A taxonomic, geographic, and temporal assessment of local birds in Portuguese ornithological collections

Bruna S. Santos; Luis M. P. Ceríaco

doi:10.3897/nhcm.2.165142

Research Article

A taxonomic, geographic, and temporal assessment of local birds in Portuguese ornithological collections

Bruna S. Santos^‡§|, Luis M. P. Ceríaco^‡|

‡ Universidade do Porto, Vairão, Portugal

§ Universidade do Porto, Porto, Portugal

| BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal

Corresponding author: Bruna S. Santos ( brunascesantos@gmail.com )

Academic editor: Marcelo Duarte

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Santos BS, Ceríaco LMP (2025) A taxonomic, geographic, and temporal assessment of local birds in Portuguese ornithological collections. Natural History Collections and Museomics 2: 1-96. https://doi.org/10.3897/nhcm.2.165142

Abstract

Since the birth of modern science, natural history collections (NHCs) have constituted the world’s largest repositories of long-term biodiversity data, making them essential to modern-day research. As part of an ongoing project aiming to review the Portuguese zoological collections in the country’s NHCs, we collected and analyzed bird data from its three major museums and two smaller collections to study coverage and representation of the taxa known from Portugal. Our results show that the collections are not taxonomically, geographically, or temporally complete and present considerable biases. Approximately 90.5% of birds are represented in the collections, but 85% are represented by fewer than 50 specimens. Geographic representation is also biased, as the country’s NHCs cover 41.7% of its territory. Thirty-one percent of the species are considered threatened or regionally extinct, yet account for only a little over one-fifth of the specimens extant in the collections. Here we present a review of the status of Portuguese bird collections, as well as a call to attention to the potential impacts on research resulting from such biases and subsequent gaps.

Key words:

Birds, collectomics, history of science, natural history collections, ornithology, specimen collecting, taxonomy

Introduction

Natural history collections (NHCs) have been the world’s repositories of biodiversity data since the 18^th century. As species disappear faster than we can describe them (Ceballos et al. 2015, 2017) and new scientific questions arise, NHCs have become unparalleled data sources and windows into the past (Holmes et al. 2016). Even so, these institutions have been dealing with several challenges, such as budget cuts (Culotta 1992; Dalton 2003; Froelich 2003; Andreone et al. 2014; Salvador and Cunha 2020) and severe gaps in the training of specialized personnel since at least the 1990s (Vivo et al. 2014; Bezerra 2015; Kemp 2015; Orr et al. 2020; Wheeler 2020). Collections without proper structure and adequate maintenance are at risk of being lost (Ceríaco et al. 2021), and in the current information era, critical data are being lost and underutilized, as specimens and their associated metadata are not being digitized in a timely manner (Peterson and Navarro-Sigüenza 2003). Although their purpose may have expanded with time, the main mission of NHCs continues to be the study, collection, preservation, and accessibility of biological specimens (Winker 1996; Andreone et al. 2022).

Paradoxically, while NHCs are once again being recognized as important research assets, the numbers of voucher depositions have hit historic lows in recent decades, halting the temporal, geographic, and taxonomic scope of the collections (Winker 1996; Gardner et al. 2014; Troudet et al. 2018; Buckner et al. 2021; Thompson et al. 2021). Rohwer et al. (2022) reported a decline of more than 50% in new accessions of amphibians, reptiles, birds, and mammals in 245 collections worldwide from 1965 to 2018. A change in societal perceptions around the collecting of live specimens has contributed to the upsurge of non-invasive sampling methods (Pauli et al. 2010; Carroll et al. 2018; Balázs 2020; Schilling et al. 2022; Byrne 2023) and observation-based occurrences (Sullivan et al. 2009; Heberling and Isaac 2018; Speed et al. 2018; Petersen et al. 2021). However, as noted by several authors, without physical specimens publicly available, replication is put at risk and research becomes impoverished (Krell and Wheeler 2014; Rocha et al. 2014; Turney et al. 2015; Ceríaco et al. 2016; Monckton et al. 2020; Thompson et al. 2021; Nachman et al. 2023).

As some of the most iconic and aesthetically appreciated groups of animals, birds have figured in NHCs since their early days (Farber 1980). As part of the initial global effort to systematically catalog and map biodiversity, bird specimens have traditionally been used as the basis of taxonomic studies—from descriptions of species, subspecies, and varieties to external morphological comparisons between populations—and to present the physical evidence of geographic distribution (Mayr and Ashlock 1991; Farber 2000; Joseph 2011). Among vertebrate collections, ornithological specimens are probably those most often used in long-term studies, whose results have become textbook examples of the importance and potential uses of scientific collections (Winker 2005). Winker (2005) noted that, as birds are excellent bioindicators, ornithological specimens are “biological filter papers” that encapsulate many of the conditions of the environment where and when the specimen was collected. Perhaps the most famous example is the study on how the synthetic pesticide DDT (dichlorodiphenyltrichloroethane) affected the thickness of bird eggshells. This correlation was first pointed out by the North American environmentalist Rachel Carson (1907–1964) in her famous book “Silent Spring” (Carson 1962) and was later supported by studies conducted by Hickey and Anderson (1968), Blus et al. (1972), and Cooke (1973) using historic and modern eggshell collections. These studies, for which ornithological collections played a critical role, eventually contributed to the ban of DDT use in the United States in 1972. A more recent example is the study on black carbon atmospheric pollution by DuBay and Fuldner (2017), in which the authors quantified the presence of black carbon in the feathers of thousands of bird specimens spanning several decades. It is not by chance that the recent concept of the “extended specimen” was coined and primarily popularized using bird specimens (Webster 2017).

A wide array of new research uses for birds and other biological specimens is rapidly developing with recent technological advancements. Natural history specimens are now being used in ways their original collectors never imagined, tackling questions on contamination (Thompson and Furness 1989; Thompson et al. 1992, 1993; Monteiro and Furness 1997; Rocque and Winker 2005), climate change (Gardner et al. 2011; Johnson et al. 2011; Bates et al. 2022), infectious diseases (Thompson et al. 2021; Keck 2023), and parasites (Fecchio et al. 2019; Harmon et al. 2019; Wood et al. 2023). The Smithsonian Institution is home to the Feather Identification Lab, which uses the collection to identify birds and other wildlife involved in aircraft strikes. It has provided answers in high-profile cases such as the “miracle on the Hudson River” (Marra et al. 2009), as well as in other areas (Dove and Wickler 2016). In this context, it is sensible to mention the scientific importance of ornithological specimens other than study skins or taxidermy mounts. Eggshell and nest collections, despite usually being less “visible” or commonly known to many users of collections, are an unparalleled source of data for a wide range of studies, as shown historically (Hickey and Anderson 1968; Blus et al. 1972; Cooke 1973) and more recently (Cassey et al. 2010; Maurer et al. 2012; Russell et al. 2013; Grealy et al. 2019; Marini et al. 2020; Bates et al. 2023).

A significant part of scientific research is dependent on museum specimens. Our knowledge of the current anthropogenic impacts on biodiversity is limited by the lack of biological data collected within this timespan (Meineke et al. 2018). Joseph (2011) highlighted this issue in the case of birds and how the incomplete nature of NHCs is hindering research by not providing the specimens needed to answer modern questions. Despite the known importance of NHCs and the need for their completeness, gaps have been reported worldwide, as early as the 1960s in the United States and Canada (Banks et al. 1973; Clench et al. 1976) with surveys conducted from 1966 onward, and in Australia and New Zealand (Gill 2006). Brazilian bird collections were assessed twice in a 10-year period (Aleixo and Straube 2007; Fontana et al. 2017), and although improvements were reported for the overall state of the collections in the most recent assessment, shortcomings such as the lack of specialized personnel and access to digital data were still not fully addressed.

Roselaar (2003) published an inventory of major European bird collections that lists seven ornithological collections in Portugal: the “Lisbon/Lisboa (MB),” formerly known as Museu Bocage and currently the Museu Nacional de História Natural e da Ciência (MUHNAC) in Lisbon; the “Lisbon/Lisboa (CZ/IICT),” currently the collections of the Instituto de Investigação Científica Tropical (IICT) housed in MUHNAC; “Coimbra (MZCoimbra),” nowadays the Museu da Ciência da Universidade de Coimbra (MCUC), in Coimbra; the “Porto Museu de História Natural – Zoologia (MZP),” now the Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP), in Porto; the “Cruz Quebrada-Dafundo Aquário Vasco da Gama,” currently the Aquário Vasco da Gama (AVG) in Dafundo; and those on the Portuguese archipelagos of the Azores and Madeira, respectively the Museu Carlos Machado (MCM) in Ponta Delgada, Azores, and the Museu Municipal do Funchal (História Natural) (MMF), in Funchal, Madeira. Of these, the first three are classified within the “A-list,” which “includes data from collections which include c. 4,000 or more bird skins for scientific study, or, for those which have less, have important numbers of skeletons, anatomical specimens, or other bird items,” while the latter are listed within the “B-list,” which encompasses the “smaller collections (…) which have fewer than c. 4,000 skins, or fewer than c. 5,000 bird items, or of which no exact information could be obtained because the questionnaire was not returned, even after repeated asking.” Using this classification and taking exclusively into account the number of Portuguese specimens, all recognized Portuguese collections would be classified under the “B-list.”

The history of MUHNAC and MCUC can be traced back to the 18^th century, when they began, respectively, as the Royal Cabinet of Natural History of Ajuda and the natural history cabinet of the Faculty of Philosophy of the University of Coimbra (Ceríaco 2021). However, while MCUC’s collection still holds specimens from its early days, MUHNAC’s zoological specimens were destroyed in the fire that devastated the museum in 1978—thus all its current ornithological holdings date from after that time or were donated later. In contrast, the collections of MHNC-UP, AVG, MCM, MMF, and IICT are more recent, with the first three being established in the second half of the 19^th century and the latter two in the first decades of the 20^th century. Currently, all these institutions hold relevant, even if small, ornithological research collections. Smaller collections, usually not accessible for research purposes, also exist in Portuguese high schools and private institutions.

Within Europe, Portugal is globally recognized as a hotspot of bird diversity. Currently, 327 species of birds, belonging to 204 genera and 23 families, are formally listed as either breeding, wintering, or migratory in Portugal (Meirinho et al. 2014; Equipa Atlas 2018; Almeida et al. 2022; Equipa Atlas 2022). This diversity is partly explained by the fact that Portugal belongs to the Iberian Peninsula biodiversity hotspot (Myers et al. 2000; Rosso et al. 2017), as well as being in the East Atlantic Flyway migratory corridor (BirdLife 2010). Regarding conservation, most species occurring in Portugal are currently classified as Least Concern according to IUCN guidelines, but 91 are threatened (Table 2). Eleven species are deemed regionally extinct for at least one population phenology (Table 2). Some species present two conservation assessments, as the red book of birds for mainland Portugal (Almeida et al. 2022) evaluated populations of different phenologies for the same species. Twenty-eight bird taxa are classified as game species by Portuguese legislation (Portaria n.° 100/2021, Ministry of Environment and Climatic Action, available at https://diariodarepublica.pt/dr/detalhe/portaria/100-2021-163012629).

Despite the interest that birds raise in society and the scientific community, the status of Portuguese collections holding birds that occur in Portuguese territory has yet to be formally assessed. As part of an ongoing project to review the specimens of Portuguese vertebrate taxa available in the main Portuguese natural history institutions (see Santos et al. 2024; Santos and Ceríaco 2025), we here assess the taxonomic, geographic, and temporal representation of Portuguese ornithological specimens in national collections. This work aims to answer the following questions: 1) Are all bird taxa occurring in Portugal represented in Portuguese collections? 2) How complete is the geographic coverage of such collections? 3) Do the current collections present temporal trends allowing them to be considered good temporal series? Furthermore, we want to understand how species of conservation concern and game species, as well as eggs and nests, are represented in collections.

Materials and methods

To assess the current situation of Portuguese birds housed in Portuguese NHCs, we consulted the collections of the country’s three main natural history museums—the Museu Nacional de História Natural e da Ciência (MUHNAC, Lisbon), the Museu da Ciência da Universidade de Coimbra (MCUC, Coimbra), and the Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP, Porto)—all part of large public universities, as well as the Aquário Vasco da Gama (AVG) collection and the Museu de História Natural do Funchal (MMF). The small IICT collection is here treated as part of MUHNAC, as its holdings were integrated into the institution in 2015. Despite its colonial (i.e., non-European) focus (see Monteiro et al. 2014, 2016, 2017), the former collection of the IICT includes a very small set of Portuguese birds assessed here, many of which were collected and donated by the first director of the institution, Fernando Frade (1898–1983). The collection of the Museu Carlos Machado (MCM, Ponta Delgada, Azores) was not assessed for this paper. Following the methodology already presented in previous studies on Portuguese herpetology and mammalogy collections (see Santos et al. 2024; Santos and Ceríaco 2025), for each collection, all available metadata associated with existing specimens were gathered, namely taxonomic identification, collecting date, and locality data. Information was compiled based on the existing internal databases, entry catalogs, and in situ review of the specimens when required. Data were standardized according to Darwin Core standards (Darwin Core 2022), and locality information was, when possible, georeferenced following Chapman and Wieczorek (2020).

Figure 1.

Bird collections of the Museu Nacional de História Natural e da Ciência (MUHNAC, Lisbon) (A), the Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP, Porto) (B), the Museu da Ciência da Universidade de Coimbra (MCUC, Coimbra) (C), and the Aquário Vasco da Gama (AVG, Dafundo) (D). Photos by Bruna S. Santos (A, D), Luis M. P. Ceríaco (B), and Marta Bento (C).

Taxonomy and nomenclature follow the most recent Portuguese atlas of breeding birds (Equipa Atlas 2022), the atlas of wintering and migrant birds (Equipa Atlas 2018), the atlas of marine birds (Meirinho et al. 2014), and the red book of birds of continental Portugal (Almeida et al. 2022). Table 1 presents the list of 327 species regarded as occurring in the country according to these references. The collections also include specimens of rare and accidental migrant species (Table 3), as well as collections of eggs and nests (Table 4). These were not included in the overall assessment and are described and analyzed separately below (see subsections “Rare or accidental species” and “Eggs and nests,” respectively). Geographic representativeness of the collections was evaluated through the mapping of extant specimens of the different taxa in comparison with their known distribution range. Species distributions were based on the maps made available through the Portuguese Atlas of Breeding Birds (Equipa Atlas 2022) and the Portuguese Atlas of Wintering and Migrant Species (Equipa Atlas 2018). Following their methodology, species occurrence data in continental Portugal were mapped based on the 10 × 10 km ETRS89/LAEA Europe grid projection. Conservation status follows Almeida et al. (2022), and the legal classification of game species was retrieved from the corresponding Portuguese legislation, Portaria n.° 100/2021.

Table 1.

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Taxonomic diversity of birds occurring in Portugal and corresponding number of existing specimens for each species in the consulted museums/collections. Asterisks (*) denote introduced species. Museums: AVG—Aquário Vasco da Gama; MMF—Museu de História Natural do Funchal; MCUC—Museu da Ciência da Universidade de Coimbra; MHNC-UP—Museu de História Natural e da Ciência da Universidade do Porto; MUHNAC—Museu Nacional de História Natural e da Ciência.

