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Woodpeckers: Distribution, conservation, and research in a global perspective


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Mikusiński, G. 2006: Woodpeckers: distribution, conservation, and research in a global perspec-tive. — Ann. Zool. Fennici 43: 86–95. The main aim of this paper is to examine and discuss the global pattern of woodpecker diversity from a conservation perspective. In addition, I review ecological traits and the conservation status of the entire family Picidae, and relate these factors to the human driven change in their habitats. Finally, I present a global overview of the research on woodpeckers in order to identify the major gaps in our knowledge which render the management of populations of these species difficult. The hotspots of woodpecker spe-cies richness identified by GIS were found in tropical and subtropical forests of South-East Asia, South and Central America, and equatorial Africa. Most of these hotspots were located in developing countries. However, almost 90% of articles published in 1985–2004 encompassed studies performed in North America and Europe, that is, in geographic areas harbouring only 17% of the global number of Picidae species.
Content may be subject to copyright.
Ann. Zool. Fennici 43: 86–95 ISSN 0003-455X
Helsinki 28 April 2006 © Finnish Zoological and Botanical Publishing Board 2006
Woodpeckers: distribution, conservation, and research in
a global perspective
Grzegorz Mikusiński
Grimsö Wildlife Research Station, Department of Conservation Biology, Swedish University of
Agricultural Sciences, SE-730 91 Riddarhyttan, Sweden (e-mail:
Received 7 Oct. 2005, revised version received 6 Mar. 2006, accepted 6 Feb. 2006
Mikusiński, G. 2006: Woodpeckers: distribution, conservation, and research in a global perspec-
tive. — Ann. Zool. Fennici 43: 86–95.
The main aim of this paper is to examine and discuss the global pattern of woodpecker
diversity from a conservation perspective. In addition, I review ecological traits and the
conservation status of the entire family Picidae, and relate these factors to the human
driven change in their habitats. Finally, I present a global overview of the research on
woodpeckers in order to identify the major gaps in our knowledge which render the
management of populations of these species difficult. The hotspots of woodpecker spe-
cies richness identified by GIS were found in tropical and subtropical forests of South-
East Asia, South and Central America, and equatorial Africa. Most of these hotspots
were located in developing countries. However, almost 90% of articles published in
1985–2004 encompassed studies performed in North America and Europe, that is, in
geographic areas harbouring only 17% of the global number of Picidae species.
The family Picidae, which encompasses wood-
peckers, wrynecks, and piculets contains ca. 216
closely related species, all with strong ties to
forest environments. Trees, snags and logs are
primary substrates providing nesting sites, shel-
ter, and food for the majority of woodpeckers
(Winkler et al. 1995). One striking feature of
woodpeckers is their ability to excavate cavi-
ties in living and dead trees (Winkler & Christie
2002). Due to this “engineering activity”, wood-
peckers have been proposed as key-stone spe-
cies in several communities with large numbers
of secondary cavity nesters (Daily et al. 1993,
Jones et al. 1995, Conner et al. 2004, Martin
et al. 2004, Ojeda 2004). Moreover, facultative
sap-consuming species are also benefited by
their drilling for phloem sap and thus their activ
ities can influence entire community structures
(Blendinger 1999, Schlatter & Vergara 2005).
Through wood excavation activities, including
nest construction and foraging, woodpeckers
play a role in wood decomposition processes
(Farris et al. 2004). It has also been suggested
that woodpeckers function as dispersal vectors
for wood-living fungi (Jackson & Jackson 2004).
It is quite apparent from these relationships that
this avian family plays a significant ecological
role in forest environments and communities.
Woodpeckers occur in all types of forest
and woodland and are found on all continents
except of Australia and Antarctica (Winkler et
al. 2005). Woodpeckers are virtually absent from
treeless landscapes such as desert, tundra and
alpine areas. In addition, most isolated islands
ANN. ZOOL. FENNICI Vol. 43 Woodpeckers: distribution, conservation and research 87
lack woodpeckers. Most woodpeckers are sed-
entary birds and are generally considered to be
relatively poor dispersers (e.g. Higuchi & Koike
1978). Blackburn et al. (1998) conducted a global
analysis and found that body mass of woodpeck-
ers was rather weakly correlated with the size of
their geographic ranges. Blackburn et al. (1998)
found that woodpecker geographic range sizes
decreased with increased woodpecker species
richness and that species living in high latitudes
tended to be more widely distributed. Interest-
ingly, the best predictor of species richness for
woodpeckers was the area of a given geographic
region; smaller regions had more woodpecker
species and more endemics.
Several species of woodpeckers have expe-
rienced dramatic population declines and range
contractions due to habitat loss and degradation
through various human activities (Winkler &
Christie 2002). This applies to large, area-sen-
sitive species such as the ivory-billed wood-
pecker (Campephilus principalis) or imperial
woodpecker (C. imperialis) that depend upon
vast areas of unmanaged forest as well as small
but highly specialised species such as the red-
cockaded woodpecker (Picoides borealis) or
South American piculets with small geographic
ranges. The effects of human activities on wood-
pecker assemblages have been observed at dif-
ferent scales. In a country-by-country analy-
sis, Mikusiński and Angelstam (1997, 1998),
reported negative population trends for most
European woodpeckers and related species rich-
ness to the degree of anthropogenic change for
this group of species. In a study from north-east-
ern Poland, species richness for woodpeckers
was positively correlated with the availability of
dead wood and large deciduous trees (Angelstam
et al. 2002) factors which are closely associ-
ated with the level of naturalness of forest stands.
Strong negative correlation between woodpecker
biomass and density with logging intensity has
been found on Borneo (Lammertink 2004) and
the Malaysian peninsula (Styring & Ickes 2001).
Conner et al. (1975) reported higher diversity of
woodpecker species in uncut versus cut wood-
lands in Virginia.
Human impact on woodpecker populations
encompasses several factors. The most obvi-
ous factor for woodpeckers as well as many
other threatened organisms — is habitat loss.
Logging of vast areas of woodland as well as
their conversion to farmland has been the major
cause of range contractions, regional extinctions,
and population declines both now and histori-
cally (Winkler & Christie 2002). Dependence of
woodpeckers on certain forest characteristics
typical for unmanaged forest (i.e. presence of
large and old trees, high structural diversity,
and presence of large quantities of dead wood)
makes them particularly susceptible to forestry
practices. Short-rotation schemes, selection of
few most productive tree species, even-age struc-
ture, removal of dead wood, active fire suppres-
sion, and replacement of native tree species with
fast-growing exotics apparently degrade habi-
tat for woodpeckers (Angelstam & Mikusiński
1994). Alternatively, in areas with traditional,
low-intensity agricultural practices as is the case
in several regions of Europe, woodpeckers may
be relatively abundant in sparsely forested land-
scapes that contain elements of forest structure
such as large, older trees (Mikusiński & Angel-
stam 1997, 1998). In these areas, the clearance
and intensified use of such semi-open landscapes
poses the main threat for woodpecker species
(Tucker & Evans 1997).
