Funding for this study was provided by the ECSIT Project: “Écosystèmes insulaires tropicaux, réponse de la faune indigène terrestre de Guadeloupe à 6 000 ans d’anthropisation du milieu”, conducted by the CNRS with financial support from the European PO-FEDER program (grant n° 2016-FED-503), Guadeloupe Regional Council, the DAC of Guadeloupe (PCR Extinctions), and the CNRS (SEEG ECSIT). We are grateful to museum curators and staff for enabling access to modern bird osteological collections: P. Sweet (AMNH); S. Guimarães and J-L. Bernos (MEC); C. Lefèvre and J. Fuchs (MNHN); M.-D. Wandhammer (MZS); J. Cooper and J. White (NHMUK); M. Peck (ROM); NR. Cannarozzi, W. Kratter, DW. Steadman and T. Webber (UF); J. Hinshaw (UMMZ); C. Gebhard, M. Florence, H. James, M. Florence and C. Milensky (USNM). We would also like to thank D. Bonissent, G. Roberts, and C. Stouvenot (DAC of Guadeloupe) for allowing access to the archaeological material; the excavation directors (M. Boudadi-Maligne, D. Cochard, S. Grouard, J.-B. Mallye, and C. Stouvenot) and teams of the sites investigated here; M. Bessou, C. Bochaton, D. Cochard, V. Conche, C. Haget, J.-B. Huchet, N. Jeantet, R. Ledevin, E. Pubert, S. Renou (PACEA) for technical assistance; B. Gravina for the English editing and constructive comments; G. Leblond and A. Levesque for ornithological information; Z. Bochenski, H. James, C. Lefèvre, A. Louchart, N. Serrand and N. Zelenkov for their suggestions which improve the quality of the manuscript.

1 - Introduction

1 The Lesser Antilles, like all the Caribbean islands of which they form the southern component, are a group of tropical oceanic islands with a highly endemic vertebrate fauna (e.g., Hedges, 1996, 2006; Ricklefs & Bermingham, 2008), including a large number of birds (Bond, 1948). This unique avian community, resulting from the position of the archipelago at the intersection of biological flows between the South American continent, the Bahamas and Greater Antilles (Lack, 1976; Bond, 1979), is fragile and threatened by several (climatic, ecological) factors that can lead to the disappearance of species (e.g., Pregill & Olson, 1981). The vulnerability of Caribbean vertebrates is illustrated by the many species that have become extinct since the end of the Pleistocene. In fact, the Caribbean Islands have experienced more mammalian extinctions than any other region on Earth (Turvey, 2009; Dávalos & Turvey, 2012), in addition to the loss of numerous squamates (e.g., Powell & Henderson, 2012; Bochaton et al., 2021a) and birds (Turvey, 2009; Steadman & Takano, 2013; Suárez & Olson, 2015). The Guadeloupe Islands form the central part of the Lesser Antilles and are currently home to 285 bird species, 81 of which are breeding populations (Levesque & Delcroix, 2019). Of these species, 8 are endemic to the Lesser Antilles and one is strictly endemic to Guadeloupe, the Guadeloupe Woodpecker Melanerpes herminieri (Benito-Espinal, 1990; Levesque & Mathurin, 2008; Eraud et al., 2009).

2 The fact that human activity is responsible for the disappearance of large terrestrial mammals has been established for some time (Steadman et al., 1984a). Several other causes potentially explaining the loss of biodiversity have also been identified, including climate change and environmental modifications (Pregill & Olson, 1981), or the reduction in land area after the last glaciation (Dávalos & Russel, 2012). The increase in the number of direct dates and detailed stratigraphic contexts has demonstrated the majority of regional extinction events to be a direct or indirect consequence of human activity (Cooke et al., 2017; Orihuela et al., 2020). Diverse societies have occupied the Caribbean since the mid-Holocene and the relative importance of human activity underlying these extinctions (overhunting, habitat transformation through cultivation, or introduction of invasive species) are yet to be explored for different subregions and faunal categories.

3 Recent studies of terrestrial vertebrates in Guadeloupe have revealed the massive extinction of endemic snakes and lizards following European colonization (Bochaton et al., 2021a). Data on rodents and bats indicates that most of the extinction/extirpation events occurred during the Holocene (Stoetzel et al., 2016; Lenoble, 2019; Goedert et al., 2020). While avian communities are also concerned by this process, no synthesis is currently available. Three endemic species are known to have become extinct (the Guadeloupe Parrot Amazona violacea, the Guadeloupe Parakeet Psittacara labati and the Lesser Antillean Macaw Ara guadeloupensis) (Du Tertre, 1654, 1667-1671; Labat, 1742; Clark, 1905; Gala & Lenoble, 2015) and 5 taxa to have been extirpated or become extremely rare (the American Flamingo Phoenicopterus ruber, the Eskimo Curlew Numenius borealis, the Black-capped Petrel Pterodroma hasitata, the Burrowing Owl Athene cunicularia guadeloupensis and the House Wren Troglodytes aedon guadeloupensis) (Labat, 1722; Barlow, 1978; Gala et al., 2020).

4 These examples show that the avifauna of Guadeloupe was equally affected by faunal turnover in the Caribbean over the last few hundred years. They also show our understanding of the islands’ past avifauna to be based essentially on observations of 17^th and 18^th century chroniclers (Breton, 1665, 1978; Du Tertre, 1654, 1667-1671; Labat, 1722, 1724, 1742) and subsequent mid-19^th century ornithological studies (Lawrence, 1879; Noble, 1916; Danforth, 1939). Until now, the archaeological and paleontological record has been underexploited in terms of documenting the past biodiversity of the Guadeloupe Islands.

5 Here we present the results of a morphometric analysis of bird remains recovered from 5 of the 6 Guadeloupe Islands and dating from the Late Pleistocene through to the Historical period. This analysis combines fossil and subfossil material from 24 archaeological and paleontological sites, providing an unprecedented dataset, especially considering the limited number of bird remains from the Guadeloupe Islands in publications from 1960 to 2018, most of which are not identified by species or genus. When access was possible, previously studied bird remains were included in our study to obtain more precise, reliable attributions.

6The analysis was designed to (a) compare the past and modern avifauna of the Guadeloupe Islands in order to highlight significant changes in avian communities since the Late Pleistocene and (b) establish a chronology for late Quaternary avian extinction and extirpation events in the Guadeloupe Islands. Finally, this data allows for an evaluation of the relative roles of environmental changes and human impact as potential causes influencing avian population dynamics.

2 - Material and methods

7 The Guadeloupe Islands are situated in the central Lesser Antilles arc. The archipelago is composed of six main islands, with the two largest ones being separated by a low mangrove swamp: the western mountainous volcanic island of Basse Terre (848 km², 16°10’N, 61°40’W) and eastern dry limestone island of Grande Terre (585 km², 16°19′03″N 61°25′43″W). The smaller volcanic islets of Les Saintes (13 km², 15°51′40″N 61°36′35″W) lie 10 km to the south of Basse Terre. The dry limestone islands of Marie Galante (158 km², 15°56′09″N 61°16′14″W), La Désirade (23 km², 16°19′06″N 61°03′07″W) and Petite-Terre (1.5 km², 16°10′15″N 61°6′55″W) are located a few kilometers to the east and southeast of Grande Terre (fig. 1). While the emergence of the Guadeloupe Islands was relatively recent, the process took many millions of years. Basse Terre was formed by a succession of volcanoes, the oldest of which are estimated to be 3 million years old (Mathieu et al., 2013), while the final emergence of the eastern limestone islands was more recent, estimated to have been half a million years old, even though several previous episodes of emergence occurred over the past 5 million years (Cornée et al., 2012). During periods of low sea levels (e.g., Late Pleistocene from 40,000 to 11,700 years ago), the islets of Petite-Terre and La Désirade were joined to the larger islands of Basse Terre and Grande Terre, forming the Guadeloupe Bank. The natural landscape of the Guadeloupe Islands is varied. Basse Terre differs from the other islands by its mountainous topography, including the highest mountain of the Lesser Antilles, the Soufrière active volcano (1467 m). The other islands in the archipelago do not exceed 300 m. Another feature of Basse Terre is vertical zonation in vegetation due to rainfall: dry forest on the coast, moist semi-evergreen forest at middle altitudes and rain forest in higher areas, while the other islands only support dry forest, dotted in places with marshes or lowland moist forest (Rousteau et al., 1994). The limestone substrate of Grande Terre, Marie Galante and La Désirade considerably influences the development of karstic features (i.e. caves, dolines and dry valleys). The Guadeloupe archipelago are the most extensive limestone islands in the Lesser Antilles and offer remarkable potential for studying the Amerindian occupation of the islands (Grouard et al., 2014) and documenting non-anthropogenic fossil-bearing deposits. Several research projects have successfully produced an inventory of caves on the Guadeloupe Islands and conducted excavations that have yielded numerous vertebrate remains (Lenoble et al., 2009; Grouard et al., 2014). Among these projects, two were dedicated to study of the past biodiversity of the islands of Guadeloupe, one for the Pleistocene period, BIVAAG (Biodiversité insulaire vertébrée ancienne des îles de Guadeloupe), and the second for the Holocene, ECSIT (Écosystèmes insulaires tropicaux, réponse de la faune indigène terrestre de Guadeloupe à 6 000 ans d’anthropisation du milieu). The fossil bone assemblages collected during these two research projects form the core of the material used in this study, complemented by published archaeological assemblages.

8 The bird remains come from 24 archaeological and paleontological sites (caves, rockshelters and open-air sites) on 5 of the 6 islands of Guadeloupe (fig. 1). The investigated sites cover periods ranging from the Late Pleistocene to the Historical period, and can be divided into 5 main periods. The first period predates the arrival of humans (PH) in Guadeloupe and groups together the Late Pleistocene (PH-Pl, from 30,000 to approximately 11,700 yBP) and early Holocene (PH-Ho, 11,700–6,000 yBP). For the following 4 periods, we used the periodization of Amerindian occupation of the Lesser Antilles and Puerto Rico proposed by Bérard (2019, table 3). The first nomadic human groups were documented in the area between Guadeloupe and Puerto Rico in the Archaic Age (AR), also known as the Preceramic, around 3,000 B.C.E (Napolitano et al., 2019). In the eastern Caribbean, this period ends with the appearance of sites assigned to the Ceramic Age (400/200 B.C.E.). We combined the Early and Middle Ceramic periods (EC-MC), up to 800 CE, and the Late and Final Ceramic (LC-FC) until the Contact period (CO). The 16^th to 17^th centuries correspond to the period during which indigenous pre-Columbian populations experienced both peaceful and hostile exchanges with European colonists (Keegan et al., 2013; Hofman et al., 2019). Beginning in the early 17^th century, European colonial populations became permanently established (Keegan & Hofman, 2017) and throughout the following period, the Colonial period (CL), the indigenous populations lost control of the islands.

9The 24 sites of interest are briefly presented below, with the context of 7 of them (Grotte des Bambous, Pointe Gros Rempart 6, Grotte Blanchard, Grotte Blanchard 2, Grotte Cadet 2, Abri Cadet 3 and Folle Anse) being detailed in Gala et al., 2020 (SI1). The archaeological collections of Sainte-Rose, Anse Sainte-Marguerite, Anse à l’Eau, Morel, Petite Rivière, Taliseronde and Folle Anse are stored in the Edgar Clerc Museum and the ones of Roseau, La Pointe de Grande Anse, Embouchure de la rivière Baillif, Anse Sainte-Marguerite, Morel, Anse à la Gourde, Pointe du Helleux; Îlet du Gosier, Petite Rivière, Folle Anse, Tourlourous, Grande Anse de Terre-de-Bas are stored in the Dépôt du Service Régional de l’Archéologie de Guadeloupe (Le Moule-Guadeloupe). Five sites are in Basse Terre (Roseau, La Pointe de Grande Anse, Cathédrale de Basse-Terre, Embouchure de la rivière Baillif, Sainte-Rose) and date from the Ceramic to Contact periods (fig. 1; tab. 1). Four are pre-Columbian sites excavated since the 1990s, and the fifth corresponds to the Barbotin collection of the Ceramic site of Sante-Rose (Dindé), without any other information. The three main occupations of Plage de Roseau (Capesterre Belle Eau) date from the Late Ceramic to Contact periods (Bochaton et al., 2021b). The radiocarbon dates from the inhabited area and cemeteries of La Pointe de Grande-Anse (Trois-Rivières) provide a chronology through the Early Ceramic and Late Ceramic periods (van den Bel & Romon, 2010). Excavations of the Amerindian midden of Cathédrale de Basse-Terre (Basse Terre) yielded a significant archaeological assemblage (malacofauna and vertebrate fauna, lithic and shell industry and a high proportion of ceramics) from the Early Ceramic period (Bonnissent & Romon., 2004). The pre-Columbian site of Embouchure de la rivière Baillif (Baillif) dates back to the Late Ceramic period (Gassies, 1996; Romon, 2001).

10 Eight sites are located in Grande Terre (Grotte de l’Anse à l’Ecu, Anse Sainte Marguerite, Grotte des Bambous, Morel, Anse à l’Eau, Anse à la Gourde, Pointe du Helleux and Îlet du Gosier) and date from the Ceramic to Historical periods. The excavations of several pre-Columbian sites near Le Moule date back to the 1950-70s (Clerc, 1965): Anse Sainte Marguerite, Morel and Anse à l’Eau. We studied material from the Late Ceramic period of Anse Sainte-Marguerite (Courtaud & Romon, 2004), the entire Ceramic period of Morel from Delpuech’s excavations in 1995 and 1999 (Delpuech et al., 1998; Hofman et al., 1999), and Anse à l’Eau (Clerc, 1964, 1968, 1970). The coastal pre-Columbian site of Anse à la Gourde (Saint Francois), on the east end of Grande Terre, was the subject of excavations from 1995 to 1999. Bird bones analyzed in this paper are from the 1995 and 1997 excavations (Delpuech et al., 1997, 2001; Hofman et al., 2001). Since the 1980s, several surveys and excavations have been carried out at the pre-Columbian site of Pointe du Helleux (Sainte-Anne) (Grouard et al., 1999). We analyzed the bird bones from the Late Ceramic period recovered during the 1997 excavations. The site of Îlet du Gosier (Gosier) was inventoried by Pierre Bodu in 1984 and subsequent surveys yielded occupations and burials dated to the Late Ceramic period (Romon, 2004). The Late Holocene natural site of Grotte de l’Anse à l’Ecu (Le Moule) was discovered in 2008 by C. Stouvenot and yielded a bone surface collection of ancient (presence of extinct taxa) and modern (historical introductions) native fauna (Lenoble et al., 2010).

11 Two sites are from La Désirade (Petite Rivière and Pointe Gros Rempart 6), dating from the Ceramic to Historical periods. The bird material of Petite Rivière (Baie-Mahault) comes from the Barbotin collection stored in the Edgar Clerc Museum and from Bodu and de Waal’s excavations (Bodu, 1984; de Waal, 2006) of this Ceramic period site.