Taxa	AVG	MMF	MCUC	MHNC-UP	MUHNAC	TOTAL
Galliformes
Common quail Coturnix coturnix	–	9	6	4	8	27
Red-legged partridge Alectoris rufa	–	5	8	9	31	53
Ring-necked pheasant Phasianus colchicus *	–	–	–	–	2	2
Grey partridge Perdix perdix	–	–	5	2	–	7
Western capercaillie Tetrao urogallus	–	–	–	–	–	0
Anseriformes
Black swan Cygnus atratus	–	–	–	–	–	0
Mute swan Cygnus olor	–	–	–	–	–	0
Greylag goose Anser anser	–	–	4	3	–	7
Common scoter Melanitta nigra	3	1	9	1	2	16
Red-breasted merganser Mergus serrator	2	–	2	2	–	6
Egyptian goose Alopochen aegyptiaca *	–	–	–	–	–	0
Common shelduck Tadorna tadorna	1	1	4	3	1	10
Ruddy shelduck Tadorna ferruginea	2	2	–	–	–	4
Marbled duck Marmaronetta angustirostris	3	–	1	–	–	4
Red-crested pochard Netta rufina	–	–	–	–	–	0
Common pochard Aythya ferina	–	2	9	4	1	16
Ferruginous duck Aythya nyroca	1	–	5	–	–	6
Tufted duck Aythya fuligula	3	–	8	6	3	20
Garganey Spatula querquedula	3	–	4	–	–	7
Northern shoveler Spatula clypeata	2	–	9	1	1	13
Gadwall Mareca strepera	–	1	–	1	–	2
Eurasian wigeon Mareca penelope	4	1	7	5	–	17
Mallard Anas platyrhynchos	2	3	4	3	5	17
Northern pintail Anas acuta	1	2	9	–	–	12
Eurasian teal Anas crecca	–	5	6	5	5	21
Podicipediformes
Little grebe Tachybaptus ruficollis	1	–	7	5	9	22
Great crested grebe Podiceps cristatus	2	–	2	1	1	6
Black-necked grebe Podiceps nigricollis	2	1	7	4	1	15
Phoenicopteriformes
Greater flamingo Phoenicopterus roseus	–	–	4	4	1	9
Columbiformes
Rock dove Columba livia	–	6	2	2	3	13
Stock dove Columba oenas	–	–	3	1	–	4
Common wood pigeon Columba palumbus	–	1	6	1	4	12
Trocaz pigeon Columba trocaz	–	29	–	–	–	29
European turtle dove Streptopelia turtur	–	5	6	4	7	22
Eurasian collared dove Streptopelia decaocto	–	1	–	–	1	2
Pterocliformes
Black-bellied sandgrouse Pterocles orientalis	–	–	5	1	–	6
Pin-tailed sandgrouse Pterocles alchata	–	–	3	–	–	3
Caprimulgiformes
Red-necked nightjar Caprimulgus ruficollis	–	–	1	1	5	7
European nightjar Caprimulgus europaeus	–	–	4	2	7	13
Alpine swift Tachymarptis melba	–	2	4	–	–	6
White-rumped swift Apus caffer	–	–	–	–	–	0
Plain swift Apus unicolor	–	4	–	–	–	4
Pallid swift Apus pallidus	–	6	2	–	1	9
Common swift Apus apus	–	1	1	3	9	14
Cuculiformes
Great spotted cuckoo Clamator glandarius	–	4	8	4	3	19
Common cuckoo Cuculus canorus	–	5	6	7	5	23
Gruiformes
Water rail Rallus aquaticus	4	1	4	4	4	17
Spotter crake Porzana porzana	2	1	5	3	2	13
Baillon’s crake Zapornia pusilla	–	1	4	1	–	6
Western swamphen Porphyrio porphyrio	–	–	5	–	1	6
Common moorhen Gallinula chloropus	3	6	6	5	16	36
Red-knobbed coot Fulica cristata	–	–	6	1	1	8
Eurasian coot Fulica atra	4	10	5	–	3	22
Demoiselle crane Anthropoides virgo	–	–	–	–	–	0
Common crane Grus grus	–	–	4	1	1	6
Otidiformes
Little bustard Tetrax tetrax	–	–	7	1	2	10
Great bustard Otis tarda	–	–	5	1	3	9
Procellariiformes
Wilson’s storm-petrel Oceanites oceanicus	–	–	–	4	–	4
White-faced storm petrel Pelagodroma marina	1	7	–	–	10	18
European storm-petrel Hydrobates pelagicus	2	1	1	–	–	4
Band-rumped storm petrel Hydrobates castro	–	33	–	1	3	37
Monteiro’s storm petrel Hydrobates monteiroi	–	1	–	–	–	1
Leach’s storm-petrel Hydrobates leucorhous	2	–	14	5	–	21
Desertas petrel Pterodroma deserta	–	30	–	–	3	33
Zino’s petrel Pterodroma madeira	–	56	–	–	1	57
Sooty shearwater Ardenna grisea	–	–	3	1	2	6
Great shearwater Ardenna gravis	–	–	3	2	1	6
Cory’s shearwater Calonectris borealis	5	25	2	2	19	53
Manx shearwater Puffinus puffinus	3	49	–	2	–	54
Sargasso shearwater Puffinus lherminieri	–	–	–	–	–	0
Balearic shearwater Puffinus mauretanicus	–	–	–	2	4	6
Bulwer’s petrel Bulweria bulwerii	3	23	–	1	7	34
Ciconiiformes
Black stork Ciconia nigra	–	–	3	1	1	5
White stork Ciconia ciconia	–	2	6	2	–	10
Pelecaniformes
Eurasian spoonbill Platalea leucorodia	1	3	4	4	1	13
Glossy ibis Plegadis falcinellus	1	1	3	3	–	8
Eurasian bittern Botaurus stellaris	3	1	8	3	–	15
Little bittern Ixobrychus minutus	1	11	9	4	3	28
Black-crowned night heron Nycticorax nycticorax	1	8	5	1	–	15
Squacco heron Ardeola ralloides	–	6	6	–	–	12
Cattle egret Bubulcus ibis	3	4	2	–	13	22
Grey heron Ardea cinerea	2	13	5	3	12	35
Purple heron Ardea purpurea	1	3	7	–	–	11
Great egret Ardea alba	–	–	–	–	–	0
Little egret Egretta garzetta	2	8	6	2	3	21
Suliformes
Northern gannet Morus bassanus	13	10	9	3	14	49
European shag Gulosus aristotelis	6	–	6	1	2	15
Great cormorant Phalacrocorax carbo	2	4	5	2	2	15
Charadriiformes
Eurasian stone-curlew Burhinus oedicnemus	–	3	6	5	3	17
Eurasian oystercatcher Haematopus ostralegus	1	2	6	5	2	16
Pied avocet Recurvirostra avosetta	5	1	5	5	3	19
Black-winged stilt Himantopus himantopus	–	8	5	7	5	25
Grey plover Pluvialis squatarola	1	2	8	4	2	17
European golden plover Pluvialis apricaria	1	3	6	3	11	24
Common ringed plove Charadrius hiaticula	2	1	6	4	–	13
Semipalmated plover Charadrius semipalmatus	–	–	–	–	–	0
Little ringed plover Charadrius dubius	3	–	6	2	6	17
Kentish plover Charadrius alexandrinus	4	1	4	3	3	15
Eurasian dotterel Eudromias morinellus	2	7	3	6	32	50
Northern lapwing Vanellus vanellus	–	4	–	1	–	5
Eurasian whimbrel Numenius phaeopus	3	4	6	8	–	21
Eurasian curlew Numenius arquata	2	2	3	5	2	14
Bar-tailed godwit Limosa lapponica	–	2	4	5	1	12
Black-tailed godwit Limosa limosa	4	7	6	4	2	23
Ruddy turnstone Arenaria interpres	3	7	6	7	2	25
Red knot Calidris canutus	3	1	5	7	1	17
Ruff Calidris pugnax	–	5	5	2	–	12
Curlew sandpiper Calidris ferruginea	–	1	5	5	1	12
Sanderling Calidris alba	1	3	5	7	1	17
Dunlin Calidris alpina	3	3	12	8	20	46
Purple sandpiper Calidris maritima	–	–	1	1	–	2
Little stint Calidris minuta	–	–	6	4	2	12
Temminck’s stint Calidris temminckii	–	–	1	–	–	1
Eurasian woodcock Scolopax rusticola	–	3	4	1	3	11
Common snipe Gallinago gallinago	–	5	4	12	11	32
Jack snipe Lymnocryptes minimus	–	5	4	3	1	13
Common sandpiper Actitis hypoleucos	1	2	4	2	4	13
Spotted redshank Tringa erythropus	–	1	–	1	–	2
Green sandpiper Tringa ochropus	1	3	5	5	–	14
Common greenshank Tringa nebularia	1	2	2	3	1	9
Common redshank Tringa totanus	3	2	6	5	2	18
Wood sandpiper Tringa glareola	–	–	4	1	1	6
Red phalarope Phalaropus fulicarius	–	2	6	3	1	12
Common buttonquail Turnix sylvaticus	–	–	5	2	–	7
Collared pratincole Glareola pratincola	3	1	2	–	1	7
Little gull Hydrocoloeus minutus	1	–	–	–	–	1
Sabine’s gull Xema sabini	–	–	–	–	–	0
Black-legged kittiwake Rissa tridactyla	3	4	7	1	6	21
Black-headed gull Larus ridibundus	6	5	4	1	4	20
Mediterranean gull Larus melanocephalus	3	–	–	–	4	7
Audouin’s gull Larus audouinii	1	–	–	–	–	1
Lesser black-backed gull Larus fuscus	2	–	7	2	11	22
European herring gull Larus argentatus	–	2	–	–	135	137
Yellow-legged gull Larus michahellis	10	13	2	–	–	25
Great black-backed gull Larus marinus	2	1	2	1	1	7
Slender-billed gull Larus genei	–	–	1	–	–	1
Ring -billed gull Larus delawarensis	–	–	–	–	–	0
Mew gull Larus canus	–	–	1	1	–	2
Sooty tern Onychoprion fuscatus	–	–	–	–	–	0
Little tern Sternula albifrons	4	–	6	11	2	23
Gull-billed tern Gelochelidon nilotica	1	–	8	2	–	11
Caspian tern Hydroprogne caspia	–	–	–	–	–	0
Whiskered tern Chlidonias hybrida	–	1	5	–	–	6
Black tern Chlidonias niger	2	1	8	3	–	14
Roseate tern Sterna dougallii	–	2	–	–	–	2
Common tern Sterna hirundo	6	2	10	2	3	23
Arctic tern Sterna paradisaea	–	–	–	–	–	0
Sandwich tern Thalasseus sandvicensis	5	–	7	1	3	16
Long-tailed jaeger Stercorarius longicaudus	–	–	–	–	–	0
Arctic jaeger Stercorarius parasiticus	2	–	1	1	–	4
Pomarine jaeger Stercorarius pomarinus	–	–	1	–	–	1
Great skua Catharacta skua	–	4	–	–	–	4
Atlantic puffin Fratercula arctica	4	9	5	2	6	26
Razorbill Alca torda	22	–	3	1	79	105
Common murre Uria aalge	10	–	7	2	18	37
Strigiformes
Barn owl Tyto alba	–	21	9	3	44	77
Little owl Athene noctua	1	–	5	2	38	46
Scops owl Otus scops	–	1	3	5	6	15
Long-eared owl Asio otus	–	–	5	3	4	12
Short-eared owl Asio flammeus	–	4	6	5	2	17
Tawny owl Strix aluco	–	–	6	1	32	39
Eurasian eagle-owl Bubo bubo	–	–	7	3	8	18
Accipitriformes
Osprey Pandion haliaetus	2	1	3	1	2	9
Black-winged kite Elanus caeruleus	–	–	3	2	5	10
European honey buzzard Pernis apivorus	–	–	–	–	2	2
Bearded vulture Gypaetus barbatus	–	–	2	–	–	2
Egyptian vulture Neophron percnopterus	–	–	7	2	–	9
Short-toed snake eagle Circaetus gallicus	–	–	4	–	4	8
Rüppell’s vulture Gyps rueppelli	–	–	–	–	–	0
Eurasian griffon vulture Gyps fulvus	–	–	8	4	5	17
Cinereous vulture Aegypius monachus	–	–	8	1	1	10
Spanish imperial eagle Aquila adalberti	–	–	2	–	–	2
Golden eagle Aquila chrysaetos	–	–	8	–	–	8
Bonelli’s eagle Aquila fasciata	–	–	6	–	2	8
Booted eagle Hieraaetus pennatus	–	–	3	3	4	10
Western marsh harrier Circus aeruginosus	–	–	8	4	2	14
Hen harrier Circus cyaneus	–	–	5	1	–	6
Montagu’s harrier Circus pygargus	–	1	6	3	3	13
Eurasian sparrowhawk Accipiter nisus	–	13	10	5	13	41
Northern goshawk Accipiter gentilis	–	–	1	2	2	5
Red kite Milvus milvus	–	–	5	2	2	9
Black kite Milvus migrans	–	–	3	1	2	6
Common buzzard Buteo buteo	–	16	8	34	26	84
Bucerotiformes
Eurasian hoopoe Upupa epops	–	8	5	2	13	28
Coraciiformes
European bee-eater Merops apiaster	–	3	5	3	17	28
European roller Coracias garrulus	–	2	6	4	6	18
Common kingfisher Alcedo atthis	2	2	5	2	21	32
Piciformes
Eurasian wryneck Jynx torquilla	–	1	3	4	1	9
Iberian green woodpecker Picus sharpei	–	–	5	4	4	13
Lesser spotted woodpecker Dryobates minor	–	–	6	1	–	7
Great spotted woodpecker Dendrocopos major	–	–	7	4	8	19
Falconiformes
Lesser kestrel Falco naumanni	–	–	1	–	7	8
Common kestrel Falco tinnunculus	–	22	3	5	22	52
Eleonora’s falcon Falco eleonorae	–	–	–	–	–	0
Merlin Falco columbarius	–	–	2	2	–	4
Eurasian hobby Falco subbuteo	–	2	8	3	–	13
Peregrine falcon Falco peregrinus	–	–	6	2	2	10
Psittaciformes
Senegal parrot Poicephalus senegalus *	–	–	–	–	–	0
Monk parakeet Myiopsitta monachus *	–	–	–	–	–	0
Blue-crowned parakeet Psittacara acuticaudatus *	–	–	–	–	–	0
Fischer’s lovebird Agarpornis fischeri *	–	–	–	–	–	0
Rose-ringed parakeet Alexandrinus krameri *	–	–	–	–	1	1
Passeriformes
Eurasian Golden Oriole Oriolus oriolus	–	4	6	5	–	15
Red-backed shrike Lanius collurio	–	–	1	–	4	5
Iberian grey shrike Lanius meridionalis	–	–	6	3	32	41
Woodchat shrike Lanius senator	–	1	6	10	34	51
Red-billed chough Pyrrhocorax pyrrhocorax	–	–	10	1	–	11
Iberian magpie Cyanopica cooki	–	–	5	3	37	45
Eurasian jay Garrulus glandarius	–	–	4	3	14	21
Eurasian magpie Pica pica	–	–	5	3	10	18
Western jackdaw Corvus monedula	–	1	3	–	4	8
Rook Corvus frugilegus	–	–	4	2	–	6
Common raven Corvus corax	–	–	6	1	–	7
Carrion crow Corvus corone	–	–	4	2	4	10
Coal tit Periparus ater	–	–	4	15	2	21
Crested tit Lophophanes cristatus	–	–	4	2	2	8
Eurasian blue tit Cyanistes caeruleus	–	–	4	6	12	22
Great tit Parus major	–	–	4	9	25	38
Eurasian penduline tit Remiz pendulinus	–	–	–	–	2	2
Dupont’s lark Chersophilus duponti	–	–	1	–	–	1
Mediterranean short-toed lark Alaudala rufescens	–	–	5	–	–	5
Calandra lark Melanocorypha calandra	–	1	6	4	–	11
Greater short-toed lark Calandrella brachydactyla	–	1	7	5	8	21
Woodlark Lullula arborea	–	–	6	10	19	35
Eurasian skylark Alauda arvensis	–	8	9	16	7	40
Thekla’s lark Galerida theklae	–	–	–	–	20	20
Crested lark Galerida cristata	–	–	5	6	11	22
Zitting cisticola Cisticola juncidis	–	–	7	3	9	19
Savi’s warbler Locustella luscinioides	–	–	–	–	2	2
Common grasshopper warbler Locustella naevia	–	–	3	2	1	6
Western olivaceous warbler Iduna opaca	–	–	–	–	–	0
Melodious warbler Hippolais polyglotta	–	–	7	2	7	16
Eurasian reed warbler Acrocephalus scirpaceus	–	–	4	–	3	7
Great reed warbler Acrocephalus arundinaceus	–	1	4	–	5	10
Aquatic warbler Acrocephalus paludicola	–	–	2	8	–	10
Sedge warbler Acrocephalus schoenobaenus	–	–	4	4	1	9
Western house martin Delichon urbicum	–	4	1	4	7	16
European red-rumped swallow Cecropis daurica	–	–	–	1	2	3
Barn swallow Hirundo rustica	–	1	2	1	18	22
Eurasian crag martin Ptyonoprogne rupestris	–	–	7	3	11	21
Sand martin Riparia riparia	–	–	4	1	–	5
Western Bonelli‘s warbler Phylloscopus bonelli	–	–	2	2	–	4
Iberian chiffchaff Phylloscopus ibericus	–	–	–	–	–	0
Common chiffchaff Phylloscopus collybita	–	1	3	7	20	31
Willow warbler Phylloscopus trochilus	–	1	3	7	2	13
Cetti’s warbler Cettia cetti	–	–	3	–	5	8
Long-tailed tit Aegithalos caudatus	–	–	5	3	16	24
Eurasian blackcap Sylvia atricapilla	–	21	5	2	27	55
Garden warbler Sylvia borin	–	–	3	–	2	5
Western Orphean warbler Curruca hortensis	–	–	2	2	–	4
Sardinian warbler Curruca melanocephala	–	–	4	–	24	28
Eastern subalpine warbler Curruca cantillans	–	–	1	–	1	2
Common whitethroat Curruca communis	–	–	9	4	2	15
Spectacled warbler Curruca conspicillata	–	3	–	–	1	4
Dartford warbler Curruca undata	–	–	5	4	5	14
Red-billed leiothrix Leiothrix lutea *	–	1	–	–	–	1
Short-toed treecreeper Certhia brachydactyla	–	–	5	2	2	9
Eurasian nuthatch Sitta europaea	–	–	7	8	7	22
Eurasian wren Troglodytes troglodytes	–	–	6	4	8	18
White-throated dipper Cinclus cinclus	–	–	6	6	5	17
Common starling Sturnus vulgaris	–	11	8	1	7	27
Spotless starling Sturnus unicolor	–	–	7	7	18	32
Crested myna Acridotheres cristatellus *	–	–	–	–	–	0
Mistle thrush Turdus viscivorus	–	–	3	5	5	13
Song thrush Turdus philomelos	–	1	1	10	23	35
Redwing Turdus iliacus	–	2	5	4	4	15
Common blackbird Turdus merula	–	23	4	11	32	70
Fieldfare Turdus pilaris	–	2	3	3	2	10
Ring ouzel Turdus torquatus	–	1	1	1	–	3
Rufous-tailed scrub robin Cercotrichas galactotes	–	1	3	–	2	6
Spotted flycatcher Muscicapa striata	–	–	5	2	3	10
European robin Erithacus rubecula	–	10	2	4	43	59
Bluethroat Luscinia svecica	–	–	5	7	–	12
Common nightingale Luscinia megarhynchos	–	–	1	2	10	13
Black redstart Phoenicurus ochruros	–	–	7	11	2	20
Common redstart Phoenicurus phoenicurus	–	3	4	3	1	11
Common rock thrush Monticola saxatilis	–	–	5	8	1	14
Blue rock thrush Monticola solitarius	–	–	9	1	2	12
Whinchat Saxicola rubetra	–	1	7	4	2	14
European stonechat Saxicola rubicola	–	1	4	15	20	40
Northern wheatear Oenanthe oenanthe	–	6	3	14	1	24
Western black-eared wheatear Oenanthe hispanica	–	–	14	9	7	30
Black wheatear Oenanthe leucura	–	–	3	2	–	5
European pied flycatcher Ficedula hypoleuca	–	2	4	–	4	10
Goldcrest Regulus regulus	–	–	2	3	–	5
Madeira firecrest Regulus madeirensis	–	3	–	–	–	7
Common firecrest Regulus ignicapilla	–	–	7	4	11	18
Alpine accentor Prunella collaris	–	–	–	2	–	2
Dunnock Prunella modularis	–	–	3	4	9	16
Yellow-crowned bishop Euplectes afer *	–	–	–	–	–	0
Black-headed weaver Ploceus melanocephalus *	–	–	–	–	–	0
Common waxbill Estrilda astrild *	–	–	–	–	20	20
Red avadavat Amandava amandava *	–	–	–	–	–	0
Scaly-breasted munia Lonchura punctulata *	–	–	–	–	–	0
Pin-tailed whydah Vidua macroura *	–	–	–	–	–	0
House sparrow Passer domesticus	–	1	3	51	156	211
Spanish sparrow Passer hispaniolensis	–	4	–	7	412	423
Eurasian tree sparrow Passer montanus	–	–	–	–	22	22
Rock sparrow Petronia petronia	–	6	5	8	6	25
Tree pipit Anthus trivialis	–	–	3	14	–	17
Meadow pipit Anthus pratensis	–	–	5	12	23	40
Water pipit Anthus spinoletta	–	–	3	5	–	8
Berthelot’s pipit Anthus berthelotii	–	7	–	–	2	9
Tawny pipit Anthus campestris	–	–	6	9	1	16
Richard’s pipit Anthus richardi	–	1	1	–	–	2
Western yellow wagtail Motacilla flava	–	1	7	26	5	39
Grey wagtail Motacilla cinerea	–	5	5	6	9	25
White wagtail Motacilla alba	–	4	7	24	14	49
Eurasian chaffinch Fringilla coelebs	–	11	4	22	60	97
Brambling Fringilla montifringilla	–	–	8	5	–	13
Hawfinch Coccothraustes coccothraustes	–	1	6	6	5	18
Azores bullfinch Pyrrhula murina	–	–	–	2	–	2
Eurasian bullfinch Pyrrhula pyrrhula	–	–	5	8	3	16
European greenfinch Chloris chloris	–	4	2	11	26	43
Common linnet Linaria cannabina	–	5	4	11	29	49
Red crossbill Loxia curvirostra	–	6	6	3	–	15
European goldfinch Carduelis carduelis	–	1	3	40	39	83
European serin Serinus serinus	–	–	5	8	22	35
Atlantic canary Serinus canaria	–	6	–	–	5	11
Eurasian siskin Spinus spinus	–	–	5	6	8	19
Corn bunting Emberiza calandra	–	–	7	7	48	62
Rock bunting Emberiza cia	–	–	5	16	9	30
Ortolan bunting Emberiza hortulana	–	–	3	4	–	7
Cirl bunting Emberiza cirlus	–	–	6	14	20	40
Yellowhammer Emberiza citrinella	–	–	–	7	3	10
Common reed bunting Emberiza schoeniclus	–	–	5	3	–	8
Snow bunting Plectrophenax nivalis	3	1	–	4	–	8
TOTAL	236	829	1251	1117	2553	5986

Table 2.

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Conservation status and percentage of representation (%) of Portuguese bird species. Conservation status: LC—Least Concern, NT—Near Threatened, VU—Vulnerable, EN—Endangered, NA—Not Applicable. For some species, there are two conservation status assessments, for breeding and wintering populations. The “Number of represented squares” column displays the number of 10 × 10 km squares for which there are museum specimens of a certain taxon versus the number of squares from which a species is known, following Equipa Atlas (2018, 2022).