The strong association that woodpeckers dis-
play with forest environments and their sensi-
tivity to structural and compositional changes
in their habitats caused by human action has
been a reason for utilizing woodpeckers in forest
and landscape management (e.g. Angelstam &
Mikusiński 1994, Diaz 1997, Hutto 1988, Jans-
son 1998, Nilsson et al. 2001, Hess & King 2002,
Lammertink 2004, Uliczka et al. 2004). Wood-
peckers may fit different concepts that recom-
mend use of surrogate species in practical con-
servation. Woodpeckers have been suggested as
indicators for forest biodiversity (Mikusiński &
Angelstam 1998, Nilsson et al. 2001). It has been
argued that since several woodpecker species
display a highly specialised selection of resources
that are typical of naturally dynamic forests (e.g.
dead wood, big and old trees) that their pres-
ence may indicate high overall biodiversity. The
empirical evidence supporting this hypothesis is
growing. Mikusiński et al. (2001) demonstrated
a strong positive relationship between the number
of woodpecker species and overall forest bird
88 Mikusiński ANN. ZOOL. FENNICI Vol. 43
species richness at the landscape scale in Poland.
The lesser spotted woodpecker (Dendrocopos
minor) has been found to be a reliable indicator of
the occurrence of avian deciduous forest special-
ists in northern Europe (Jansson 1998, Roberge &
Angelstam 2006). Also tree species diversity was
found to be positively correlated with the number
of woodpecker species observed in one study
performed in south-central Sweden (Angelstam
1990). Martikainen et al. (1998) found high num-
bers of threatened wood-species beetles in areas
with white-backed woodpeckers (Dendrocopos
leucotos) in Finland and Russian Karelia.
Woodpeckers have also been proposed as
indicators of structural diversity and ecological
quality of forest habitats and as umbrella/focal
species for forest and landscape management
(McClelland & McClelland 1999, Derleth et al.
2000, Angelstam et al. 2002, Pakkala et al. 2002,
Lammertink 2004). Hess and King (2002) used
the pileated woodpecker (Dryocopus pileatus)
as focal species of mature forest for conserva-
tion planning in North Carolina. Two species
of woodpeckers have been employed as focal
species in designing a forest habitat network
in northern Italy (Bani et al. 2002). The habi-
tat requirements of the three-toed woodpecker
(Picoides tridactylus) have been used to propose
the quantitative snag targets for the conifer-
ous forest management in Europe (Bütler et
al. 2004). In the state of Mississippi, the suc-
cessful restoration of pine-grassland habitat for
red-cockaded woodpeckers resulted in increased
diversity of bird community (Wood et al. 2004).
Moreover, woodpeckers have been used to pre-
dict the impact of forest management on wild-
life habitats (Cox & Engstrom 2001, Marzluff
2002). Angelstam et al. (2004) proposed several
woodpeckers as focal species for the assessment
of forest habitat networks in Europe applying
habitat suitability modelling. In summary, wood-
peckers are an important systematic group from
forest and landscape management perspectives
because of their sensitivity to forestry and other
anthropogenic impacts on the forest environ-
ment. It remains to be seen whether they will be
used to their full capacity as tools in forest biodi-
versity management in a global perspective.
In this paper I examine and discuss the
global pattern of woodpecker diversity from a
conservation perspective. I also review some
ecological traits and the conservation status of
the entire family Picidae and relate them to the
human driven change in their habitats. Finally,
I present a global overview of the research on
woodpeckers in order to identify the major gaps
in our knowledge.
Material and methods
Analyses in this study were performed at the
global scale. I created a spatially-explicit Geo-
graphic Information System (GIS) database on
the distribution of the breeding ranges of 216
Picidae species, including woodpeckers, wry-
necks and piculets. The source of information on
geographic ranges of species was taken from dis-
tribution maps published in Winkler and Christie
(2002). Spatial analyses were performed using
ArcView 3.2 (ESRI 2000). A set of global maps
provided with this software was used as back-
ground information during the process of digi-
talisation. In addition, information on the global
distribution of major vegetation zones was pro-
vided with this software. The spatial resolution
of the created database corresponded to a grid
with cell dimensions latitude ¥ longitude.
Breeding ranges of all species were manually
digitised into this grid and then summarised spa-
tially using the overlay function in vector format.
Borders between adjacent cells with the same
number of woodpecker species were then dis-
solved. This resulted in a vector-based map with
polygons with values corresponding to 1–25 spe-
cies present.
Information on woodpeckers listed into dif-
ferent threat categories according to IUCN clas-
sification as well as on the numbers of wood-
pecker species present in different countries was
extracted from BirdLife International homepage
in the Search for Species mode (www.birdlife.
org/datazone/species/index.html). The data on
woodpecker’s body size and their diet was assem-
bled from Winkler and Christie (2002).
The literature survey for articles on wood-
peckers has been performed using Wildlife and
Ecology Studies Worldwide database (NISC
2005). I searched for articles that contained
at least one of following words in the title
ANN. ZOOL. FENNICI Vol. 43 Woodpeckers: distribution, conservation and research 89
(translated title): woodpecker(s), wryneck(s),
piculet(s), flicker(s), sapsucker(s), flameback(s),
goldenback(s), yellownape(s). I covered a 20-
year period (1985–2004). All titles and abstracts
(when available) were reviewed and the follow-
ing parameters were recorded in spread sheet
format: species, year of publication, country,
continent, first author and type of study (ecologi-
cal, behavioural, faunistic and biogeographic,
other). Publications concerning two or more spe-
cies of woodpeckers were recorded as “several
species” instead of species names. All references
that appeared two or more times in the database
were manually removed.
The map illustrating the global pattern in spe-
cies richness within family Picidae pointed to
the regions of southeast Asia and northern and
central parts of South America as species rich-
ness hotspots with at least 15 species present at
the regional level (Fig. 1). In particular, hotspots
were found in Myanmar, Thailand, Vietnam,
Cambodia, Malaysia and Indonesia in Asia, as
well as in Brazil, Colombia, Peru, Ecuador and
Suriname in South America. Equatorial Africa
and Central America also had relatively high
global levels of woodpecker species richness.
Among different major vegetation zones with
woody vegetation, moist broadleaved tropical
and subtropical forests had, on average, the high-
est number of woodpecker species (Fig. 2). Most
occurrences of high woodpecker species rich-
ness were found to be in economically develop-
ing countries (Fig. 3).
Ants, other arthropods, fruits and berries
were among the most commonly observed food
items taken by woodpeckers (Winkler & Chris-
Fig. 1. Global pattern of
species richness in family
Tropical and subtr.
moist broad-leaf
forests (1440)
Tropical and subtr.
grasslands, savannas,
and shrublands (1386)
Tropical and subtr.
dry broad-leaf
forests (524)
Temperate grasslands,
savannas, and
shrublands (1065)
coniferous forests
broad-leaf and
mixed forests (1319)
scrub (151)
Deserts and xeric
shrublands (887)
Boreal forest/
Mean number of species
Fig. 2. Mean number of woodpecker species in major
woody vegetation types. Number of 1 ¥ 1 geographic
degree plots representing each type in parentheses.