12 Eight sites are located on Marie Galante (Taliseronde, Grotte du Morne Rita, Tourlourous, Grotte Blanchard, Grotte Blanchard 2, Grotte Cadet 2, Abri Cadet 3, Folle Anse) and span the period from the Late Pleistocene to present. The bird bones from the Ceramic site of Taliseronde come from the Barbotin collection (Barbotin, 1970; Emond, 1980; Haviser, 1997). Part of this bird assemblage is stored in the Florida Museum and was studied by Wing (Wing & Reitz, 1982), and was not analyzed in our study.

13Three additional sites yielded bird remains: the pre-Columbian sites of Tourlourous (Capesterre de Marie Galante) and Folle Anse (Grand-Bourg de Marie Galante), and the Late Holocene natural site of Grotte du Morne Rita (Capesterre de Marie Galante). The material in our study derives from the Early Ceramic period of Tourlourous (Barbotin, 1970; Etrich, 2001; Colas et al., 2002; Fourteau, 2009; Serrand, 2016) and from the old (Early Ceramic) and new excavations (Late Ceramic) of Folle Anse that document an archaeological sequence spanning from the Early to Late Ceramic period (Barbotin, 1970, 1991; Lenoble, 2019; Goedert et al., 2020). Several remains reported by Grouard (2001a) from Chenorkian’s excavations (1999a,b), and by Williams and Steadman (2001) and Olson and López (2008) from the excavations by Barbotin, are not examined in this study.

14The bird bones collected on the surface during Bodu’s excavation of Grotte du Morne Rita and stored in the Dépôt du Service Régional de l’Archéologie de Guadeloupe (Bodu, 1985) were included and testify to the more recent frequentation of the cave by seabirds during the Contact and Colonial periods. A single archaeological site is on Les Saintes islets (Grande-Anse de Terre-de-Bas) and dates from the Ceramic period (Hofman, 1995; Grouard, 2001a). In addition to analyzing the collections of the above-mentioned sites, we also updated and re-examined the bird remains from 11 sites: Embouchure de la rivière Baillif, Morel, Petite Rivière, Folle Anse, Grande-Anse de Terre-de-Bas (Grouard, 2001a, 2007), Anse à la Gourde, (Grouard, 2001a; 2001b), Pointe du Helleux (Grouard, 2001a; 2001c), Pointe Gros Rempart 6 (Boudadi-Maligne et al., 2015), Tourlourous (Grouard, 2016), Grotte Blanchard 2 (Mallye et al., 2018) and Abri Cadet 3 (Stouvenot et al., 2014).

15 Seven radiocarbon dates were obtained from bird bones discovered on 5 sites (Grotte des Bambous, Anse à l’Eau, Pointe Gros Rempart 6, Grotte Blanchard 2, Grotte du Morne-Rita). Five of the dated samples concern remains of Audubon’s Shearwater Puffinus lherminieri, which were calibrated using the Marine20 curve (Heaton et al., 2020) with respect to the diet of these birds using a regional mean value of -146 ± 114 yrs as a correction for the marine reservoir effect (DiNapoli et al., 2021). The other two dates were obtained on bones of the Yellow-crowned Night-Heron Nyctanassa violacea and Burrowing Owl Athene cunicularia and calibrated using the Intcal20 terrestrial curve (Reimer et al., 2020).

16 The bird remains were identified by direct comparison with modern specimens from the skeletal collections of the PACEA laboratory, University of Bordeaux (Lenoble et al., 2019; Guimaraes & Lenoble, 2019), as well as the American Museum of Natural History of New York (AMNH), Musée Edgar Clerc of Le Moule (MEC), Muséum national d’Histoire naturelle of Paris (MNHN), Natural History Museum of London (NHMUK), Royal Ontario Museum of Toronto (ROM), University of Florida-Florida Museum of Natural History of Gainesville (UF), University of Michigan Museum of Zoology of Ann Arbor (UMMZ) and the Smithsonian Institution - National Museum of Natural History of Washington (USNM). Details concerning the comparative material are provided in the Supplemental Material (Appendix A). For comparative purposes, we also included several fossils from Cueva Catedral and Cueva Clara (Wetmore, 1922), Burma Quarry (Pregill et al., 1994), Folle Anse (Wing, 1973; Wing & Reitz, 1982; Olson & Lopez, 2008), Montserrat (UF collection, unpublished material), Tyto noeli from Barbuda (USNM 359244) observed during our study session at the Smithsonian Institution and University of Florida. Specimens from Cathédrale de Basse-Terre were identified in collaboration with D. Steadman (UF). We use the term “fossil” to refer to bone from paleontological or archaeological sites and distinguish it from “modern” specimens from osteological collections.

17 The number of identified specimens (NISP) and the minimum number of individuals (MNI) were recorded for each stratigraphic unit of the site (Appendix B), taking into account bone lateralization, age (based on bone texture: juvenile, subadult, adult) and sex (presence of medullary bone).

18 The 3D surface models of the various fossil, subfossil and modern specimens were produced using microtomography with a lCTscanner (GE Phoenix vjtomejx s) at the PLACAMAT Platform of the University of Bordeaux. We selected paleontological and archaeological bone remains that were fragile or difficult to identify in order to facilitate comparisons with osteological collections and several anatomical elements from rare and endangered species for comparison with our sample.

19 The taxonomy follows Clements et al. (2019). The anatomical terminology is after Baumel and Witmer (1993) with some exceptions. Bone measurements were taken with a digital caliper to the nearest 0.1 mm and follow those described by von den Driesch (1976). In the case of 3D models, the measurements were derived with the Avizo® and Meshlab® software packages. Morphometric data on some elements of different groups of birds (Wetmore, 1922; Olson & Hilgartner, 1982; Steadman & Takano, 2013; Steadman et al., 1984b; Tellkamp, 2005; Tomek & Bochenski, 2009) was also integrated into this study.

20 A review of previous accounts (historical sources, museum specimens, past and modern ornithological data, fossil distribution in the Lesser Antilles) of bird families identified during our study provides an overview of current data for avian communities in the region. We included early accounts of travelers from the 17^th and 18^th centuries, such as Breton’s dictionary (1665) and his reports in Relations de l’île de la Guadeloupe (1978), Du Tertre’s Histoire génerale des Antilles (1667-1671), Labat’s Nouveau voyage aux isles de l’Amérique (1722) and Allan de Cramant’s Voyage d’Allan de Cramant à la Martinique et à la Guadeloupe, 1732 (2010), as well as all available historical sources with the mentions of the birds and their exploitation.

21 The collections inherited from the era of bird collecting during the first half of the 19^th century and subsequent ornithological studies, as well as lists and publications of Guadeloupe birds by Lawrence (1879), Noble (1916) and Danforth (1939), complemented by Bond’s observations in the early versions of his checklist (1945), also contain useful information. The status and range of birds in the Guadeloupe Islands and West Indies were reported following Levesque and Delcroix (2019, 2021) and Raffaele et al. (2020). Finally, we combined this information with data from the STOC Program (“Suivi Temporel des Oiseaux Communs”) that produces indices (or indicators) of annual changes from 2014 to 2019 in the abundance of various species of common birds in Guadeloupe (Levesque et al., 2020). We classified the taxa into five categories: 1) globally extinct; 2) locally extirpated -- no longer exist in Guadeloupe Islands; 3) locally extirpated but still exist elsewhere on one or more islands of Guadeloupe; 4) introduced; 5) extant.

3 - Results

22 Of a total of 4,584 bird remains, more than half (NISP = 2,457) were identified to order, family, genus or species. At least 55 taxa belonging to a minimum of 330 individuals representing 26 families and 16 orders were identified in one or more layers of the studied sites (tabs. 1 & 2; Appendix B). Most sites date to the Ceramic period and the largest number of remains are from the Marie Galante sites that produced evidence from the Late Pleistocene to the Historical period (fig. 2).

23 We report the results of fossil and subfossil bird bones grouped by family. Shearwaters, petrels (Procellariidae) and songbirds (Mimidae, Turdidae, Thraupidae, Tyrannidae and Icteridae) are the most frequent taxa, followed by pigeons and doves (Columbidae). We also identified waterfowl (Anatidae), pheasants (Phasianidae), nightjars and hummingbirds (Caprimulgidae and Trochilidae), wading birds (Rallidae, Charadriidae, Scolopacidae, Ardeidae), pelicans (Pelecanidae), gulls (Laridae), tropicbirds (Phaethontidae), boobies (Sulidae), diurnal and nocturnal raptors (Pandionidae, Accipitridae, Falconidae, Tytonidae, Strigidae), woodpeckers (Picidae) and parrots (Psittacidae). For specimens that were difficult to identify to taxon, we specify the specific characters that allow them to be referred to a species or genus. The results concerning Strigiformes fossil and subfossil remains (Tytonidae and Strigidae) are assessed in the light of previous investigations (Gala et al., 2020; 2022).

3.1 - Anseriformes – Anatidae

24This family is represented by 2 genera (NISP = 12) and 7 individuals from 6 sites distributed across all the islands of Guadeloupe and dating from the Ceramic to Colonial periods (tab. 2, Appendix B).

25Four incomplete bones from Folle Anse are similar in size and shape to the West Indian Whistling-Duck Dendrocygna arborea and were tentatively assigned to it. In order to confirm this attribution, these bones must be compared with other reference specimens of congeneric species that are similar in size to Dendrocygna arborea: the Fulvous Whistling Duck Dendrocygna bicolor and Black-bellied Whistling-Duck Dendrocygna autumnalis. An omal part of coracoid from Cathédrale de Basse-Terre does not present features that allow it to be assigned to species (Dendrocygna sp.). The Whistling-Duck was previously identified in pre-Columbian assemblages based on a femur labelled Dendrocygna arborea and determined by Alexander Wetmore. The bone is exhibited at the Edgar Clerc Museum and comes from an unknown Ceramic site on the Atlantic coast of Grande Terre (Clerc, 1965).

26 A fragmented scapula from Roseau was referred to the genus Anser. Six remains of an unidentified Anatidae were reported from this latter site and 4 other sites belonging to the Colonial (Pointe Gros Rempart 6) and Ceramic periods (Cathédrale de Basse-Terre, Anse à l’Eau and Grande-Anse de Terre-de-Bas).

27Since the discovery of the Americas, Anseriformes have been described as abundant in the West Indies and as being consumed by Amerindian groups in Guadeloupe, assuming that the report of geese bones in Caribbean pots by Martyr d’Anghiera (1907). According to Bénito-Espinal (1978), the Whistling-Duck is also the nocturnal species that the chroniclers named Vigeon (Breton, 1978; Du Tertre, 1667-1671) or Vingeon (Labat, 1724), like a Dendrocygna. The West Indian Whistling-Duck was extirpated from several islands during the modern period (Raffaele et al., 2020). It relies on freshwater, brackish, and marine wetlands for feeding and roosting (Sorenson et al., 2004), and was not known to be nesting currently in Guadeloupe (Bénito-Espinal & Hautcastel, 2003), until a recent report of its presence in Terre-de-Bas, Les Saintes by Levesque and Sorenson (2012) (fig. 3).

28Except for the Wetmore specimen from an unspecified location on Grande Terre, these Whistling-Duck and geese bones constitute the first mentions of fossil Anseriformes in Guadeloupe. A few bones belonging to this order have also been identified in the Lesser Antilles: Dendrocygna arborea, Dendrocygna autumnalis, Dendrocygna sp., Aythya collaris and Neochen barbadiana nov. sp. on Barbados (Brodkorb, 1964; Wing, 1991); Anas bahamensis on Antigua (Pregill et al., 1994); and Neochen cf. jubata on Barbuda (Pregill et al., 1994).

3.2 - Galliformes - Phasianidae

29Very few remains of Galliformes (NISP = 4) have been found in the paleontological and archaeological record of Guadeloupe. They come from 3 sites on Grande Terre dating from the Ceramic (Anse à la Gourde and Îlet du Gosier) to Colonial periods (Grotte des Bambous, Cochard et al., 2019; Goedert et al., 2020; cf. tab. 2, Appendix B).

30 A sternal part of right coracoid from Grotte des Bambous (Bam-O-01-2014) was assigned to the turkey Meleagris sp., based on the characters described by Tomek and Bochenski (2009). The turkey was introduced in the mid-17^th century (Anonyme, 1660; Du Tertre, 1667-1771). The 3 specimens (a humerus and 2 ulnae) of juvenile Galliformes do not show any specific similarities to the modern Phasianidae, Numididae and Odontophoridae specimens in the PACEA laboratory collection (Appendix A). The ulnae from Anse à la Gourde and Îlet du Gosier are, respectively, larger and smaller than any of the Galliformes specimens examined. The humerus from Îlet du Gosier could fit with turkey, but more osteological material (e.g., Cracidae was not available for comparison) is needed to improve the taxonomic resolution.

31A single example of remains of an unidentified Phasianidae was previously reported from Basse Terre from a colonial level in Gare Maritime, but was not revised in our study (Tomadini et al., 2014).

3.3 - Columbiformes - Columbidae

32Columbidae is one of the most successful bird families in the world, but also one of the most threatened (Walker, 2007). The excellent flying abilities afforded by larger wings and flight muscles (Hartman, 1961) enabled this family to colonize oceanic islands. Currently, 22 species belonging to 7 genera are listed for the Caribbean region (Baptista et al., 1997; Raffaele et al., 2020). Their biogeographic value and the common occurrence of remains from this family in the Caribbean archaeological and paleontological record makes the Columbidae particularly interesting. In order to improve species-level identification of Columbidae bone remains, a morphometrical analysis of 8 postcranial elements was conducted on modern specimens belonging to the 5 genera that represent most of the indigenous taxa of the West Indies (Gala et al., 2021). This study identified 42 diagnostic characters and five size classes (large and small pigeons, large, medium and small doves).

33More than two hundred bone remains (NISP = 216) from 17 sites assigned to the pre-human to Colonial periods belong to this family, among which 4 genera (23 individuals of Patagioenas, 10 of Columbina, 15 of Geotrygon and 14 of Zenaida) have been identified in the palaeontological and archaeological record (tab. 2, Appendix B). Their presence is well documented in the sites of Basse Terre (NISP = 95) from the Ceramic to Contact periods (Roseau, Cathédrale de Basse-Terre and Embouchure de la rivière Baillif) and less so on Les Saintes (NISP = 5) at the Ceramic site of Grande-Anse de Terre de Bas. On the other islands, bone remains from this family have been identified at 6 sites of Grande Terre (NISP = 25) from the Archaic to Colonial periods (Grotte de l’Anse à l’Ecu, Anse Sainte Marguerite, Grotte des Bambous, Morel, Anse à l’Eau and Anse à la Gourde), 2 sites of La Désirade (NISP = 17) from the Ceramic to Colonial periods (Petite Rivière and Pointe Gros Rempart 6) and 5 sites of Marie Galante (NISP = 76) from pre-human to Ceramic periods (Taliseronde, Tourlourous, Grotte Blanchard, Abri Cadet 3, and Folle Anse).

34Based on a recent morphometrical analysis (Gala et al., 2021), we were able to refer 74 bones to species: 3 species of Patagioenas (the Scaly-naped Pigeon P. squamosa, White-crowned Pigeon P. leucocephala and Plain Pigeon P. inornata), 2 of Geotrygon (Ruddy Quail-Dove G. montana and Bridled Quail-Dove G. mystacea) 1 of Zenaida (Zenaida Dove Z. aurita) and the only species representing the genus Columbina (Common Ground-Dove C. passerina). For the other 58 bones, as the absence of diagnostic characters and/or the fragmentary nature of the specimens did not allow attribution to a species, the specimens were assigned to pigeons (Patagioenas squamosa/inornata and Patagioenas sp.) and doves (Geotrygon cf. mystacea, Geotrygon sp., Zenaida cf asiatica, Zenaida cf. aurita and Zenaida sp.) based on their size. Other traits characteristic of this family were identified on 84 bone remains (Columbidae indet.). The fragmentary nature and lack of distinguishing features of a particular species or genus made it difficult to assign these remains to specific size classes.