Taxa	Conservation status	Game species	Number of represented squares	Percentage of representation (%)	Map
Galliformes
Common quail Coturnix coturnix	LC	Yes	11/625	1.8	Fig. 8A
Red–legged partridge Alectoris rufa	LC	Yes	23/839	2.7	Fig. 8C
Ring–necked pheasant Phasianus colchicus	NA	Yes	1/51	2.0	Fig. 8B
Grey partridge Perdix perdix	RE	No	–	–	–
Western capercaillie Tetrao urogallus	RE	No	–	–	–
Anseriformes
Black swan Cygnus atratus	NA	No	–	–	–
Mute swan Cygnus olor	NA	No	–	–	–
Greylag goose Anser anser	VU	No	2/31	6.5	Fig. 8D
Common scoter Melanitta nigra	VU	No	8/46	17.4	Fig. 8E
Red–breasted merganser Mergus serrator	EN	No	3/11	27.3	Fig. 8F
Egyptian goose Alopochen aegyptiaca	NA	No	–	–	–
Common shelduck Tadorna tadorna	NT/LC	No	6/44	13.6	Fig. 8G
Ruddy shelduck Tadorna ferruginea	NA	No	1/13	7.7	Fig. 8H
Marbled duck Marmaronetta angustirostris	NA/NA	No	2/4	50	Fig. 8I
Red–crested pochard Netta rufina	VU/NT	No	–	–	–
Common pochard Aythya ferina	EN/VU	Yes	9/82	11.0	Fig. 9A
Ferruginous duck Aythya nyroca	NA/NA	No	1/22	4.5	Fig. 8J
Tufted duck Aythya fuligula	NA/VU	Yes	9/45	20.0	Fig. 9B
Garganey Spatula querquedula	NA/LC	No	2/14	14.3	Fig. 9C
Northern shoveler Spatula clypeata	VU/LC	Yes	4/123	3.3	Fig. 9D
Gadwall Mareca strepera	LC/LC	Yes	1/187	0.5	Fig. 9E
Eurasian wigeon Mareca penelope	LC	Yes	7/36	19.4	Fig. 9F
Mallard Anas platyrhynchos	LC	Yes	5/679	0.7	Fig. 9G
Northern pintail Anas acuta	LC	Yes	3/31	9.7	Fig. 9H
Eurasian teal Anas crecca	LC	Yes	13/115	11.3	Fig. 9I
Podicipediformes
Little grebe Tachybaptus ruficollis	LC	No	10/409	2.4	Fig. 10A
Great crested grebe Podiceps cristatus	LC	No	3/234	1.3	Fig. 10B
Black–necked grebe Podiceps nigricollis	NA/VU	No	9/43	20.9	Fig. 10C
Phoenicopteriformes
Greater flamingo Phoenicopterus roseus	NA/LC	No	4/56	7.1	Fig. 10D
Columbiformes
Rock dove Columba livia	DD	Yes	4/897	0.4	Fig. 10E
Stock dove Columba oenas	VU/DD	Yes	2/45	4.4	Fig. 10F
Common wood pigeon Columba palumbus	LC	Yes	5/1014	0.5	Fig. 10G
Trocaz pigeon Columba trocaz	LC	No	6/12	50	Fig. 10H
European turtle dove Streptopelia turtur	LC	Yes	10/732	1.4	Fig. 11A
Eurasian collared dove Streptopelia decaocto	LC	No	2/991	0.2	Fig. 10J
Pterocliformes
Black–bellied sandgrouse Pterocles orientalis	EN	No	3/37	8.1	Fig. 10I
Pin–tailed sandgrouse Pterocles alchata	CR	No	1/7	14.3	Fig. 10K
Caprimulgiformes
Red–necked nightjar Caprimulgus ruficollis	LC	No	5/236	2.1	Fig. 11B
European nightjar Caprimulgus europaeus	LC	No	10/344	2.9	Fig. 11C
Alpine swift Tachymarptis melba	NT	No	4/89	4.5	Fig. 11D
White–rumped swift Apus caffer	VU	No	–	–	–
Plain swift Apus unicolor	NA	No	2/17	11.8	Fig. 11E
Pallid swift Apus pallidus	LC	No	4/391	1.0	Fig. 11F
Common swift Apus apus	LC	No	7/784	0.9	Fig. 11G
Cuculiformes
Great spotted cuckoo Clamator glandarius	NT	No	15/212	7.1	Fig. 11H
Common cuckoo Cuculus canorus	LC	No	12/688	1.7	Fig. 11I
Gruiformes
Water rail Rallus aquaticus	LC	No	8/122	6.6	Fig. 12A
Spotter crake Porzana porzana	NA	No	8/11	72.7	Fig. 12B
Baillon’s crake Zapornia pusilla	NA	No	4/7	57.1	Fig. 12C
Western swamphen Porphyrio porphyrio	NT	No	3/45	6.7	Fig. 12D
Common moorhen Gallinula chloropus	LC	Yes	19/569	3.3	Fig. 12E
Red–knobbed coot Fulica cristata	RE/NA	No	5/12	41.7	Fig. 12F
Eurasian coot Fulica atra	LC/LC	Yes	11/280	3.9	Fig. 12H
Demoiselle crane Anthropoides virgo	RE	No	–	–	–
Common crane Grus grus	RE/VU	No	4/33	12.1	Fig. 12G
Otidiformes
Little bustard Tetrax tetrax	CR	No	5/123	4.1	Fig. 12I
Great bustard Otis tarda	EN	No	7/65	10.8	Fig. 13A
Procellariiformes
Wilson’s storm–petrel Oceanites oceanicus	–	No	–	–	–
White–faced storm petrel Pelagodroma marina	EN	No	2/2	100	Fig. 13B
European storm–petrel Hydrobates pelagicus	–	No	4/12	33	Fig. 13C
Band–rumped storm petrel Hydrobates castro	VU	No	6/12	50	Fig. 13D
Monteiro’s storm petrel Hydrobates monteiroi	VU	No	1/4	25	Fig. 13E
Leach’s storm–petrel Hydrobates leucorhous	–	No	7/10	70	Fig. 13F
Desertas petrel Pterodroma deserta	VU	No	2/4	50	Fig. 13G
Zino’s petrel Pterodroma madeira	EN	No	1/3	33.3	Fig. 13H
Sooty shearwater Ardenna grisea	–	No	4/10	40	Fig. 13I
Great shearwater Ardenna gravis	–	No	4/12	33.3	Fig. 13J
Cory’s shearwater Calonectris borealis	EN	No	13/76	17.1	Fig. 13K
Manx shearwater Puffinus puffinus	LC	No	8/34	23.5	Fig. 14A
Sargasso shearwater Puffinus lherminieri	NT	No	–	–	–
Balearic shearwater Puffinus mauretanicus	CR	No	3/24	12.5	Fig. 14B
Bulwer’s petrel Bulweria bulwerii	LC	No	8/9	88.9	Fig. 14C
Ciconiiformes
Black stork Ciconia nigra	EN	No	3/150	2	Fig. 14E
White stork Ciconia ciconia	LC	No	4/596	0.7	Fig. 14D
Pelecaniformes
Eurasian spoonbill Platalea leucorodia	LC/LC	No	5/130	3.8	Fig. 14F
Glossy ibis Plegadis falcinellus	LC	No	3/62	4.8	Fig. 14G
Eurasian bittern Botaurus stellaris	NA/CR	No	8/14	57.1	Fig. 14I
Little bittern Ixobrychus minutus	VU	No	16/84	19.0	Fig. 14J
Black–crowned night heron Nycticorax nycticorax	NT	No	6/59	10.2	Fig. 15A
Squacco heron Ardeola ralloides	EN/EN	No	3/27	11.1	Fig. 14H
Cattle egret Bubulcus ibis	VU	No	15/389	3.9	Fig. 15B
Grey heron Ardea cinerea	LC	No	12/559	2.1	Fig. 15C
Purple heron Ardea purpurea	VU	No	4/98	4.1	Fig. 15D
Great egret Ardea alba	NA/NT	No	–	–	–
Little egret Egretta garzetta	LC	No	10/326	3.1	Fig. 15E
Suliformes
Northern gannet Morus bassanus	LC	No	22/68	32.4	Fig. 15G
European shag Gulosus aristotelis	EN	No	5/42	11.9	Fig. 15F
Great cormorant Phalacrocorax carbo	NA/LC	No	8/441	1.8	Fig. 15H
Charadriiformes
Eurasian stone–curlew Burhinus oedicnemus	VU	No	9/217	4.1	Fig. 16A
Eurasian oystercatcher Haematopus ostralegus	RE/VU	No	7/41	17.1	Fig. 16B
Pied avocet Recurvirostra avosetta	NT/LC	No	6/50	12	Fig. 16C
Black–winged stilt Himantopus himantopus	LC	No	11/239	4.6	Fig. 16E
Grey plover Pluvialis squatarola	NT	No	9/61	14.8	Fig. 16D
European golden plover Pluvialis apricaria	LC	Yes	13/128	10.2	Fig. 16G
Common ringed plover Charadrius hiaticula	LC	No	6/87	6.9	Fig. 16F
Semipalmated plover Charadrius semipalmatus	–	No	–	–	–
Little ringed plover Charadrius dubius	LC	No	9/306	2.9	Fig. 16H
Kentish plover Charadrius alexandrinus	VU/EN	No	7/109	6.4	Fig. 16J
Eurasian dotterel Eudromias morinellus	NA/LC	No	4/12	33.3	Fig. 16I
Northern lapwing Vanellus vanellus	–	No	32/326	9.8	Fig. 17A
Eurasian whimbrel Numenius phaeopus	NT	No	9/69	13.0	Fig. 17B
Eurasian curlew Numenius arquata	EN	No	7/43	16.3	Fig. 17C
Bar–tailed godwit Limosa lapponica	LC	No	5/42	11.9	Fig. 17E
Black–tailed godwit Limosa limosa	LC	No	11/67	16.4	Fig. 17D
Ruddy turnstone Arenaria interpres	VU	No	10/81	12.3	Fig. 17F
Red knot Calidris canutus	EN	No	7/37	18.9	Fig. 17G
Ruff Calidris pugnax	VU	No	6/31	19.4	Fig. 17H
Curlew sandpiper Calidris ferruginea	EN	No	7/42	16.7	Fig. 17I
Sanderling Calidris alba	LC	No	10/81	12.3	Fig. 18A
Dunlin Calidris alpina	LC	No	15/79	19.0	Fig. 18B
Purple sandpiper Calidris maritima	EN	No	2/9	22.2	Fig. 18C
Little stint Calidris minuta	EN	No	7/42	16.7	Fig. 18D
Temminck’s stint Calidris temminckii	–	No	1/8	12.5	Fig. 18E
Eurasian woodcock Scolopax rusticola	DD	Yes	6/73	8.2	Fig. 18G
Common snipe Gallinago gallinago	CR/LC	Yes	14/242	5.8	Fig. 18H
Jack snipe Lymnocryptes minimus	DD	Yes	8/15	53.3	Fig. 19A
Common sandpiper Actitis hypoleucos	VU/NT	No	9/426	2.1	Fig. 18F
Spotted redshank Tringa erythropus	EN	No	2/28	7.1	Fig. 19B
Green sandpiper Tringa ochropus	NT	No	9/249	3.6	Fig. 19C
Common greenshank Tringa nebularia	NT	No	5/84	6.0	Fig. 19D
Common redshank Tringa totanus	CR/VU	No	10/71	14.1	Fig. 19E
Wood sandpiper Tringa glareola	–	No	4/18	22.2	Fig. 19F
Red phalarope Phalaropus fulicarius	–	No	6/11	54.5	Fig. 19G
Common buttonquail Turnix sylvaticus	RE	No	–	–	–
Collared pratincole Glareola pratincola	VU	No	2/57	3.5	Fig. 19H
Little gull Hydrocoloeus minutus	–	No	1/15	6.7	Fig. 19I
Sabine’s gull Xema sabini	–	No	–	–	–
Black–legged kittiwake Rissa tridactyla	VU	No	11/16	68.8	Fig. 19J
Black–headed gull Larus ridibundus	DD/LC	No	10/216	4.6	Fig. 20A
Mediterranean gull Larus melanocephalus	LC	No	1/65	1.5	Fig. 19K
Audouin’s gull Larus audouinii	VU	No	–	–	–
Lesser black–backed gull Larus fuscus	VU/LC	No	9/241	3.7	Fig. 20B
European herring gull Larus argentatus	–	No	5/228	2.2	Fig. 20D
Yellow–legged gull Larus michahellis	LC	No	7/27	25.9	Fig. 20F
Great black–backed gull Larus marinus	–	No	2/38	5.3	Fig. 20C
Slender–billed gull Larus genei	–	No	1/10	10.0	Fig. 20E
Ring –billed gull Larus delawarensis	–	No	–	–	–
Mew gull Larus canus	–	No	2/22	9.1	Fig. 20G
Sooty tern Onychoprion fuscatus	VU	No	–	–	–
Little tern Sternula albifrons	VU	No	8/55	14.5	Fig. 20H
Gull–billed tern Gelochelidon nilotica	VU	No	3/69	4.3	Fig. 20I
Caspian tern Hydroprogne caspia	VU	No	–	–	–
Whiskered tern Chlidonias hybrida	CR	No	3/16	18.8	Fig. 20J
Black tern Chlidonias niger	–	No	7/20	35	Fig. 20K
Roseate tern Sterna dougallii	LC	No	–	–	–
Common tern Sterna hirundo	EN	No	9/123	7.3	Fig. 21A
Arctic tern Sterna paradisaea	–	No	–	–	–
Sandwich tern Thalasseus sandvicensis	LC	No	9/81	11.1	Fig. 21C
Long–tailed jaeger Stercorarius longicaudus	–	No	–	–	–
Arctic jaeger Stercorarius parasiticus	–	No	3/15	20.0	Fig. 21B
Pomarine jaeger Stercorarius pomarinus	–	No	1/10	10.0	Fig. 21E
Great skua Catharacta skua	LC	No	2/23	8.7	Fig. 21D
Atlantic puffin Fratercula arctica	DD	No	11/14	78.6	Fig. 21F
Razorbill Alca torda	NT	No	14/34	41.2	Fig. 21G
Common murre Uria aalge	CR/NT	No	11/15	73.3	Fig. 21H
Strigiformes
Barn owl Tyto alba	NT	No	34/396	8.6	Fig. 21I
Little owl Athene noctua	LC	No	31/650	4.8	Fig. 21J
Scops owl Otus scops	VU	No	11/229	4.8	Fig. 22A
Long–eared owl Asio otus	VU	No	9/143	6.3	Fig. 22B
Short–eared owl Asio flammeus	VU	No	10/45	22.2	Fig. 22C
Tawny owl Strix aluco	LC	No	25/550	4.5	Fig. 22D
Eurasian eagle–owl Bubo bubo	NT	No	11/247	4.5	Fig. 22E
Accipitriformes
Osprey Pandion haliaetus	CR/VU	No	7/73	9.6	Fig. 22F
Black–winged kite Elanus caeruleus	LC	No	5/413	1.2	Fig. 22G
European honey buzzard Pernis apivorus	NT	No	–	–	–
Bearded vulture Gypaetus barbatus	RE	No	–	–	–
Egyptian vulture Neophron percnopterus	EN	No	4/81	4.9	Fig. 22H
Short–toed snake eagle Circaetus gallicus	NT	No	7/631	1.1	Fig. 22I
Rüppell’s vulture Gyps rueppelli	NA	No	–	–	–
Eurasian griffon vulture Gyps fulvus	LC	No	8/186	4.3	Fig. 22J
Cinereous vulture Aegypius monachus	EN	No	4/79	5.1	Fig. 22K
Spanish imperial eagle Aquila adalberti	CR	No	2/33	6.1	Fig. 22L
Golden eagle Aquila chrysaetos	EN	No	6/109	5.5	Fig. 23A
Bonelli’s eagle Aquila fasciata	VU	No	4/238	1.7	Fig. 23B
Booted eagle Hieraaetus pennatus	LC	No	6/656	0.9	Fig. 23C
Western marsh harrier Circus aeruginosus	NT/NT	No	5/225	2.2	Fig. 23D
Hen harrier Circus cyaneus	CR/EN	No	3/157	1.9	Fig. 23E
Montagu’s harrier Circus pygargus	EN	No	6/259	2.3	Fig. 23F
Eurasian sparrowhawk Accipiter nisus	LC	No	17/612	2.8	Fig. 23G
Northern goshawk Accipiter gentilis	LC	No	5/364	1.4	Fig. 23H
Red kite Milvus milvus	CR/LC	No	5/240	2.1	Fig. 23I
Black kite Milvus migrans	LC	No	3/668	0.4	Fig. 23J
Common buzzard Buteo buteo	LC	No	29/968	3.0	Fig. 23L
Bucerotiformes
Eurasian hoopoe Upupa epops	LC	No	11/888	1.2	Fig. 24A
Coraciiformes
European bee–eater Merops apiaster	LC	No	9/700	1.3	Fig. 23K
European roller Coracias garrulus	CR	No	11/40	27.5	Fig. 24B
Common kingfisher Alcedo atthis	LC	No	17/625	2.7	Fig. 24C
Piciformes
Eurasian wryneck Jynx torquilla	LC	No	5/251	2.0	Fig. 24D
Iberian green woodpecker Picus sharpei	LC	No	9/678	1.3	Fig. 24E
Lesser spotted woodpecker Dryobates minor	LC	No	5/415	1.2	Fig. 24F
Great spotted woodpecker Dendrocopos major	LC	No	10/899	1.1	Fig. 24G
Falconiformes
Lesser kestrel Falco naumanni	EN	No	6/52	11.5	Fig. 24H
Common kestrel Falco tinnunculus	VU	No	18/737	2.4	Fig. 24I
Eleonora’s falcon Falco eleonorae	RE	No	–	–	–
Merlin Falco columbarius	EN	No	3/48	6.3	Fig. 24J
Eurasian hobby Falco subbuteo	VU	No	4/221	1.8	Fig. 24K
Peregrine falcon Falco peregrinus	VU	No	4/254	1.6	Fig. 25A
Psittaciformes
Senegal parrot Poicephalus senegalus	NA	No	–	–	–
Monk parakeet Myiopsitta monachus	NA	No	–	–	–
Blue–crowned parakeet Psittacara acuticaudatus	NA	No	–	–	–
Fischer’s lovebird Agarpornis fischeri	NA	No	–	–	–
Rose–ringed parakeet Alexandrinus krameri	NA	No	–	–	–
Passeriformes
Eurasian Golden Oriole Oriolus oriolus	LC	No	8/698	1.1	Fig. 25B
Red–backed shrike Lanius collurio	VU	No	2/52	3.8	Fig. 25C
Iberian grey shrike Lanius meridionalis	VU	No	23/673	3.4	Fig. 25D
Woodchat shrike Lanius senator	VU	No	22/584	3.8	Fig. 25E
Red–billed chough Pyrrhocorax pyrrhocorax	VU	No	5/41	12.2	Fig. 25F
Iberian magpie Cyanopica cooki	LC	No	17/542	3.1	Fig. 25G
Eurasian jay Garrulus glandarius	LC	No	14/897	1.6	Fig. 25H
Eurasian magpie Pica pica	LC	Yes	10/713	1.4	Fig. 25I
Western jackdaw Corvus monedula	EN	No	5/85	5.9	Fig. 25J
Rook Corvus frugilegus	RE	No	–	–	–
Common raven Corvus corax	LC	No	5/552	0.9	Fig. 25K
Carrion crow Corvus corone	LC	Yes	7/884	0.8	Fig. 25L
Coal tit Periparus ater	LC	No	9/535	1.7	Fig. 25M
Crested tit Lophophanes cristatus	LC	No	5/764	0.7	Fig. 25N
Eurasian blue tit Cyanistes caeruleus	LC	No	12/958	1.3	Fig. 26A
Great tit Parus major	LC	No	25/973	2.6	Fig. 26B
Eurasian penduline tit Remiz pendulinus	NA/NT	No	1/26	3.8	Fig. 26C
Dupont’s lark Chersophilus duponti	RE	No	–	–	–
Mediterranean short–toed lark Alaudala rufescens	EN	No	1/5	20.0	Fig. 26D
Calandra lark Melanocorypha calandra	NT	No	6/128	4.7	Fig. 26E
Greater short–toed lark Calandrella brachydactyla	LC	No	9/171	5.3	Fig. 26F
Woodlark Lullula arborea	LC	No	18/873	2.1	Fig. 26G
Eurasian skylark Alauda arvensis	LC	No	16/489	3.3	Fig. 26I
Thekla’s lark Galerida theklae	LC	No	15/476	3.2	Fig. 26H
Crested lark Galerida cristata	LC	No	16/655	2.4	Fig. 26J
Zitting cisticola Cisticola juncidis	LC	No	10/855	1.2	Fig. 26K
Savi’s warbler Locustella luscinioides	VU	No	2/41	4.9	Fig. 26L
Common grasshopper warbler Locustella naevia	–	No	3/45	6.7	Fig. 26M
Western olivaceous warbler Iduna opaca	DD	No	–	–	–
Melodious warbler Hippolais polyglotta	LC	No	10/837	1.2	Fig. 26N
Eurasian reed warbler Acrocephalus scirpaceus	LC	No	5/274	1.8	Fig. 26O
Great reed warbler Acrocephalus arundinaceus	LC	No	4/177	2.3	Fig. 27A
Aquatic warbler Acrocephalus paludicola	EN	No	2/3	66.7	Fig. 27B
Sedge warbler Acrocephalus schoenobaenus	–	No	5/29	17.2	Fig. 27C
Western house martin Delichon urbicum	LC	No	7/924	0.8	Fig. 27E
European red–rumped swallow Cecropis daurica	LC	No	3/913	0.3	Fig. 27D
Barn swallow Hirundo rustica	LC	No	13/976	1.3	Fig. 27F
Eurasian crag martin Ptyonoprogne rupestris	LC	No	10/606	1.7	Fig. 27G
Sand martin Riparia riparia	LC	No	3/518	0.6	Fig. 27H
Western Bonelli’s warbler Phylloscopus bonelli	LC	No	3/326	0.9	Fig. 27I
Iberian chiffchaff Phylloscopus ibericus	LC	No	–	–	–
Common chiffchaff Phylloscopus collybita	NA/LC	No	18/693	2.6	Fig. 27J
Willow warbler Phylloscopus trochilus	–	No	6/357	1.7	Fig. 27K
Cetti’s warbler Cettia cetti	LC	No	4/869	0.5	Fig. 27L
Long–tailed tit Aegithalos caudatus	LC	No	12/842	1.4	Fig. 27M
Eurasian blackcap Sylvia atricapilla	LC	No	25/1052	2.4	Fig. 28A
Garden warbler Sylvia borin	VU	No	3/112	2.7	Fig. 28B
Western Orphean warbler Curruca hortensis	NT	No	2/114	1.8	Fig. 28C
Sardinian warbler Curruca melanocephala	LC	No	21/945	2.2	Fig. 28D
Eastern subalpine warbler Curruca cantillans	LC	No	2/427	0.5	Fig. 28E
Common whitethroat Curruca communis	LC	No	5/294	1.7	Fig. 28G
Spectacled warbler Curruca conspicillata	NT	No	2/93	2.2	Fig. 28H
Dartford warbler Curruca undata	LC	No	9/698	1.3	Fig. 28F
Red–billed leiothrix Leiothrix lutea	NA	No	–	–	–
Short–toed treecreeper Certhia brachydactyla	LC	No	6/929	0.6	Fig. 28I
Eurasian nuthatch Sitta europaea	LC	No	14/754	1.9	Fig. 28J
Eurasian wren Troglodytes troglodytes	LC	No	10/939	1.1	Fig. 28K
White–throated dipper Cinclus cinclus	NT	No	8/136	5.9	Fig. 28L
Common starling Sturnus vulgaris	LC	Yes	10/254	3.9	Fig. 29A
Spotless starling Sturnus unicolor	LC	No	17/975	1.7	Fig. 29B
Crested myna Acridotheres cristatellus	NA	No	–	–	–
Mistle thrush Turdus viscivorus	LC	Yes	9/649	1.4	Fig. 29C
Song thrush Turdus philomelos	LC	Yes	18/753	2.4	Fig. 29D
Redwing Turdus iliacus	LC	Yes	7/203	3.4	Fig. 29E
Common blackbird Turdus merula	LC	No	33/1068	3.1	Fig. 29F
Fieldfare Turdus pilaris	LC	Yes	4/51	7.8	Fig. 29G
Ring ouzel Turdus torquatus	NT	No	2/22	9.1	Fig. 29H
Rufous–tailed scrub robin Cercotrichas galactotes	EN	No	3/27	11.1	Fig. 29I
Spotted flycatcher Muscicapa striata	NT	No	5/328	1.5	Fig. 29J
European robin Erithacus rubecula	LC	No	36/985	3.7	Fig. 30A
Bluethroat Luscinia svecica	NA/LC	No	1/63	1.6	Fig. 30B
Common nightingale Luscinia megarhynchos	NA/LC	No	6/864	0.7	Fig. 30C
Black redstart Phoenicurus ochruros	LC	No	8/877	0.9	Fig. 30D
Common redstart Phoenicurus phoenicurus	LC	No	5/289	1.7	Fig. 30E
Common rock thrush Monticola saxatilis	EN	No	5/40	12.5	Fig. 30F
Blue rock thrush Monticola solitarius	LC	No	2/312	0.6	Fig. 30G
Whinchat Saxicola rubetra	EN	No	5/156	3.2	Fig. 30H
European stonechat Saxicola rubicola	LC	No	22/962	2.3	Fig. 30I
Northern wheatear Oenanthe oenanthe	LC	No	6/388	1.5	Fig. 30J
Western black–eared wheatear Oenanthe hispanica	VU	No	13/257	5.1	Fig. 30K
Black wheatear Oenanthe leucura	CR	No	5/28	17.9	Fig. 30L
European pied flycatcher Ficedula hypoleuca	–	No	4/466	0.9	Fig. 31A
Goldcrest Regulus regulus	LC	No	2/93	2.2	Fig. 31B
Madeira firecrest Regulus madeirensis	LC	No	2/13	15.4	Fig. 31C
Common firecrest Regulus ignicapilla	LC	No	10/619	1.6	Fig. 31E
Alpine accentor Prunella collaris	EN	No	1/17	5.9	Fig. 31F
Dunnock Prunella modularis	LC	No	13/443	2.9	Fig. 31G
Yellow–crowned bishop Euplectes afer	NA	No	–	–	–
Black–headed weaver Ploceus melanocephalus	NA	No	–	–	–
Common waxbill Estrilda astrild	NA	No	9/641	1.4	Fig. 31D
Red avadavat Amandava amandava	NA	No	–	–	–
Scaly–breasted munia Lonchura punctulata	NA	No	–	–	–
Pin–tailed whydah Vidua macroura	NA	No	–	–	–
House sparrow Passer domesticus	LC	No	60/1041	5.8	Fig. 31H
Spanish sparrow Passer hispaniolensis	LC	No	16/324	4.9	Fig. 31I
Eurasian tree sparrow Passer montanus	NT	No	9/530	1.7	Fig. 32A
Rock sparrow Petronia petronia	LC	No	9/440	2.0	Fig. 31J
Tree pipit Anthus trivialis	LC	No	3/171	1.8	Fig. 32B
Meadow pipit Anthus pratensis	LC	No	22/640	3.4	Fig. 32C
Water pipit Anthus spinoletta	EN/LC	No	4/102	3.9	Fig. 32D
Berthelot’s pipit Anthus berthelotii	LC	No	4/17	23.5	Fig. 32E
Tawny pipit Anthus campestris	LC	No	7/225	3.1	Fig. 32F
Richard’s pipit Anthus richardi	–	No	2/10	20.0	Fig. 32G
Western yellow wagtail Motacilla flava	LC	No	9/212	4.2	Fig. 32H
Grey wagtail Motacilla cinerea	LC	No	13/755	1.7	Fig. 32K
White wagtail Motacilla alba	LC	No	18/961	1.9	Fig. 32I
Eurasian chaffinch Fringilla coelebs	LC	No	41/1052	3.9	Fig. 33A
Brambling Fringilla montifringilla	LC	No	4/106	3.8	Fig. 32J
Hawfinch Coccothraustes coccothraustes	LC	No	11/366	3.0	Fig. 33C
Azores bullfinch Pyrrhula murina	VU	No	1/4	25.0	Fig. 33B
Eurasian bullfinch Pyrrhula pyrrhula	LC	No	9/255	3.5	Fig. 33E
European greenfinch Chloris chloris	LC	No	29/1011	2.9	Fig. 33F
Common linnet Linaria cannabina	LC	No	20/976	2.0	Fig. 33D
Red crossbill Loxia curvirostra	VU/DD	No	5/45	11.1	Fig. 33G
European goldfinch Carduelis carduelis	LC	No	34/1037	3.3	Fig. 34A
European serin Serinus serinus	LC	No	16/976	1.6	Fig. 34B
Atlantic canary Serinus canaria	LC	No	4/90	4.4	Fig. 34C
Eurasian siskin Spinus spinus	LC	No	9/288	3.1	Fig. 34D
Corn bunting Emberiza calandra	LC	No	33/782	4.2	Fig. 34E
Rock bunting Emberiza cia	LC	No	18/518	3.5	Fig. 34F
Ortolan bunting Emberiza hortulana	VU	No	4/74	5.4	Fig. 34G
Cirl bunting Emberiza cirlus	LC	No	26/757	3.4	Fig. 34H
Yellowhammer Emberiza citrinella	EN	No	2/21	9.5	Fig. 34I
Common reed bunting Emberiza schoeniclus	EN/VU	No	5/65	7.7	Fig. 34J
Snow bunting Plectrophenax nivalis	–	No	4/9	44.4	Fig. 34K

Table 3.

