in Grey Parrots underwent
several periods of embargo in Ghana after 1967
until, in May 1980, a ban on exports was imposed
owing to suspicions that the species was being used
to smuggle diamonds (D
andliker 1992). Bans and
revocations were repeated multiple times between
1980 and 1989, until the last ban, still in force
owing to lack of data for establishing an export
quota, was effected in 1994 (CITES 2012). CITES-
reported exports from Ghana averaged 3047
3234 (sd) per annum between 1976 and 1990.
Reported export numbers were highly variable in
these 15 years, ranging from 0 in 1976 to 9585 in
1978 and 8680 in 1985 (http://trade.cites.org/).
CITES-reported exports averaged just two per
annum in the years 19912012.
During NNDAseldwork, ve recently caught
Grey Parrots were found for sale in three markets
two in separate stalls at Kantamanto, two at
Kumasi and one at Assin-Fosu. The traders con-
cerned stated that these ve individuals had been
bought from trappers in the previous few months.
All 26 dealers who gave an opinion reported that
the trade collapsed many years previously, and
that current supply is based on opportunistic trap-
ping by villagers. These dealers kept large stocks of
parrots in their market aviaries in the mid- to late
1990s, and had an abundant supply from trappers.
According to seven dealers, current (2014) prices
were variable (mean =700 621 sd Ghana
Cedis; approximately US$230) but two dealers
stated that Grey Parrots could sell for 10002000
Ghana Cedis (approximately US$330660) if
bought by expatriates. These prices contrast
strongly with those from the early 1990s, when
birds were bought from trappers for US$812, and
passed on from dealer to exporter usually for less
than US$20 (D
andliker 1992).
No currently operating parrot trappers could be
found for interview. Of 23 former trappers inter-
viewed, 17 had become farmers and only one
admitted having trapped a Grey Parrot in the pre-
vious year. Two others had trapped Grey Parrots
in the previous 5 years but 15 stated that they had
not caught any in the previous decade. Nine trap-
pers stated that their livelihoods from Grey Parrot
trapping became unsustainable in the mid-1990s
when parrot populations declined, and many of
their fellow trappers emigrated to other countries
including C^
ote dIvoire, Liberia and even DRC to
ply their trade. Besides Grey Parrots, interviewees
indicated that during the 1980s they trapped other
species including Red-headed Lovebird Agapornis
pullarius, Red-fronted Parrot Poicephalus gulielmi,
West African Pied Hornbill Lophoceros semifascia-
tus, Western Long-tailed Hornbill Horizocerus
albocristatus, Yellow- and Black-casqued Hornbills
Ceratogymna elata and Ceratogymna atrata, Afri-
can Green-pigeon Treron calvus and Great Blue
Turaco Corythaeola cristata (taxonomy follows del
Hoyo & Collar 2014).
DISCUSSION
Grey Parrot populations in Ghana have declined
catastrophically over the past 20 years. This
Table 2. Responses of interviewees concerning the main fac-
tors they considered to be drivers of declines in Ghanas Grey
Parrots. Responses are split according to whether the factor
was identied as acting alone or with other factors.
Factor Alone With other factors Total
Hunting for food
(H)
7 (1.7) 29 (2.3)
Trapping for
trade (T)
211 (49.8) 172(+F); 30(+L);
41(+FL)
463 (36.8)
Felling farm
trees (F)
178 (42.0) 172(+T); 127(+L);
41(+TL)
533 (42.3)
Mechanized
logging (L)
28 (6.6) 127(+F); 30(+T) 234 (18.6)
Figures are shown for combinations of factors (with other fac-
tors) only if numbers are greater than 20. Figures in parenthe-
ses are the percentage of times a particular response was
given either alone or along with other factors.