35Two bones from layer 5 of Grotte Blanchard, a right coracoid (GB-2014-O-76) and a left carpometacarpus (GB-2014-O-77), were previously assigned to the large White-winged Dove Zenaida asiatica (Gala et al., 2016). However, osteological study of a modern specimen (Gala et al., 2021) assigned these two bones to Z. cf. asiatica for the carpometacarpus (Appendices C-D) and to Z. aurita for the coracoid (Appendices E-G). The carpometacarpus specimen falls within the size range of large Z. asiatica and 4 characters out of 11 described for this bone fit with this species unambiguously, while the seven others (dorsal margin of the os metacarpale minus near trochlea carpalis; development of trochlea carpalis; distal dorsal margin of trochlea carpalis; curve of the facies articularis ulnocarpalis; processus extensorius; ventral margin of the os metacarpale minus; symphysis metacarpalis distalis) do not distinguish it from the morphologically similar Z. aurita. The four diagnostic characters are 1) the alignment of the ventral margin of the facies articularis ulnocarpalis and the ventral margin of the proximal portion of the os metacarpale minus in the proximal area of the bone; 2) the shaft of the major metacarpal towards the processus alularis is very slightly bowed and does not slope up as markedly (Olson & Hilgartner 1982); 3) the facies articularis digitalis major is closer to the facies articularis digitalis minor; and 4) the dorsal margin between the distal part of the sulcus tendineus and tuberosity of the os metacarpale majus is narrow and concave (fig. 4).

36Columbina passerina, G. mystacea (endemic to the West Indies) and Z. aurita are currently common birds in the Guadeloupe Islands (Levesque & Delcroix, 2019). The other species (P. squamosa, P. leucocephala, G. montana, and Z. asiatica) are uncommon or rare, while P. inornata is not known to occur in the Lesser Antilles. The 5 remains of P. inornata from Cathédrale de Basse-Terre are the first evidence for this species in the Guadeloupe Islands and Lesser Antilles (fig. 3). Currently, the Plain Pigeon is restricted to the Greater Antilles where it is an endemic species (Raffaele et al., 2020). In addition, the fossil from Grotte Blanchard tentatively assigned to Zenaida cf. asiatica would be the first evidence (fig. 3) of this species in the Lesser Antilles paleontological record. This dove species inhabits dry and poorly forested environments of the Bahamas and Greater Antilles (Raffaele et al., 2020), and was reported as nesting in Guadeloupe ten years ago (Levesque, 2013).

37It should be noted that birds belonging to this family of mostly frugivorous arboreal birds are threatened by human impact via environmental degradation, hunting and the introduction of predator species (Walker, 2007). The most affected species are the regionally endemic Patagioenas leucocephala (Delcroix et al., 2016), P. inornata, both near-threatened species in IUCN 2020, and Patagioenas squamosa. The latter species was well represented in our study, was abundant in the 17^th century (Breton, 1665; Du Tertre, 1667-1671; Labat, 1724), and is the main game bird in Guadeloupe (Lawrence, 1879; Noble, 1916). The process of rarefaction of pigeons in the 19^th and 20^th centuries (Ballet, 1895; Noble, 1916) nearly led to the disappearance of this species from La Désirade in the 1950s (Pinchon, 1976) and it is found today mainly in the forests of Basse Terre (Feldmann, 1998; Levesque & Lartiges, 2000). G. monatana is also present essentially in the forests of Basse Terre, where it is uncommon and localized (Feldmann, 1998; Levesque & Lartiges, 2000). On the other hand, and unlike today, doves were rare when Europeans settled Guadeloupe, while pigeons were abundant (Breton, 1665). The recent inversion of the ratio between pigeons and doves is documented by ornithological data, with a high number of observations of doves (Zenaida aurita) and limited reports of pigeons (Levesque et al., 2020). In addition, our results show that two additional extirpations occurred in the past. They concern Zenaida cf. asiatica, represented by a bone identified in a pre-human layer from Marie Galante, and Patagioenas inornata, limited remains of which have been identified in a Ceramic period site on Basse Terre. During the Ceramic period, after Procellariidae, Columbidae was the second most abundant family in the studied archaeological material. Reports of Amerindian groups hunting pigeons and doves go back a long way. Sir Monpilliers, who ran aground in Guadeloupe in 1635, reported that the crew of the wrecked ship was taken in and fed by the Amerindians, who gave them, among other things, ramiers (a French word for pigeon) to eat (Monpilliers, 1654: 57). This report is in agreement with the West Indian archaeological data which shows Columbidae to be the most represented species in middens of this period (Grouard, 2001a; Wing, 2001; Newsom & Wing, 2004).

38We provide a list of all columbid fossil records in the Lesser Antilles in the supplemental material (Appendix H). Bone remains have previously been identified at Abri Cadet 3 on Marie Galante and on numerous other islands, mainly to the north of Guadeloupe (Anguilla, Tintamarre, St Martin, St Kitts, Nevis, St Eustache, Barbuda, Antigua, Saba and Montserrat), and on 2 islands in the southern Lesser Antilles (Martinique and Tobago).

3.4 - Caprimulgiformes - Caprimulgidae

39Three bird bones from this family, belonging to 2 individuals, were found in 2 sites from 2 islands (Grande Terre and Marie Galante) (tab. 2, Appendix B). The first individual is represented by a single bone discovered in a layer of Grotte de Bambous dating from the Archaic to Colonial periods, and the other 2 bones of the second individual were recovered from a pre-human layer (8) of Grotte Blanchard.

40 The Grotte des Bambous specimen was compared to the carpometacarpi of Antillean Nighthawk Chordeiles gundlachii and Common Nighthawk C. minor. These two species are similar in size and morphology. For this reason, it was difficult to assign the fossil specimen to one particular species. The measurements of 9 modern specimens show a relatively weak separation between smaller C. minor and larger C. gundlachii (Appendix I), and the distal measurement of the Bambous specimen is compatible with both species. Based on morphological characters, the carpometacarpus is compatible with that of C. gundlachii in 3 characters (distal caudal edge caudally projected, shape of distal shaft and large distal part of spatium intermetacarpale) and C. minor in 2 characters (approximately straight ventral edge of facies articularis digitalis major and large facies articularis digitali minor). While two of these characters were used by Steadman and Takano (2013) to distinguish 5 genera of Caprimulgidae, we only observed interspecific differences for these two characters within Chordeiles, leading us to assign this specimen tentatively to Chordeiles gundlachii.

41 The mandibles and humerus from Grotte Blanchard belong to a large individual of Nightjar Antrostomus. The shape of epicondylus ventralis and processus supracondylaris dorsalis allow us to assign it to this genus (fig. 5). The left distal humerus is smaller than that of Antrostomus carolinensis and larger than that of A. vociferus and A. cayennensis (Appendix I). Due to the fragmented nature of the available material, we failed to assign the specimen to species.

42 Chordeiles gundlachii currently nests in Guadeloupe (Levesque & Delcroix, 2019); however, this is a recent phenomenon, with the appearance of the first individuals reported in the early 2000s (Levesque, 2001) and reported nesting since 2013-2014 only (Perlut & Levesque, 2020). Despite the difficulty in observing this species, the nightjar is nevertheless part of the West Indian avifauna, present in the different islands of the Greater and Lesser Antilles (Bond, 1936). In Guadeloupe, the bird is historically known as Héleux, and a cove on the southern coast of Grande Terre is named after it. It is under this term that Allan de Cramant reported the hunting of a specimen from the island of Petite Terre in 1732 (Allan de Cramant, 2010). The vague description provided by the author does not allow the species to be identified. However, one account suggests that the historically-present species was of the genus Antrostomus, not Chordeiles. This assumption is based on a representation on a vellum entitled “Engoulevent de la Guadeloupe” showing a Caprimulgiformes with red and black striped feathers (fig. 5), a character in agreement with a member of the genus Antrostomus (MNHN, Direction des Bibliothèques, Collection des vélins, Portefeuille 79, folio 06). The wording “natural size” on the vellum suggests an early 19^th century depiction (C. Lim, in litteris), which is consistent with the mention of a nightjar of the genus Caprimulgus on the list of birds of Guadeloupe by L’Herminier (Lawrence, 1879). No additional Caprimulgidae were recorded in the 19^th and 20^th century ornithological studies (Lawrence, 1879; Noble, 1916; Danforth, 1939; Terborgh et al., 1978; Benito-Espinal & Hautcastel, 2003), suggesting that the historical form of Nightjar in Guadeloupe disappeared during the 19^th century (fig. 3). Fossil remains of Chordeiles gundlachii occur only in Antigua (Pregill et al., 1994).

3.5 - Caprimulgiformes - Trochilidae

43Although they have a unique morphology, the bones of this family are rarely recognized in the fossil or archaeological record due to their very small size. The 2 hummingbird specimens from pre-human layers of 2 sites located on Marie Galante Island are the only examples of this family in the studied material (tab. 2, Appendix B). Based on several characters, the humerus from layer 5/6 of Abri Cadet 3 was assigned to the endemic species of Purple-throated Carib Eulampis jugularis or the Green-throated Carib Eulampis holosericeus. The humerus is smaller than that of Eulampis holosericeus and shares all the traits of the proximal end and dorsal edge of crista deltopectoralis, except for the shape of the tuberculum dorsale. This latter feature is more pointed and similar to that of Eulampis jugularis. The carpometacarpus from layer 9 of Grotte Blanchard needs to be reexamined and compared with more Trochilidae species in order to assign it to a genus or species.

44 Two large species of hummingbirds of the genus Eulampis and one small species, the Antillean Crested Hammingbird Orthorhynchus cristatus that is currently present in the Guadeloupe Islands (Levesque & Delcroix, 2019), are adapted to the different ecological conditions of each island (Lack, 1973). Eulampis holosericeus occurs mainly in coastal scrub forest and Eulampis jugularis in montane forest. Orthorhynchus cristatus frequents both habitats in Guadeloupe (Terborgh et al., 1978). The 3 species inhabit the forests of Marie Galante today (Benito-Espinal, 1990) and Eulampis jugularis is found in the wet, shaded forest of the ravines in the center of the island (eBird, 2021).

45 While our material is limited to the pre-human period, historical accounts provide little information about this family during the Colonial period. Adoption of the hunting of hummingbirds by early colonists was reported by French chroniclers as a Caribbean heritage (Breton, 1665; Du Tertre, 1667-1671; Pinchon, 1976) and provides evidence of human exploitation (e.g., feather use) of these birds since the Amerindian times.

46Eulampsis sp. has been mentioned at the Ceramic period site of Pointe du Helleux on Guadeloupe (Grouard, 2001a) and was not revised in the course of our study. Eulampsis holosericeus and Orthorhynchus cristatus occur on Barbuda (Pregill et al., 1994). Remains of this latter species have also been found on Antigua (Pregill et al., 1988, 1994).

3.6 - Gruiformes - Rallidae

47This family is well represented on all the islands except Les Saintes, with 19 individuals from 10 sites (NISP = 44) corresponding to 4 taxa, ranging from the Ceramic to Colonial periods (tab. 2, Appendix B).

48 Two small rail specimens from the Historical period on La Désirade (Pointe Gros Rempart 6) and on Marie Galante (Abri Cadet 3) were compared with the two tiny rails that have been recorded elsewhere in the West Indies (Olson, 1977; Pregill et al., 1988): the Yellow-breasted Crake Hapalocrex flaviventer (previously known as Porzana flaviventer or Poliolimnas flaviventer) and the Black Rail Laterallus jamaicensis.

49 The specimen from La Désirade is a distal tarsometatarsus (PGR6-2010-O-125) which, despite the smaller size (Appendix J) and more rounded lateral margin compared to the modern specimens, can be related to Hapalocrex flaviventer on the basis of 2 characters: a trochlea metatarsi III that projects slightly distally from the trochlea metatarsi IV in plantar view; and a trochlea metatarsi II that projects more plantarly than trochlea metatarsi IV in distal view (fig. 6). The specimen from Marie Galante is an incomplete coracoid (AC3-2019-O-1446), smaller than actual specimens of Laterallus jamaicensis (Appendix J) but which nevertheless shows the distinctive features of this taxon: the shape of the omal end and the corpus of bone (fig. 7). For both Hapalocrex flaviventer and Laterallus jamaicensis, the comparison was made between numerical bone model and fossil specimen, without it being possible to achieve a direct comparison between fossil and present-day specimen, for which reason and in spite of the distinctive characters observed, only a precautionary assignment cf. Hapalocrex flaviventer and Laterallus cf. jamaicensis is proposed.

50Bone remains of two large-sized rails, the Common Gallinule Gallinula galeata and the Purple Gallinule Porphyrio martinica, were documented from several sites and islands (tab. 2). Among the 9 remains of Gallinula galeata, one humerus from the Ceramic site of Îlet du Gosier is very small (Appendix J) but presents all the characters typical of this species. The other 3 incomplete bones from the Ceramic site of Tourlourous and from the Ceramic and Colonial layers of Pointe Gros Rempart 6 have been tentatively attributed to Gallinula galeata, as they differ in some morphological characters from modern specimens, but are similar in size.

51 The six bone remains of Porphyrio martinica were recovered from 2 sites on Basse Terre and La Désirade and date to the Ceramic and Colonial periods: Cathédrale de Basse-Terre and Pointe Gros Rempart 6. Three fragmentary bones from this latter site (scapula, humerus and tarsometatarsus) were attributed to the Purple Gallinule based on size (Appendix J) and the occurrence of the following osteological characters: large acromion of the scapula; processus supracondylaris dorsalis positioned away from the condylus dorsalis and straight dorsal margin of humerus; and similar shape of the trochlea metatarsi II on the tarsometatarsus (fig. 8).

52 Rallidae is the only family of the Gruiformes order in Guadeloupe. Amongst them, Gallinula galeata was very common in historical times (Du Tertre, 1667-1671) and remains so today (Levesque & Delcroix, 2019, 2021), while Porphyrio martinica became rarer since historical times (Danforth, 1939; Pinchon, 1976), to the point of being only sporadically present on La Désirade (Levesque & Delcroix, 2021) or very occasionally seen in the north of Basse Terre (e-bird, 2021). The two small rails are not known to occur in Guadeloupe (fig. 3): the Yellow-breasted Crake is a local but uncommon permanent resident in Cuba, Jamaica, and Puerto Rico, while the Black Rail is rare or vagrant in the West Indies (Raffaele et al., 2020).

53Rallids (Gallinula galeata and Porphyrio martinica) are well-represented in the Amerindian sites of the West Indies (Wing, 2001), but their rarefaction over the centuries is most likely due to their hunting by the first inhabitants of the islands and subsequent European colonists, as well as the loss or degradation of their habitat. An example in our study is provided by Porphyrio martinica, whose remains have been found in the southern part of Basse Terre where this bird is no longer found.