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Taxonomic diversity of rare and accidental migrant birds occurring in Portugal and corresponding number of existing specimens for each species in the consulted museum/collection. Museums: AVG—Aquário Vasco da Gama, MMF—Museu de História Natural do Funchal, MCUC—Museu da Ciência da Universidade de Coimbra, MHNC-UP—Museu de História Natural e da Ciência da Universidade do Porto, and MUHNAC—Museu Nacional de História Natural e da Ciência.

Taxa	AVG	MMF	MCUC	MHNC-UP	MUHNAC	Total
Anseriformes
Greater white-fronted goose Anser albifrons		1		1		2
Bean goose Anser fabalis			3			3
Pink-footed goose Anser brachyrhynchus		1				1
Surf scoter Melanitta perspicillata		1				1
Green-winged teal Anas carolinensis		1				1
Ring-necked duck Aythya collaris		1				1
Brant Branta bernicla			2			2
Common goldeneye Bucephala clangula				1		1
Smew Mergus albellus					1	1
Common merganser Mergus merganser					1	1
Common eider Somateria mollissima	1					1
Gruiformes
Corn crake Crex crex		10	6		2	18
Little crake Zapornia parva				1		1
Procellariiformes
Northern fulmar Fulmarus glacialis		1			1	2
Little shearwater Puffinus assimilis/baroli		28		1	1	30
Gaviiformes
Red-throated loon Gavia stellata			2	4	1	7
Black-throated loon Gavia arctica			1	2		3
Common loon Gavia immer	1	1			3	5
Charadriiformes
Eurasian dotterel Charadrius morinellus				1		1
Slender-billed curlew Numenius tenuirostris		1				1
White-rumped sandpiper Calidris fuscicollis		1				1
Broad-billed sandpiper Calidris falcinellus				1		1
Pectoral sandpiper Calidris melanotos				1		1
Upland sandpiper Bartramia longicauda		2				2
Red-necked phalarope Phalaropus lobatus			1			1
Caspian gull Larus cachinnans		5				5
Glaucous gull Larus hyperboreus		3		1		4
Little auk Alle alle		2			7	9
Thick-billed murre Uria lomvia					1	1
Accipitriformes
Lesser spotted eagle Clanga pomarina			1			1
Falconiformes
Gyrfalcon Falco rusticolus				1		1
Red-footed falcon Falco vespertinus		1				1
Passeriformes
Alpine chough Pyrrhocorax graculus				4		4
Bohemian waxwing Bombycilla garrulus					2	2
Wood warbler Phylloscopus sibilatrix		1				1
Eurasian treecreeper Certhia familiaris				9		9
Wallcreeper Tichodroma muraria			1			1
Desert wheatear Oenanthe deserti		1				1
Rosy starling Sturnus roseus				2		2
European rock pipit Anthus petrosus			3	3		6
Twite Linaria flavirostris				1		1
TOTAL	2	62	20	34	20	138

Table 4.

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Eggs and nest specimens extant in the consulted museum/collection. Asterisks (*) denote introduced species. Museums: AVG—Aquário Vasco da Gama, MMF—Museu de História Natural do Funchal, MCUC—Museu da Ciência da Universidade de Coimbra, MHNC-UP—Museu de História Natural e da Ciência da Universidade do Porto, and MUHNAC—Museu Nacional de História Natural e da Ciência.

Taxa	Eggs					Nests				Total
Taxa	AVG	MMF	MCUC	MHNC-UP	MUHNAC	MMF	MCUC	MHNC-UP	MUHNAC	Total
Common quail Coturnix coturnix	–	6	44	7	–	–	–	–	–	57
Red–legged partridge Alectoris rufa	–	7	21	8	19	–	2	–	–	57
Ring–necked pheasant Phasianus colchicus *	–	–	–	–	–	–	–	–	–	0
Grey partridge Perdix perdix	–	–	7	–	–	–	–	–	–	7
Western capercaillie Tetrao urogallus	–	–	–	–	–	–	–	–	–	0
Black swan Cygnus atratus	–	–	–	1	–	–	–	–	–	1
Mute swan Cygnus olor	–	–	–	1	–	–	–	–	–	1
Greylag goose Anser anser	–	–	–	–	–	–	–	–	–	0
Common scoter Melanitta nigra	–	–	–	–	–	–	–	–	–	0
Red–breasted merganser Mergus serrator	–	–	–	–	–	–	–	–	–	0
Egyptian goose Alopochen aegyptiaca *	–	–	–	–	–	–	–	–	–	0
Common shelduck Tadorna tadorna	–	–	–	–	7	–	–	–	–	7
Ruddy shelduck Tadorna ferruginea	–	–	–	–	–	–	–	–	–	0
Marbled duck Marmaronetta angustirostris	–	–	–	–	–	–	–	–	–	0
Red–crested pochard Netta rufina	–	–	–	–	–	–	–	–	–	0
Common pochard Aythya ferina	–	–	–	–	–	–	–	–	–	0
Ferruginous duck Aythya nyroca	–	–	–	–	–	–	–	–	–	0
Tufted duck Aythya fuligula	–	–	–	–	–	–	–	–	–	0
Garganey Spatula querquedula	–	–	–	–	–	–	–	–	–	0
Northern shoveler Spatula clypeata	–	–	–	–	–	–	–	–	–	0
Gadwall Mareca strepera	–	–	–	–	–	–	–	–	–	0
Eurasian wigeon Mareca penelope	–	–	–	–	–	–	–	–	–	0
Mallard Anas platyrhynchos	–	–	13	8	–	–	2	–	–	23
Northern pintail Anas acuta	–	–	–	–	–	–	–	–	–	0
Eurasian teal Anas crecca	–	–	–	–	–	–	–	–	–	0
Little grebe Tachybaptus ruficollis	–	–	6	5	–	–	1	–	–	12
Great crested greb Podiceps cristatus	–	–	–	–	–	–	–	–	–	0
Black–necked greb Podiceps nigricollis	–	–	–	–	–	–	–	–	–	0
Greater flaming Phoenicopterus roseus	–	–	–	–	1	–	–	–	–	1
Rock dove Columba livia	–	4	–	–	–	–	–	–	–	4
Stock dove Columba oenas	–	–	–	–	–	–	–	–	–	0
Common wood pigeon Columba palumbus	–	–	4	2	–	–	2	–	–	8
Trocaz pigeon Columba trocaz	–	1	–	–	–	–	–	–	–	1
European turtle dove Streptopelia turtur	–	1	10	4	–	–	5	3	–	23
Eurasian collared dove Streptopelia decaocto	–	–	–	–	–	–	–	–	–	0
Black–bellied sandgrouse Pterocles orientalis	–	–	6	–	–	–	–	–	–	6
Pin–tailed sandgrouse Pterocles alchata	–	–	3	–	–	–	–	–	–	3
Red–necked nightjar Caprimulgus ruficollis	–	–	–	2	–	–	–	–	–	2
European nightjar Caprimulgus europaeus	–	–	5	2	–	–	–	–	–	7
Alpine swift Tachymarptis melba	–	–	–	–	–	–	–	–	–	0
White–rumped swift Apus caffer	–	–	–	–	–	–	–	–	–	0
Plain swift Apus unicolor	–	–	–	–	–	–	–	–	–	0
Pallid swift Apus pallidus	–	–	–	–	1	–	–	–	–	1
Common swift Apus apus	–	–	6	2	–	–	2	1	–	11
Great spotted cuckoo Clamator glandarius	–	–	–	–	–	–	–	–	–	0
Common cuckoo Cuculus canorus	–	–	4	–	–	–	–	–	–	4
Water rail Rallus aquaticus	–	–	–	7	–	–	–	–	–	7
Spotter crake Porzana porzana	–	–	–	–	–	–	–	–	–	0
Baillon’s crake Zapornia pusilla	–	–	–	–	–	–	–	–	–	0
Western swamphen Porphyrio porphyrio	–	–	–	–	–	–	–	–	–	0
Common moorhen Gallinula chloropus	–	–	13	25	7	–	2	–	1	48
Red–knobbed coot Fulica cristata	–	–	–	–	–	–	–	–	–	0
Eurasian coot Fulica atra	–	–	2	–	–	–	–	–	–	2
Demoiselle crane Anthropoides virgo	–	–	–	–	–	–	–	–	–	0
Common crane Grus grus	–	–	–	–	–	–	–	–	–	0
Little bustard Tetrax tetrax	–	–	12	–	–	–	–	–	–	12
Great bustard Otis tarda	–	–	6	–	–	–	–	–	–	6
Wilson’s storm–petrel Oceanites oceanicus	–	–	–	–	–	–	–	–	–	0
White–faced storm petrel Pelagodroma marina	–	8	–	–	2	–	–	–	–	10
European storm–petrel Hydrobates pelagicus	–	–	–	–	–	–	–	–	–	0
Band–rumped storm petrel Hydrobates castro	–	4	–	–	–	–	–	–	–	4
Monteiro’s storm petrel Hydrobates monteiroi	–	–	–	–	–	–	–	–	–	0
Leach’s storm–petrel Hydrobates leucorhous	–	–	–	–	–	–	–	–	–	0
Desertas petrel Pterodroma deserta	–	19	–	–	–	–	–	–	–	19
Zino’s petrel Pterodroma madeira	–	45	–	–	–	–	–	–	–	45
Sooty shearwater Ardenna grisea	–	–	–	–	–	–	–	–	–	0
Great shearwater Ardenna gravis	–	–	–	–	–	–	–	–	–	0
Cory’s shearwater Calonectris borealis	–	16	1	1	41	–	–	–	–	59
Manx shearwater Puffinus puffinus	3	2	–	–	–	–	–	–	–	5
Sargasso shearwater Puffinus lherminieri	–	–	–	–	–	–	–	–	–	0
Balearic shearwater Puffinus mauretanicus	–	–	–	–	–	–	–	–	–	0
Bulwer’s petrel Bulweria bulwerii	–	18	–	–	16	–	–	–	–	34
Black stork Ciconia nigra	–	–	7	5	1	–	1	–	–	14
White stork Ciconia ciconia	–	–	8	3	6	–	2	–	–	19
Eurasian spoonbill Platalea leucorodia	–	–	–	–	–	–	–	–	–	0
Glossy ibis Plegadis falcinellus	–	–	–	–	–	–	–	–	–	0
Eurasian bittern Botaurus stellaris	–	–	–	–	–	–	–	–	–	0
Little bittern Ixobrychus minutus	–	–	13	5	–	–	3	–	–	21
Black–crowned night heron Nycticorax nycticorax	–	–	–	–	–	–	–	–	–	0
Squacco heron Ardeola ralloides	–	–	–	–	–	–	–	–	–	0
Cattle egret Bubulcus ibis	–	–	–	–	–	–	–	–	–	0
Grey heron Ardea cinerea	–	–	–	–	–	–	–	–	–	0
Purple heron Ardea purpurea	–	–	–	5	–	–	–	–	–	5
Great egret Ardea alba	–	–	–	–	–	–	–	–	–	0
Little egret Egretta garzetta	–	–	–	8	–	–	–	–	–	8
Northern gannet Morus bassanus	–	–	–	–	–	–	–	–	–	0
European shag Gulosus aristotelis	–	–	9	–	–	–	2	–	–	11
Great cormorant Phalacrocorax carbo	–	–	–	–	–	–	–	–	–	0
Eurasian stone–curlew Burhinus oedicnemus	–	–	5	8	–	–	–	–	–	13
Eurasian oystercatcher Haematopus ostralegus	–	–	–	8	–	–	–	–	–	8
Pied avocet Recurvirostra avosetta	–	–	–	7	–	–	–	–	–	7
Black–winged stilt Himantopus himantopus	–	–	–	11	1	–	–	–	–	12
Grey plover Pluvialis squatarola	–	–	–	–	–	–	–	–	–	0
European golden plover Pluvialis apricaria	–	–	–	–	–	–	–	–	–	0
Common ringed plover Charadrius hiaticula	–	–	12	–	–	–	–	–	–	12
Semipalmated plover Charadrius semipalmatus	–	–	–	–	–	–	–	–	–	0
Little ringed plover Charadrius dubius	–	–	3	3	–	–	–	–	–	6
Kentish plover Charadrius alexandrinus	–	3	–	3	–	–	–	–	–	6
Eurasian dotterel Eudromias morinellus	–	–	–	–	–	–	–	–	–	0
Northern lapwing Vanellus vanellus	–	–	–	–	–	–	–	–	–	0
Eurasian whimbrel Numenius phaeopus	–	–	–	–	–	–	–	–	–	0
Eurasian curlew Numenius arquata	–	–	–	–	–	–	–	–	–	0
Bar–tailed godwit Limosa lapponica	–	–	–	–	–	–	–	–	–	0
Black–tailed godwit Limosa limosa	–	–	–	–	–	–	–	–	–	0
Ruddy turnstone Arenaria interpres	–	–	–	–	–	–	–	–	–	0
Red knot Calidris canutus	–	–	–	–	–	–	–	–	–	0
Ruff Calidris pugnax	–	–	–	–	–	–	–	–	–	0
Curlew sandpiper Calidris ferruginea	–	–	–	–	–	–	–	–	–	0
Sanderling Calidris alba	–	–	–	–	–	–	–	–	–	0
Dunlin Calidris alpina	–	–	–	–	–	–	–	–	–	0
Purple sandpiper Calidris maritima	–	–	–	–	–	–	–	–	–	0
Little stint Calidris minuta	–	–	–	–	–	–	–	–	–	0
Temminck’s stint Calidris temminckii	–	–	–	–	–	–	–	–	–	0
Eurasian woodcock Scolopax rusticola	–	2	–	–	–	–	–	–	–	2
Common snipe Gallinago gallinago	–	–	–	4	–	–	–	1	–	5
Jack snipe Lymnocryptes minimus	–	–	–	–	–	–	–	–	–	0
Common sandpiper Actitis hypoleucos	–	–	–	4	–	–	–	–	–	4
Spotted redshank Tringa erythropus	–	–	–	4	–	–	–	–	–	4
Green sandpiper Tringa ochropus	–	–	–	–	–	–	–	–	–	0
Common greenshank Tringa nebularia	–	–	–	–	–	–	–	–	–	0
Common redshank Tringa totanus	–	–	–	4	–	–	–	–	–	4
Wood sandpiper Tringa glareola	–	–	–	–	–	–	–	–	–	0
Red phalarope Phalaropus fulicarius	–	–	–	–	–	–	–	–	–	0
Common buttonquail Turnix sylvaticus	–	–	–	–	–	–	–	–	–	0
Collared pratincole Glareola pratincola	–	–	–	2	–	–	–	–	–	2
Little gull Hydrocoloeus minutus	–	–	–	–	–	–	–	–	–	0
Sabine’s gull Xema sabini	–	–	–	–	–	–	–	–	–	0
Black–legged kittiwake Rissa tridactyla	–	–	–	–	–	–	–	–	–	0
Black–headed gull Larus ridibundus	–	–	–	–	–	–	–	–	–	0
Mediterranean gull Larus melanocephalus	–	–	–	–	–	–	–	–	–	0
Audouin’s gull Larus audouinii	–	–	–	–	–	–	–	–	–	0
Lesser black–backed gull Larus fuscus	–	–	–	–	–	–	–	–	–	0
European herring gull Larus argentatus	2	15	–	10	–	–	–	–	–	27
Yellow–legged gull Larus michahellis	–	–	8	–	49	–	3	–	–	60
Great black–backed gull Larus marinus	2	–	–	–	–	–	–	–	–	2
Slender–billed gull Larus genei	–	–	–	–	–	–	–	–	–	0
Ring –billed gull Larus delawarensis	–	–	–	–	–	–	–	–	–	0
Mew gull Larus canus	–	–	–	–	–	–	–	–	–	0
Sooty tern Onychoprion fuscatus	–	–	–	–	–	–	–	–	–	0
Little tern Sternula albifrons	–	–	–	3	–	–	2	–	–	5
Gull–billed tern Gelochelidon nilotica	–	–	–	2	–	–	–	–	–	2
Caspian tern Hydroprogne caspia	–	–	–	–	–	–	–	–	–	0
Whiskered tern Chlidonias hybrida	–	–	–	–	–	–	–	–	–	0
Black tern Chlidonias niger	–	–	–	–	–	–	–	–	–	0
Roseate tern Sterna dougallii	–	–	–	–	–	–	–	–	–	0
Common tern Sterna hirundo	–	5	–	5	–	–	–	–	–	10
Arctic tern Sterna paradisaea	–	–	–	–	–	–	–	–	–	0
Sandwich tern Thalasseus sandvicensis	–	–	6	–	–	–	–	–	–	6
Long–tailed jaeger Stercorarius longicaudus	–	–	–	–	–	–	–	–	–	0
Arctic jaeger Stercorarius parasiticus	–	–	–	–	–	–	–	–	–	0
Pomarine jaeger Stercorarius pomarinus	–	–	–	–	–	–	–	–	–	0
Great skua Catharacta skua	–	–	–	–	–	–	–	–	–	0
Atlantic puffin Fratercula arctica	–	–	–	–	–	–	–	–	–	0
Razorbill Alca torda	–	–	–	1	–	–	–	–	–	1
Common murre Uria aalge	7	–	5	1	–	–	–	–	–	13
Barn owl Tyto alba	–	1	4	–	2	–	–	–	–	7
Little owl Athene noctua	–	–	6	–	2	–	–	–	–	8
Scops owl Otus scops	–	–	12	–	–	–	–	–	–	12
Long–eared owl Asio otus	–	–	–	–	–	–	–	–	–	0
Short–eared owl Asio flammeus	–	–	–	–	–	–	–	–	–	0
Tawny owl Strix aluco	–	–	–	8	–	–	–	–	–	8
Eurasian eagle–owl Bubo bubo	–	–	3	2	–	–	–	–	–	5
Osprey Pandion haliaetus	–	–	–	–	–	–	–	–	–	0
Black–winged kite Elanus caeruleus	–	–	–	–	–	–	–	–	–	0
European honey buzzard Pernis apivorus	–	–	–	4	–	–	–	–	–	4
Bearded vulture Gypaetus barbatus	–	–	–	–	–	–	–	–	–	0
Egyptian vulture Neophron percnopterus	–	–	–	9	–	–	–	–	–	9
Short–toed snake eagle Circaetus gallicus	–	–	–	6	–	–	–	–	–	6
Rüppell’s vulture Gyps rueppelli	–	–	–	–	–	–	–	–	–	0
Eurasian griffon vulture Gyps fulvus	–	–	1	2	–	–	1	–	–	4
Cinereous vulture Aegypius monachus	–	–	1	–	–	–	–	–	–	1
Spanish imperial eagle Aquila adalberti	–	–	3	–	–	–	1	–	–	4
Golden eagle Aquila chrysaetos	–	–	4	–	–	–	1	–	–	5
Bonelli’s eagle Aquila fasciata	–	–	–	–	–	–	–	–	–	0
Booted eagle Hieraaetus pennatus	–	–	–	12	–	–	–	–	–	12
Western marsh harrier Circus aeruginosus	–	–	3	8	–	–	1	–	–	12
Hen harrier Circus cyaneus	–	–	1	3	–	–	–	–	–	4
Montagu’s harrier Circus pygargus	–	–	–	3	–	–	–	1	–	4
Eurasian sparrowhawk Accipiter nisus	–	3	9	–	–	1	2	–	–	15
Northern goshawk Accipiter gentilis	–	–	–	–	–	–	–	–	–	0
Red kite Milvus milvus	–	–	5	6	–	–	3	–	–	14
Black kite Milvus migrans	–	–	–	10	–	–	–	–	–	10
Common buzzard Buteo buteo	–	2	8	31	–	–	2	–	–	43
Eurasian hoopoe Upupa epops	–	1	6	27	–	–	–	1	–	35
European bee–eater Merops apiaster	–	–	15	5	–	–	–	–	–	20
European roller Coracias garrulus	–	–	–	–	9	–	–	–	–	9
Common kingfisher Alcedo atthis	–	–	10	7	–	–	–	–	–	17
Eurasian wryneck Jynx torquilla	–	–	1	–	–	–	–	1	–	2
Iberian green woodpecker Picus sharpei	–	–	25	5	–	–	1	–	–	31
Lesser spotted woodpecker Dryobates minor	–	–	3	–	–	–	–	–	–	3
Great spotted woodpecker Dendrocopos major	–	–	–	8	–	–	–	–	–	8
Lesser kestrel Falco naumanni	–	–	–	9	54	–	–	–	–	63
Common kestrel Falco tinnunculus	–	4	15	21	16	–	3	–	1	60
Eleonora’s falcon Falco eleonorae	–	–	–	–	–	–	–	–	–	0
Merlin Falco columbarius	–	–	–	–	–	–	–	–	–	0
Eurasian hobby Falco subbuteo	–	–	–	–	–	–	–	–	–	0
Peregrine falcon Falco peregrinus	–	–	–	–	–	–	–	–	–	0
Senegal parrot Poicephalus senegalus*	–	–	–	–	–	–	–	–	–	0
Monk parakeet Myiopsitta monachus *	–	–	–	–	–	–	–	–	–	0
Blue–crowned parakeet Psittacara acuticaudatus*	–	–	–	–	–	–	–	–	–	0
Fischer’s lovebird Agarpornis fischeri *	–	–	–	–	–	–	–	–	–	0
Rose–ringed parakeet Alexandrinus krameri *	–	–	–	–	–	–	–	–	–	0
Eurasian Golden Oriole Oriolus oriolus	–	–	6	5	–	–	3	1	–	15
Red–backed shrike Lanius collurio	–	–	–	1	–	–	–	1	–	2
Iberian grey shrike Lanius meridionalis	–	–	–	–	–	–	–	–	–	0
Woodchat shrike Lanius senator	–	–	26	21	–	–	6	1	–	54
Red–billed chough Pyrrhocorax pyrrhocorax	–	–	4	–	–	–	1	–	–	5
Iberian magpie Cyanopica cooki	–	–	11	6	–	–	2	1	–	20
Eurasian jay Garrulus glandarius	–	–	28	16	4	–	6	2	1	57
Eurasian magpie Pica pica	–	–	15	22	–	–	3	–	–	40
Western jackdaw Corvus monedula	–	–	–	–	64	–	–	–	–	64
Rook Corvus frugilegus	–	–	–	–	–	–	–	–	–	0
Common raven Corvus corax	–	–	–	–	–	–	–	–	–	0
Carrion crow Corvus corone	–	–	8	–	–	–	2	2	–	12
Coal tit Periparus ater	–	–	–	–	–	–	–	–	–	0
Crested tit Lophophanes cristatus	–	–	8	10	–	–	2	1	–	21
Eurasian blue tit Cyanistes caeruleus	–	–	6	11	–	–	1	1	–	19
Great tit Parus major	–	–	18	15	–	–	3	1	2	39
Eurasian penduline tit Remiz pendulinus	–	–	–	–	–	–	–	–	1	1
Dupont’s lark Chersophilus duponti	–	–	–	–	–	–	–	–	–	0
Mediterranean short–toed lark Alaudala rufescens	–	–	–	–	–	–	–	–	–	0
Calandra lark Melanocorypha calandra	–	–	4	7	–	–	1	–	–	12
Greater short–toed lark Calandrella brachydactyla	–	–	3	7	–	–	1	2	–	13
Woodlark Lullula arborea	–	–	16	16	–	–	4	1	–	37
Eurasian skylark Alauda arvensis	–	–	4	11	–	–	1	–	–	16
Thekla’s lark Galerida theklae	–	–	–	–	–	–	–	–	–	0
Crested lark Galerida cristata	–	–	13	22	–	–	3	1	–	39
Zitting cisticola Cisticola juncidis	–	–	2	4	–	–	2	2	1	11
Savi’s warbler Locustella luscinioides	–	–	–	–	–	–	–	–	–	0
Common grasshopper warbler Locustella naevia	–	–	–	–	–	–	–	–	–	0
Western olivaceous warbler Iduna opaca	–	–	–	–	–	–	–	–	–	0
Melodious warbler Hippolais polyglotta	–	–	13	19	–	–	4	2	1	39
Eurasian reed warbler Acrocephalus scirpaceus	–	–	4	3	–	–	1	–	1	9
Great reed warbler Acrocephalus arundinaceus	–	–	17	24	–	–	3	–	–	44
Aquatic warbler Acrocephalus paludicola	–	–	–	–	–	–	–	–	–	0
Sedge warbler Acrocephalus schoenobaenus	–	–	–	–	–	–	–	–	–	0
Western house martin Delichon urbicum	–	–	8	3	–	–	3	–	–	14
European red–rumped swallow Cecropis daurica	–	–	–	–	–	–	–	–	–	0
Barn swallow Hirundo rustica	–	–	14	8	2	–	3	2	–	29
Eurasian crag martin Ptyonoprogne rupestris	–	–	–	–	–	–	–	–	–	0
Sand martin Riparia riparia	–	–	–	3	–	–	–	–	–	3
Western Bonelli‘s warbler Phylloscopus bonelli	–	–	–	6	–	–	–	–	–	6
Iberian chiffchaff Phylloscopus ibericus	–	–	–	–	–	–	–	–	–	0
Common chiffchaff Phylloscopus collybita	–	–	13	11	–	–	4	2	–	30
Willow warbler Phylloscopus trochilus	–	–	–	5	–	–	–	–	–	5
Cetti’s warbler Cettia cetti	–	–	4	–	–	–	1	–	–	5
Long–tailed tit Aegithalos caudatus	–	–	19	–	–	–	3	–	–	22
Eurasian blackcap Sylvia atricapilla	–	5	32	2	–	4	8	2	1	54
Garden warbler Sylvia borin	–	–	–	–	–	–	–	–	–	0
Western Orphean warbler Curruca hortensis	–	–	–	8	–	–	–	1	–	9
Sardinian warbler Curruca melanocephala	–	–	3	5	–	–	1	1	4	14
Eastern subalpine warbler Curruca cantillans	–	–	–	3	–	–	–	–	–	3
Common whitethroat Curruca communis	–	–	18	15	–	–	5	3	–	41
Spectacled warbler Curruca conspicillata	–	–	–	–	–	1	–	–	–	1
Dartford warbler Curruca undata	–	–	4	8	–	–	1	1	–	14
Red–billed leiothrix Leiothrix lutea *	–	–	–	–	–	1	–	–	–	1
Short–toed treecreeper Certhia brachydactyla	–	–	7	–	–	–	2	–	–	9
Eurasian nuthatch Sitta europaea	–	–	7	15	–	–	–	–	–	22
Eurasian wren Troglodytes troglodytes	–	–	17	15	–	–	8	3	4	47
White–throated dipper Cinclus cinclus	–	–	8	–	–	–	3	–	–	11
Common starling Sturnus vulgaris	–	–	–	–	–	–	–	–	–	0
Spotless starling Sturnus unicolor	–	–	6	5	10	–	1	–	–	22
Crested myna Acridotheres cristatellus *	–	–	–	–	–	–	–	–	–	0
Mistle thrush Turdus viscivorus	–	–	16	11	–	–	5	2	–	34
Song thrush Turdus philomelos	–	–	–	–	–	–	–	–	–	0
Redwing Turdus iliacus	–	–	–	–	–	–	–	–	–	0
Common blackbird Turdus merula	–	6	31	19	–	7	7	4	8	82
Fieldfare Turdus pilaris	–	–	–	–	–	–	–	–	–	0
Ring ouzel Turdus torquatus	–	–	–	–	–	–	–	–	–	0
Rufous–tailed scrub robin Cercotrichas galactotes	–	–	9	–	–	–	3	–	–	12
Spotted flycatcher Muscicapa striata	–	–	4	–	–	–	1	–	–	5
European robin Erithacus rubecula	–	4	20	7	–	4	5	2	–	42
Bluethroat Luscinia svecica	–	–	–	–	–	–	–	–	–	0
Common nightingale Luscinia megarhynchos	–	–	16	25	–	–	4	–	1	46
Black redstart Phoenicurus ochruros	–	–	2	11	–	–	1	2	1	17
Common redstart Phoenicurus phoenicurus	–	–	–	18	–	–	–	–	–	18
Common rock thrush Monticola saxatilis	–	–	1	–	–	–	1	–	–	2
Blue rock thrush Monticola solitarius	–	–	1	3	–	–	1	2	–	7
Whinchat Saxicola rubetra	–	–	–	17	–	–	–	–	–	17
European stonechat Saxicola rubicola	–	–	30	19	–	–	8	1	2	60
Northern wheatear Oenanthe oenanthe	–	–	–	2	–	–	–	1	–	3
Western black–eared wheatear Oenanthe hispanica	–	–	17	5	–	–	4	–	–	26
Black wheatear Oenanthe leucura	–	–	1	–	–	–	1	–	–	2
European pied flycatcher Ficedula hypoleuca	–	–	–	4	–	–	–	–	–	4
Goldcrest Regulus regulus	–	–	–	–	–	–	–	–	–	0
Madeira firecrest Regulus madeirensis	–	–	–	–	–	–	–	–	–	0
Common firecrest Regulus ignicapilla	–	3	–	–	–	5	–	–	–	8
Alpine accentor Prunella collaris	–	–	–	–	–	–	–	–	–	0
Dunnock Prunella modularis	–	–	3	9	–	–	1	4	–	17
Yellow–crowned bishop Euplectes afer *	–	–	–	–	–	–	–	–	–	0
Black–headed weaver Ploceus melanocephalus *	–	–	–	–	–	–	–	–	–	0
Common waxbill Estrilda astrild *	–	–	–	–	–	–	–	–	1	1
Red avadavat Amandava amandava *	–	–	–	–	–	–	–	–	–	0
Scaly–breasted munia Lonchura punctulata *	–	–	–	–	–	–	–	–	–	0
Pin–tailed whydah Vidua macroura *	–	–	–	–	–	–	–	–	–	0
House sparrow Passer domesticus	–	–	27	32	1	–	4	2	1	67
Spanish sparrow Passer hispaniolensis	–	2	–	4	6	1	–	–	–	13
Eurasian tree sparrow Passer montanus	–	–	–	1	–	–	–	–	–	1
Rock sparrow Petronia petronia	–	4	4	1	–	2	1	–	–	12
Tree pipit Anthus trivialis	–	–	–	–	–	–	–	–	–	0
Meadow pipit Anthus pratensis	–	–	–	–	–	–	–	–	–	0
Water pipit Anthus spinoletta	–	–	–	–	–	–	–	–	–	0
Berthelot’s pipit Anthus berthelotii	–	2	–	–	–	1	–	–	–	3
Tawny pipit Anthus campestris	–	–	18	–	–	–	4	–	–	22
Richard’s pipit Anthus richardi	–	–	–	–	–	–	–	–	–	0
Western yellow wagtail Motacilla flava	–	–	–	8	–	–	–	–	–	8
Grey wagtail Motacilla cinerea	–	4	7	3	–	3	2	1	–	20
White wagtail Motacilla alba	–	–	9	4	–	–	2	1	–	16
Eurasian chaffinch Fringilla coelebs	–	5	21	13	–	2	5	2	1	49
Brambling Fringilla montifringilla	–	–	–	–	–	–	–	–	–	0
Hawfinch Coccothraustes coccothraustes	–	–	–	–	–	–	–	–	–	0
Azores bullfinch Pyrrhula murina	–	–	–	–	–	–	–	–	–	0
Eurasian bullfinch Pyrrhula pyrrhula	–	–	–	–	–	–	–	–	–	0
European greenfinch Chloris chloris	–	–	11	27	–	1	4	3	2	48
Common linnet Linaria cannabina	–	–	26	18	–	1	6	1	1	53
Red crossbill Loxia curvirostra	–	–	–	–	–	–	–	–	–	0
European goldfinch Carduelis carduelis	–	–	39	8	1	–	14	3	4	69
European serin Serinus serinus	–	–	–	13	–	–	–	6	2	21
Atlantic canary Serinus canaria	–	2	19	–	–	2	6	–	–	29
Eurasian siskin Spinus spinus	–	–	–	–	–	–	–	–	–	0
Corn bunting Emberiza calandra	–	–	14	20	–	–	4	1	–	39
Rock bunting Emberiza cia	–	–	11	10	–	–	3	3	–	27
Ortolan bunting Emberiza hortulana	–	–	–	–	–	–	–	–	–	0
Cirl bunting Emberiza cirlus	–	–	28	25	–	–	7	–	–	60
Yellowhammer Emberiza citrinella	–	–	–	–	–	–	–	–	–	0
Common reed bunting Emberiza schoeniclus	–	–	–	–	–	–	–	–	–	0
Snow bunting Plectrophenax nivalis	–	–	–	–	–	–	–	–	–	0
Total	14	204	1125	1036	322	36	240	83	42	3102