Bars denote 95% confidence intervals of the mean.
0 5000 10000 15000 20000 25000 30000 35000 40000 45000
Gross Domestic Product in 2003 (USD)
Number of species
Fig. 3. Number of woodpecker species in countries with
different Gross National Product per capita according
to UNEP (2005). Only countries with area > 5000 km
were included (n = 135).
90 Mikusiński ANN. ZOOL. FENNICI Vol. 43
tie 2002; Fig. 4). Clearly, well-described and
known species had much higher numbers of
foods listed. Interestingly, this source indicates
that the diets of 37 species (17%) are virtually
unknown to science.
In total, 25 species of Picidae are listed by
IUCN as threatened or near threatened in the
following categories: CR = 3 species, VU =
7 species, NT = 15 species (Table 1). Almost
half of them (12 species) are considered as
restricted-range species (i.e. with geographic
range < 50 000 km
). Most of the endangered
species occur in areas with generally high wood-
pecker species richness. The percentage of spe-
cies in different size classes in threatened (near
threatened) species and non-threatened catego-
ries is illustrated in Fig. 5. The proportions of
threatened (near threatened) species in differ-
ent size classes were dissimilar from those of
the remaining woodpecker species ( χ
= 11.4,
df = 5, p = 0.044). In particular size class 10–
15 cm (containing mostly piculets) and the larg-
est woodpeckers were relatively more abundant
among threatened (near threatened) species in
comparison with the remaining species. Habitat
loss and degradation due to forest logging and
removal of wood are the major threats indicated
for all listed species. In some cases, hunting
and collecting (e.g. imperial woodpecker), forest
Woodboring beetles
Other arthropod
Fruits & berries
Seeds & nuts
Woodpecker species (%)
Fig. 4. Diet of Picidae according to species accounts in
Winkler and Christie (2002).
Table 1. Species listed by IUNC as threatened or near threatened. Categories: Critically Endangered (CR), Vulner
able (V), Near Threatened (NT). Regions: Africa (AF), Asia (AS), Latin America (LA), North America (NA).
Species Category Region Comments
Rusty-necked piculet (Picumnus fuscus) NT LA restricted-range species
Speckle-chested piculet (Picumnus steindachneri ) VU LA restricted-range species
Tawny piculet (Picumnus fulvescens) NT LA restricted-range species
Ochraceous piculet (Picumnus limae) VU LA restricted-range species
Mottled piculet (Picumnus nebulosus) NT LA
Guadeloupe woodpecker (Melanerpes herminieri ) NT LA restricted-range species
Red-headed woodpecker (Melanerpes erythrocephalus) NT NA
Knysna woodpecker (Campethera notata) NT AF restricted-range species
Stierling’s woodpecker (Dendropicos stierlingi ) NT AF
Sulu woodpecker (Dendrocopos ramsayi ) VU AS restricted-range species
Arabian woodpecker (Dendrocopos dorae) VU AS
Red-cockaded woodpecker (Picoides borealis) VU NA
Chocó woodpecker (Veniliornis chocoensis) NT LA restricted-range species
Yellow-browed woodpecker (Piculus aurulentus) NT LA
Fernandina’s flicker (Colaptes fernandinae) VU LA restricted-range species
Helmeted woodpecker (Dryocopus galeatus) VU LA restricted-range species
Black-bodied woodpecker (Dryocopus schulzi ) NT LA
Andaman woodpecker (Dryocopus hodgei ) NT AS restricted-range species
Guayaquil woodpecker (Campephilus gayaquilensis) NT LA
Imperial woodpecker (Campephilus imperialis) CR LA probably extinct
Ivory-billed woodpecker (Campephilus principalis) CR NA < 50 individuals
Red-collared woodpecker (Picus rabieri ) NT AS
Olive-backed woodpecker (Dinopium rafflesii ) NT AS
Okinawa woodpecker (Sapheopipo noguchii ) CR AS restricted-range species
Buff-necked woodpecker (Meiglyptes tukki ) NT AS
ANN. ZOOL. FENNICI Vol. 43 Woodpeckers: distribution, conservation and research 91
fires and hurricanes (e.g. red-cockaded wood-
pecker) were considered to be additional causes
of population declines.
In total, 1252 articles on woodpeckers pub-
lished between 1985 and 2004 were found in
Wildlife and Ecology Studies Worldwide data-
base. These included 1050 single-species articles
that encompassed 65 species. Table 2 provides
the list of the 15 species with highest numbers of
articles published. I found a great disproportion
in the number of woodpecker species occurring
in Europe and North America and the number of
articles published about these species (Fig. 6).
The global pattern of species diversity in Pici-
dae generally follows the well-known latitudinal
gradient of decreasing species richness from
lower to higher latitudes (Rosenzweig 1995,
Gaston 2000). This trend confirms an earlier
finding at a much more coarse resolution by
Blackburn et al. (1998). Identified hotspots of
woodpecker diversity largely overlapped with
distribution of hotspots of avian species richness
(Orme et al. 2005). In the case of woodpeck-
ers, the prerequisite for high species richness
appears to be presence of vast areas of woodland
with high structural and compositional diversity.
These environments are found mostly in tropi-
cal and subtropical moist broadleaved forests of
< 10 10–15 15–20 20–25 25–30 30–35 > 35
Body length (cm)
Woodpecker species (%)
Not threatened (%)
Threatened (%)
Fig. 5. Body length in globally threatened (near threat-
ened) vs. unthreatened species of woodpeckers.
Table 2. Species with highest number of articles pub
lished between 1985 and 2004 according to Wildlife &
Ecology Studies Worldwide database.
Species Number of articles
Red-cockaded woodpecker
(Picoides borealis) 301
Great spotted woodpecker
(Dendrocopos major) 62
Northern flicker
(Colaptes auratus) 61
Pileated woodpecker
(Dryocopus pileatus) 58
Black woodpecker
(Dryocopus martius) 55
Acorn woodpecker
(Melanerpes formicivorus) 54
Middle spotted woodpecker
(Dendrocopos medius) 40
White-backed woodpecker
(Dendrocopos leucotos) 36
Red-bellied woodpecker
(Melanerpes carolinus) 27
Three-toed woodpecker
(Picoides tridactylus) 24
Red-headed woodpecker
(Melanerpes erythrocephalus) 23
Yellow-bellied Sapsucker
(Sphyrapicus varius) 22
(Jynx torquilla) 21
Downy woodpecker
(Picoides pubescens) 20
Lewis’s woodpecker
(Melanerpes lewis) 16
Fig. 6. Number of articles found in Wildlife & Ecology
Studies Worldwide database (NISC 2005) and number
of woodpeckers in different regions of the world.