54About twenty fragmented specimens of an unidentified Rallidae were recorded from the Ceramic to Colonial period at Pointe Gros Rempart 6 and Roseau (fig. 9) and found from the Ceramic period layers of Folle Anse, Taliseronde and Tourlourous.

55The archeological remains tentatively assigned to Laterallus jamaicensis would be the first evidence of this species in the Lesser Antilles. No more than one species of Rallidae from each island was identified in the other Lesser Antilles islands: Hapalocrex flaviventer on Antigua (Steadman et al., 1984a; Pregill et al., 1988; 1994); Gallinula galeata on Marie Galante (Wing & Reitz, 1982), on Montserrat (Steadman et al., 1984b) and on Sint Eustatius (Van der Klif, 1992); Porphyrio martinica on Barbados (Wing, 1991), on Martinique (Allaire, 1977), on Antigua (Wing et al., 1968; Steadman et al., 1984a; Pregill et al., 1994; Healy et al., 2001, 2005) and on St Kitts (Hoffman, 1973). Other rails were identified on Barbados and Barbuda: Fulica podagrica (Brodkorb, 1964) and Rallus limicola (Pregill et al., 1994). Rallidae remains have been recorded on Saint Martin (Grouard, 2004).

56About twenty fragmented specimens of an unidentified Rallidae were recorded from the Ceramic to Colonial period at Pointe Gros Rempart 6 and Roseau (fig. 9) and found from the Ceramic period layers of Folle Anse, Taliseronde and Tourlourous.

3.7 - Charadriiformes - charadriidae

57Charadriidae is one of the families belonging to the Charadriiformes order in our sample, and is represented in the Historical period by 2 individuals from La Désirade (Pointe Gros Rempart 6) (tab. 2, Appendix B). A complete carpometacarpus differs in some morphological characters from modern specimens of Semipalmated Plover Charadrius semipalmatus, but is similar in size, while a proximal humerus is smaller than that of this species and similar in shape.

58Today, at least 7 species of Charadriidae are present in La Désirade and on the other islands of Guadeloupe, but only Charadrius semipalmatus is a very common migrant bird (Levesque & Delcroix, 2019, 2021). These bones are the first fossil occurrences of Charadriidae in the Lesser Antilles.

3.8 - Charadriiformes - scolopacidae

59Only a few Scolopacidae were recorded in our sample and they represent a second family of Charadriiformes (NISP = 11). They were recovered from the pre-human layer of Grotte Blanchard on Marie Galante and in layers dating from the Ceramic to Colonial periods of Pointe Gros Rempart 6 on La Désirade (tab. 2, Appendix B). Six specimens from Pointe Gros Rempart 6 agree in shape and size with the Semipalmated Sandpiper Calidris pusilla, while 2 specimens from Grotte Blanchard were tentatively assigned to this species. The other 3 remains of an unidentified Scolopacidae (medium and small size) have also been recorded from this site. Currently, among the Scolopacidae, Calidris pusilla is a very common wintering migrant bird in Guadeloupe Islands (Levesque & Delcroix, 2019, 2021).

60The material described here represents the first fossil occurrence of the Calidris genus on the Guadeloupe Islands. Calidris melanotos has been identified on Antigua (Steadman et al., 1984a; Pregill et al., 1988, 1994). Other Scolopacidae remains belonging to Burhinus bistriatus have been found on Barbuda (Pregill et al., 1994), while Limnodromus griseus has been mentioned on Tobago and on Martinique (Steadman & Jones, 2006; Fraser, 1981).

3.9 - Charadriiformes – laridae

61Laridae is the third family of the Charadiiformes order represented by 2 genera among fossils of Guadeloupe (Sternula and Sterna) discovered in 2 sites, one on Grande Terre (Anse à la Gourde) and the other on La Désirade (Pointe Gros Rempart 6), dating from the Ceramic to Colonial periods (tab. 2, Appendix B). The coracoid and humerus of a small Laridae from Pointe Gros Rempart 6 are similar in size, but do not agree in all characters with those of modern specimens of the Least Tern Sternula antillarum, so were only tentatively referred to this taxon. Three bones from Anse à la Gourde and one specimen from Pointe Gros Rempart 6 were referred to the genus Sterna. Four remains of an unidentified small- and large-sized Laridae have been recorded in the Ceramic layers of 2 sites on La Désirade (Pointe Gros Rempart 6 and Petite-Rivière).

62 Today, Sternula antillarum is common on La Désirade (Levesque & Delcroix, 2021) and nesting sites are present on the island (Gomes et al., 2014). Four species of the genus Sterna are uncommon or rare in the Guadeloupe Islands: Roseate Tern Sterna dougallii, Common Tern Sterna hirundo, Arctic Tern Sterna paradisaea and Forster’s Tern Sterna forsteri (Levesque & Delcroix, 2019). Except for Sterna forsteri, the other 3 species have also been observed at La Désirade (Levesque & Delcroix, 2021).

63 The chroniclers report very significant historical exploitation of this family, with the nesting colonies located in Saintes “decimated” by settlers who caught them with sticks. Father Breton (1978: 32) records that “around Guadeloupe there are many small, very pleasant islands. Les Saintes are the most beautiful, where even the savages still have some cotton gardens. The inhabitants of Guadeloupe go there to hunt and shoot turtles. They hunt tuërou, a kind of mauve, with a stick, killing them by the thousands, which they salt and dry in the form of a dried herring”. Moreover, Father Du Tertre provided much the same information for the Amerindians, who filled their pirogues with birds as they passed over the islets where the birds breed (1667‑1671 vol. 2: 274).

64The bones of Sternula and Sterna are the first fossil occurrences of these 2 genera in the Lesser Antilles. Leucophaeus atricilla is the only Laridae previously identified on Guadeloupe (Grouard, 2001a), Antigua (Wing et al., 1968; Pregill et al., 1994; Healy et al., 2005) and St Kitts (Wing & Scudder, 1983). Bone remains of Anous stolidus have been found on Nevis (Wing, 2001).

3.10 - Charadriiformes – family, genus and species indeterminate

65 It should be noted that no Charadriiformes remains have been found in Basse Terre or Les Saintes. Seventeen bones have been recorded on Marie Galante (in 2 layers from the pre-human to Ceramic periods of Grotte Blanchard and in the Ceramic layer of Abri Cadet 3), on Grande Terre (from Archaic to Colonial periods in the 3 sites of Anse à l’Eau, Pointe du Helleux and Grotte des Bambous) and on La Désirade (Pointe Gros Rempart 6). Due to the absence of diagnostic characters on these remains and the substantial variety of species belonging to this order, it was not possible to refer them to family level.

3.11 - Phaethontiformes - phaethontidae

66Nine remains have been referred to this family from the Ceramic layers of 2 sites, one on Grande Terre (Anse à l’Eau) the other on La Désirade (Petite-Rivière) (tab. 2, Appendix B). Six specimens from Anse à l’Eau can be attributed to the Red-billed Tropicbird Phaethon aethereus. The 3 incomplete bones from Petite-Rivière show some morphological similarities with the skull and ulna of this species but differ in their smaller size, which is closer to the White-tailed Tropicbird Phaethon lepturus. These bones were assigned to the genus Phaethon.

67 Today, this seabird family is represented on the Guadeloupe Islands by 2 species: Phaethon aethereus which is more common than Phaethon lepturus, also found on La Désirade (Levesque & Delcroix, 2019, 2021; Raffaele et al., 2020). Cory (1886) only reported Phaethon aethereus on this island.

68The 2 species - Red-billed Tropicbird and White-tailed Tropicbird - have been recorded elsewhere in the Lesser Antilles, on Antigua (Wing et al., 1968; Pregill et al., 1994). Furthermore, unidentified bones of a Phaethontidae have been reported from Sint Eustatius (Van der Klif, 1992; Wing, 2001).

3.12 - Procellariiformes - Procellariidae

69The Procellariidae is one of the best represented bird families in our sample, comprising 960 remains belonging to 92 individuals recovered from 19 sites spread across all the Guadeloupe Islands and dating from the pre-human to Colonial periods (tab. 2, Appendices B and K). The most abundant species is Audubon’s Shearwater Puffinus lherminieri (NISP = 625), found in 15 sites. On Marie Galante, the earliest evidence for this bird comes from the pre-human layers of Grotte Blanchard, Grotte Cadet 2 and Abri Cadet 3, and the taxon is present up to the Colonial period at Abri Cadet 3, Grotte Blanchard 2 and Grotte du Morne Rita. On the same island, shearwaters are also well represented in the Amerindian site of Tourlourou.

70 In order to understand the origin of the bone accumulations of Puffinus lherminieri in one of the sites which provided the highest number of remains of this bird, Abri Cadet 3, we conducted a preliminary taphonomic analysis of the bones. The results revealed a near total absence of digestion traces, the presence of mainly adult individuals, and a spatial association with artefacts from the archaic period which corresponds to the arrival of the first humans to the island. The abundance of bone remains in the archaic period (layer 4, NISP = 116) suggests that puffins were mainly accumulated by humans (Lenoble et al., 2019). This question deserves further analysis and the evidences of human processing of shearwater bones (cut marks, burning marks, fractures) are being currently investigated by the authors.

71 The caves on the island were home to shearwater colonies until the end of the 19^th century, at least discontinuously. On Marie Galante, at Grotte Blanchard 2, the nests of this species have also been documented up until at least the end of the 19^th century (Mallye et al., 2018). The high proportion of juvenile bones at Grotte Cadet 2, Blanchard and Morne Rita also suggests that Puffinus lherminieri nested near Grotte Blanchard 2, from the pre-human to Colonial periods. Three dates obtained on the ulnae of Puffinus lherminieri from Grotte du Morne Rita (1354-1862 CE (765 ± 30 BP, Lyon-15445 (SacA-54107); 1391-1892 CE (745 ± 30 BP, Lyon-15446 (SacA-54108); 1413-1905 CE (720 ± 30 BP, Lyon-15447 (SacA-54109)) and a tarsometatarsus from Grotte Blanchard 2 (1359-1875 CE (760 ± 30 BP, Lyon-9693 (SacA-31250)) provide evidence for the presence of this species on Marie Galante throughout the Historical period. In Grotte du Morne Rita, the Amerindian occupation of the caves did not preclude their later reoccupation by birds.

72 On the other islands, juvenile individuals of Puffinus lherminieri are represented by a few remains on Grande Terre, at the site of Grotte de l’Anse à l’Ecu. This small cave at the foot of the cliff is accessible and the bird bones are associated with those of extinct reptiles (Bochaton et al., 2021a) dating from Colonial period or earlier, as confirmed by a date obtained on a Puffinus lherminieri humerus (1455-1932 CE (680 ± 30 BP, Lyon-14288 (SacA-50617)). Other remains of this species have been identified in 2 sites of La Désirade that date from Ceramic to Colonial periods, and from the Ceramic period site of Les Saintes.

73 Two additional Procellaridae taxa are attested to on Basse Terre at the Ceramic to Contact period site of Roseau: Shearwater Calonectris sp. and Black-capped Petrel Pterodroma hasitata. The right mandibular ramus of the Shearwater is similar in the position of the foramina neurovascolaria to that of Cory’s Shearwater Calonectris diomedea, but is too fragmentary to be referred to a species.

74 Of the 2 remains belonging to Pterodroma hasitata, the ulna (ROS-2002-O-88; fig. 9) is comparable to modern specimens and an ulna that A. Wetmore identified as Pterodroma hasitata from Ceramic sites of the Grande Terre Atlantic coast (Clerc, 1965). The proximal end measurements are slightly larger than those of comparative specimens (Appendix L). However, 3 ulnae (ROS-2001-O-44, ROS-O-69 and ROS-2002-O-80) from the same site and an ulna from Îlet du Gosier (IG-2003-O-554) were cautiously attributed to this species, being similar in size (Appendix L) but with poorly preserved extremities. For the same reasons, a carpometacarpus and a coracoid from Roseau were referred as cf. Pterodroma hasitata.

75 A large number of remains of an unidentified Procellariidae (NISP = 159) were recovered from several Ceramic period sites of 5 islands, among which some sites not mentioned above on Basse Terre (La Pointe de Grande Anse and Embouchure de la rivière Baillif), on Grande Terre (Anse Sainte Marguerite) and on Marie Galante (Folle Anse and Taliseronde).

76 Today, the genus Calonectris is represented by 2 species on the Guadeloupe Islands, C. diomedea, a fairly common migrant bird in the Bahamas and Guadeloupe (Raffaele et al., 2020), and Cape Verde Shearwater C. edwardsii, an occasional visitor (Levesque & Delcroix, 2019). The Roseau specimen could possibly be attributable to the first species. Puffinus lherminieri is currently observed throughout almost all the islands of Guadeloupe, except for La Désirade where it is less common (Levesque & Delcroix, 2019, 2021), but its nesting in Guadeloupe remains uncertain, and could potentially be on the most inaccessible cliffs (Levesque & Mathurin, 2008). Our study shows that the past nesting of this bird was much more widespread than in modern times, as previously assumed by Mallye et al. (2018), including on terrestrial sites and in accessible caves. As confirmed by the dates from Grotte du Morne Rita, the presence of these birds is attested to until the Historical period at the latest.

77 The exact causes leading to the abandonment of the nesting site are not precisely known, but several factors are identified: the introduction of mammal species, such as the black rat (over the last five centuries) or mongoose (in the early 20^th century), or human exploitation following the arrival of the first European colonists in the region (in the early 17^th century) (Pinchon, 1976; Pregill et al., 1994; Lorvelec et al., 2007; Atkinson, 1985; Towns et al., 2006; Jones et al., 2008; Goedert et al., 2020). The high occurrence of this taxon in the middens of pre-Columbian villages suggests that Amerindians hunted this species. Although extremely rare in the archaeological sites of Guadeloupe and the Lesser Antilles, Pterodroma hasitata has lost its principal nesting zones, like Puffinus. This pelagic seabird is one of the most critically endangered species in the Caribbean (UICN), and previously bred in Guadeloupe until the middle of the 18^th or the early 19^th century (Lafresnaye, 1844; Feilden, 1889; Goetz et al., 2012; Levesque & Yésou, 2018) (fig. 3). The main cause of its disappearance is human overhunting from the beginning of colonization (Du Tertre, 1667-1671; Froger, 1698; Labat, 1722) (tab. 3). This practice lasted until the beginning of the 19^th century (e.g., Léonard, 1797; Longin, 1848). Pterodroma hasitata became rare in Guadeloupe towards the end of the first half of the 19^th century, according to the account of Collardeau, the mayor of St Claude (Lawrence, 1891). Ober himself was unable to collect any specimens (Lawrence, 1879). Noble indicates that he learned from Father Lowinsky that Pterodroma hasitata were no longer heard on the island following the earthquake of 1843 (Noble, 1916), although the last to be seen in Guadeloupe was captured in St Claude in the second half of the 19^th century (Lawrence, 1891). It should be noted that the petrel was so appreciated by the French that it led to trade in the species from Dominica (Atwood, 1791). The collapse of petrel populations in the first part of the 19^th century followed two centuries of overhunting by colonists.