Results

A total of 5,986 specimens with associated data, prepared either as study skins or taxidermy mounts and belonging to 297 species, were recorded (Table 1). Additionally, 138 specimens from 41 rare or accidental species are extant in the collections (Table 3), as well as 2,701 eggs and 401 nests from 170 taxa (Table 4). Detailed results are provided in the following subsections.

Taxonomic representation

Of the 327 species known to occur regularly in Portugal, 297 are represented in the country’s NHCs. Among the non-represented taxa are three regionally extinct species—the western capercaillie Tetrao urogallus, the demoiselle crane Anthropoides virgo, and Eleonora’s falcon Falco eleonorae—as well as several exotic species (see “Exotic species” section below). Most species extant in the examined collections are resident breeders in Portugal or wintering species (Tables 1, 2). In addition, the collections also house some more elusive species for which there are fewer records (see “Rare or accidental species” section below; Table 3). Distribution maps were not produced for the following species due to lack of spatial data from specimens: Wilson’s storm petrel Oceanites oceanicus, Audouin’s gull Larus audouinii, the roseate tern Sterna dougallii, the European honey buzzard Pernis apivorus, and the red-billed leiothrix Leiothrix lutea. No maps were produced for some species with a conservation status of regionally extinct, despite the existence of spatial data from specimens, as the atlas did not provide historically known distributions. This is the case for the grey partridge Perdix perdix, the rook Corvus frugilegus, the common buttonquail Turnix sylvaticus, and Dupont’s lark Chersophilus duponti.

Analyzing taxonomic representation within each institution shows MCUC’s collection as the most taxonomically diverse, with 256 of 327 species represented (78.3%), while holding 20.9% of all existing specimens extant in the assessed Portuguese NHCs (Table 1). MHNC-UP’s collection covers 70.9% (232 of 327) of occurring taxa, corresponding to 18.7% of all assessed specimens, while MUHNAC’s collection represents 65.1% (213 of 327) of species, corresponding to 42.6% of specimens. MMF’s collection covers 45.6% (149 of 327) of species, corresponding to 13.8% of specimens, and AVG’s collection represents 23.9% (78 of 327) of taxa, corresponding to 3.9% of specimens.

Two passerine species—the Spanish sparrow Passer hispaniolensis (423 specimens) and the house sparrow Passer domesticus (211 specimens)—and a gull, the yellow-legged gull Larus michahellis (137 specimens), are the three most represented species in the collections, making up 12.9% of all specimens. Approximately 85% of extant taxa in the collections are represented by 50 or fewer specimens, with only 18 species represented by more (Table 1). A little over 30% of taxa (116) are represented by 10 or fewer specimens. Order Passeriformes is the most numerous in the collections, both in species represented (116) and in number of extant specimens (2,909); it is followed by order Charadriiformes with 58 represented species and 951 specimens and order Accipitriformes with 20 represented species and 242 specimens (Fig. 2). More detailed descriptions are presented below.

Figure 2.

Proportion of Portuguese bird specimens extant in the country’s NHCs by taxonomic order. The pie chart order is alphabetical and clockwise.

Order Galliformes

Five species are known from Portugal, all from the family Phasianidae. Of these, only the western capercaillie Tetrao urogallus is not represented in the collections. The remaining four species account for a total of 89 specimens in the collections. The red-legged partridge Alectoris rufa is the most represented species, with 53 specimens that cover 2.7% of its known distribution (Fig. 8C), followed by the common quail Coturnix coturnix with 27 specimens (Fig. 8A; 1.8%), the gray partridge Perdix perdix with seven specimens, and the ring-necked pheasant Phasianus colchicus with two specimens (Fig. 8B; 2%). Both P. perdix and T. urogallus are considered regionally extinct (Almeida et al. 2022).

Order Anseriformes

Twenty species, all from the family Anatidae, occur in Portugal. Sixteen of them are represented in the collections by a total of 178 specimens. The Eurasian teal Anas crecca is the most represented, with 21 specimens covering 11.3% of its known distribution (Fig. 9I); followed by the tufted duck Aythya fuligula with 20 specimens covering 20% of its known distribution (Fig. 9B); the Eurasian wigeon Mareca penelope and the mallard Anas platyrhynchos with 17 specimens each (Fig. 9F; 19.4% and Fig. 9G; 0.7%, respectively); the common scoter Melanitta nigra and the common pochard Aythya ferina with 15 specimens each (Fig. 8E; 17.4% and Fig. 9A; 11%, respectively); the northern shoveler Spatula clypeata with 13 specimens (Fig. 9D; 3.3%); the northern pintail Anas acuta with 12 specimens (Fig. 9H; 9.7%); the common shelduck Tadorna tadorna with 10 specimens (Fig. 8G; 13.6%); the greylag goose Anser anser and the garganey Spatula querquedula with seven specimens each (Fig. 8D; 6.5% and Fig. 9C; 14.3%, respectively); the red-breasted merganser Mergus serrator and the ferruginous duck Aythya nyroca with six specimens each (Fig. 8F; 27.3% and Fig. 8J; 4.5%, respectively); the marbled duck Marmaronetta angustirostris and the ruddy shelduck Tadorna ferruginea with four specimens each (Fig. 8I; 50% and Fig. 8H; 7.7%, respectively); and the gadwall Mareca strepera with one specimen (Fig. 9E; 0.5%). The four species not represented in the collections are the black swan Cygnus atratus, the mute swan Cygnus olor, the Egyptian goose Alopochen aegyptiaca, and the red-crested pochard Netta rufina.

Order Podicipediformes

Three species from the family Podicipedidae occur in Portugal, and all are present in the collections, represented by a total of 43 specimens. The little grebe Tachybaptus ruficollis is the most represented species, with 22 specimens (Fig. 10A; 2.4%), followed by the black-necked grebe Podiceps nigricollis with 15 specimens (Fig. 10C; 20.9%) and the great crested grebe Podiceps cristatus with six specimens (Fig. 10B; 1.3%).

Order Phoenicopteriformes

Only one species occurs in the country, the greater flamingo, Phoenicopterus roseus. It is represented in the collections by nine specimens that cover 7.1% of its known distribution (Fig. 10D).

Order Columbiformes

Six species, all from the family Columbidae, are found in Portugal and are represented in its NHCs by 82 specimens. The trocaz pigeon Columba trocaz is the most represented in the collections, with 29 specimens that cover 50% of its known distribution (Fig. 10H), followed by the European turtle dove Streptopelia turtur with 22 specimens (Fig. 11A; 1.4%), the rock dove Columba livia with 13 specimens (Fig. 10E; 0.4%), the common wood pigeon Columba palumbus with 12 specimens (Fig. 10G; 0.5%), the stock dove Columba oenas with four specimens (Fig. 10F; 4.4%), and the Eurasian collared dove Streptopelia decaocto with two specimens (Fig. 10J; 0.2%).

Order Pterocliformes

Two species from the family Pteroclidae are known to occur in Portugal, and both are represented in the collections by a total of nine specimens. The black-bellied sandgrouse Pterocles orientalis is represented by six specimens (Fig. 10I; 8.1%), while the pin-tailed sandgrouse Pterocles alchata is represented by three specimens (Fig. 10K; 14.3%).

Order Caprimulgiformes

Seven species from the families Caprimulgidae and Apodidae are known from Portugal. Of those, six are represented in the collections by a total of 53 specimens. Both Caprimulgidae species are recorded in the NHCs: the European nightjar Caprimulgus europaeus with 13 specimens (Fig. 11C; 2.9%) and the red-necked nightjar Caprimulgus ruficollis with seven specimens (Fig. 11B; 2.1%). Four of the five species of the family Apodidae are present in the collections: 14 specimens of the common swift Apus apus (Fig. 11G; 0.9%), nine specimens of the pallid swift Apus pallidus (Fig. 11F; 1%), six specimens of the alpine swift Tachymarptis melba (Fig. 11D; 4.5%), and four specimens of the plain swift Apus unicolor (Fig. 11E; 11.8%). No specimens of the white-rumped swift Apus caffer are found in the collections.

Order Cuculiformes

Two species from the family Cuculidae occur in the country and are represented in its collections by 42 specimens. The common cuckoo Cuculus canorus is represented by 23 specimens (Fig. 11I; 1.7%), while the great spotted cuckoo Clamator glandarius is represented by 19 specimens (Fig. 11H; 7.1%).

Order Gruiformes

Nine species from the families Rallidae and Gruidae are found in Portugal. Eight of those are represented in the collections by 114 specimens, but the collections hold no specimens of the demoiselle crane Anthropoides virgo, regionally extinct in the country (Almeida et al. 2022). The family Rallidae is represented by seven species: the common moorhen Gallinula chloropus with 36 specimens (Fig. 12E; 3.3%), the Eurasian coot Fulica atra with 12 specimens (Fig. 12H; 3.9%), the water rail Rallus aquaticus with 17 specimens (Fig. 12A; 6.6%), the western swamphen Porzana porzana with 13 specimens (Fig. 12B; 72.7%), the red-knobbed coot Fulica cristata with eight specimens (Fig. 12F; 41.7%), and the spotted crake Porphyrio porphyrio and Baillon’s crake Zapornia pusilla with six specimens each (Fig. 12D; 6.7% and Fig. 12C; 57.1%, respectively). The common crane, Grus grus, is the only representative of the family Gruidae in the collections, with six specimens (Fig. 12G; 12.1%).

Order Otidiformes

Two species from the family Otididae occur in the country and are represented in the collections by 19 specimens. The little bustard Tetrax tetrax is represented by 10 specimens (Fig. 12I; 4.1%), while the great bustard Otis tarda is represented by nine specimens (Fig. 13A; 10.8%).