92 Mikusiński ANN. ZOOL. FENNICI Vol. 43
south-east Asia and South America (Figs. 1 and
2). Temperate and boreal forest vegetation zones
have a moderate number of species present at
regional level.
In a global perspective, arthropods clearly
dominate diets in Picidae (Fig. 4). However,
woodpecker diets also include vegetable items
such as fruit, nuts, berries and sap. I found a
relatively low proportion of species that spe-
cialised in wood-boring beetles and their larvae
(< 30% of species), the food often perceived
as most typical for woodpeckers. Interest-
ingly, ants seem to be an important ingredient
of woodpecker diets world-wide. An obvious
weakness of this analysis is that the data used
are not quantitative and thus it is difficult to
know the importance of different food items
for lesser-known species. Detailed studies on
food of European woodpecker species indicate
significant regional and seasonal variations in
diets (Cramp 1985). Information on the diet
of the great majority of woodpecker species
comes not from comprehensive foraging stud-
ies or stomach contents analyses but from more
or less incidental observations. Askins (1983)
compared foraging behaviour of 11 species of
woodpeckers in Guatemala, Maryland, and Min-
nesota and found a similar degree of specialisa-
tion in both tropical and temperate species. I
argue that with more knowledge on foraging
habits of woodpeckers we are going to see, not
only a broader spectrum of different food items
utilised by particular species, but also a clearer
pattern of seasonal and regional specialisation
and the importance of animal versus vegetarian
ingredients in diets.
Judging from the proportion of woodpecker
species listed by IUCN (11.5%), the global con-
servation status of this group of birds is not
extreme. Some other broadly distributed groups
of forest birds like parrots (Psittacidae), horn-
bills (Bucerotidae), or nuthatches (Sittidae) have
higher proportions of threatened species (40%,
39% and 27%, respectively), while others like
tits (Paridae) or wrens (Troglodytidae) have
lower proportions of threatened species (5.5%
and 9.2%, respectively). High proportions of
threatened species among parrots and hornbills
may be partly explained by the fact that these
families are nearly exclusive to the tropics where
the fastest habitat destruction is occurring. In
addition, many parrot species are small island
Does the relatively low proportion of offi-
cially listed species mean that woodpeckers as
a group of forest specialists are not particularly
threatened by recent global intensification of log-
ging and other human-related changes affecting
forest environments? A closer look at the geo-
graphic locations of the hotspots of woodpecker
species richness may suggest the opposite. Most
hotspots are located in areas with tropical and
subtropical moist forests as the major vegeta-
tion types. Since the majority of these areas
have been recently subjected to intensive clear-
ing operations the amount and quality of avail-
able woodpecker habitat has declined drastically.
Estimated cumulative forest loss in Malaysia,
Myanmar, Thailand and Vietnam, countries that
harbour the majority of Asian woodpecker diver-
sity hotspots, amount to 39%, 55%, 71% and
68%, respectively (FAO 2000, Pahari & Murai
1999). These figures illustrate loss of original
forest cover but are silent on the structural and
compositional changes in managed forests that
negatively affect the quality of woodpecker habi-
tats (e.g. Lammertink 2004).
Similarly, forest loss in the South American
hotspots is drastic. In the case of Brasilian Ama-
zonia, the tropical forest area remained a largely
intact area until the inauguration of the Transa-
mazon Highway in 1970, but the pace of forest
clearing has been rapid since then (Fearnside
2005). Also, in Colombia, Peru, and Ecuador the
deforestation has been a major land-use change
affecting entire regions (Pahari & Murai 1999).
In an area of over 3000 km
located at the border
between Colombia and Ecuador within one
of the woodpecker diversity hotspots Viña
et al. (2004) estimated with the use of satellite
images the deforestation rate during a 23-yr
period at 43% and 22% on the Colombian and
Ecuadorian sides of the border, respectively.
The establishment of protected areas (PA) in
woodpecker diversity hotspots is not necessarily
securing the maintenance of their natural habi-
tats. In protected areas in south-western Borneo
(Kalimantan) for example, the lowland forest
cover declined by 56% (29 000 km
) from 1985
to 2001 (Curran et al. 2004).
ANN. ZOOL. FENNICI Vol. 43 Woodpeckers: distribution, conservation and research 93
I predict that a clear positive relationship
between the deforestation rate and human popu-
lation density (Pahari & Murai 1999), consid-
ered together with high annual human popula-
tion growth (> 1.5%) in the woodpecker-richest
countries, may lead to a rapid increase in the
number of globally threatened species of Picidae
in the near future. Cincotta et al. (2000) found
that human population growth in global biodi-
versity hotspots is substantially above that of
developing countries in general, and predicted
that human-induced changes negatively affecting
biodiversity are likely to continue in these areas.
Regional extinctions and population contractions
of woodpecker species related to socio-economic
development resulting in forest loss and degra-
dation have been described from temperate and
subtropical zones (e.g. Pettersson 1984, Virkkala
et al. 1993, Jackson 1994, Mikusiński & Angel-
stam 1997, 1998). Since the rapid deforestation
of areas with highest woodpecker diversity in
the tropics is a relatively recent phenomenon, we
have to consider a potentially large unrealised
“extinction debt” (sensu Tilman et al. 1994)
as an important conservation issue in the near
Some features of woodpecker distribution
patterns and biology make them especially prone
to rapid declines due to increase of human popu-
lation. First, woodpecker species occurring in
low latitudes tend to have relatively small geo-
graphic ranges (Blackburn et al. 1998) and there-
fore several range restricted species may be more
susceptible to regional forest loss. Second, many
woodpecker species are specialised foragers that
do not attain high population densities but have
relatively large home-ranges, and therefore a
viable forest population will need relative large
areas with a network of functional habitats. The
situation of the largest Camphepilus species is a
good illustration of this problem. Third, due to
their incompatibility with many forestry prac-
tices concerning the lack of big and old trees, of
dead wood, and the reduction of compositional
and structural diversity, woodpeckers may be
unable to thrive in managed forests and forest
The literature survey on woodpeckers pro-
vided striking results concerning the dispropor-
tionate number of studies performed on species
in temperate versus tropical zones. Fifteen North
American and European species of woodpeck-
ers (7% of Picidae) accounted for 70% of all
single-species articles published in the years
1985–2004. I found a total of 33 articles (less
than 3% of the total) concerning woodpeckers in
Latin America where over half of all woodpecker
species occur. Even if we take into account inac-
cessibility of some local, regional, and national
studies in the database used, the geographic
distribution of woodpecker studies seems to be
far from representative. The lack of knowledge
on diets of 37 species in the most recent and in-
depth global accounts on woodpecker life-histo-
ries provided by Winkler and Christie (2002) is
a good illustration of this problem. What kinds
of studies on woodpeckers are needed to be per-
formed and reported to the international commu-
nity? Most importantly, more reports on wood-
pecker assemblages from the areas identified as
continental or regional hotspots are required.