78The specimen from Roseau is the first mention of a fossil Calonectris in the Lesser Antilles. In addition to the ulna identified by Wetmore from the Ceramic sites of the eastern coast of Grande Terre, remains belonging to Pterodroma hasitata were listed by Grouard at Embouchure de la rivière Baillif (2001a), and by Wetmore for Martinique (1952), where it was supposed to have been extirpated in the pre-Columbian period according to Pinchon (1976). A few bones have been found on St. Kitts and on Antigua (Wing, 1973; Wing, et al., 1968).

79 Several authors have reported remains of other procellarids (Puffinus lherminieri and Puffinus sp.) in archaeological and paleontological sites on Marie Galante (Honoré, 2014; Stouvenot et al., 2014). Generally, remains of Puffinus lherminieri are common in archaeological sites of the Lesser Antilles, and it probably provided a seasonal food resource for Amerindian groups (Wing, 2001) on Grenada (Frandrich, 1990), Martinique (Grouard, 2013), Barbuda (Watters et al., 1984), Antigua (Wing et al., 1968; Steadman et al., 1984a), Saba (Hofman & Hoogland, 2003), Anguilla (Watters, 1991), and Saint Martin (Grouard, 2004, 2005).

80 The fossil records of paleontological sites on most of these islands also indicate an extensive distribution of Puffinus lherminieri during the pre-human period: on Barbuda (Pregill et al., 1994), Antigua (Pregill et al., 1988, 1994), and Anguilla (Cope, 1883; Pregill et al., 1994). Puffinus lherminieri also occurred on Barbados, as demonstrated by fossil remains identified in natural sites on this island (Brodkorb, 1964). Bone remains of Puffinus sp. have been found on Antigua (Pregill et al., 1994; Healy et al., 2005) and Saba (Wing, 2001).

3.13 - Suliformes - Sulidae

81The Sulidae are represented by 4 individuals (NISP = 4) of the genus Sula from 4 sites on Basse Terre (Sainte Rose), La Désirade (Petite-Rivière) and Marie Galante (Tourlourous and Folle Anse), all dating from the Ceramic period (tab. 2, Appendix B). The 2 coracoids from Sainte Rose (Dindé) and Petite-Rivière are similar in size to the Red-footed Booby Sula sula, but differ by a slenderer facies articularis humeralis in the Sainte Rose specimen, and by a more rounded labrum internum coracocoidei in the Petite-Rivière specimen. Comparisons with modern coracoids of the Brown Booby Sula leucogaster reveal no specific similarities. To confirm the species determination, a direct comparison with larger coracoids of Masked Booby Sula dactylatra is needed. Identifying the other two bones (a maxilla and scapula) from Tourlourous and Folle Anse in Marie Galante was more difficult due to their fragmentary nature and they were therefore referred simply to the genus Sula.

82 The Red-footed Booby Sula sula is currently uncommon in the Guadeloupe Islands and is found occasionally on La Désirade (Levesque & Delcroix, 2019, 2021). The other 2 species of the same genus are present today: one is common (Brown Booby Sula leucogaster), the other rare (Masked Booby Sula dactylatra).

83 Allan de Cramant hunted boobies and pelicans in the salt ponds of Petite-Terre (La Désirade) during the first half of the 18^th century (Allan de Cramant, 2010: 92), although hunting of these birds probably dates back to the Ceramic period. Boobies, shearwaters, pigeons, purple and common gallinules are the most common birds found in archaeological sites of the West Indies. The boobies, like shearwaters, are thought to have been captured at the nest, being oceanic birds that approach the coast only for nesting, although no medullary and immature bones have been found providing evidence of nesting (Wing, 2001). The few bone remains of boobies from the Guadeloupe Islands do not support this hypothesis, as all the individuals are adults and none bears medullary bone.

84Several specimens of the genus Sula (Sula sp.) have been found on Antigua (Wing et al., 1968; Pregill et al., 1994), Nevis (Wing, 2001), Sint Eustatius (Van der Klif, 1992), St Kitts (Wing & Scudder, 1983) and Saba (Hofman & Hoogland, 1991; Wing, 2001).

3.14 - Pelecaniformes - Pelecanidae

85The Pelecaniformes order is represented in our sample by 2 families: Pelecanidae and Ardeidae. Only one left phalanx 2 digit III from the Ceramic and Contact period deposits of Roseau (ROS-O-45) was assigned to Pelecanidae (tab. 2, Appendix B). The Roseau specimen has a similar tuberculum extensorium (Livezey and Zusi, 2006) to that of the Brown Pelican Pelecanus occidentalis, but the trochleae articulares are slenderer.

86 Pelecanus occidentalis is the only species of this family reported from the modern period in the Guadeloupe Islands (Levesque & Delcroix, 2019). This common migrant bird inhabits calm coastal waters (Raffaele et al., 2020).

87A small number of fossil remains of Pelecanus occidentalis and Pelecanus sp. have been found on Grenada (Frandrich, 1990) and Antigua (Murphy & Healy, 1997), respectively.

3.15 - Pelecaniformes - Ardeidae

88Except for Grande Terre, bone remains of Herons and Egrets (NISP = 62) have been identified from 7 sites on 4 islands of Guadeloupe and date from the pre-human to Colonial periods (tab. 2, Appendix B). The 30 remains assigned to the Yellow-crowned Night-Heron Nyctanassa violacea come from the levels spanning the pre-human to Colonial periods of sites on La Désirade and Marie Galante (Pointe Gros Rempart 6, Taliseronde, Grotte Blanchard, Grotte Blanchard 2). A pedal phalanx from layer 3 of Pointe Gros Rempart 6 attests to the presence of this species at La Désirade during the Ceramic period (1301-1407 CE; 600 ± 30 BP, Lyon-14478 (SacA-51014)).

89 The remains of the Egret Egretta sp. are less abundant in the early Ceramic period of Taliseronde, as are the Great Blue Heron Ardea herodias in the late Ceramic and Contact deposits of Roseau. The Taliseronde specimen (MEC-D-16.13.57.5) has similar characters to the modern femurs of the Little Egret Egretta garzetta and Snowy Egret Egretta thula (tuberculum musculi gostrocnemialis lateralis, impressiones ansae musculi iliofibularis, trochlea fibularis, crista iliofibularis, condyli lateralis and medialis). In fact, the femur is similar in size to that of Nyctanassa violacea, which differs in the shape of distal part of the bone. In order to attribute this specimen to species, a comparison with a larger sample of species of the genus Egretta present in the West Indies is necessary: the Little Blue Heron Egretta caerulea, Tricolored Heron Egretta tricolor and Reddish Egret Egretta rufescens (Raffaele et al., 2020).

90 The phalanx 1 digit I from layer 1 of Roseau (ROS-O-96) was assigned to the Great Blue Heron Ardea herodias on the basis of its shape and size which differ from those of Pelecanidae from the same layer. The remaining 30 specimens have not been assigned to either genus or species but nevertheless indicate the presence of Ardeidae (large size) in the Ceramic and Colonial periods at Cathédrale de Basse-Terre and Grande-Anse de Terre de Bas. This material mainly comprises phalanges and difficult-to-attribute vertebrae, as well as long bones that do not present species-specific diagnostic features.

91 At present, Nyctanassa violacea is a resident breeding species in the Guadeloupe Islands, while the migrant Ardea herodias is an uncommon winter visitor (Levesque & Delcroix, 2019, 2021). Egretta thula is the most common species (resident breeding) among the Egretta genus. The other 3 migrant species are uncommon (E. garzetta, E. caerulea and E. tricolor) and Egretta rufescens is very rare in the Lesser Antilles (Raffaele et al., 2020).

92 Among the ornithological studies of the 19^th and 20^th centuries, the list of the Guadeloupean birds produced by L’Herminier reports 6 species of Ardea (Lawrence, 1879). In addition to the mention of A. herodias and N. violacea (=A. cayennensis), 2 egrets are listed: Egretta caerulea (=A. caerulea) and E. garzetta (=A. garzetta).

93The remains of Nyctanassa violacea were found on Marie Galante (Mallye et al., 2018), La Désirade (Boudadi-Maligne et al., 2015) and on other Lesser Antilles islands: Tobago (Steadman & Jones, 2006) and Antigua (Wing et al., 1968; Pregill et al., 1994). The largest number of genera and species attributed to this family (including Nyctanassa violacea) have been reported for Antigua: Butorides striata, Egretta sp., Nycticorax cf. nycticorax, Nycticorax sp. (Wing et al., 1968; Pregill et al., 1994). Moreover, specimens of Nycticorax nycticorax were found on Sint Eustatius (Van der Klif, 1992) and St Kitts (Wing & Scudder, 1983), 2 species of Ardea on Saint Martin, A. herodias (Grouard, 2004) and A. alba on Tobago (Steadman & Jones, 2006).

3.16 - Accipitriformes - Pandionidae

94In our analysis, only one remain (distal humerus) of Osprey Pandion haliaetus from the Ceramic period site of Tourlourous in Marie Galante was identified (tab. 2, Appendix B). This wintering migrant bird occurs throughout the West Indies but is uncommon in the Lesser Antilles (Levesque & Delcroix, 2019; Raffaele et al., 2020). Pandion haliaetus is on the USNM list of several bird skins from Guadeloupe, presumably collected before 1885 by Guesde (Danforth, 1939). Very few remains have been found in the Lesser Antilles islands: Grenada (Frandrich, 1990) and Sint Eustatius (Van der Klif, 1992).

3.17 - Accipitriformes - Accipitridae

95Like the Pandionidae, bone remains attributed to the Accipitridae family are rare in our sample (NISP = 3) and restricted to the Ceramic period on 2 islands: Grande Terre and La Désirade (tab. 2, Appendix B). A reliable morphological comparison (incisura tibialis, pons suprotendineus, condylus medialis and incisura intercondylaris) can be made between the right tibiotarsus from the site of Anse à l’Eau (MEC-I-16.3.802) and a large Accipitridae, Swainson’s Hawk Buteo swainsoni, although it is slightly larger (Appendix M). Among the congeneric species, the tibiotarsus is comparatively larger and smaller, respectively, than modern specimens of Red-tailed Hawk Buteo jamaicensis and Broad-winged Hawk Buteo platypterus. Several morphological differences with these specimens were also observed. Two specimens from Pointe Gros Rempart 6 (coracoid and claw) have little diagnostic value beyond the family level.

96 The Anse à l’Eau specimen represents the first occurrence of Buteo swainsoni in the Lesser Antilles (fig. 3). This bird currently frequents coastal habitats and is an irregular and generally uncommon and local migrant in Cuba (Raffaele et al., 2020). Two others Buteo species occur only occasionally in the Guadeloupe Islands: Buteo jamaicensis and Buteo platypterus (Levesque & Delcroix, 2019).

97An unidentified Accipitridae was found on Marie Galante (Stouvenot et al., 2014). Fossil bones of Buteo swainsoni are very rare in the Caribbean and have only been mentioned on Abaco (Steadman & Franklin, 2015). Conversely, the remains of 2 Buteo species have been identified in the Lesser Antilles: Buteo jamaicensis on Barbados (Brodkorb, 1964); Buteo platypterus on Martinique (Allaire, 1977) and on Antigua (Pregill et al., 1994). Additional Accipitridae bones attributed to Buteogallus urubitinga were found on Tobago (Steadman & Jones, 2006).

3.18 - Strigiformes - Tytonidae

98The only Tytonidae remains identified in the course of our study were a distal penultimate phalanx (ROS-O-61) from the Ceramic to Contact period deposits of Roseau (tab. 2, Appendix B). The morphometric analysis of this fragment is consistent with a pedal phalanx from an extinct giant Barn Owl Tyto sp. (Gala et al., 2022).

99 Currently, no tytonid owls inhabit the Guadeloupe Islands. A small endemic tytonid occurs on the neighboring island, the Lesser Antillean barn owl Tyto insularis, whose distribution is restricted to the southern Lesser Antilles islands (Dominica, St. Vincent, Grenadines islands and Grenada) (Ridgway 1914; Wetmore & Swales 1931; Weick 2006). Four extinct species of giant tytonid owls (genus Tyto) inhabited the Antilles and Bahamian Archipelago during different periods (Wetmore, 1922, 1937; Arredondo 1972a, 1972b; Suárez and Olson, 2015, 2020). Until now, the only fossil evidence of a giant Barn Owl in the Lesser Antilles has been described in Barbuda during the late Quaternary as Tyto neddi (Steadman & Hilgartner, 1999), subsequently reattributed to Tyto noeli by Suárez and Olson (2015). This species, which is the smallest extinct giant Barn Owl, is represented by 6 remains, including several pedal phalanges that are not homologous with the Roseau specimen. Therefore, even if it is impossible to assign the Roseau specimen to Tyto noeli, this discovery nevertheless suggests that a large tytonid inhabited the Lesser Antilles up to the European colonization of the Caribbean (fig. 3). This giant owl became extinct along with, and potentially as a consequence of, the disappearance of endemic species of the medium-sized rodents on which they preyed (Gala et al., 2022), a hypothesis that is consistent with the regional context (Olson, 1978; Steadman & Hilgartner, 1999; Suárez & Olson, 2015, 2020).

100The Roseau specimen is the first mention of a giant barn owl in Guadeloupe and the second recorded in the Lesser Antilles, alongside the fossil specimens from Barbuda.

3.19 - Strigiformes - Strigidae

101The Burrowing Owl Athene cunicularia is a small Strigiformes that inhabited the Guadeloupe Islands (Grande Terre, Marie Galante and probably La Désirade) from the pre-human to Colonial periods (Gala et al., 2020) (fig. 3). About one hundred remains (NISP = 101) from 7 sites, representing 21 individuals, could be attributed to this small owl (Grotte des Bambous, Pointe Gros Rempart 6, Grotte Blanchard, Grotte Blanchard 2, Grotte Cadet 2, Abri Cadet 3, Folle Anse; tab. 2, Appendix B). Among these owl remains, 30 were only tentatively assigned to the Burrowing Owl (cf. A. cunicularia) due to the fact that different morphometric characters or their degree of fragmentation preclude reliable attribution to this species, as in the case of 2 specimens from Pointe Gros Rempart 6 and Folle Anse (tarsometatarsus and scapula), which are the only strigid remains from these sites.

102 Several Pleistocene and early Holocene fossils represent the oldest evidence of A. cunicularia in the Guadeloupe archipelago, before the first human settlements. They come from 3 neighboring sites on Marie Galante (fig. 2 and see tab. 2 in Gala et al., 2020). Evidence of the mid-Holocene (Archaic period) distribution of this species in Marie Galante comes from Grotte Blanchard and Grotte Cadet 2 (from Archaic to Colonial periods). Several remains confidently or tentatively referred to this species were found from the Ceramic to Colonial period layers of 3 sites on this island (Grotte Blanchard 2, Abri Cadet 3, Folle Anse).

103 Further data on the distribution and chronology of the Burrowing Owl on the other islands, Grande Terre and La Désirade, is provided by over 20 owl remains from the Archaic to Colonial periods. A direct radiocarbon date obtained on an ulna from Grotte des Bambous provides evidence for the presence of this owl on Grande Terre throughout the Contact period and possibly during the first decades of European colonization (1445–1631 CE (385 ± 30 BP, Lyon-14109, SacA49751)).