Order Procellariiformes

Fifteen species from the families Oceanitidae, Hydrobatidae, and Procellariidae occur in Portugal and are represented in the collections by 334 specimens. The family Oceanitidae is represented by the white-faced storm petrel Pelagodroma marina with 18 specimens (Fig. 13B; 100%) and Wilson’s storm-petrel Oceanites oceanicus with four specimens. The family Hydrobatidae is represented by four species: the band-rumped storm petrel Hydrobates castro with 37 specimens (Fig. 13D; 50%), Leach’s storm-petrel Hydrobates leucorhous with 21 specimens (Fig. 13F; 70%), the European storm-petrel Hydrobates pelagicus with four specimens (Fig. 13C; 33%), and Monteiro’s storm petrel Hydrobates monteiroi with one specimen (Fig. 13E; 25%). The family Procellariidae is represented by nine species, of which eight are represented in the collections: Zino’s petrel Pterodroma madeira with 57 specimens (Fig. 13H; 33.3%), Puffinus puffinus with 54 specimens (Fig. 14A; 23.5%), Cory’s shearwater Calonectris borealis with 53 specimens (Fig. 13K; 17.1%), Bulwer’s petrel Bulweria bulwerii with 34 specimens (Fig. 14C; 88.9%), the Desertas petrel Pterodroma deserta with 30 specimens (Fig. 13G; 50%), the sooty shearwater Ardenna grisea with six specimens (Fig. 13I; 40%), the great shearwater Ardenna gravis with six specimens (Fig. 13J; 33.3%), and the Balearic shearwater Puffinus mauretanicus with six specimens (Fig. 14B; 12.5%). The sargasso shearwater Puffinus lherminieri is the only species not represented in the NHCs.

Order Ciconiiformes

Two species from the family Ciconiidae occur in the country and are represented in its collections by 13 specimens. The white stork Ciconia ciconia is represented by 10 specimens (Fig. 14D; 0.7%), while the black stork Ciconia nigra is represented by five specimens (Fig. 14E; 2%).

Order Pelecaniformes

Eleven species, two from the family Threskiornithidae and nine from the family Ardeidae, occur in Portugal and are represented in the collections by 122 specimens. Only the great egret Ardea alba is not represented in the country’s NHCs. The family Threskiornithidae is represented by 13 specimens of the Eurasian spoonbill Platalea leucorodia (Fig. 14F; 3.8%) and eight specimens of the glossy ibis Plegadis falcinellus (Fig. 14G; 4.8%). The family Ardeidae is represented in the collections by eight species: the gray heron Ardea cinerea with 35 specimens (Fig. 15C; 2.1%), the little bittern Ixobrychus minutus with 28 specimens (Fig. 14J; 19%), the cattle egret Bubulcus ibis with 22 specimens (Fig. 15B; 3.9%), the little egret Egretta garzetta with 21 specimens (Fig. 15E; 3.1%), the Eurasian bittern Botaurus stellaris with 15 specimens (Fig. 14I; 57.1%), the black-crowned night heron Nycticorax nycticorax with 15 specimens (Fig. 15A; 10.2%), the squacco heron Ardeola ralloides with 12 specimens (Fig. 14H; 11.1%), and the purple heron Ardea purpurea with 11 specimens (Fig. 15D; 4.1%).

Order Suliformes

One species from the family Sulidae and two from the family Phalacrocoracidae occur in Portugal and are represented in the collections by 79 specimens. The northern gannet Morus bassanus is the sole representative of the family Sulidae, with 49 specimens (Fig. 15G; 32.4%). The family Phalacrocoracidae is represented by the European shag Gulosus aristotelis with 15 specimens (Fig. 15F; 11.9%) and the great cormorant Phalacrocorax carbo with 15 specimens (Fig. 15H; 1.8%).

Order Charadriiformes

Sixty-seven species from 10 families occur in Portugal, with 60 species represented in the collections by 1,092 specimens. The family Burhinidae is represented by the Eurasian stone-curlew Burhinus oedicnemus with 17 specimens (Fig. 16A; 4.1%). The family Haematopodidae is represented by the Eurasian oystercatcher Haematopus ostralegus with 16 specimens (Fig. 16B; 17.1%). The family Recurvirostridae is represented by the black-winged stilt Himantopus himantopus with 25 specimens (Fig. 16E; 4.6%) and the pied avocet Recurvirostra avosetta with 19 specimens (Fig. 16C; 12%). The family Charadriidae is represented by 141 specimens of seven species: the northern lapwing Vanellus vanellus with 50 specimens (Fig. 17A; 9.8%); the European golden plover Pluvialis apricaria with 24 specimens (Fig. 16G; 10.2%); the little ringed plover Charadrius dubius with 17 specimens (Fig. 16H; 2.9%); the gray plover Pluvialis squatarola with 17 specimens (Fig. 16D; 14.8%); the Kentish plover Charadrius alexandrinus with 15 specimens (Fig. 16J; 6.4%); the common ringed plover Charadrius hiaticula with 13 specimens (Fig. 16F; 6.9%); and the Eurasian dotterel Eudromias morinellus with five specimens (Fig. 16I; 33.3%). No specimens of the semipalmated plover Charadrius semipalmatus are present in the country’s NHCs. The family Scolopacidae is represented by 344 specimens of 23 species, including the dunlin Calidris alpina with 46 specimens (Fig. 18B; 19%), the common snipe Gallinago gallinago with 36 specimens (Fig. 18H; 5.8%), the ruddy turnstone Arenaria interpres with 25 specimens (Fig. 17F; 12.3%), the black-tailed godwit Limosa limosa with 23 specimens (Fig. 17D; 16.4%), the Eurasian whimbrel Numenius phaeopus with 21 specimens (Fig. 17B; 13%), the common redshank Tringa totanus with 18 specimens (Fig. 19E; 14.1%), the red knot Calidris canutus with 17 specimens (Fig. 17G; 18.9%), the sanderling Calidris alba with 17 specimens (Fig. 18A; 12.3%), the Eurasian curlew Numenius arquata with 14 specimens (Fig. 17C; 16.3%), the green sandpiper Tringa ochropus with 14 specimens (Fig. 19C; 3.6%), the jack snipe Lymnocryptes minimus with 13 specimens (Fig. 19A; 53.3%), the common sandpiper Actitis hypoleucos with 13 specimens (Fig. 18F; 2.1%), the bar-tailed godwit Limosa lapponica with 12 specimens (Fig. 17E; 11.9%), the ruff Calidris pugnax with 12 specimens (Fig. 17H; 19.4%), the curlew sandpiper Calidris ferruginea with 12 specimens (Fig. 17I; 16.7%), the little stint Calidris minuta with 12 specimens (Fig. 18D; 16.7%), the red phalarope Phalaropus fulicarius with 12 specimens (Fig. 19G; 54.5%), the Eurasian woodcock Scolopax rusticola with 11 specimens (Fig. 18G; 8.2%), the common greenshank Tringa nebularia with nine specimens (Fig. 19D; 6%), the wood sandpiper Tringa glareola with six specimens (Fig. 19F; 22.2%), the purple sandpiper Calidris maritima with two specimens (Fig. 18C; 22.2%), the spotted redshank Tringa erythropus with two specimens (Fig. 19B; 7.1%), and Temminck’s stint Calidris temminckii with one specimen (Fig. 18E; 12.5%). The family Turnicidae is represented by the regionally extinct common buttonquail Turnix sylvaticus with seven specimens. The family Glareolidae is represented by the collared pratincole Glareola pratincola with seven specimens (Fig. 19H; 3.5%). The family Laridae is represented by 23 species, five of which are not represented in the collections: Sabine’s gull Xema sabini, the ring-billed gull Larus delawarensis, the sooty tern Onychoprion fuscatus, the Caspian tern Hydroprogne caspia, and the Arctic tern Sterna paradisaea. The remaining 18 species are represented as follows: the yellow-legged gull Larus michahellis with 137 specimens (Fig. 20D; 2.2%); the European herring gull Larus argentatus with 25 specimens (Fig. 20F; 12.5%); the little tern Sternula albifrons with 23 specimens (Fig. 20H; 14.5%); the common tern Sterna hirundo with 23 specimens (Fig. 21A; 7.3%); the lesser black-backed gull Larus fuscus with 22 specimens (Fig. 20B; 3.7%); the black-legged kittiwake Rissa tridactyla with 21 specimens (Fig. 19J; 68.8%); the black-headed gull Larus ridibundus with 20 specimens (Fig. 20A; 4.6%); the sandwich tern Thalasseus sandvicensis with 16 specimens (Fig. 21C; 11.1%); the black tern Chlidonias niger with 14 specimens (Fig. 20K; 35%); the gull-billed tern Gelochelidon nilotica with 11 specimens (Fig. 20I; 4.3%); the Mediterranean gull Larus melanocephalus with seven specimens (Fig. 19K; 1.5%); the great black-backed gull Larus marinus with seven specimens (Fig. 20C; 5.3%); the whiskered tern Chlidonias hybrida with six specimens (Fig. 20J; 18.8%); the common gull Larus canus with two specimens (Fig. 20G; 9.1%); the roseate tern Sterna dougallii with two specimens; the little gull Hydrocoleus minutus with one specimen (Fig. 19I; 6.7%); Audouin’s gull Larus audouinii with one specimen; and the slender-billed gull Larus genei with one specimen (Fig. 20E; 10%). The family Stercorariidae is represented by four species, one of which is not present in the collections—the long-tailed jaeger Stercorarius longicaudus—while the other three are: the Arctic jaeger Stercorarius parasiticus with four specimens (Fig. 21B; 20%), the great skua Catharacta skua with four specimens (Fig. 21D; 8.7%), and the pomarine jaeger Stercorarius pomarinus with one specimen (Fig. 21E; 10%). The family Alcidae is represented by three species: the razorbill Alca torda with 105 specimens (Fig. 21G; 41.2%), the common murre Uria aalge with 37 specimens (Fig. 21H; 73.3%), and the Atlantic puffin Fratercula arctica with 26 specimens (Fig. 21F; 78.6%).

Order Strigiformes

Seven species from the families Tytonidae and Strigidae are represented by 224 specimens in the collections. The family Tytonidae is represented by a single species, the barn owl Tyto alba, with 77 specimens that account for 8.6% of its known distribution (Fig. 21I). The family Strigidae is represented by six species: the little owl Athene noctua with 46 specimens (Fig. 21J; 4.8%); the tawny owl Strix aluco with 39 specimens (Fig. 22D; 4.5%); the Eurasian eagle-owl Bubo bubo with 18 specimens (Fig. 22E; 4.5%); the short-eared owl Asio flammeus with 17 specimens (Fig. 22C; 22.2%); the Eurasian scops owl Otus scops with 15 specimens (Fig. 22A; 4.8%); and the long-eared owl Asio otus with 12 specimens (Fig. 22B; 6.3%).

Order Accipitriformes

Twenty-one species from the families Pandionidae and Accipitridae are known from Portugal, with 20 represented in the collections by 273 specimens. The family Pandionidae is solely represented by the osprey Pandion haliaetus, with nine specimens (Fig. 22F; 9.6%). The family Accipitridae is represented by 20 species, of which one is not found in the collections, the regionally extinct Rüppell’s vulture Gyps rueppelli. The following species are represented: the common buzzard Buteo buteo with 84 specimens (Fig. 23L; 3%); the Eurasian sparrowhawk Accipiter nisus with 41 specimens (Fig. 23G; 2.8%); the Eurasian griffon vulture Gyps fulvus with 17 specimens (Fig. 22J; 4.3%); the western marsh harrier Circus aeruginosus with 14 specimens (Fig. 23D; 2.2%); Montagu’s harrier Circus pygargus with 13 specimens (Fig. 23F; 2.3%); the black-winged kite Elanus caeruleus with 10 specimens (Fig. 22G; 1.2%); the cinereous vulture Aegypius monachus with 10 specimens (Fig. 22K; 5.1%); the booted eagle Hieraaetus pennatus with 10 specimens (Fig. 23C; 0.9%); the Egyptian vulture Neophron percnopterus with nine specimens (Fig. 22H; 4.9%); the red kite Milvus milvus with nine specimens (Fig. 23I; 2.1%); the short-toed snake eagle Circaetus gallicus with eight specimens (Fig. 22I; 1.1%); the golden eagle Aquila chrysaetos with eight specimens (Fig. 23A; 5.5%); Bonelli’s eagle Aquila fasciata with eight specimens (Fig. 23B; 1.7%); the hen harrier Circus cyaneus with six specimens (Fig. 23E; 1.9%); the black kite Milvus migrans with six specimens (Fig. 23J; 0.4%); the northern goshawk Accipiter gentilis with five specimens (Fig. 23H; 1.4%); and two specimens each of the European honey buzzard Pernis apivorus, the bearded vulture Gypaetus barbatus, and the Spanish imperial eagle Aquila adalberti (Fig. 22L; 6.1%).

Order Bucerotiformes

The Eurasian hoopoe Upupa epops is the single representative of the order in Portugal and is represented by 28 specimens (Fig. 24A; 1.2%).

Order Coraciiformes

Three species, from the families Meropidae, Coraciidae, and Alcedinidae, are represented in the collections by 78 specimens. The family Meropidae is represented solely by the European bee-eater Merops apiaster with 28 specimens (Fig. 23K; 1.3%). The family Coraciidae is represented by the European roller Coracias garrulus with 18 specimens (Fig. 24B; 27.5%). The family Alcedinidae is represented by the common kingfisher Alcedo atthis with 32 specimens (Fig. 24C; 2.7%).

Order Piciformes

Four species from the family Picidae are found in Portugal and are represented in the collections by 48 specimens: the great spotted woodpecker Dendrocopos major with 19 specimens (Fig. 24G; 1.1%); the Iberian green woodpecker Picus sharpei with 13 specimens (Fig. 24E; 1.3%); the Eurasian wryneck Jynx torquilla with nine specimens (Fig. 24D; 2%); and the lesser spotted woodpecker Dryobates minor with seven specimens (Fig. 24F; 1.2%).

Order Falconiformes

Six species, all from the family Falconidae, are found in Portugal, five of which are represented in the collections by 87 specimens. Eleonora’s falcon Falco eleonorae is considered regionally extinct and is not present in the country’s NHCs. The remaining species are represented as follows: the common kestrel Falco tinnunculus with 52 specimens (Fig. 24I; 2.4%); the Eurasian hobby Falco subbuteo with 13 specimens (Fig. 24K; 1.8%); the peregrine falcon Falco peregrinus with 10 specimens (Fig. 25A; 1.6%); the lesser kestrel Falco naumanni with eight specimens (Fig. 24H; 11.5%); and the merlin Falco columbarius with four specimens (Fig. 24J; 6.3%).

Order Psittaciformes

Five species of the family Psittacidae occur in Portugal, all resulting from human introduction (see Table 1). Only the rose-ringed parakeet Alexandrinus krameri is extant in the collections, represented by a single specimen.

Order Passeriformes

One hundred twenty-five species, split between 32 families, are known to occur in Portugal. Of these, 116 are represented in the NHCs by 2,909 specimens. The family Oriolidae is solely represented by the Eurasian golden oriole, Oriolus oriolus, with 15 specimens that cover 1.1% of its known distribution (Fig. 25B). The family Laniidae is represented by three species: the woodchat shrike Lanius senator with 51 specimens (Fig. 25E; 3.8%); the Iberian gray shrike Lanius meridionalis with 41 specimens (Fig. 25D; 3.4%); and the red-backed shrike Lanius collurio with five specimens (Fig. 25C; 3.8%). The family Corvidae is represented by eight species: the Iberian magpie Cyanopica cooki with 45 specimens (Fig. 25G; 3.1%); the Eurasian jay Garrulus glandarius with 21 specimens (Fig. 25H; 1.6%); the Eurasian magpie Pica pica with 18 specimens (Fig. 25I; 1.4%); the red-billed chough Pyrrhocorax pyrrhocorax with 11 specimens (Fig. 25F; 12.2%); the carrion crow Corvus corone with 10 specimens (Fig. 25L; 0.8%); the western jackdaw Corvus monedula with eight specimens (Fig. 25J; 5.9%); the common raven Corvus corax with seven specimens (Fig. 25K; 0.9%); and the rook Corvus frugilegus with six specimens. The family Paridae is represented by four species: the great tit Parus major with 38 specimens (Fig. 26B; 2.6%); the Eurasian blue tit Cyanistes caeruleus with 22 specimens (Fig. 26A; 1.3%); the coal tit Periparus ater with 21 specimens (Fig. 25M; 1.7%); and the crested tit Lophophanes cristatus with eight specimens (Fig. 25N; 0.7%). The family Remizidae is represented by a single species, the Eurasian penduline tit Remiz pendulinus, with two specimens (Fig. 26C; 3.8%). The family Alaudidae is represented by eight species: the Eurasian skylark Alauda arvensis with 40 specimens (Fig. 26I; 3.3%); the woodlark Lullula arborea with 35 specimens (Fig. 26G; 2.1%); the crested lark Galerida cristata with 22 specimens (Fig. 26J; 2.4%); the greater short-toed lark Calandrella brachydactyla with 20 specimens (Fig. 26F; 5.3%); Thekla’s lark Galerida theklae with 20 specimens (Fig. 26H; 3.2%); the Calandra lark Melanocorypha calandra with 11 specimens (Fig. 26E; 4.7%); the Mediterranean short-toed lark Alaudala rufescens with five specimens (Fig. 26D; 20%); and Dupont’s lark Chersophilus duponti with one specimen. The family Cisticolidae is represented by one species, the zitting cisticola Cisticola juncidis, with 19 specimens (Fig. 26K; 1.2%). The family Locustellidae is represented by two species: the common grasshopper warbler Locustella naevia with six specimens (Fig. 26M; 6.7%) and Savi’s warbler Locustella luscinioides with two specimens (Fig. 26L; 4.9%). The family Acrocephalidae is represented by six species, five of which are extant in the collections: the melodious warbler Hippolais polyglotta with 16 specimens (Fig. 26N; 1.2%); the aquatic warbler Acrocephalus paludicola with 10 specimens (Fig. 27B; 66.7%); the great reed warbler Acrocephalus arundinaceus with 10 specimens (Fig. 27A; 2.3%); the sedge warbler Acrocephalus schoenobaenus with nine specimens (Fig. 27C; 17.2%); and the Eurasian reed warbler Acrocephalus scirpaceus with seven specimens (Fig. 26O; 1.8%). No specimen of the western olivaceous warbler Iduna opaca exists in the collections. The family Hirundinidae is represented by five species: the barn swallow Hirundo rustica with 22 specimens (Fig. 27F; 1.3%); the Eurasian crag martin Ptyonoprogne rupestris with 21 specimens (Fig. 27G; 1.7%); the western house martin Delichon urbicum with 16 specimens (Fig. 27E; 0.8%); the sand martin Riparia riparia with five specimens (Fig. 27H; 0.6%); and the European red-rumped swallow Cecropis daurica with three specimens (Fig. 27D; 0.3%). The family Phylloscopidae is represented by four species, three present in the collections: the common chiffchaff Phylloscopus collybita with 31 specimens (Fig. 27J; 2.6%); the willow warbler Phylloscopus trochilus with 13 specimens (Fig. 27K; 1.7%); and the western Bonelli’s warbler Phylloscopus bonelli with four specimens (Fig. 27I; 0.9%). No specimens of the Iberian chiffchaff Phylloscopus ibericus are extant in the collections. The families Scotocercidae and Aegithalidae are represented by a sole species each, the former by Cetti’s warbler Cettia cetti with eight specimens (Fig. 27L; 0.5%), and the latter by the long-tailed tit Aegithalos caudatus with 24 specimens (Fig. 27M; 1.4%). The family Sylviidae is represented by eight species: the Eurasian blackcap Sylvia atricapilla with 55 specimens (Fig. 28A; 2.4%); the Sardinian warbler Curruca melanocephala with 28 specimens (Fig. 28D; 2.2%); the common whitethroat Curruca communis with 15 specimens (Fig. 28G; 1.7%); the Dartford warbler Curruca undata with 14 specimens (Fig. 28F; 1.3%); the garden warbler Sylvia borin with five specimens (Fig. 28B; 2.7%); the western orphean warbler Curruca hortensis with four specimens (Fig. 28C; 1.8%); the spectacled warbler Curruca conspicillata with four specimens (Fig. 28H; 2.2%); and Curruca cantillans with two specimens (Fig. 28E; 0.5%). The families Leiothrichidae, Certhiidae, Sittidae, Troglodytidae, and Cinclidae are all represented by a single specimen each: the red-billed leiothrix Leiothrix lutea; the short-toed treecreeper Certhia brachydactyla with nine specimens (Fig. 28I; 0.6%); the Eurasian nuthatch Sitta europaea with 22 specimens (Fig. 28J; 1.9%); the Eurasian wren Troglodytes troglodytes with 18 specimens (Fig. 28K; 1.1%); and the white-throated dipper Cinclus cinclus with 17 specimens (Fig. 28L; 5.9%). The family Sturnidae is represented by three species, of which the crested myna Acridotheres cristatellus is not in the collections: the spotless starling Sturnus unicolor with 32 specimens (Fig. 29B; 1.7%) and the common starling Sturnus vulgaris with 27 specimens (Fig. 29A; 3.9%). The family Turdidae is represented by six species: the common blackbird Turdus merula with 70 specimens (Fig. 29F; 3.1%); the song thrush Turdus philomelos with 35 specimens (Fig. 29D; 2.4%); the mistle thrush Turdus viscivorus with 15 specimens (Fig. 29C; 1.4%); the redwing Turdus iliacus with 13 specimens (Fig. 29E; 3.1%); the fieldfare Turdus pilaris with 10 specimens (Fig. 29G; 7.8%); and the ring ouzel Turdus torquatus with three specimens (Fig. 29H; 9.1%). The family Muscicapidae is represented by 15 species: the European robin Erithacus rubecula with 59 specimens (Fig. 30A; 3.7%); the European stonechat Saxicola rubicola with 40 specimens (Fig. 30I; 2.3%); the western black-eared wheatear Oenanthe hispanica with 30 specimens (Fig. 30K; 5.1%); the northern wheatear Oenanthe oenanthe with 24 specimens (Fig. 30J; 1.5%); the black redstart Phoenicurus ochruros with 20 specimens (Fig. 30D; 0.9%); the common rock thrush Monticola saxatilis with 14 specimens (Fig. 30F; 12.5%); the whinchat Saxicola rubetra with 14 specimens (Fig. 30H; 3.2%); the common nightingale Luscinia megarhynchos with 13 specimens (Fig. 30C; 0.7%); the bluethroat Luscinia svecica with 12 specimens (Fig. 30B; 1.6%); the blue rock thrush Monticola solitarius with 12 specimens (Fig. 30G; 0.6%); the common redstart Phoenicurus phoenicurus with 11 specimens (Fig. 30E; 1.7%); the spotted flycatcher Muscicapa striata with 10 specimens (Fig. 29J; 1.5%); the European pied flycatcher Ficedula hypoleuca with 10 specimens (Fig. 31A; 0.9%); the rufous-tailed scrub robin Cercotrichas galactotes with six specimens (Fig. 29I; 11.1%); and the black wheatear Oenanthe leucura with five specimens (Fig. 30L; 17.9%). The family Regulidae is represented by three species: the common firecrest Regulus ignicapilla with 18 specimens (Fig. 31E; 1.6%); the Madeira firecrest Regulus madeirensis with seven specimens (Fig. 31C; 15.4%); and the goldcrest Regulus regulus with five specimens (Fig. 31B; 2.2%). The family Prunellidae is represented by two species: the dunnock Prunella modularis with 16 specimens (Fig. 31G; 2.9%) and the alpine accentor Prunella collaris with two specimens (Fig. 31F; 5.9%). Two species are known from the family Ploceidae, the yellow-crowned bishop Euplectes afer and the black-headed weaver Ploceus melanocephalus, but neither is represented in the collections. The family Estrildidae is represented by three species, but only one is found in the collections—the common waxbill Estrilda astrild with 20 specimens (Fig. 31D; 1.4%)—while the red avadavat Amandava amandava and the scaly-breasted munia Lonchura punctulata are not represented. The family Viduidae is represented by a single species, the pin-tailed whydah Vidua macroura, for which no specimens exist in the collections. The family Passeridae is represented by four species: the Spanish sparrow Passer hispaniolensis with 423 specimens (Fig. 31I; 4.9%); the house sparrow Passer domesticus with 211 specimens (Fig. 31H; 5.8%); the rock sparrow Petronia petronia with 25 specimens (Fig. 31J; 2%); and the Eurasian tree sparrow Passer montanus with 22 specimens (Fig. 32A; 1.7%). The family Motacillidae is represented by nine species: the white wagtail Motacilla alba with 49 specimens (Fig. 32I; 1.9%); the meadow pipit Anthus pratensis with 40 specimens (Fig. 32C; 3.4%); the western yellow wagtail Motacilla flava with 39 specimens (Fig. 32H; 4.2%); the gray wagtail Motacilla cinerea with 25 specimens (Fig. 32K; 1.7%); the tree pipit Anthus trivialis with 17 specimens (Fig. 32B; 1.8%); the tawny pipit Anthus campestris with 16 specimens (Fig. 32F; 3.1%); Berthelot’s pipit Anthus berthelotii with nine specimens (Fig. 32E; 23.5%); the water pipit Anthus spinoletta with eight specimens (Fig. 32D; 3.9%); and Richard’s pipit Anthus richardi with two specimens (Fig. 32G; 20%). The family Fringillidae is represented by 12 species: the Eurasian chaffinch Fringilla coelebs with 97 specimens (Fig. 33A; 3.9%); the European goldfinch Carduelis carduelis with 83 specimens (Fig. 34A; 3.3%); the common linnet Linaria cannabina with 49 specimens (Fig. 33D; 2%); the European greenfinch Chloris chloris with 43 specimens (Fig. 33F; 2.9%); the European serin Serinus serinus with 35 specimens (Fig. 34B; 1.6%); the Eurasian siskin Spinus spinus with 19 specimens (Fig. 34D; 3.1%); the hawfinch Coccothraustes coccothraustes with 18 specimens (Fig. 33C; 3%); the Eurasian bullfinch Pyrrhula pyrrhula with 16 specimens (Fig. 33E; 3.5%); the red crossbill Loxia curvirostra with 15 specimens (Fig. 33G; 11.1%); the brambling Fringilla montifringilla with 13 specimens (Fig. 32J; 3.8%); the Atlantic canary Serinus canaria with 11 specimens (Fig. 34C; 4.4%); and the Azores bullfinch Pyrrhula murina with two specimens (Fig. 33B; 25%). The family Emberizidae is represented by six species: the corn bunting Emberiza calandra with 62 specimens (Fig. 34E; 4.2%); the cirl bunting Emberiza cirlus with 40 specimens (Fig. 34H; 3.4%); the rock bunting Emberiza cia with 30 specimens (Fig. 34F; 3.5%); the yellowhammer Emberiza citrinella with 10 specimens (Fig. 34I; 9.5%); the common reed bunting Emberiza schoeniclus with eight specimens (Fig. 34J; 7.7%); and the ortolan bunting Emberiza hortulana with seven specimens (Fig. 34G; 5.4%). The family Calcariidae is represented by a sole species, the snow bunting Plectrophenax nivalis, with eight specimens (Fig. 34K; 44.4%).