These should cover the conservation status and
habitat requirements of particular species, their
sensitivity to human impact, and their ecological
role in the ecosystems. Good examples of such
studies are those provided by Styring and Ickes
(2001), Lammertink (2004) and Schlatter and
Vergara (2005).
Winkler and Christie (2002) identified gaps
in our knowledge on life-history traits and the
conservation status of particular species. The
keystone role of woodpeckers as excavators in
communities with secondary cavity users and
with species that feed on sap or invertebrates
exposed by woodpeckers should be assessed.
The knowledge on habitat requirements of such
key-stone species are necessary for conservation
planning in both primeval and managed forests.
It is possible that several woodpecker species in
the low latitudes will be useful focal/umbrella
species. In terms of woodpecker conservation,
it seems that emphasis should be put on larger
species. The large species tend to be more often
threatened (Fig. 5) due to their large area require-
ments, their specialisation on foraging substrate,
and their need for trees with large dimensions for
nesting and roosting.
The woodpeckers are an important part of
forest ecosystems on five continents. This group
includes highly specialised species that are
94 Mikusiński ANN. ZOOL. FENNICI Vol. 43
threatened by the recent and future deforestation
of tropical and subtropical areas. At the same
time our knowledge on the majority of species
is quite limited. I urge all woodpecker research
ers to design and perform studies that can fill
the gaps in our knowledge. Such studies will
not only illuminate our understanding of wood
pecker biology and ecology, but will also con
tribute to the maintenance of forest biodiversity
Many thanks to Martjan Lammertink, Jean-Michel Roberge
and Scott Brainerd for their comments on earlier drafts of
this paper.
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This article is also available in pdf format at
... Most woodpeckers are forest dependent and are considered ecosystem engineers because of their role as excavators of cavities, which are then exploited by a diversity of birds, mammals and insects (Martin et al., 2004;Tremblay et al., 2015). They are considered particularly vulnerable to forest degradation and fragmentation due to their requirement for nesting and foraging trees (Mikusiński, 2006;Lammertink, 2014), and are considered good indicators of general bird diversity (Drever et al., 2008;Kumar et al., 2011). Although different species may have specific habitat requirements, there is evidence that management practices that favor woodpeckers (e.g., tree species richness, presence of old trees, and dead trees) may also benefit bird diversity (Drever and Martin, 2010). ...
... Home ranges of Robust Woodpeckers were smaller with a higher density of dead trees, a resource that this species uses for foraging , excavation of nests , and roosting by occupying old woodpecker nest cavities (unpublished data). Several studies on woodpeckers have shown that standing dead (snags) and decaying trees are important structural attributes in foraging grounds (Villard, 1994;Müller et al., 2009;Nappi et al., 2015;Tingley et al., 2014;Tingley et al., 2020), and that unlogged forests contain a higher amount of these resources (Mikusiński 2006;Lindenmayer et al., 2012). Although our results do not support the latter, since the availability of dead trees did not differ between unlogged and logged forests, they do support the importance of dead wood for this species, since a greater availability of dead trees for foraging, nesting, and roosting resulted in a smaller home range sizes. ...
Understanding the habitat requirements of forest-dependent species, such as woodpeckers, is crucial for effective management strategies in environments facing significant anthropogenic pressures. During 2013–2019, we investigated the variation in the availability of forest structural attributes in two unlogged forests, one forest with active selective logging, and one forest 33 years after it had been logged, in the Upper Paran´a Atlantic Forest in Misiones, Argentina. Then, we studied the selection of the forest attributes by co-occurring populations of Helmeted Woodpecker (Celeus galeatus), Lineated Woodpecker (Dryocopus lineatus), and Robust Woodpecker (Campephilus robustus), along with their correlation to variations in home range sizes of woodpeckers. We found that the density of potential nest trees of the three woodpecker species, the density of potential Helmeted Woodpecker roost trees, the basal area ha-1 of live trees ≥ 30 cm DBH, the density of live large trees ≥ 30 cm DBH, and the volume of dead wood ≥ 30 cm diameter differed among study sites. Home range sizes of Helmeted Woodpeckers decreased with higher potential roost tree density and greater understory bamboo cover. The home range sizes of Robust Woodpeckers decreased with a higher density of dead trees, while home range sizes of Lineated Woodpeckers were not influenced by any of the variables we measured. The three woodpecker species selected diameter classes and tree species for foraging and nesting disproportionally to their availability. Helmeted Woodpecker preferred guatambú (Balfourodendron riedelianum) over other trees for roosting. We recommend considering key attributes associated with smaller home range sizes of the threatened Helmeted Woodpecker in sustainable forest management, as these attributes are also important for the other two woodpecker species. To ensure the availability of the resources selected by these woodpeckers, we recommend increasing the legal minimum cutting diameter to 60 cm for grapia (Apuleia leiocarpa), guatambú, rabo duro (Lonchocarpus leucanthus) and palo de canga (Symplocos uniflora), and to 50 cm for laurel negro (Nectandra megapotamica) and laurel amarillo (N. lanceolata). We found similar quantities of most of the tree resources we measured in unlogged forests and a forest undergoing a natural recovery for 33 years after the cessation of selective logging, and we recommend that more logged forests be restored in this manner.
... O liczebności dzięciołów najczęściej decydują wiek, struktura i bogactwo gatunkowe drzewostanu oraz ilość martwego drewna (Wesołowski & Tomiałojć 1986, Walankiewicz et al. 2002, Kosiński et al. 2018. Wąski zakres tolerancji ekologicznej dzięciołów wykorzystywany jest w ocenie różnorodności biologicznej oraz jakości siedlisk leśnych, szczególnie zasobności drzewostanów w zamierające drzewa lub martwe drewno (Mikusiński & Angelstam 1998, Mikusiński et al. 2001, Wübbenhorst & Südbeck 2003, Mikusiński et al. 2006, Kajtoch & Figarski 2014. Cechą wyróżniającą dzięcioły jest zdolność do drążenia dziupli, które mogą być wykorzystywane jako miejsca gniazdowania przez tzw. ...
... dzięcioł białogrzbiety, spełniają funkcję osłonową w ochronie organizmów związanych z martwym drewnem, np. saproksylicznych chrząszczy, mszaków i porostów (Mikusiński 2006. ...