104 The result of this study shows that the Burrowing Owl was present in Guadeloupe on Grande Terre and La Désirade until the Colonial period. For this latter island, this is a first mention of the species. More widely, the review of past occurrences of the Burrowing Owl in the Lesser Antilles indicates anthropic impact rather than climate change as the driving force behind the renewal of Caribbean avian communities since the Pleistocene (Gala et al., 2020).

105 During the Historical period, 2 now-extinct forms of A. cunicularia were present in the Lesser Antilles: a small form A. c. amaura (Lawrence, 1878) on the islands of Nevis and Antigua, and a larger form A. c. guadeloupensis (Ridgway, 1874) on Marie Galante. Eight specimens of this latter form were known to occur on the Guadeloupe Islands until at least the end of the 19^th century (Noble, 1916; Danforth, 1939; Deignan, 1961).

106 Native raptor populations in Guadeloupe and the Caribbean have declined significantly since European colonization. Seventeenth-century historical sources suggest that their distribution was wider than suggested by 19^th century collections. In fact, both Breton (1665) and Du Tertre (1667-1671) attested to the presence of a small-sized Strigidae on the main islands of Guadeloupe during the first decades of French colonization, a bird in which an Athene cunicularia can be recognized unambiguously according to the details given by Du Tertre (Gala et al., 2020). The causes of the disappearance of this species are a matter of debate, and our results challenge the previous hypothesis of a relict distribution of the Burrowing Owl as a consequence of climatically-induced modifications of the Caribbean environment since the last glaciation (Pregill & Olson, 1981). The disappearance of the owl involved a species rarefaction stage. Pregill and Olson’s hypothesis predicts habitat fragmentation and the abundance of remains from paleontological sites to be coincident with the Pleistocene-Holocene transition, whereas our data shows that this fragmentation and rarefaction is much more recent, coincident with the human occupation of the islands. This rarefaction came early, before the middle of the 19^th century, as shown by Ober’s failure to obtain this bird during his visit to Guadeloupe in 1877. This is proof of the extreme rarity if not the disappearance of this bird from the islands of Guadeloupe at this time. Several ornithologists (Noble, Bond, Danforth) have proposed that the disappearance of this Caribbean owl was linked to the introduction of the mongoose. However, the early character of this rarefaction is better explained by a modification of the environment, especially as the introduction of the mongoose only occurred at the very end of the 19^th century (Lorvellec et al., 2021). Habitat degradation linked to increasing human settlement of the island during historical periods therefore appears the most parsimonious explanation for the regional extinction of this species.

107Our results confirm the presence of Athene cunicularia in the fossil record of Marie Galante, which was previously suspected based on 2 remains from the Archaic layers of Abri Cadet 3 (Sierpe, 2011; Stouvenot et al., 2014) and from the Ceramic Age site of Tourlourous (Colas et al., 2002; Grouard, 2016). The study of archaeological bones also highlights the presence of this species in St Martin during the Ceramic period (Gala et al., 2020). Fossils have been found on several islands, from the Pleistocene in Barbuda (Pregill et al., 1994) to the Ceramic period on Sint Eustatius (Van der Klif, 1992) and Antigua (Wing et al., 1968; Steadman et al., 1984a; Pregill et al., 1988; Healy et al., 2001).

3.20 - Piciformes - Picidae

108The 8 remains belonging to the scapular girdle, wing and leg of a single individual of Guadeloupe Woodpecker Melanerpes herminieri from the Archaic to Colonial periods on Grande Terre (Grotte des Bambous) are the only evidence of this family in the material that was studied (tab. 2, Appendix B).

109 The distribution of this endemic woodpecker of Guadeloupe during the modern period is restricted to six different habitat types, ranging from the seasonal evergreen second-growth forest and rainforest of Basse Terre to the mangrove, swamp and semi-deciduous forests of Grande Terre (Villard et al., 2010).

110 Maynard (1898) limited the occurrence of Melanerpes herminieri to Basse Terre, while other past reports have also described this species as being common on the western part of Basse Terre and rare on Grande Terre (Noble, 1916; Danforth, 1939; Pinchon, 1976; Short 1982). However, recent ornithological lists reflect the current expansion of this species on Grande Terre, where it occupies the central and western part of the island (Levesque et al., 2020; Villard et al., 2010). On the other hand, it remains absent from the eastern part of Grande Terre where the fossil specimen comes from.

111 The presence of this species at Grotte des Bambous therefore testifies to a wider distribution in the past, extending to the eastern coast of Grande Terre. The most important known factor affecting woodpecker populations, and in particular the Guadeloupe Woodpecker, is deforestation and habitat fragmentation (Villard & Rousteau, 1998).

112These specimens constitute the first fossil record of Melanerpes herminieri for the Lesser Antilles. Several remains from the Greater Antilles and Bahamas were previously assigned to the genus Melanerpes and 3 species: M. portoricensis in Puerto Rico (Wetmore, 1922), M. striatus in Hispaniola (Steadman & Takano, 2013) and M. superciliaris in the Bahamas (Wetmore, 1937; Brodkorb, 1959; Olson & Hilgartner, 1982).

3.21 - Falconiformes - Falconidae

113A limited number of remains of diurnal raptors belonging to the order Falconiformes (NISP = 5 and 3 individuals) were identified in 2 sites from the pre-human to Colonial periods on Marie Galante and Grande Terre (tab. 2, Appendix B). Falconidae remains from Marie Galante were recovered from the pre-human layers 5 and 2 of Grotte Blanchard. Four specimens have been attributed to the American Kestrel Falco sparverius. One additional bone (ulna) found in a more recent deposit (probably from the late Ceramic to Colonial period) at the Grotte de l’Anse à l’Ecu in Grande Terre has been tentatively referred to this species.

114 Currently, Falco sparverius is common in the Guadeloupe Islands (Levesque & Delcroix, 2019) where it inhabits dry, open lowlands, as well as the periphery of mountain forests (Raffaele et al., 2020). Its presence has been documented since the 19^th century. L’Herminier, Ober and Wynch collected this species from both Grande Terre and Basse Terre (Lawrence, 1879; Cory, 1891) and Richardson from Marie Galante and La Désirade (Cory, 1886).

115Falconidae remains have previously been identified at a small number of sites: Falco sp. on Basse Terre (Tomadini et al., 2014); Falco sparverius on Barbuda (Pregill et al., 1994) and on Anguilla (Pregill et al., 1994).

3.22 - Psittaciformes - Psittacidae

116Psittacidae remains are rare in the studied sample (n = 3) and belong to 2 genera from 3 sites dating from the pre-human to Contact period (tab. 2, Appendix B). Two remains from sites on Basse Terre dating to the Ceramic and Contact periods represent one or two species of Amazona Parrot (Amazona sp.). The distal humerus from layer 1 of Roseau (Bochaton et al., 2021b) (fig. 9) was compared to 2 large Lesser Antillean parrots (St Lucia Parrot Amazona versicolor from the island of the same name and the Red-Necked Parrot A. arausiaca from Dominica) and 3 smaller parrots from the Greater Antilles (Cuban Parrot Amazona leucocephala and Black-billed Parrot Amazona agilis) and Central America (Red-crowned Parrot Amazona viridigenalis). The Roseau specimen falls within the size range of Lesser Antillean parrots but is smaller than the Amazona versicolor and larger than the humerus of A. arausiaca (Appendix N). However, the specimen was not compared with the large St Vincent Parrot A. guildingii and the Imperial Parrot A. imperialis. The humerus of A. guildingii should be slightly larger than Roseau specimen, and too large to be compatible with A. imperialis.

117 An incomplete tarsometatarsus from US 3005 of Cathédrale de Basse-Terre is too fragmentary for species-level identification. The smallest breadth of the corpus (Sc) is larger than that of A. arausiaca and places this specimen within the Lesser Antillean parrots (Appendix N). Based on its size, this bone may be associated with the above-mentioned humerus and could therefore represent a single species of Amazona that lived or was kept in captivity on Basse Terre during the pre-Columbian and Contact periods.

118 In the course of the initial study of bird bones from Grotte Blanchard, we cautiously assigned a talon of a large macaw from the pre-human period (layer 5) to the extinct Lesser Antilles Macaw (Ara guadeloupensis) described by Clark in 1905, based on its size and geographic arguments (Gala & Lenoble, 2015) (fig. 3). In addition to the large macaw, three Psittacidae are listed as inhabiting the Guadeloupe Islands in historical times, with all becoming extinct by the 18^th or 19^th centuries. Fossil and/or historic accounts also indicate that a Guadeloupe Parakeet Psittacara labati (Labat, 1742) and a Guadeloupe Parrot Amazona violacea (Du Tertre, 1654) occurred on these islands.

119 Another macaw was described by Rothschild (1905, 1907), the Violet Macaw Anodorhynchus purpurascens. A recent study (Lenoble, 2015) showed that this species never existed, however, and that the description actually corresponds to Amazona violacea. The persistence of the Guadeloupe Parrot has been reliably established for the 18^th century (Labat, 1722, 1724, 1742; Brisson, 1760) and suggested by L’Herminier’s list in the first part of the 19^th century (Lawrence, 1879). Two bones from the Ceramic site of Folle Anse on Marie Galante (not examined in this study) were assigned to A. violacea, with the assumption that this species and A. imperialis referred to the same parrot (Olson & López, 2008). The first bone is a tibiotarsus comparable to the endemic A. imperialis from Dominica (Williams & Steadman, 2001). The second one is an ulna, referred first to an Ara by Steadman and Willams (2001) and re-examined by Olson and López (2008), who referred it to A. imperialis, based on its size and robustness.

120 Historical sources indicate that various other species of Psittacidae may have inhabited the Caribbean at the time of the discovery of the Americas; however, their validity is subject to debate (Du Tertre, 1667-1671; Ridgway, 1915; Greenway, 1967; Snyder et al., 1987; Butler, 1992; Williams & Steadman, 2001; Olson & López, 2008; Wiley & Kirwan, 2013).

121 While several historical sources report Caribbean groups having kept parrots in their villages (Anonyme, 1660; Monpilliers, 1654; Las Casas, 1875-1876), there is no mention of whether these birds were native or exchanged between villages and islands. The abundance of Psittacidae at the time of Columbus, as reported by several documents, declined over the next 3 centuries (Wiley & Kirwan, 2013). The causes underlying the disappearance of these birds were mostly anthropogenic (e.g., hunting, trade, habitat loss), a hypothesis that is difficult to test with the studied material. Furthermore, Psittacidae were an appreciated game species (Du Tertre, 1654, 1667-1671; Labat, 1742; Chanvalon, 1763) and hunted because they damaged crops.

122In addition to the remains of two extinct species on Marie Galante and the Amazona bones from Basse Terre, Grouard (2001a) assigned a bone from the Ceramic period site of Petite Rivière on La Désirade to the Amazona genus. Based on its size (smaller than that of an Orange-winged Parrot Amazona amazonica) the specimen could not be identified as a large Amazona and would seem to belong to a smaller species of Greater Antillean parrot. Unfortunately, the bone is now lost (S. Grouard, personal communication) and thus not revised in the course of this study.

123 In the Lesser Antilles, fossil bones of Ara sp. were found in Montserrat (Williams & Steadman, 2001) and of Aratinga sp. in Barbuda (Pregill et al., 1994). Bones of Amazona come from sites on 4 islands: Puerto Rican Parrot Amazona vittata on Barbuda (Williams & Steadman, 2001) and Antigua (Wing et al., 1968; Steadman et al., 1984a; Pregill et al., 1994), the Orange-winged Amazon Amazona amazonica on Tobago (Steadman & Jones, 2006) and possibly the St. Lucia Parrot Amazona versicolor on St. Lucia (Williams & Steadman, 2001).

3.23 - Passeriformes - Tyrannidae

124Six specimens from 2 sites dating from the pre-human to Colonial periods on 2 islands have been identified as Tyrannidae, based on their characters and size (tab. 2, Appendix B). Among the 5 specimens from the pre-human layers (3, 5, 6 and 8) of Grotte Blanchard on Marie Galante tentatively assigned to the genus Elaenia, only one is complete (wing phalanx). The sternum from the Archaic to Colonial periods of Grande Terre (Grotte des Bambous) is too small for Elaenia, but can be assigned to Tyrannidae based on the shape of the rostrum sterni. Two small Tyrannidae species are currently present in Guadeloupe, the Lesser Antillean Flycatcher Myiarchus oberi and the Lesser Antillean Pewee Contopus latirostris. As these birds are yet to be included in the osteological collections consulted in the course of this study, the specimen is impossible to assign to a species.

125 Currently, the genus Elaenia is represented in the Guadeloupe Islands by the Caribbean Elaenia E. martinica, a very common bird (Levesque & Delcroix, 2019) that inhabits woodlands, scrub, and forests (Raffaele et al., 2020). Furthermore, larger Tyrannidae such as the Gray Kingbird Tyrannus dominicensis are very common in the archipelago, while the Lesser Antillean Flycatcher Myiarchus oberi and Fork-tailed Flycatcher Tyrannus savanna are uncommon or rare, respectively. Additional similar-sized or smaller Tyrannidae than Elaenia martinica are uncommon or occasional: the Lesser Antillean Pewee Contopus latirostris, Eastern Wood-Pewee Contopus virens and Willow/Alder Flycatcher Empidonax traillii/alnorum (Levesque & Delcroix, 2019).

126 Together with the Grey Kingbird Tyrannus dominicensis, Elaenea martinica figures in the list of Cory (1891), while only Tyrannus dominicensis and Tyrannus sp. are recorded in the list of Lawrence (1879).

127If the identification of Grotte Blanchard specimens is confirmed, it will be the first mention of Elaenea martinica in the fossil record of the Lesser Antilles. Three different fossils of Tyrannidae have been identified on 3 islands in the Lesser Antilles: Cnemotriccus fuscatus on Tobago (Steadman & Jones, 2006); Myiarchus cf. oberi on Barbuda (Pregill et al., 1994); and Tyrannus dominicensis on Antigua (Steadman et al., 1984a; Pregill et al., 1994).

3.24 - Passeriformes - Mimidae

128Mimidae is the best represented Passeriformes family in our sample, comprising 84 remains belonging to 31 individuals. The material was recovered from 10 sites spread across all the Guadeloupe Islands, except Les Saintes, and dates from the pre-human to Colonial periods (tab. 2, Appendix B).

129 This family is one of the best-known examples of avian radiation in the West Indies (Hunt et al., 2001). The substantial inter-specific and inter-generic variability of this family and the difficulty in identifying remains to species have driven a recent study focusing on the carpometacarpus morphological variation of West Indies Mimidae species (Jeantet et al., 2021). This skeletal element was explored using a combination of geometric morphometrics and conventional osteology. A modern reference collection of carpometacarpi was built to assess 12 Mimidae remains from paleontological and archaeological sites. Morphometric data collected from over 50 modern carpometacarpi belonging to 7 genera and 9 species of Caribbean mimid birds revealed morphological differences of the entire or distal portion of the carpometacarpus separating the Antillean endemic from pan-continental clades. Inter-genera and inter-specific differences were documented for both clades, especially within the genus Mimus. Several characters successfully distinguish the different Mimidae species. Combining morphological characters with size data also produced more reliable species determinations. Scaly-breasted Thrasher Allenia fusca and Pearly-eyed Thrasher Margarops fuscatus are distinguished by their large size while Grey Catbird Dumetella carolinensis is characterized by its small size, with the other species being intermediate in size.