© 2015 British OrnithologistsUnion
Grey Parrot declines 87
decline is certainly in excess of 90%, and could be
as high as 99%, as evidenced by the near-total loss
of the major roosts known in 1992 (D
andliker
1992). This said, Grey Parrots were recorded in
one quarter of surveyed squares, and small num-
bers remain over quite large areas of the country
a pattern shown elsewhere in Africa of signicant
numerical contraction with little or no distribu-
tional change (e.g. Kori Bustard Ardeotis kori;
Senyatso et al. 2013). Results of our study largely
concur with a recently published Ghana bird atlas
(Dowsett-Lemaire & Dowsett 2014), which found,
at a coarse scale, that Grey Parrots were still pre-
sent in very small numbers across much of the
southeast of the country. Three- to ve-day visits
to eight forest reserves in 20082009 yielded
sightings of single pairs (three reserves), single
birds (three reserves) and no birds (two reserves).
Encounter rates of 0.15 individuals per hour
(around one group per 23 days) of targeted
searching over wide areas indicate the enormity of
the decline since previous studies, which recorded
encounter rates of around two individuals per hour
in the early 1990s (D
andliker 1992). Although
comparisons of encounter rates presented in differ-
ent studies are problematic, these ndings are fur-
ther reinforced by intervieweesperceptions and a
general lack of observations.
Causes of decline
We identify four factors that condently can be
regarded as having contributed to the decline of
the Grey Parrot in Ghana: trade, overall forest
reduction, silvicultural practice and farmland tim-
ber harvesting.
The CITES trade database has Grey Parrot
imports originating from Ghana totalling 67 259
birds for the period 197690 (www.unep-wcmc-
apps.org/citestrade). Allowing for domestic con-
sumption, which does not appear in CITES trade
gures, high pre-export mortality (variable, but
averaging around 50% from capture to market:
Clemmons 2003), and the absence of data on pre-
sumably signicant illegal trade in ofcial reports
(Pain et al. 2006), the real number taken from the
wild in Ghana in this period is probably far higher
and plausibly well over twice as many. D
andliker
(1992) estimated annual offtake levels in excess of
10 000, and up to 20 000 birds during years of
maximum trade, between the early 1970s and late
1980s. The heavy trade in the 1970s and 1980s
presumably contributed signicantly to population
declines well before D
andlikers surveys. However,
most striking is the fact that in the years 1991
2012 when trade was outlawed and Ghanas
CITES-reported exports of Grey Parrots totalled
just 35 individuals (two per annum), the popula-
tion in the country still declined by 95%. As inter-
viewed trappers still held large numbers of Grey
Parrots in the mid-1990s, this illegal trade must
surely have contributed to the post-1990 declines
that we report. However, because some trappers
judged the trade unsustainable from the mid-
1990s, the bulk of the illegal trade may have
occurred in the years immediately after 1992. It is
also possible that heavy capture of juveniles in the
decades preceding 1992 had left the Grey Parrot
population with a highly skewed age structure.
Parrots are long-lived (Forshaw & Cooper 1989)
and there may have been a signicant time lag
between the perturbation and the resultant decline
of the species (Marsden & Pilgrim 2003, Hylander
& Ehrlen 2013).
Logging and farming are major causes of forest
loss and alteration in Ghana, compounded by poor
forest management, re and mining (Benhin &
Barbier 2004). Ghanas human population
increased from 8.5 million in 1970 to 24.2 million
in 2010, at rates well over 3% per annum
(National Population Council of Ghana 2006,
2011, Ghana Statistical Service 2012). This rapid
population growth has shortened fallow periods
and increased demand for land (Donkor & Vlosky
2003). At the time of D
andlikers survey in 1992,
forest cover totalled 74 480 km
2
, whereas by 2010
it had fallen by a third to 49 400 km
2
(http://rain-
forests.mongabay.com/deforestation/2000/Ghana.
htm). This clearly represents a signicant reduc-
tion of Ghanas carrying capacity for Grey
Parrots.