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Abstrakt: W pracy przedstawiono wyniki inwentaryzacji dzięciołów Picidae przeprowadzonej z użyciem stymulacji głosowej w roku 2020 w centralnej części Lasów Suchedniowskich na Wyżynie Kieleckiej. Cenzusem objęto fragment lasów, głównie jodłowo-bukowych oraz sosnowych, o powierzchni 2940 ha. Zagęszczenie dzięcioła dużego Dendrocopos major oszacowano na powierzchni próbnej o powierzchni 365 ha. Na badanym obszarze za lęgowe uznano sześć gatunków dzięciołów związanych z wnętrzem zwartych lasów mieszanych. Dla każdego z nich oszacowano zagęszczenie krajobrazowe, a dla gatunków stenotopowych także zagęszczenie ekologiczne. Poszczególne gatunki gniazdowały w zagęszczeniach: dzięcioł duży – 0,8 ter./10 ha, dzięcioł czarny Dryocopus martius – 5,1 ter./10 km2 powierzchni całkowitej, dzięcioł białogrzbiety D. leucotos – 0,7 ter./10 km2 powierzchni całkowitej i 0,4 ter./km2 lasów liściastych, dzięcioł średni Dendrocoptes medius – odpowiednio 5,8–6,1 ter./10 km2 i 3,3–3,5 ter./km2, dzięcioł zielonosiwy Picus canus – 3,5 ter./10 km2 i 0,7 ter./km2 i dzięciołek Dryobates minor – 2,0–2,4 ter./10 km2 i 1,2–1,3 ter./km2. Wysoka różnorodność gatunkowa dzięciołów w Lasach Suchedniowskich oraz obecność gatunków stenotopowych, w tym dzięcioła białogrzbietego, wskazuje na zadowalający stan zachowania lasów oraz ich znaczną zasobność w martwe oraz zamierające drzewa. Słowa kluczowe: dzięcioły, Picidae, dzięcioł białogrzbiety, Dendrocopos leucotos, drzewostany jodłowo-bukowe, Lasy Suchedniowskie, liczebność Woodpecker Picidae numbers in the Suchedniowskie Forest. Abstract: The paper presents results of counts of woodpeckers Picidae conducted in 2020 in the central part of the Suchedniowskie Forests located on Kielce Upland. The study area of ca 2,940 ha was covered mainly by fir-beech and pine forests. Sound stimulation was used to detect the presence of woodpeckers. The density of Great Spotted Woodpecker Dendrocopos major was estimated on a survey plot of 365 ha. The density of stenotopic species was calculated per a total study area (landscape density) and per the area of one the preferred forest type (ecological density). We found six species of woodpeckers associated with the interior of mixed forests. The densities of woodpeckers were as follows: GreatSpotted Woodpecker – 0.8 ter./10 ha, Black Woodpecker Dryocopus martius – 5.1 ter./10 km2 of the total area, White-backed Woodpecker Dendrocopos leucotos – 0.7 ter./10 km2 of the total area and 0.4 ter./km2 of deciduous forests, Middle Spotted Woodpecker Dendrocoptes medius – respectively 5.8–6.1 ter./10 km2 and 3.3–3.5 ter./km2, Grey-headed Woodpecker Picus canus – 3.5 ter./10 km2 and 0.7 ter./km2, and Lesser Spotted Woodpecker Dryobates minor – 2.0–2.4 ter./10 km2 and 1.2–1.3 ter./km2. High diversity of woodpeckers in the Suchedniowskie Forests and the presence of stenotopic species, including the White-backed Woodpecker, suggest a relatively good conditions of these stands associated with large amounts of dead wood. Słowa kluczowe: woodpeckers, Picidae, White-backed Woodpecker, Dendrocopos leucotos, firbeech forests, the Suchedniowskie Forests, numbers
... Woodpecker resources can be defined both in terms of suitable nesting locations (cavities that provide protection from the elements and from predation) and food availability (Bonnot et al., 2009;Conner et al., 1994;Mikusiński, 2006). These resources have been shown to be directly linked to woodpecker nest site location and the distance they travel during foraging bouts (Conner, 1981;Lorenz et al., 2016;Wiktander et al., 2001). ...
... In terms of food availability, an increase in population size of one insect taxa can sometimes be mirrored by similar increases in other insect taxa, usually correlating with changing environmental conditions and available resources (Brooks et al., 2012;Koivula, 2011;Lehnert et al., 2013). (Mikusiński, 2006). Therefore, woodpeckers and secondary cavities nesters may have an interwoven relationship with food resources as the woodpecker acts as an ecosystem engineer by constructing cavities in areas with increased foraging opportunities for both taxa. ...
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Abstract Ecosystem engineers alter, and can be influenced in turn by, the ecosystems they live in. Woodpeckers choose foraging and nesting sites based, in part, on food availability. Once abandoned, these cavities, particularly within areas of high forage, may be crucial to secondary cavity‐nesting birds otherwise limited by cavities formed through decay. Our study examined factors that influence the nesting success of primary cavity nesters and the subsequent impact on secondary cavity‐nesting birds. Using 5 years of point count data, we monitored the outcomes of cavity‐nesting birds in South Texas. We used logistic‐exposure models to predict daily survival rates based on cavity metrics and used woodpecker foraging trends and insect surveys to determine if nesting where woodpeckers actively forage benefits secondary cavity‐nesting birds. Both woodpeckers and secondary cavity nesters shared predictors of daily survival; nests were more successful in cavities with small openings in minimally decayed trees. All secondary cavity nesters had higher probabilities of success when nesting in an abandoned woodpecker cavity, opposed to ones formed by decay. Woodpeckers tended to forage in areas with higher‐than‐average levels of the insect orders Coleoptera, Hymenoptera, and Orthoptera, and secondary cavity nesters had higher rates of success when nesting in these areas. Our results suggest abandoned woodpecker cavities may be constructed in a way that directly benefit secondary cavity nesters. Additionally, we suggest an interplay between these ecosystem engineers, food availability, and secondary cavity nesters: Woodpeckers engineer superior nesting cavities in areas where food is more abundant, and the resultant cavities in areas of high forage may benefit local secondary cavity nesters. Our findings indicate that there is still much to be explored in the role of ecosystem engineers, and how they influence local communities on multiple trophic levels.
... At the population level, most woodpecker species are neither too rare nor too common, largely resident, and fairly conspicuous due to their loud calls, colourful plumage and tree drumming behaviour (Drever et al. 2008;Mikusiński et al. 2001). This makes woodpeckers easily identifiable across their range so amateur birdwatchers and field personnel may be trained in their species-level identification (Angelstam and Mikusiński 1994;Mikusiński 2006). All these characteristics make woodpeckers easier to monitor and hence effective candidates for forest indicator species from a management point of view. ...
... The hotspots of woodpecker diversity are in the tropical and sub-tropical countries of South-east Asia, Equatorial Africa and South and Central America. Yet more than half of the published studies on woodpeckers are from North America and Europe, which account for only 17% of Picidae species (Mikusiński 2006). Woodpeckers are largely under studied in tropical countries, even those where forests face high levels of threat. ...