130 The 12 fossil carpometacarpi allowed us to confirm the presence of Allenia fusca at Grotte Cadet 2 and Brown Trembler Cinclocerthia ruficauda at Grotte Blanchard, also based on other anatomical elements. In addition, this study based on carpometacarpi increases the variety of mimids present in the pre-human and Archaic deposits from 2 sites of Marie Galante that are not currently present in the Guadeloupe avifauna: Dumetella carolinensis at Abri Cadet 3 and 2 large-sized specimens at Grotte Blanchard (similar in size to Allenia but with morphological characters of Margarops) (fig. 3) (Jeantet et al., 2021). The remains attributed to the latter two taxa are rare (1 for Dumetella and 2 for the large Mimidae species), while bone remains of Cinclocerthia ruficauda (NISP = 27) and Margarops fuscatus (NISP = 20) are more abundant, being the best represented Mimidae taxa in our sample, followed by Allenia fusca (NISP = 10), Bahama Mockingbird Mimus gundlachii (NISP = 1) and Mimus sp. (NISP = 3) (tab. 2). Several remains can be attributed to Cinclocerthia ruficauda (fig. 10, Appendix O) and 2 specimens from Abri Cadet 3 that did not display all diagnostic characters were tentatively referred to this species.

131The genus Mimus is represented by 4 remains from Grotte des Bambous, but only a tibiotarsus presents diagnostic characters that allow it to be identified to species (Mimus gundlachii) compared to all other congeneric specimens: a very high crest attachment (fig. 11; Appendix P). The other specimens (2 coracoids and a femur) do not exhibit any distinguishing features that might support a determination to species level (Mimus sp.). Unidentified Mimidae remains (NISP = 18) were found in several of the above-mentioned sites (from pre-human to Colonial period) and the early Ceramic levels of Folle Anse.

132The fossil record documents 5 species, 3 of which currently inhabit Guadeloupe and are endemic to the Lesser Antilles: Allenia fusca, Margarops fuscatus and Cinclocerthia ruficauda. The other two species, Dumetella carolinensis and Mimus gundlachii, are absent from the Guadeloupe Islands (Levesque & Delcroix, 2019; Raffaele et al., 2020) (fig. 3). Allenia fusca and Margarops fuscatus inhabit the forests and woodlands of the Lesser Antilles and the swamp forests of Grande Terre, whereas Cinclocerthia ruficauda is a rainforest bird that lives at higher elevations on Basse Terre (Terborgh et al., 1978; Cody, 2005; Raffaele et al., 2020). This latter species is not present any longer on the dry islands of La Désirade and Marie Galante, nor in the eastern part of Grande Terre (Levesque et al., 2020; Levesque & Delcroix, 2021; e-bird, 2021) (fig. 3). Dumetella carolinensis is a common migrant and non-breeding resident in the Bahamas, Cuba, Cayman Islands, San Andrés, and Providencia (Raffaele et al., 2020). Mimus gundlachii is endemic to Bahamas and the Greater Antilles and inhabits semi-arid scrub and woodlands (Raffaele et al., 2020).

133Concerning Cinclocerthia ruficauda, the difficulty lies in establishing the precise origin of specimens recorded in Guadeloupe (Grande Terre and Basse Terre) during 19^th century ornithological studies. In fact, L’Herminier, Ober and Wynch collected this species from Grande Terre and/or from Basse Terre (Lawrence, 1879; Cory, 1891). However, we know that Cinclocerthia ruficauda was collected by Richardson from Grande Terre (Cory, 1886). The rarefaction of this bird on Grande Terre was already evident by the beginning of 20^th century or at the end of the Historical period (Noble, 1916; Danforth, 1939) following the contraction of forested areas.

134 Even in the cases of Dumetella carolinensis and Mimus gundlachii, although documented by very few remains, our data evidences specimens in the fossil record up to the Historical period from islands (Marie Galante, Désirade) or part of an island (eastern Grande Terre) where the species does not currently occur, thus indicating a reduction in its distribution in Colonial times. The causes of the decline of these species are probably more related to the loss of habitat. Several are forest birds (C. ruficauda) for which deforestation is the main cause of habitat reduction, while another xeric-adapted species (M. gundlachii) was “once more widely distributed but has apparently retreated with the loss of xeric habitat” (Pregill & Olson, 1981, p. 82).

135The specimens of Dumetella carolinensis and Mimus gundlachii are the first mentions of these species in the Guadeloupe Islands. No fossils of Dumetella carolinensis are mentioned for the other Lesser Antilles islands and Mimus gundlachi remains have also been identified on Barbuda (Pregill et al., 1994).

136 Prior to this study, Margarops sp. was the only mimidae identified in the Guadeloupe archaeological record (Grouard, 2001a). Fossils of Margarops fuscatus have been recovered from Montserrat (Steadman et al., 1984b), Barbuda (Pregill et al., 1994) and Saba (Wing, 2001), as well as from Antigua (Steadman et al., 1984a; Pregill et al., 1988; Pregill et al., 1994), and the Anguilla Bank: Anguilla, St Martin and Tintamarre (Watters, 1991; Pregill et al., 1994; Wing, 1995; Newsom & Wing, 2004; Bochaton et al., 2020). Except on Saba and the islands of the Anguilla bank, bone remains of Cinclocerthia ruficauda were found on the same islands as Margarops. Bone remains of Allenia fusca have also been also identified on Montserrat (Steadman et al., 1984b), Barbuda (Pregill et al., 1994) and Saba (Wing, 2001).

3.25 - Passeriformes - Turdidae

137Turdidae is represented by 18 remains corresponding to 8 individuals that can primarily be attributed to the Forest Thrush Turdus lherminieri (NISP = 13) (tab. 2, Appendix B). Its presence is attested to during the pre-human period on Marie Galante (Grotte Blanchard) and in 4 sites spread across all the Guadeloupe Islands except Les Saintes and dating from the Archaic to the Historical period (Grotte Blanchard 2, Roseau, Grotte des Bambous and Pointe Gros Rempart 6). Other occurrences of Turdidae have been reported from the Pleistocene layer (5) of Grotte Blanchard (unidentified Turdidae) and in the Ceramic period levels of Cathédrale de Basse-Terre (Turdus cf. lherminieri) (tab. 2).

138 Twelve bones are in good agreement with the comparative material of Turdus lherminieri; however, identification of an almost complete coracoid from Grotte de Bambous (BAM-2014-O-28) is more problematic. The omal part of the bone where the facies articularis clavicularis (fig. 12) is broken makes it impossible to observe the discriminant character identifying the genus Turdus, according to Steadman and Franklin (2017). However, the specimen is robust (Appendix Q) and the similar facies articularis humeralis and position of the hole in lateral view allow it to be attributed to Turdus lherminieri.

139The characters and large size of the 3 specimens from Cathédrale de Basse-Terre suggest they belong to Turdus lherminieri, but a direct comparison is needed to confirm this identification. The 2 femora proximal ends from Grotte Blanchard (GB-2013-O-207 and GB-2013-O-208) are very similar in shape to the Turdidae specimens from the reference collection.

140 Turdus lherminieri is endemic to the Lesser Antilles and currently inhabits most of the tropical forests on Grande Terre and Basse Terre (Eraud et al., 2013; Parashuram et al., 2015). The modern distribution of this species does not include the east side of Grande Terre, La Désirade or Marie Galante (Levesque et al., 2020; Levesque & Delcroix, 2021; e-bird, 2021). Except for the bones from Basse Terre, the past presence of Turdus lherminieri at the Grotte des Bambous on the eastern coast of Grande Terre and in the dry forest of Marie Galante and La Désirade, suggests a contraction of its range on Grande Terre, in addition to its extirpation from La Désirade and Marie Galante (fig. 3).

3.26 - Passeriformes - Icteridae

141Only 2 Icteridae remains were found in the caves on Marie Galante (tab. 2, Appendix B). The most complete specimen (GC2-2014-O-3), a nearly complete left humerus from the pre-human layer (4-5) of Grotte Cadet 2 (fig. 13), was compared with 10 modern and 6 fossil specimens (7 species) of Icteridae. The shape of the ventral margin of the crista bicipitalis, epicondylus ventralis and processus supracondilaris dorsali fits well with all the comparative specimens.

142 The fossil humerus from Marie Galante is larger than that of Icterus oberi from Montserrat, I. portoricensis and Agelaius xanthomus from Cueva Catedral (Wetmore, 1922) and the modern specimen of Baltimore Oriole I. galbula. This specimen is smaller than that of Venezuelan Troupial I. icterus, and much smaller than Carib Grackle Quisqualus lugubris (Appendix R). Finally, we could not assign the Blanchard Cave fossil specimen to a particular species, nor could we rule out the possibility that it might not fit one of the present-day species, because we could not compare it to some West Indian species that were not represented in our comparison corpus (e.g., Icterus bonana from Martinique or Icterus laudabilis from Ste Lucia).

143 The distal humerus from a pre-human layer (5) of Grotte Blanchard appears much larger than the one from Grotte Cadet 2, but its fragmentary nature makes it impossible to be sure that it belongs to the same species. Furthermore, differences in the general proportions do not rule out the possibility that it is from the same species.

144 Except for the Carib Grackle Quiscalus lugubris, members of the Icteridae family are uncommon or rare in Guadeloupe: Bobolink Dolichonyx oryzivorus, Orchard Oriole Icterus spurius and I. galbula (Levesque & Delcroix, 2019). Today, another passerine bird of the New World family Icteridae, the Shiny Cowbird Molothrus bonariensis, is present in the West Indies. This bird is native to South America and Trinidad and Tobago, and their range is increasing towards the north (Cruz et al., 1989).

145 However, the diversity of Icteridae species may have been greater in the past. The Bobolink Dolichonyx oryzivorus, a small species of Icteridae that inhabits open environments, is listed by Pinchon for Guadeloupe (Pinchon & Bon Saint-Côme, 1951) and also from the beginning of the 19^th century on the list of L’Herminier under the name of Emberiza oryzivora (Lawrence, 1879). Above all, an icterid of comparable size to the Antillean orioles, reputed to come from Guadeloupe, was collected by Moreau de Jonnès and sent to the Museum of Paris in 1814, according to the year on the base plate (catalog number 2010-429) (Lesson, 1831). The bird was named Pendulinus rufigaster by Vieillot (1816) then “Troupiale enfumé” Bananivorus rufigaster by Bonaparte (1854), and was thought to represent a species in its own right, distinct from the other West Indian orioles, according to these two authors. According to Hellmayr (1937), on the other hand, the bird was not different from the Martinican form, Icterus bonana, which he considered synonymous with Bananivorus rufigaster. The hypothesis of an oriole species specific to Guadeloupe was subsequently abandoned, although it should be noted that a specimen of “Troupiale enfumé” from Guadeloupe was inventoried by the NHMUK in the course of the 19^th century (Gray, 1870).

146 In the Lesser Antilles, the Icteridae depicts a pattern whereby the main islands either sheltered or were home to now rare endemic species, to which can be added regionally undifferentiated species (Molothrus, Dolichonyx, Quiscalus) resulting from a recent settlement from the American continent. Currently, the relevant elements for testing whether these bones represent an extinct species would be extant skeletons of the endemic species of the nearby islands (Martinique, St. Lucia and Montserrat). At the very least, the fossil bones described in this study or identified by Wing in Marie Galante are evidence of the past presence of an Icteridae in Guadeloupe (fig. 3).

147Remains of Icteridae indet. have previously been identified at Taliseronde on Marie Galante (Wing & Reitz, 1982) and St Kitts (Wing & Scudder, 1983).

3.27 - Passeriformes - Thraupidae

148This family is represented by 3 genera and 4 individuals (NISP = 11) from 3 sites on 3 islands, dated from the pre-human to Colonial periods (tab. 2, Appendix B).

149 Except for the two remains of the Bananaquit Coereba flaveola and Lesser Antillean Saltator Saltator albicollis from the Archaic to Colonial period layers of Pointe Gros Rempart 6 and Grotte des Bambous, the remaining specimens of Antillean Bullfinch Loxigilla noctis and Saltator albicollis come from the pre-human layers of Grotte Blanchard. Saltator albicollis and Loxigilla noctis are endemic to the Lesser Antilles and Coereba flaveola is a very common permanent resident today throughout the West Indies (Raffaele et al., 2020).

150The specimen of Coereba flaveola from La Désirade and the 5 remains of Saltator albicollis from Grande Terre and Marie Galante are the first evidence of these species from the Guadeloupe Islands and Lesser Antilles. Fossils of Loxigilla noctis have been found on Barbuda (Pregill et al., 1994), Antigua (Steadman et al., 1984a; Pregill et al., 1994) and Tobago (Steadman & Jones, 2006). Another species of this family, the Black-faced Grassquit Melanospiza bicolor, was found on Anguilla (Pregill et al., 1994) and an unidentified Thraupidae on St Kitts (Wing & Scudder, 1983).

3.28 - Passeriformes - family, genus and species indeterminate

151Numerous passerine bones (856) that require further analysis come primarily from the lower layers (pre-human and Archaic periods) of the Marie Galante caves (Grotte Blanchard, Grotte Cadet 2 and Abri Cadet 3), alongside a few remains from other sites dating from the early Ceramic (Taliseronde and Folle Anse) to Colonial periods (Grotte Blanchard 2) (tab. 2, Appendix B). On the other islands, Passeriformes remains are present throughout the Ceramic period until the Contact and Colonial periods (Roseau, Cathédrale de Basse-Terre, Grotte de l’Anse à l’Ecu, Anse Sainte Marguerite, Grotte des Bambous, Morel, Anse à la Gourde, Pointe du Helleux and Pointe Gros Rempart 6).

4 - Discussion

152Just a few years ago, our understanding of avian communities in the Guadeloupe Islands was based on studies of bones from Amerindian sites, along with descriptions and reports from historical sources. Archaeological excavations carried out from the 1950s to 1980s in Guadeloupe yielded small quantities of bones, allowing the first determinations of birds in the archaeological record. E. Clerc had entrusted a small sample of bones collected from sites in the east of Grande Terre to A. Wetmore, who identified Puffins lherminieri, Dendrocygna arborea and Pterodroma hasitata (Clerc, 1964, 1968). A sample of bones from the sites of Folle Anse and Taliseronde on Marie Galante was given to E. Wing by Father Barbotin (Barbotin, 1970; Wing, 1973; Wing & Reitz, 1982). This led to identification of additional taxa determined to either family (Threskiornithidae, Icteridae) or species (Gallinula galeata, Patagioenas squamosa, Zenaida aurita). This sample also contained two Psittacidae bones which may have represented the Guadeloupean form of Amazona (Williams & Steadman, 2001; Olson & López, 2008).

153More recently, data concerning the avifauna from the Ceramic period has benefited from the study of assemblages recovered during the 19^th and 20^th centuries as part of S. Grouard’s doctoral thesis (2001a), and then from analysis during subsequent rescue excavations. This work documented the presence of Puffins lherminieri, Ardea herodias, seabirds (Fregata magnificens, Leucophaeus cf. atricilla, Sula leucogaster), a small parrot (Amazona sp.), several columbids (Patagioenas sp., Geotrygon cf. montana, Zenaida cf. aurita) and passerines (cf. Margarops, Turdus sp.). Most of the identifications still remained at the supra-species levels and thus only partially reflect past avian biodiversity during the pre-Columbian periods.