Temporal patterns

Collecting date records are known from approximately 89.7% of the specimens. The collecting efforts in all assessed NHCs are historically inconsistent at both institutional and national levels. MUHNAC holds the largest collection among the assessed institutions (Figs 3, 4). Its current numbers reflect a troubled history, as the museum lost nearly its entire vertebrate collection in a fire in 1978 (Saldanha 1978). Pre-1978 specimens extant today were mostly donated later by researchers and museum personnel, including those incorporated from the collections of the former IICT. The growth observed just after the fire during the 1980s and 1990s (Fig. 3) reflects the efforts to rebuild the collection, in large part conducted by ornithologist and preparator António A. Soares (dates of birth and death unknown) and other associated researchers. Since then, new records have been scarce and mostly prepared from specimens collected in the vicinity of the museum or from opportunistic donations.

This pattern is in line with that observed for both amphibians and reptiles (Santos et al. 2024) and mammals (Santos and Ceríaco 2025). In comparison, MCUC holds the largest number of 19^th-century specimens—close to a thousand—as that also corresponds to the period of greatest growth in the collection (Figs 3, 4). MCUC has registered no new records since the late 1920s. These specimens are listed in the catalogue “Aves de Portugal” (Catálogo da Colecção do Museu Zoológico de Coimbra), published by Themido (1933). MHNC-UP’s collection was constructed chiefly in the early 1900s (Fig. 3), due in large part to the contributions of João Alves dos Reis Júnior (1871–1939), one of the most prolific Portuguese ornithologists (Catry et al. 2010). Most of these specimens correspond to those cited by Reis Júnior in his “Catálogo Sistemático e Analítico das Aves de Portugal” (Reis Júnior 1931). Other entries after his time in the institution were much rarer, and it has received no new Portuguese specimens since the late 1970s. AVG’s collection presents a larger number of specimens lacking temporal data when compared to the other assessed NHCs. Despite the lack of precise data, many of its specimens were most likely collected in the late 19^th and early 20^th centuries, given that they were captured as part of King Carlos I’s hunting parties and oceanographic campaigns. These specimens have been held at AVG since 1935 (see Ceríaco 2021). King Carlos I had a keen interest in the natural world and, aside from undertaking several expeditions, also published multiple manuscripts on the birds of Portugal, including the “Catalogo Illustrado das Aves de Portugal” [Illustrated Catalog of the Birds of Portugal] (Catry et al. 2010). Later collections from the 1960s onward (Fig. 3) correspond to sporadic donations made to the institution. Despite the lower number of specimens accessioned per year, the MMF collection presents the most consistent collecting time series among all analyzed institutions, with non-stop records since the 1930s, when it was originally founded (Fig. 3). The recent increase in specimens (and taxa) has also been reported by Zino et al. (1995), Romano et al. (2010), and Correia-Fagundes et al. (2021). Overall, the patterns observed follow those previously reported by Santos et al. (2024) and Santos and Ceríaco (2025) for the herpetology and mammalogy collections, respectively, held in the assessed NHCs.

Figure 3.

Recorded collections of Portuguese birds per year from the 1810s to 2020, for the MUHNAC + IICT, MCUC, MHNC-UP, AVG, and MMF collections. Only specimens with available collecting date data were used in this analysis. Note that the scales differ in each graphic representation. The most recent MCUC record dates from the late 1920s. MUHNAC + IICT only reflects currently extant specimens and does not account for those existing in the collection before the 1978 fire. Egg and nest specimens were not used in this analysis.

Figure 4.

Aggregated growth of the MUHNAC + IICT (pink), MCUC (yellow), MHNC-UP (blue), AVG (green), and MMF (purple) bird collections. Only specimens with available collecting date data were used in this analysis. MUHNAC + IICT only reflects currently extant specimens and does not account for those existing in the collection before the 1978 fire. Egg and nest specimens were not used in this analysis.

Geographic coverage

The overall countrywide collecting effort is geographically biased (Fig. 5). Regions around each museum’s location (Lisbon, Porto, Coimbra, Funchal) have historically been the object of more surveys and present an easily identifiable pattern (Fig. 5). Other collection hotspots can be found in the vicinity of Castelo Branco and the Berlengas and Madeira archipelagos, as well as in some areas of the Alentejo and Oeste e Vale do Tejo regions (Fig. 5). Nearly 1,500 localities were recorded, although approximately 70% were represented by a single specimen.

Taking into account collecting efforts per institution, this bias is further exacerbated (Fig. 6). AVG specimens were collected mostly in the Greater Lisbon region, with a few other records scattered throughout the country. MMF specimens are almost exclusively from the Madeira archipelago, with a few records from the Azores region and none from continental Portugal. MCUC specimens originate from the surroundings of Coimbra and the Beiras region, as well as the northern part of Portugal, with additional records further south. MHNC-UP specimens exhibit a stronger bias toward northern Portugal and the vicinity of the institution in the Porto district. MUHNAC covers a much broader area, including both mainland and insular regions of Madeira and the Azores. As mentioned, much of the collecting effort after the 1978 fire was carried out by António A. Soares, as well as other associated researchers who occasionally delivered new material. Although these patterns do not exactly correspond to those reported for amphibians and reptiles (Santos et al. 2024) and mammals (Santos and Ceríaco 2025) of Portugal, the results still reveal geographic bias.

Figure 5.

Map of collecting localities for all collections. Increasing circle size and color intensity correspond to the number of records for each location. Egg and nest specimens were not used in this analysis.

Dividing the country into a 10 × 10 km grid following the method used by Equipa Atlas (2022), we obtained a quantitative overview of the geographic coverage of all collections (Fig. 7). Overall, the assessed collections cover 41.7% of Portugal. MUHNAC covers the highest percentage of all institutions, with 33.4%, followed by MCUC with 7.3%, MHNC-UP with 6.9%, AVG with 3.1%, and MMF with 2.1%. Detailed accounts for each species, using the same grid and comparing museum specimens with the species’ known distributions, are presented in Figs 8–34. Species’ known distributions were retrieved from the most recent Portuguese atlas of breeding taxa (Equipa Atlas 2022) and wintering and migrating species (Equipa Atlas 2018). These cover the whole range of each taxon, combining known breeding, wintering, and migrating ranges when applicable. Whenever museum specimen data fell outside of known distributions, we accounted for those records as additional distribution data and added them to the previously known range in order to evaluate the percentage of coverage that museums represent.

Figure 6.

Map of collecting localities for all collections: AVG (green), MMF (purple), MCUC (yellow), MHNC-UP (blue), and MUHNAC + IICT (pink). Egg and nest specimens were not used in this analysis.

Only a single taxon, the white-faced storm petrel Pelagodroma marina, has the totality of its known distribution covered in the collections, as the species is found only in the Madeira archipelago (Fig. 13B). The next best represented species in the collections are Bulwer’s petrel Bulweria bulwerii with 88.9% (Fig. 14C), the Atlantic puffin Fratercula arctica with 78.6% (Fig. 21F), the common murre Uria aalge with 73.3% (Fig. 21H), and the spotted crake Porzana porzana (Fig. 12B) with 72.7% of its known distribution represented in collections (Table 2). Additionally, 11 other taxa have 50% or more of their known distributions represented in the collections (Table 2). All these species have very restricted distributions.

Figure 7.

Map of collecting localities for all collections, presented on a 10 × 10 km grid following Equipa Atlas (2022). Egg and nest specimens were not used in this analysis.

Of the species represented in the collections, 249 have 20% or less of their distribution represented. Of those, 201 have 10% or less, and 160 have less than 5%. Among the latter are some of the most ubiquitous species in the country: the Eurasian collared dove Streptopelia decaocto with 0.2% of its known distribution covered (Fig. 10J), the European red-rumped swallow Cecropis daurica with 0.3% (Fig. 27D), and the rock dove Columba livia (Fig. 10E) and the black kite Milvus migrans with 0.4% (Fig. 23J).

Figure 8.

Distribution maps of A. Coturnix coturnix; B. Phasianus colchicus; C. Alectoris rufa; D. Anser anser; E. Melanitta nigra; F. Mergus serrator; G. Tadorna tadorna; H. Tadorna ferruginea; I. Marmaronetta angustirostris, and J. Aythya ferina, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 9.

Distribution maps of A. Aythya nyroca; B. Aythya fuligula; C. Spatula querquedula; D. Spatula clypeata; E. Mareca strepera; F. Mareca penelope; G. Anas platyrhynchos; H. Anas acuta, and I. Anas crecca, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 10.

Distribution maps of A. Tachybaptus ruficollis; B. Podiceps cristatus; C. Podiceps nigricollis; D. Phoenicopterus roseus; E. Columba livia; F. Columba oenas; G. Columba palumbus; H. Columba trocaz; I. Pterocles orientalis; J. Streptopelia decaocto, and K. Pterocles alchata, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 11.

Distribution maps of A. Streptopelia turtur; B. Caprimulgus ruficollis; C. Caprimulgus europaeus; D. Tachymarptis melba; E. Apus unicolor; F. Apus pallidus; G. Apus apus; H. Clamator glandarius, and I. Cuculus canorus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 12.

Distribution maps of A. Rallus aquaticus; B. Porzana porzana; C. Zapornia pusilla; D. Porphyrio porphyrio; E. Gallinula chloropus; F. Fulica cristata; G. Grus grus; H. Fulica atra, and I. Tetrax tetrax, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 13.

Distribution maps of A. Otis tarda; B. Pelagodroma marina; C. Hydrobates pelagicus; D. Hydrobates castro; E. Hydrobates monteiroi; F. Hydrobates leucorhous; G. Pterodroma deserta; H. Pterodroma madeira; I. Ardenna grisea; J. Ardenna gravis, and K. Calonectris borealis, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 14.

Distribution maps of A. Puffinus puffinus; B. Puffinus mauretanicus; C. Bulweria bulwerii; D. Ciconia ciconia; E. Ciconia nigra; F. Platalea leucorodia; G. Plegadis falcinellus; H. Ardeola ralloides; I. Botaurus stellaris, and J. Ixobrychus minutus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 15.

Distribution maps of A. Nycticorax nycticorax; B. Bubulcus ibis; C. Ardea cinerea; D. Ardea purpurea; E. Egretta garzetta; F. Gulosus aristotelis; G. Morus bassanus, and H. Phalacrocorax carbo, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 16.

Distribution maps of A. Burhinus oedicnemus; B. Haematopus ostralegus; C. Recurvirostra avosetta; D. Pluvialis squatarola; E. Himantopus himantopus; F. Charadrius hiaticula; G. Pluvialis apricaria; H. Charadrius dubius; I. Eudromias morinellus, and J. Charadrius alexandrinus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 17.

Distribution maps of A. Vanellus vanellus; B. Numenius phaeopus; C. Numenius arquata; D. Limosa limosa; E. Limosa lapponica; F. Arenaria interpres; G. Calidris canutus; H. Calidris pugnax, and I. Calidris ferruginea, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 18.

Distribution maps of A. Calidris alba; B. Calidris alpina; C. Calidris maritima; D. Calidris minuta; E. Calidris temminckii; F. Actitis hypoleucos; G. Scolopax rusticola, and H. Gallinago gallinago, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 19.

Distribution maps of A. Lymnocryptes minimus; B. Tringa erythropus; C. Tringa ochropus; D. Tringa nebularia; E. Tringa totanus; F. Tringa glareola; G. Phalaropus fulicarius; H. Glareola pratincola; I. Hydrocoloeus minutus; J. Rissa tridactyla, and K. Larus melanocephalus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 20.

Distribution maps of A. Larus ridibundus; B. Larus fuscus; C. Larus marinus; D. Larus michahellis; E. Larus genei; F. Larus argentatus; G. Larus canus; H. Sternula albifrons; I. Gelochelidon nilotica; J. Chlidonias hybrida, and K. Chlidonias niger, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 21.

Distribution maps of A. Sterna hirundo; B. Stercorarius parasiticus; C. Thalasseus sandvicensis; D. Catharacta skua; E. Stercorarius pomarinus; F. Fratercula arctica; G. Alca torda; H. Uria aalge; I. Tyto alba, and J. Athene noctua, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 22.

Distribution maps of A. Otus scops; B. Asio otus; C. Asio flammeus; D. Strix aluco; E. Bubo bubo; F. Pandion haliaetus; G. Elanus caeruleus; H. Neophron percnopterus; I. Circaetus gallicus; J. Gyps fulvus; K. Aegypius monachus, and L. Aquila adalberti, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 23.

Distribution maps of A. Aquila chrysaetos; B. Aquila fasciata; C. Hieraaetus pennatus; D. Circus aeruginosus; E. Circus cyaneus; F. Circus pygargus; G. Accipiter nisus; H. Accipiter gentilis; I. Milvus milvus; J. Milvus migrans; K. Merops apiaster, and L. Buteo buteo, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 24.

Distribution maps of A. Upupa epops; B. Coracias garrulus; C. Alcedo atthis; D. Jynx torquilla; E. Picus sharpei; F. Dryobates minor; G. Dendrocopos major; H. Falco naumanni; I. Falco tinnunculus; J. Falco columbarius, and K. Falco subbuteo, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 25.

Distribution maps of A. Falco peregrinus; B. Oriolus oriolus; C. Lanius collurio; D. Lanius meridionalis; E. Lanius senator; F. Pyrrhocorax pyrrhocorax; G. Cyanopica cooki; H. Garrulus glandarius; I. Pica pica; J. Corvus monedula; K. Corvus corax; L. Corvus corone; M. Periparus ater, and N. Lophophanes cristatus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 26.

Distribution maps of A. Cyanistes caeruleus; B. Parus major; C. Remiz pendulinus; D. Alaudala rufescens; E. Melanocorypha calandra; F. Calandrella brachydactyla; G. Lullula arborea; H. Galerida theklae; I. Alauda arvensis; J. Galerida cristata; K. Cisticola juncidis; L. Locustella luscinioides; M. Locustella naevia; N. Hippolais polyglotta, and O. Acrocephalus scirpaceus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 27.

Distribution maps of A. Acrocephalus arundinaceus; B. Acrocephalus paludicola; C. Acrocephalus schoenobaenus; D. Cecropis daurica; E. Delichon urbicum; F. Hirundo rustica; G. Ptyonoprogne rupestris; H. Riparia riparia; I. Phylloscopus bonelli; J. Phylloscopus collybita; K. Phylloscopus trochilus; L. Cettia cetti, and M. Aegithalos caudatus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 28.

Distribution maps of A. Sylvia atricapilla; B. Sylvia borin; C. Curruca hortensis; D. Curruca melanocephala; E. Curruca cantillans; F. Curruca undata; G. Curruca communis; H. Curruca conspicillata; I. Certhia brachydactyla; J. Sitta europaea; K. Troglodytes troglodytes, and L. Cinclus cinclus, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 29.

Distribution maps of A. Sturnus vulgaris; B. Sturnus unicolor; C. Turdus viscivorus; D. Turdus philomelos; E. Turdus iliacus; F. Turdus merula; G. Turdus pilaris; H. Turdus torquatus; I. Cercotrichas galactotes, and J. Muscicapa striata, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 30.

Distribution maps of A. Erithacus rubecula; B. Luscinia svecica; C. Luscinia megarhynchos; D. Phoenicurus ochruros; E. Phoenicurus phoenicurus; F. Monticola saxatilis; G. Monticola solitarius; H. Saxicola rubetra; I. Saxicola rubicola; J. Oenanthe oenanthe; K. Oenanthe hispanica, and L. Oenanthe leucura, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 31.

Distribution maps of A. Ficedula hypoleuca; B. Regulus regulus; C. Regulus madeirensis; D. Estrilda astrild; E. Regulus ignicapilla; F. Prunella collaris; G. Prunella modularis; H. Passer domesticus; I. Passer hispaniolensis, and J. Petronia petronia, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 32.

Distribution maps of A. Passer montanus; B. Anthus trivialis; C. Anthus pratensis; D. Anthus spinoletta; E. Anthus berthelotii; F. Anthus campestris; G. Anthus richardi; H. Motacilla flava; I. Motacilla alba; J. Fringilla montifringilla, and K. Motacilla cinerea, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 33.

Distribution maps of A. Fringilla coelebs; B. Pyrrhula murina; C. Coccothraustes coccothraustes; D. Linaria cannabina; E. Pyrrhula pyrrhula; F. Chloris chloris, and G. Loxia curvirostra, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Figure 34.

Distribution maps of A. Carduelis carduelis; B. Serinus serinus; C. Serinus canaria; D. Spinus spinus; E. Emberiza calandra; F. Emberiza cia; G. Emberiza hortulana; H. Emberiza cirlus; I. Emberiza citrinella; J. Emberiza schoeniclus, and K. Plectrophenax nivalis, comparing museum records (black dots) with the species’ known extent in Portugal (pink).

Exotic species

Several species have been introduced in Portugal as a result of the wildlife pet trade, in conditions dissimilar to historic introductions (Abellán et al. 2016). Wild populations of these taxa are now found in the country, with some species well established, namely the rose-ringed parakeet Alexandrinus krameri, the monk parakeet Myiopsitta monachus, the common waxbill Estrilda astrild, and the red avadavat Amandava amandava (Baquero et al. 2021), while others are rapidly expanding their known distribution (Equipa Atlas 2022). Given the fact that most of these invasive episodes are considerably recent, only four exotic species are represented in the collections: the ring-necked pheasant Phasianus colchicus with two specimens, the rose-ringed parakeet Alexandrinus krameri and the red-billed leiothrix Leiothrix lutea with a single specimen each, and the common waxbill Estrilda astrild with 20 specimens.