Full-text available
Woodpeckers (family Picidae) are a specialised group of insectivores that are sensitive to forest degradation and fragmentation. We evaluated the woodpecker taxa as potential indicators of habitat quality and forest bird diversity in temperate moist hardwood forests (1500–2400 m asl), a threatened biome in the Western Himalaya. 74 forest sites, selected to represent a gradient of anthropogenic use, were surveyed for birds, vegetation structure and proportion of land under dense forest in the surrounding landscape. Individual woodpecker species were observed to quantify their foraging niche preferences. We found that forest sites with higher woodpecker richness were also rich in all other bird species. Further, the richness and abundance of woodpeckers and all other birds were affected by similar habitat variables. Four out of the eight woodpecker species occurring in the study area were found to fit our habitat models suitably, with canopy cover, tree density, and forest proportion proving to be important variables. Behavioural observations showed that the same four woodpecker species significantly preferred larger and taller trees for foraging. Given the difficulty of directly monitoring forest characteristics and total bird diversity over large landscapes, consistent monitoring of sensitive woodpecker species can provide answers to both. Further, as woodpeckers are moderately common and conspicuous birds, areas with high woodpecker diversity can be easily identified and prioritised for conservation.
... Nonetheless, aside from a few exceptions, the foraging ecology of woodpeckers in tropical/subtropical regions that harbour a high picid diversity has not been studied in great detail (Mikusiński 2006). Parts of Asia, like north-western India that supports 15 species (of which ca. 12 are sympatric), are well suited to assess foraging niche partitioning among woodpeckers (Styring 2002). ...
... In the area we identified eight study sites representing the structural and compositional diversity of the forests, and established transects of 2.0 km length. At each site we conducted field surveys for woodpeckers by traversing the marked transects on foot at a steady pace of 1-2 km/h, with each site being surveyed 20 times during 2006(Kumar et al. 2011, 2014. ...
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The sub-Himalayan dipterocarp forests of subtropical region in northwest India support 17 species of woodpecker. From a conservation perspective it is necessary to assess the ecological requirements of woodpeckers in these biologically diverse landscapes, which are experiencing habitat modification and decline in some woodpeckers. We studied the foraging niche differentiation among ten sympatric woodpeckers in the sub-Himalayan forests of northwest India: Dendrocopos canicapillus, D. macei, Picus chlorolophus, P. xanthopygaeus, P. canus, Dinopium shorii, D. benghalense, Chrysophlegma [Picus] flavinucha, Chrysocolaptes lucidus, and Mulleripicus pulverulentus. We examined the foraging site preferences of individual woodpecker species and explored the role of inter-specific differences in foraging behaviour as a possible mechanism for their coexistence. Observations on foraging woodpeckers were taken vis-a-vis the following niche dimensions: diameter of the foraging tree, height of the foraging bird, type of substrate on which it was foraging, vertical position with respect to canopy, condition of the forage tree and condition of the substrate. Distinct preferences were evident among species in their foraging tree diameters, foraging heights, vertical positions, and choice of substrate type, while preference for dead substrates was not an important distinguishing factor. Species that overlapped in one dimension generally segregated along other dimensions. Niche segregation in forage tree diameter was associated with body weight. Based on their overall niche overlaps species could broadly be differentiated into groups according to body size viz. small (3 species), medium (3 species), large (3 species), and very large (1 species). Our study demonstrates that differentiation in foraging may be a likely mechanism for coexistence of sympatric woodpeckers in this region. Given that larger species prefer larger substrates, removal of mature trees could affect their abundance, and homogenisation of stand structure could lead to impoverishment of woodpecker diversity.
... A lack of published radiotracking studies of woodpeckers from subtropical or tropical areas reflects the general paucity of ecological studies of woodpeckers from these regions. This stands in contrast to high woodpecker species diversity in (sub)tropical regions and high pressures from deforestation and logging that woodpeckers there experience, and the resulting clear need for detailed information on their ecological requirements (Mikusiński 2006, Lammertink 2014, Vergara-Tabares et al. 2018. Moreover, few radio-tracking studies of woodpeckers, anywhere in the world, have reported on tracking of multiple species in the same study sites, even though such work will yield the clearest understanding of interspecific differences in area use in response to forestry practices. ...
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Home range mapping studies of birds inform about area requirements and responses to land management as reflected by home range sizes and by resource selection within home ranges. Tracking studies of woodpeckers (Picidae) so far have been concentrated in temperate regions. In the subtropical Atlantic Forest of northeast Argentina, we assessed interspecific differences in home range sizes in old-growth forests and selectively logged forests of Helmeted Woodpecker (Celeus galeatus, a globally threatened species), Lineated Woodpecker (Dryocopus lineatus), and Robust Woodpecker (Campephilus robustus). Helmeted Woodpecker had larger breeding home ranges in selectively logged forests, averaging 105 ± 39 ha for pairs, versus 60 ± 13 ha in old-growth forests. Lineated Woodpecker breeding home ranges of pairs averaged 56 ± 22 ha, and those of Robust Woodpecker 43 ± 22 ha, with no differences between forest types. Helmeted Woodpeckers had an unusual separation between the home ranges of males and females in breeding pairs, with a mean area overlap of only 8% ± 9% near the nest tree, resulting in large home ranges for pairs. Helmeted Woodpecker and Robust Woodpecker individuals that were followed into the post-breeding stage had marked expansions of their home range sizes relative to breeding home ranges. To place our findings with Atlantic Forest woodpeckers in perspective we reviewed whether woodpecker home range sizes increase with latitude and body mass globally. For 29 populations of 22 woodpecker species, a power regression model with these factors explained 24.1% of variation in breeding home range sizes, with 17.1% of variation explained by latitude alone. Woodpecker species with larger home ranges than predicted values were three species of North American woodpeckers of coniferous forests, as well as the Helmeted Woodpecker. Our results of smaller home ranges in old-growth forests for the Helmeted Woodpecker affirm an association of this species with such forests. We urge the conservation of the few remaining tracts of old-growth Atlantic Forest and more restoration of logged forests to mature conditions.
... El carpintero negro Campephilus magellanicus es un ave que pertenece al orden Piciforme La familia Picidae abarca a más de 216 especies en el mundo, todas vinculadas con ambientes boscosos (Mikusiński 2006). Estas aves trepan árboles, anidar en ellos y extraen larvas de insectos de la madera (Fuchs et al. 2007) y han sido reconocidos como únicos por sus especializaciones morfológicas para excavar en árboles (Benz et al. 2006). ...
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The Black woodpecker ( Dryocopus martius , L. 1758) is the largest primary cavity digger in western Europe. Its cavities represent an essential microhabitat for many other forest species but the landscape factors that influence the cavity digging by the Black woodpecker are poorly known and rarely been quantified at the scale of the vital range. We used cavity maps by foresters and naturalists to build a large (2689 cavity bearing trees) database distributed over 11 sites in France. Based on this and on a set of pseudo-absence data, we analysed the effects of stand composition and landscape features at three different scales around each plot corresponding to a forest management unit (10ha), the core (100ha) and extended (250ha) vital range scales. We showed that landscape shape index and forest composition (mixed forests) had significant effects but that the magnitude varied across the three scales. The Black woodpecker tend to avoid conifer-dominated stands to dig cavities, and also prefer homogeneous forest landscapes with low edge densities. These results vary with precipitation and slope with stronger effects of landscape in wetter climates and higher slopes. Although forest management rarely modifies the landscape in western Europe, a better understanding of the features that influence cavity digging by the Black woodpecker may help to better integrate their conservation at the management planning scale. Our results also show the importance to maintain mixed broadleaf-conifer forests as well as connected forest landscapes to favour features that benefit forest biodiversity at the large scale.