Reduction in habitat quality, especially by the
removal of large trees, is another important factor
in parrot declines (e.g. Manning et al. 2013). Par-
rots typically nest in very large trees (e.g. Marsden
& Jones 1997, Monterrubio-Rico & Enkerlin-Hoe-
ich 2003), and Grey Parrots are no exception
(Clemmons 2003, Valle 2015). In the 1970s and
1980s, the practice of salvage loggingwas proba-
bly particularly detrimental to Ghanas Grey Par-
rots, as this permitted the unlimited felling of
large trees as a means of cleansingforests of over-
mature timber, regarded as undesirable under
national forest management policy (Hawthorne &
© 2015 British OrnithologistsUnion
88 N. N. D. Annorbah, N. J. Collar & S. J. Marsden
Abu-Juam 1995, Dykstra et al. 1996, Benhin &
Barbier 2004).
Licensed logging of individual trees in agricul-
tural areas evidently further damaged Grey Parrot
populations, particularly as 70% of interviewees
reported the species nesting outside forests. Over
50% of timber harvests in Ghana during the per-
iod 197292 were from outside reserves, with
farmers having no legal rights to timber trees on
their land and receiving little or no compensation
from loggers for crop damage during felling (Sar-
gent et al. 1994). Crucially, this circumstance
remains a disincentive to retain trees on farmland
and contributes heavily to the pre-emptive
destruction of large trees outside forest reserves
(Ruf 2011).
The status of Grey and Timneh Parrots
across West Africa
More than 6 months of dedicated searching within
likely areas of Ghana, including visits to roosts
which previously held 7001200 individuals,
yielded just a handful of Grey Parrot sightings.
Undoubtedly, the species has declined precipi-
tously, and is now extremely rare across Ghana. It
is likely that the status of the Grey Parrot and its
recently split sister species the Timneh Parrot is
similar across West Africa (west of Cameroon):
both species have been virtually eliminated from
wider landscapes, and exist in reasonable numbers
at very few locations. Such sites include the
Bijag
os Islands, Guinea-Bissau (Martin et al. 2014),
and perhaps parts of the Gola Forest in Liberia
and Sierra Leone, where ocks of around 70 Tim-
neh Parrots have recently been recorded (NNDA
pers. data, C. Showers in litt. 2013). Interestingly,
Liberia appears to have retained its larger animals
relatively well (Tweh et al. 2015), and further
work might nd small parrot populations else-
where in the country. However, densities else-
where within the range of the Timneh Parrot
appear extremely low (Martin et al. 2014). For
example, around 32 km of distance sampling line
transects and 38 h of dedicated encounter rate sur-
vey within Parc National dAzagny and other likely
areas in Cote dIvoire yielded no records of the
species (Marsden et al. 2015). The situation for
Grey Parrot in West Africa is quite possibly worse,
with the species being probably extinct in Togo,
and very rare right across Nigeria (McGowan
2001, Green et al. 2007, Martin et al. 2014).
Absence of evidence that any Psittacus popula-
tions in the region are healthy must surely now
preclude trade from West Africa, and calls into
question whether there should be any further
trade in much of mainland Central Africa. The
Red List classication of both Grey Parrot and
especially the much smaller-ranged Timneh Parrot
clearly requires re-evaluating.
NNDAs PhD studentship was generously funded by the
Loro Parque Fundaci
on. We thank the many local guides
who assisted with eldwork, as well as E. Brown and,
especially, S. Kenyenso for his immense contribution by
way of dedicating nearly 2 years of his time to inter-
views and eldwork. NNDA also thanks L. Holbech, D.
Attuquayeo and A. Asamoah for their constant encour-
agement and practical eld advice. We are grateful to
the Forestry Commission of Ghana for granting access to
forest reserves and national parks. We thank Gottlieb
D
andliker for his help with aspects of his original report,
and two anonymous referees for valuable comments on
the manuscript.
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SUPPORTING INFORMATION
Additional Supporting Information may be found
in the online version of this article:
Appendix S1. Counts of Grey Parrots at roosts
identied by D
andliker (1992) in Ghanas high
forest zone, and repeat counts from the current
study.
Appendix S2. Structured questionnaire showing
list of questions and possible answers administered
during surveys in the study area.
Appendix S3. GPS coordinates for markets sur-
veyed.
© 2015 British OrnithologistsUnion
Grey Parrot declines 91
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