Rare or accidental species

Some rare wintering, passage migrant, and accidental species can also be found in Portugal from time to time (Catry et al. 2010; Equipa Atlas 2018). The assessed collections have records of 41 such species (Table 3), for a total of 138 skin and taxidermy mounts. The MMF collection houses a specimen of the slender-billed curlew Numenius tenuirostris, collected in 1990 in “Santa Cruz.” This species is considered “Critically Endangered,” with less than 50 individuals estimated to still exist globally (BirdLife International 2018c).

Eggs and nests

A total of 2,701 eggs of 166 species and 401 nests from 95 species are currently available in the studied collections (Fig. 35, Table 4). MCUC’s collection is the richest in numbers, with 1,127 eggs from 107 taxa and 240 nests from 80 taxa in total, followed by the MHNC-UP with 968 eggs and 82 nests from 113 and 46 taxa, respectively. MUHNAC, MMF, and AVG’s collections are much smaller: MUHNAC amasses 311 eggs from 22 taxa and 22 nests from 12 taxa, while MMF has 204 eggs from 31 species and 40 nests from 15 taxa, and AVG has 12 eggs from 3 species. The most represented species in egg collections are the western jackdaw Corvus monedula with 64, the lesser kestrel Falco naumanni with 63, and the house sparrow Passer domesticus with 60. When it comes to nests, the common blackbird Turdus merula is the most represented with 26, followed by the European goldfinch Carduelis carduelis with 21 and the Eurasian blackcap Sylvia atricapilla with 15. Most of the egg specimens were collected in the late 1880s when the MCUC collection grew the most, with the next peak occurring in the 1930s and 1940s with the MHNC-UP collection (Fig. 36). Due to the 1978 fire, the MUHNAC collection only contains more recent records with a clear peak in the 2010s (Fig. 36). Unlike the other collections, MMF has had a more stable tradition in collecting egg and nest specimens, with continued growth from when the museum was established to the current day, where new material is still being accessioned (Fig. 36).

Figure 35.

Examples of eggs from the MHNC-UP collection (A) and nests from the MUHNAC collection (B).

Figure 36.

Recorded collections of Portuguese bird eggs (red solid line) and nests (green dashed line) per year from the 1870s to 2025, for the MUHNAC + IICT, MCUC, MHNC-UP, and MMF. Only specimens with available collecting date data were used in this analysis. Note that the scales differ in each graphic representation. The most recent MCUC record dates from the late 1950s. MUHNAC + IICT only reflects currently extant specimens and does not account for those existing in the collection before the 1978 fire.

Game species

Twenty-eight game species from six orders are known from Portugal. All taxa are represented in the collections (Table 1) by a total of 517 specimens, approximately 8.6% of bird specimens in Portuguese NHCs (Fig. 37). Thirteen species are known breeders in Portugal, while eleven winter in the country, and three others are known from both breeding and wintering populations. The ring-necked pheasant Phasianus colchicus was introduced in the country for hunting purposes and does not have viable reproducing populations in the wild as of now (Equipa Atlas 2022). Despite the presence of all taxa in the collections, almost all species are represented by less than 50 specimens. The red-legged partridge Alectoris rufa is the most represented, with 53 specimens (2.7% of known distribution), followed by the common moorhen Gallinula chloropus with 36 (3.3%) and the song thrush Turdus philomelos with 35 (2.4%). When it comes to known distributions, the best represented game species are the jack snipe Lymnocryptes minutus with 53.3% (Fig. 19A), the tufted duck Aythya fuligula with 20% (Fig. 9B), and the Eurasian wigeon Mareca penelope with 19.4% (Fig. 9F). All three of these species have quite restricted distributions.

Figure 37.

Proportion of specimens belonging to non-game and game species, per order. Detailed numbers per species are presented.

Conservation status

The most recent red list of bird species from Portugal accounts only for the taxa found in the country’s mainland (Almeida et al. 2022). Species known only as passage migrants were not evaluated. Conservation status for species occurring primarily in the Azores and Madeira archipelagos was retrieved from the IUCN Red List (BirdLife International 2018a, 2018b, 2018d, 2018e, 2018f, 2021a, 2021b, 2021c, 2021d, 2021e, 2021f, 2021g, 2022, 2024a, 2024b). For some taxa, two distinct conservation statuses are given, according to the assessed population phenology, either breeding or wintering (Table 2). Around 60% of all assessed species are classified as “Least Concern” or “Near Threatened,” and more than a quarter have at least one population phenology considered threatened, that is, with an attributed status of “Vulnerable” (43 species), “Endangered” (34 species), or “Critically Endangered” (14 species) (Fig. 38). Eight species are classified as “Regionally Extinct,” while the breeding population in continental Portugal for three others is considered extinct—the red-knobbed coot Fulica atra, the common crane Grus grus, and the Eurasian oystercatcher Haematopus ostralegus. The latter two have wintering populations deemed “Vulnerable.” When it comes to the number of specimens, the collections currently hold 431 specimens of “Endangered” taxa, 176 specimens of “Critically Endangered” taxa, and 53 specimens of the eleven “Regionally Extinct” taxa.

Figure 38.

Proportion of specimens belonging to non-threatened (soft yellow), threatened (red tones), and no data (grey) taxa. Specimens are classified in categories of increasing danger, from “Least Concern” to “Regionally Extinct.” Species for which no assessment exists for Portugal, are “Data Deficient” or for which the assessment was “Not Applicable” are grouped within “No Data.”

Discussion

With a total of 6,124 specimens, between the more common and the rarer or accidental species, the collections of Portuguese birds in Portuguese NHCs are considerably small, especially when comparing them to other European and international counterparts (Roselaar 2003). This number increases if we include the number of eggs and nests, but it still amounts to a small total, especially considering that it combines all specimens from all the assessed institutions. However, this is in line with what is known for the collections of other Portuguese vertebrate taxa in national NHCs—amphibians and reptiles combined have a similar number (4,950 fide Santos et al. 2024), and mammals, while slightly more (7,402 fide Santos and Ceríaco 2025), still house fewer than 10,000 specimens. Despite close to 90% of the occurring avian taxa being represented in the collections (297 out of 327 species), the overall collection numbers per species are considerably low. Only 1.3% of the represented species have more than 100 specimens in the collections, while the majority (93.9%) have 50 or fewer, and a considerable percentage (39.3%) have 10 or fewer specimens. The country is not fully sampled, with nearly 60% of its area lacking a single recorded specimen. The sampling effort is also geographically biased toward the areas around major cities and some protected areas. The vast majority of the species represented, 82.5%, have less than 20% of their known distribution covered in the collections, with only 4% having more than 50%. Most collections also do not form any kind of time series, as collecting events were not continuous and reflect institutional history rather than concerted efforts to create such time series. The single exception appears to be MMF, which has kept a stable and continuous collecting practice since its establishment in the early decades of the twentieth century. Researcher interest and opportunistic donations have been the backbone of collection growth, rather than continued and sustained institutional collecting policies. Overall, Portuguese ornithological collections are not taxonomically complete, are geographically biased, and do not provide robust time series for any of the species. This is also the case for the egg and nest collections, for which nearly half of the occurring species do not have either eggs or nests represented in the NHCs. Those that do are represented by very low numbers and were collected from only a few distinct localities, altogether failing to make up any relevant time series. This overall scenario is similar to that presented by other vertebrate groups in Portuguese NHCs (Santos et al. 2024; Santos and Ceríaco 2025).

Apart from Portuguese material, the country’s NHCs hold a considerable number of non-Portuguese and especially non-European specimens. This reflects Portuguese colonial history and its historical ties with countries such as Brazil. While MUHNAC ornithological collections are almost entirely based on Portuguese specimens, the majority of IICT ornithological collections, held and managed by MUHNAC, originate from the former Portuguese colonies in Africa and Asia and are represented by approximately 5,600 specimens (Monteiro et al. 2014, 2016, 2017; Figueira 2017). MCUC collections hold approximately 1,930 specimens from other regions, namely from the Afrotropics (575; Themido 1938a, 1938b), the Neotropics (1,093; Themido 1938b), and other parts of the world (267; Themido 1938b). Among these, close to 120 were donated by the Museu Bocage and 26 by King Pedro V in the last decades of the nineteenth to early twentieth century. The Museu Bocage material includes important specimens collected by the Portuguese explorer José d’Anchieta (1832–1897) in Angola and used by José Vicente Barbosa du Bocage (1823–1907) in his various works on Angolan ornithology (Ceríaco 2021). MHNC-UP has one of the largest ornithological collections in the country, of approximately 5,200 non-Portuguese specimens, with around 2,000 of those originating from tropical regions (Gonçalves et al. 2016). It is worth highlighting the impressive collection of South American birds belonging to the collection of José Teixeira da Silva Braga Júnior (1844–1904), held today at MHNC-UP, including a collection of hummingbirds (Anonymous 1905; Lopes et al. 2021). MHNC-UP also hosts a considerable collection of African birds, some donated by the Museu Bocage in the early 1900s, also including Anchieta specimens, a collection of Guinea-Bissau specimens (Madureira 1978), and the recent deposition of the holotype of the recently described Otus bikegila Melo, Freitas, Verbelen, Costa, Pereira, Fuchs, Sangster, Correia, Lima, and Crottini, 2022, endemic to Príncipe Island, Gulf of Guinea (Melo et al. 2022). AVG ornithological collections are almost exclusively based on Portuguese specimens.

A comparison between the overall number of specimens in Portuguese NHCs and those of other European countries of similar size reveals a stark difference. In the case of Austria, even if accounting only for the bird skins extant at the Naturhistorisches Museum Wien, Roselaar (2003) reported 104,300 Austrian specimens, roughly five times more than all Portuguese collections combined, with an annual growth of up to 1,000 specimens per year. More recent numbers indicate a growth of approximately 500 specimens per year, with the collection reaching 130,000 specimens (Naturhistorisches Museum Wien 2025). This includes a large donation of almost 20,000 scientific skins in 2022 by the British Harrison Institute, all currently digitized (Vohland et al. 2025). Before its destruction in 1956, the Magyar Nemzeti Muzeum, in Budapest, Hungary, housed approximately 70,000 skins, with a current collection of 12,500 reported by Roselaar (2003). As for the Czech Republic, the Národní Muzeum, Zoologické oddělení, alone reported 22,000 skins in Roselaar (2003), with a present collection of 35,000 skins and 1,500 skeletons (National Museum 2025). Looking to neighboring Spain, according to Roselaar (2003), 7,000 skins and approximately 1,300 skeletons of Spanish birds were catalogued at the Museu de Zoologia de Barcelona (currently Museu de Ciències Naturals de Barcelona, MCNB), 19,000 skins and 6,000 skeletons at the Museo Nacional de Ciencias Naturales (MNCN), and 22,000 skins and 1,000 skeletons at the Estacíon Biológica de Doñana (EBD). Updated numbers report 12,863 bird specimens from the MCNB (Museu de Ciències Naturals de Barcelona 2025), 30,000 from the MNCN (Museo Nacional de Ciencias Naturales 2025), and 33,000 bird specimens from EBD (Estación Biológica Doñana – CSIC 2025). Even when compared to its former colonies in Africa, Portuguese ornithological collections fall short—in the ornithological collection housed in the Instituto Superior de Educação do Lubango (ISCED-Lubango), Huíla Province, southwestern Angola, more than 40,000 Angolan birds are preserved (Mills et al. 2010), almost double the Portuguese specimens available in the assessed NHCs.

Portugal has had a longstanding tradition of hunting, reflected today in the high number of bird game species. Nevertheless, these make up less than 10% of the overall specimens available and are geographically poorly represented (see section above). Ubiquitous species like the rock dove Columba livia or the common wood pigeon Columba palumbus, known from all of continental Portugal and both archipelagos (the latter currently extinct in Madeira), are represented by 13 and 12 specimens, respectively. In both cases, this accounts for less than 1% of their known distributions and makes them both the most poorly geographically represented game species. Taxidermy mounts were at one point a common trophy from recreational hunting (pers. obs.) and can still be found throughout the country in hunting clubs and small private collections, but they are surprisingly scarce in NHCs. These numbers are even lower than those observed for mammalian game species, which make up 17% of mammal specimens in the Portuguese NHCs (Santos and Ceríaco 2025).

Given their popularity as gaming trophies and expositive taxidermies, additional bird specimens exist in other institutions in the country, such as regional hunting museums, private collections, and high school biology laboratories or museums (Fig. 39). Gomes (2017) provided an overview of the importance of natural history collections in Portuguese high schools, but a systematic cataloguing of these collections remains to be done. The Museu da Caça de Vila Viçosa (Vila Viçosa, southern Portugal), a hunting museum, also has some ornithological collections that are currently being catalogued (Perestrelo et al. in prep.). While many of these specimens may lack basic data, many do not, and given the small number of specimens of some species available in the “main” museums, these specimens could represent a considerable increase in the material available for those taxa. They can also provide additional geographic records or dates of collecting, which would contribute to closing some of the existing gaps and biases. The importance of such small and sometimes “hidden” collections has been shown to be particularly significant for the case of rare and threatened species (Casas-Marce et al. 2012).

Figure 39.

Bird collections of the Hunting Museum of Vila Viçosa (A) and the Military School of Lisbon. Photos by Sofia Perestrelo (A) and Luis M. P. Ceríaco (B).

Ethical concerns and social reactions against specimen collection are particularly strong for birds and mammals. While the debate regarding collecting specimens is decades old, it was recently reignited by discussions generated by several highly impactful papers, such as those by Minteer et al. (2014), Rocha et al. (2014), Byrne (2023), and Nachman et al. (2023). An iconic case was the recent collection of a single male specimen of the Guadalcanal moustached kingfisher Actenoides boungainvillei excelsus in the Solomon Islands by American Museum of Natural History researchers (Filardi 2015). Although the species is classified in the IUCN non-threatened category “Least Concern” (BirdLife International 2002) and the collection of a single male specimen “would not affect the population’s success” given that the state of the population and the habitat was “substantial and healthy enough” (Sweet in Silber 2015), its collection caused a global public outcry, which forced both the collector and the AMNH to issue public statements (Filardi 2015). Cases like this put additional societal pressure on researchers who use and collect bird specimens for research purposes, making it more difficult for them to collect new material and partly explaining the sharp decline in the addition of new specimens to Portuguese NHCs.

Despite the current scenario, the gaps found in the assessed collections could be mitigated through targeted collecting efforts of poorly represented species and/or poorly sampled regions. The 1978 fire depleted the already incomplete collection of the country’s largest natural history museum and further deepened existing gaps. A considerable part of what exists today in MUHNAC’s ornithological collection reflects the post-fire collecting efforts established by the then institutional policy (Almaça 2000) and carried out by the institution’s preparator, who had previously published several catalogues of the collection (Soares 1970, 1971). Following his retirement, all new accessions were made sporadically by associated researchers and invited curators. Overall, no collecting campaigns to fill known gaps in NHCs have occurred in the twenty-first century in the country. This is, however, not a modern issue, as even before the 1978 fire, a considerable part of Portuguese zoologists were focused on projects regarding the biodiversity of the then Portuguese overseas territories in Africa, and collecting Portuguese fauna was not a research priority in most institutions.

Besides active collecting, other sources of specimens could contribute to the enrichment of Portuguese ornithological collections, such as road-killed specimens or individuals that died under the care of recovery centers. Pinto et al. (2020) recorded more than 2,500 road-killed birds of just six species (European goldfinch Carduelis carduelis, blue tit Cyanistes caeruleus, Eurasian blackcap Sylvia atricapilla, house sparrow Passer domesticus, corn bunting Emberiza calandra, and tawny owl Strix aluco) between January 2005 and December 2013 on selected roads in southern Portugal. These 2,500 specimens correspond to nearly half of all available specimens of Portuguese birds in Portuguese museums (6,124). While some of these specimens would likely be in a less-than-ideal state for preservation as good scientific specimens, many could have been salvaged and accessioned in collections. There are over a dozen wild animal recovery centers across the country (managed by either local NGOs or governmental authorities) that receive several injured birds every month (pers. obs.). Some of the birds that unfortunately die are currently not being deposited in scientific collections either.

This situation is considerably difficult to explain but surely has roots in the disassociation between researcher and conservationist communities and the natural history museums in Portugal (as already noted by Ceríaco et al. 2021; Santos et al. 2024; Santos and Ceríaco 2025), as well as the lack of curatorial and technical staff in Portuguese museums. Of the four institutions analyzed in this study, only MUHNAC has a full-time taxidermist on staff. The lack of curators and collection managers not only results in the absence of specific growth strategies for ornithological collections but also in an overall deficiency of the resources needed to deal with the accession and preparation of new material. The current staff is insufficient, and part of the curatorial duties are carried out by invited researchers (non-staff) who do not have full-time presence in the institutions. Therefore, most efforts are concentrated on immediate curatorial practices. Such a severe lack of staff and growth strategies may also create distrust among potential researchers who need to deposit specimens in Portuguese institutions, many of whom end up depositing them in international collections instead.

Specimen-based research is at the core of conservation practices, and time series are crucial to investigating the rapid changes impacting biodiversity worldwide. Good coverage provided by NHCs reflects good baseline data to tackle these issues. Zimova et al. (2023) studied changes in bird morphology using two large datasets: one with more than 70,000 Field Museum specimens from 52 species collected from 1978 to 2016, and another with a little over 15,000 specimens from 77 species collected as part of the Biological Dynamics of Forest Fragments Project in Brazil. Such projects would be impossible to conduct using only Portuguese NHCs, as they do not have even a fraction of the numbers or time series. This not only affects the science that could be conducted in Portuguese NHCs but also limits the possibilities of studying several aspects of Portuguese bird fauna. Peterson et al. (1998) conducted a geographic analysis of Mexican bird specimens in 26 institutions from Mexico, the United States, Canada, and several European collections, retrieving about 221,000 specimens, and reported that the representation of Mexican birds was uneven and inadequate. The IUCN Red List of Threatened Species requires assessment of past and present data per its base criteria to classify population size trends. As NHCs are the repositories of such data, they are fundamental to species conservation policies. Targeted field censuses can mitigate the lack of more recent records in collections, but historical data rely heavily on museum specimens. Although citizen science and observation records have proved useful (Fu et al. 2024), they are also incomplete and biased (Callaghan et al. 2021) and do not provide physical vouchers that can be reassessed in the future.

Conclusion

Similarly to what has been observed for other taxonomic groups (Santos et al. 2024; Santos and Ceríaco 2025), the bias and incompleteness of Portuguese birds in Portuguese NHCs present serious consequences for the research community. These biases hinder, if not completely prevent, several types of studies, namely those that require robust geographic and temporal series, such as those related to pollutants, parasites, and pathogens (DuBay and Fuldner 2017; Harmon et al. 2019; Thompson et al. 2021; Wood et al. 2023). These institutions require urgent and stable investment, the allocation of resources toward hiring and training new professionals, and rebuilding the bridge between museums and their scientific community. They should also aim to foster the digitization of metadata and its availability and to increase their number of specimens through regular field collecting. A debate on the future of Portuguese ornithological collections is urgently needed.

Acknowledgements

We thank all collections managers and curators who provided access and data from the collections under their care: Ana Cristina Rufino from the Museu da Ciência da Universidade de Coimbra, Maria Judite Alves from the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa, Francisco Calado from the Aquário Vasco da Gama, and Manuel Biscoito and Carolina Ornelas from the Museu de História Natural do Funchal. Data on the ornithological collections of the Museu Nacional de História Natural e da Ciência da Universidade do Porto were provided in 2015 by its former curator of zoology, Luzia de Sousa. We are especially grateful to Carlos Godinho (LabOr, Universidade de Évora) for providing the shapefiles with the current known distribution of Portuguese birds used in the Portuguese Atlas of Breeding Birds and the Portuguese Atlas of Wintering and Migrant Species. Luis Fábio Silveira and Marcelo Duarte from the Museu de Zoologia da Universidade de São Paulo provided important comments and revisions on earlier versions of this manuscript. The authors are members of the NATHIST—Natural History, Collections & Taxonomy research group at CIBIO-InBIO, which provided institutional support for this study.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

Bruna S. Santos is funded by the Fundação para a Ciência e a Tecnologia (FCT) grant 2021.06659.BD..

Author contributions

Both authors have contributed equally.

Author ORCIDs

Bruna S. Santos https://orcid.org/0000-0002-3798-2664

Luis M. P. Ceríaco https://orcid.org/0000-0002-0591-9978

Data availability

All of the data that support the findings of this study are available in the main text.

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﻿Abstract

Key words:

﻿Introduction

﻿Materials and methods

﻿Results

﻿Taxonomic representation

﻿Order Galliformes

﻿Order Anseriformes

﻿Order Podicipediformes

﻿Order Phoenicopteriformes

﻿Order Columbiformes

﻿Order Pterocliformes

﻿Order Caprimulgiformes

﻿Order Cuculiformes

﻿Order Gruiformes

﻿Order Otidiformes

﻿Order Procellariiformes

﻿Order Ciconiiformes

﻿Order Pelecaniformes

﻿Order Suliformes

﻿Order Charadriiformes

﻿Order Strigiformes

﻿Order Accipitriformes

﻿Order Bucerotiformes

﻿Order Coraciiformes

﻿Order Piciformes

﻿Order Falconiformes

﻿Order Psittaciformes

﻿Order Passeriformes

﻿Temporal patterns

﻿Geographic coverage

﻿Exotic species

﻿Rare or accidental species

﻿Eggs and nests

﻿Game species

﻿Conservation status

﻿Discussion

﻿Conclusion

﻿Acknowledgements

﻿Additional information

Conflict of interest

Ethical statement

Use of AI

Funding

Author contributions

Author ORCIDs

Data availability

﻿References

Abstract

Introduction

Materials and methods

Results

Taxonomic representation

Order Galliformes

Order Anseriformes

Order Podicipediformes

Order Phoenicopteriformes

Order Columbiformes

Order Pterocliformes

Order Caprimulgiformes

Order Cuculiformes

Order Gruiformes

Order Otidiformes

Order Procellariiformes

Order Ciconiiformes

Order Pelecaniformes

Order Suliformes

Order Charadriiformes

Order Strigiformes

Order Accipitriformes

Order Bucerotiformes

Order Coraciiformes

Order Piciformes

Order Falconiformes

Order Psittaciformes

Order Passeriformes

Temporal patterns

Geographic coverage

Exotic species

Rare or accidental species

Eggs and nests

Game species

Conservation status

Discussion

Conclusion

Acknowledgements

Additional information

References