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We documented the foraging activities of woodpeckers on selected trees in an established conservation-oriented management study in five oak-dominated forests in Hungary. We examined the tree species preference of woodpeckers as a group and the impact of specific tree characteristics on the habitat use of woodpeckers. We estimated the percentage of visible foraging signs on the trunks and upper limbs of selected trees through the winter and early spring of 2019-2020. Based on the Jacobs' index, woodpeckers preferred oak species for foraging and most foraging signs were on limbs rather than trunks. Foraging signs on trunks were more frequent on those of larger diameters and greater heights. It was also found that the lower the tree, the greater the effect of its diameter on the occurrence of signs.
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The Magellanic Woodpecker (Campephilus magellanicus) is a poorly known species endemic of the Austral Temperate Forests of South America, where it is a potential keystone habitat modifier. Here, I summarize data on the social and breeding biology of this woodpecker, based on 22 active nests located from 1998–2002 in forests from northwestern Argentine Patagonia. Woodpeckers normally traveled in pairs or family parties. In late Austral winter, one to three cavities were selected for completion at each territory. Breeding occurred between mid- to late spring and early to mid-summer, and took about 65 days. Monogamous parents shared duties in nest excavation, incubation and young rearing. Egg length (SD) measured 34.130.79mmand egg breadth 23.910.67 mm, and incubation took roughly 20 days. Nestlings were altricial and remained at the nest for about 45 days. Clutch size was one, occasionally two eggs, and one nestling was produced at all successful nests. Young remained with their family group for up to 2 years or more, and were fed by adults, who normally bred every second year. Nest re-use, nest predation and helpers at the nest were not recorded. Holes were placed (SD) 8.843.71m high and were 32.35.32cmdeep. Entrances (SD) were 8.920.46cmwide and 15.592.54cmhigh and mostly oval in shape. Peculiarities of the breeding biology and social behaviour of this species are discussed in the light of patterns common to picids, especially Campephilus spp.
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In the Monte desert of South America, the overall supply of water and food for birds decreases in the dry, sold season (June through September). During this period the White-fronted Woodpecker (Melanerpes cactorum) drills holes in the trunks and branches of Prosopis flexuosa and feeds on the exuded sap. Other bird species, diverse in ecological attributes and taxonomic affinities, take advantage of this resource which otherwise would be rarely available. Sap is a major constituent of the diet of the White-fronted Woodpecker and 11 other bird species, and sap feeding comprises between 16% to 83% of foraging observations made during June and July. Aggression by White-fronted Woodpeckers significantly reduced the time smaller bird species spent feeding on sap, indicating that White-fronted Woodpeckers actively compete for this resource. Other bird species profit from having access to a resource rich in water and sugar.
Woodpeckers are a widespread and popular family of birds. They inhabit virtually all forest and woodlands of the tropics, subtropics and temperate zones. This comprehensive guide to the woodpecker group should aid identification by plumage and voice, and gives scientifically accurate information about their habitats. It covers 198 species worldwide, apart from Australia.
The pileated woodpecker (Dryocopus pileatus) is of special interest to wildlife managers; it requires large trees for nesting, and its abandoned excavations are used by many birds and other small animals for nesting, roosting, hiding, and feeding. Prior to our study, little had been published on pileated woodpecker habitat in the northern Rocky Mountains. From 1973 through 1995, we located nest and roost trees of pileated woodpeckers in northwestern Montana forests dominated by western larch (Larix occidentalis) and Douglas-fir (Pseudotsuga menziesii). Nests (113 in 97 trees) were in western larch (n=52), ponderosa pine (Pinus ponderosa, n=18), black cottonwood (Populus trichocarpa, n=15), trembling aspen (Populus tremuloides, n=7), western white pine (Pinus monticola, n=3), grand fir (Abies grandis, n=1), and Douglas-fir (n=1). Nest-tree diameter-at-breast-height averaged 73 cm, and height averaged 29 m. Roost trees (n=40) were similar to nest trees, but had more cavity entrances and higher basal area of surrounding forest. Nest trees and roost trees typically were snags (81% and 78%, respectively) with broken tops (77% in both). Old-growth stands containing western larch were common nesting sites for pileated woodpeckers. Old-growth ponderosa pine, black cottonwood, and trembling aspen were locally important, but their distribution was more restricted. Compared to other nest-tree species in Montana, undecayed larch wood is hard, making excavation difficult for woodpeckers. Heartwood decay, which softens the wood, becomes more prevalent as a forest matures and was characteristic of western larch nest trees. In the northern Rocky Mountains, the pileated woodpecker has been used too broadly and simplistically as a management indicator of old growth. A more realistic strategy would nurture western larch old growth, defined ecologically, as an indicator of high-quality nesting habitat for pileated woodpeckers. Large trees, logs, snags, carpenter ants (Camponotus spp.), and heartwood decay are intrinsic components of 'healthy' old growth that sustains pileated woodpeckers.
The ability to predict the patch occurrence of a deciduous forest bird guild through an indicator species was tested using published data. Except for the indicator, the Long-tailed Tit (LT), the guild consisted of the Marsh Tit (MT), the Blue Tit (BT) and the Lesser Spotted Woodpecker (LW). All single species, as well as the entire guild, occurred more often like LT than expected by chance. Over all, 86% of the guild occurred in concurrence with LT. When LT was present the mean number of guild species was 3.52 and when absent 0.79. The occurrence pattern of the guild did not change when one multi-sampled region in Sweden was compared with patches scattered throughout four countries, whereas concurrence was more significant in studies from the boreal zone than from the nemoral zone. The guild showed a nested structure in patch occupancy, where the occurrence of LW strongly predicted that of the entire guild. However, because of the better overall statistical agreement in relation to the guild, the Long-tailed Tit, or rather, its known thresholds regarding habitat fragmentation, are suggested as suitable indicators for the occurrence of this guild.
Interactions between organisms are a major determinant of the distribution and abundance of species. Ecology textbooks (e.g., Ricklefs 1984, Krebs 1985, Begon et al. 1990) summarise these important interactions as intra- and interspecific competition for abiotic and biotic resources, predation, parasitism and mutualism. Conspicuously lacking from the list of key processes in most text books is the role that many organisms play in the creation, modification and maintenance of habitats. These activities do not involve direct trophic interactions between species, but they are nevertheless important and common. The ecological literature is rich in examples of habitat modification by organisms, some of which have been extensively studied (e.g. Thayer 1979, Naiman et al. 1988).