154The chronological record was extended at the beginning of the 2000 by the discovery of natural or paleontological sites on Marie Galante (Lenoble et al., 2009). An initial study of the Abri Cadet 3 site revealed a more varied record of taxa in the Mid Holocene, including Columbidae, small passerines, and also Falconiformes and Strigiformes (Stouvenot et al., 2014). In total, 13 taxa were recognized, half of them determined to species.

155Our recent work in the Guadeloupe Islands first highlighted a form of endemic macaw, Puffinus lherminieri, Nyctanassa violacea, Athene cunicularia and a giant Barn Owl (Gala & Lenoble, 2015; Mallye et al., 2018; Gala et al., 2020, 2022), while the present study identified other 50 bird taxa in the fossil record for a total of 55 taxa, more than half of which were determined to species level (n = 29). This computation does not include bird remains identification by other authors, in a dozen of sites considered in our work (see Appendix S for the details of the results of the re-examination of bird remains studied by other authors). Analysis of assemblages from natural sites predating the first human settlement of the islands (Grotte Blanchard, Grotte de Cadet 2) and contemporary to pre-Columbian island occupation (Abri Cadet 3, Grotte Blanchard 2, Pointe Gros Rempart 6 and Grotte des Bambous) provides material from a long chronology. Improved by historical sources, this data documents the main trends in the evolution of the Guadeloupe bird community through time.

156A preliminary remark regarding interpretation of the data is that the different types of sites studied do not provide directly comparable information. Concerning natural sites, occasional use by humans cannot be excluded; although sedimentation and accumulation of bone remains mainly reflect non-human and post-depositional processes. The deposits at these sites may have been built up over a long period and potentially contain remains linked to various accumulation agents and processes. However, for the most part, bird bones from these sites attest to an attritional mortality profile related to the use of cavities by nesting birds (example of the Puffinus from Grotte Blanchard 2, cf. Mallye et al., 2018) and above all, the accumulation of bird remains by raptors (Bailon et al., 2015; Boudadi-Maligne et al., 2015). Conversely, the studied material also includes collections recovered during the excavation of Amerindian middens, where the accumulation of bones is assumed to be of human origin.

157These different contexts provide comparable information in terms of the number of species: 22 taxa for sites before human settlement (Late Pleistocene and early Holocene), 30 for Amerindian sites, and 26 for contemporary natural sites (Late Holocene). However, these differences may induce biases in the accumulation of bones, such as under-representation of small taxa in the Amerindian sites due to their being less sought after by these populations or because their consumption did not preserve bones. In addition, excavation methods for these two types of sites may differ. While excavations of natural sites carried out in the last decade included systematic use of a fine mesh during wet sieving, sieving was not systematic during excavation of archaeological sites, particularly during earlier fieldwork, and may have favored the recovery of large bones (Payne, 1975).

158Whatever the reason, this differential representation of taxa according to body mass is evident when the species recognized at the different site types are plotted according to size (fig. 14). Birds weighing less than 100 g (i.e., small-to-medium sized forest birds) are almost exclusively represented in natural sites, whereas birds weighing 300 g or more, including large seabirds, Accipitriformes, or Anseriformes, come mostly from archaeological deposits (fig. 15). This size difference should be kept in mind when comparing data from these different sources.

159Among the 55 taxonomic determinations, most are the first mention for the Guadeloupe or Lesser Antilles fossil records. Overall, forest birds represent more than half of the taxa determined, with Passeriformes represented by the highest number of taxa, followed by the Columbiformes, Caprimulgiformes (nightjars and hummingbirds), diurnal raptors (Accipitriformes and Falconiformes), with some orders represented by only 1 or 2 species (Strigiformes, Piciformes and Psittaciformes).

160These forest birds are completed by waterbirds (Gruiformes, Pelecaniformes and Anseriformes), seabirds (Procellariiformes, Phaetontiformes and Suliformes) and waders (Charadriiformes). This assemblage depicts considerable variability and complementarity between its different components. However, it represents only a fraction of the current Guadeloupe avian community. This is particularly apparent when the number of nesting species in the dataset is compared to that of current avifauna. Forty-five taxa represented in the fossil record are species that nest in the Caribbean, while the list of current Guadeloupe bird community records 81 (Levesque & Delcroix, 2019).

4.1 - Recomposition of Avian communities

161Published data for Guadeloupe bird communities suggests that the number of extinct and extirpated species is scarce. Three Psittacidae are listed as extinct in the Guadeloupe avifauna and one, Athene cunicularia, as extirpated (Levesque & Delcroix, 2019). Two other species, Numenius borealis and the Guadeloupean form of Troglodytes aedon, are suspected to be (globally) extinct and extirpated, respectively. Historical sources nevertheless suggest that other birds may also be concerned: the possibility of an endemic icterid, Pendilunus rufigaster (Vieillot, 1816; Hellmayr, 1937), and a Caribbean form of nightjar represented by a vellum preserved in the Muséum national d’Histoire naturelle. While the exact number of extinct taxa remains difficult to estimate, this data suggests it is not significant.

162Analyses of the archaeological remains, however, nuance this first impression and point to more substantial recomposition of the avian community. Additionally, this change takes several forms: extinction, local disappearance or extirpation, reduction in habitat size within the same island or rarefaction expressed by a reduction in the number of individuals forming a population, or by loss of nesting status or the arrival of new taxa in an impoverished native bird community. When an extirpation event is considered as the disappearance of a single taxon from one of the main islands of Guadeloupe (Basse Terre and Grande Terre counted as one island, La Désirade, Marie Galante, Les Saintes), 33 local disappearances are observable, 4 of which are extinctions (the 3 Psittacidae and the giant form of the Barn Owl; see the “Chronology of extinction and extirpation” section below). This assessment combines the fossil record with historical evidence for the birds of Guadeloupe.

163In terms of orders, among those identified in our work, these disappearance events reveal forest birds, to be the most affected, primarily Passeriformes (Mimidae and Turdidae) and Psittaciformes (Psittacidae), followed by marine birds, in this case Procellariiformes (fig. 16).

164Among the species not identified by traditional ornithological sources are Mimus gundlachii identified at Grotte des Bambous, a giant owl form identified at the Amerindian site of Roseau (Gala et al., 2022), Patagioenas inornata at the site of Cathédrale de Basse-Terre or the small forms of rail similar to Hapalocrex flaviventer and Laterallus jamaicensis dated to the Historical period from Pointe Gros Rempart 6 and Abri Cadet 3, respectively. In addition, species still present in Guadeloupe have disappeared from certain islands, such as Turdus lherminieri and Cinclocerthia ruficauda, which have been recognized on La Désirade and Marie Galante, where they are now absent.

165Except for the Psittacidae, which are known to have been of interest to human populations for a very long time, ground-nesting birds have long been identified as a fragile ecological group in the face of human-induced modifications of island ecosystems, first and foremost the introduction of mammals, whether carnivores (cat, mongoose) or not (rat). This bird category is represented by Puffinus lherminieri and Pterodroma hasitata, the Caprimulgidae, and Athene cunicularia, as well as the Rallidae. Together they account for just over a third of all disappearance events (n = 12/33). Avian community recomposition thus involves a much greater diversity of species than just ground-nesting species, particularly forest birds.

166The past presence of Turdus lherminieri, Cinclocerthia ruficauda and Melanerpes herminieri at the Grotte des Bambous on the eastern coast of Grande Terre documents a reduction in the habitat of these species. The combination of these 3 species is remarkable because they are all forest species and have comparable distributions restricted to the rain forest of Basse Terre and the swamp forest of western Grande Terre. Their past occurrence therefore indicates a reduction in their habitat in human-modified (deforested) sectors of Grande Terre, which is congruent with the disappearance of thrush and trembler from the other dry islands (Marie Galante and La Désirade). An analysis of the Grotte Bambous faunal spectrum shows that non-avian forest species were also affected by the emergence of new environmental conditions (e.g., Lesser Antillean Tree Bat Ardops nichollsi and Guadeloupe big brown bat Eptesicus guadeloupensis) (Cochard et al., 2019).

167Several species were abundant in the past, presumably forming breeding populations, such as Dendrocygna (Du Tertre, 1667-1671). The Caribbean form, Dendrocygna arborea, is a largely sedentary breeding species. Its past presence on Amerindian sites suggests that it nested in Guadeloupe, which is not the case today, with the exception of an attempt observed on Les Saintes in the early 2010s (Levesque & Sorenson, 2012).

168Historical sources also show a decrease in some populations of what were abundant forest birds in the 17^th century, as in the case of Patagioenas squamosa (Breton, 1665; Du Tertre, 1667-1671; Labat, 1724). The ornithological observations reported by the STOC-Guadeloupe project show that this bird has become rare today and is confined to the canopy of the high forest (Levesque et al., 2020). On the other hand, Zenaida aurita, which is very common today throughout the archipelago, would have been rare at the time of the first European settlement of Guadeloupe, as reflected in the text of Father Breton, who was present on the island from the first years of its colonization (Breton, 1665).

169The list of species represented in the fossil record includes a small number of forest birds considered to be recent colonists among the Caribbean avifauna. According to Ricklefs & Bermingham (2001), these recent colonists show little genetic divergence between the different island populations and a continuous or near-continuous distribution in the Lesser Antilles. Their presence in the archipelago may thus be more recent than 0.55 to 0.75 Ma, which distinguishes them from the old colonists that are now either endemic (e.g., Turdus lherminieri or Melanerpes herminieri) or native taxa that expanded their range recently to include the Lesser Antilles (e.g., Cinclocerthia ruficauda or Margarops fuscatus). The recent colonists represent a large proportion of current forest birds (orders Columbiformes, Caprimulgiformes, Psittaciformes, Piciformes, and Passeriformes) and form the large and fairly generalist populations that are well represented in open environments (Ricklefs, 2010). Except for the columbids, that are considered as recent colonist in reason of the continuing migration through the islands as well as for their association with drier habitats of recent glacial periods (Ricklefs & Bermingham, 2004), these “new arrivals” are thought to represent the last stage of settlement of the Lesser Antillean avifauna. A small number of these “new arrivals” are taxa whose range extension to include the West Indies has been reported in the ornithological literature, such as the Spectacled Thrush Turdus nudigenis (Bond, 1945; Levesque, 1997) or the Tropical Mockingbird Mimus gilvus (Bond, 1963; Guth, 1971). Most represent taxa whose presence is attested to as early as the first ornithological lists of the 19^th century (e.g., the Grey Kingbird Tyrannus dominicensis, the Mangrove Cuckoo Coccyzus minor, the Carib Grackle Quiscalus lugubris, cf. Lawrence, 1879).

170When the columbids are excluded, the assignment of birds in the Guadeloupe fossil record to one of three categories, “Young” (recent colonists), “Spread” (old colonists that have recently spread within the Lesser Antilles) and “Old” (colonists that are endemic to individual islands within the Lesser Antilles) shows the recent colonists to be under-represented in the fossil record, especially when the distribution of these groups is compared with the current avifauna of Guadeloupe (fig. 17). In addition, the recent colonist identified in the fossil record, namely the Bananaquit Coereba flaveola occurs in sites that are contemporary with the first human presence on the archipelago as well as the Antillean Nighthawk, Chordeiles gundlachii, if confirmed the identification. Thus, none of the recent colonists are identified in the fossil record before the earliest human settlement. This absence is unexpected as arid, open habitats that from favorable conditions for these new arrivals prevailed during the Late Pleistocene (Royer et al., 2017). This absence thus suggests that part if not all of these new taxa may reflect a very recent event, coincident with human settlement of the island. According to this interpretation, these new arrivals represent an additional expression of human-induced turnover in avian community by substituting the disappeared, extirpated, and habitat-reduced species highlighted in this work.

4.2 - Chronology of extinction and extirpation events

171Over the past few decades, the estimated date of extinction/extirpation of bird species in the Lesser Antilles in particular, and the Caribbean in general, has been based on cultural (archaeological) and non-cultural (paleontological) contexts, as well as on the degree of bone surface mineralization. In fact, a significant portion of the avifauna has been considered to be of Pleistocene age without supporting evidence (Pregill & Olson, 1981; Pregill et al., 1994), and climatically induced changes that emerged at the Pleistocene-Holocene transition (PHT, ca. 11.7 ka) have long been associated with vertebrate community turnover (Pregill & Olson, 1981). However, analysis of the fossil content of the Burma Quarry site, in addition to the Ceramic sites on Antigua, has refined this model, highlighting the disappearance of several taxa during the Holocene (Steadman et al., 1984a; Pregill et al., 1988). The relative contribution of these climatic events to renewal of Caribbean avifauna thus remains unclear.

172The timing of extinction/extirpation events of birds from the Guadeloupe Islands has been established based on direct radiometric dates obtained on bird bones or other components of the assemblage of the sites or even sediments (see presentation of the sites). For the archaeological sites, the cultural material also makes it possible to include them in the regional archeo-chrono-stratigraphy and to subsequently assign a terminus post-quem for these extinction events (e.g., Orihuela et al. 2020). Completed with information from historical or ornithological sources, this data provides the Last Appearance Datum for a large number of species (LAD; representing the last time a species was mentioned in the fossil record or historical sources) (fig. 18). Five extinction events have a Pleistocene LAD, all of which concern Marie Galante, the only island to have produced Pleistocene fossil material. Four of these five LADs are compatible with the PHT because of their chronological proximity to this transition.

173A Caribbean form of Nightjar (Antrostomus sp.) occurs only in the early part of the fossil record and is represented by a small number of remains (n = 2). The presence of these two bones in the lowermost levels of Grotte Blanchard may thus reflect the randomness of the chronological distribution of a taxon that is rare in the fossil record. It is therefore difficult to rule out the possibility that this taxon might have persisted over a longer period of time. In fact, the presence of a Caribbean form of Nightjar on the neighboring island of Guadeloupe is suggested by one specimen from the Historical period.

174A similar situation can be seen for 2 of the 4 taxa with an LAD close to the PHT. The persistence of a local form of Icteridae in Guadeloupe is suggested by the icterid specimen collected by Moreau de Joannes in the early 19^th century, while the presence of the Ara is clearly attested in Guadeloupe during historical times by the accounts of chroniclers. Furthermore, the fact that these taxa occur only in Pleistocene layers from sites on Marie Galante may simply reflect the more diversified faunal spectrum of natural sites from the pre-human period compared to those of the following periods.

175In addition, the PHT changes are characterized by increased precipitation favoring the extension of forest and closed landscapes. The taxa whose disappearance may have been induced by these environmental changes are therefore primarily the taxa of arid or open habitats. Zenaida asiatica is the only species in this category. This bird is one of the recent colonists of the Lesser Antilles avifauna, where it is becoming a regular breeding species (Levesque, 2013), favored by the emergence of more open landscapes and aridification as a result of deforestation. The possible past presence of this species in the fossil record may reflect favorable conditions for this species in the Pleistocene, with an extirpation event resulting from the extension of forests in the Holocene.

176Furthermore, our data suggests that most extinction or extirpation events over the last millennia are contemporary with human presence in the archipelago. This pattern is observable for Basse Terre, Grande Terre and La Désirade, where this pattern may reflect the absence of a fossil record before the earliest arrival of humans. The same can be observed on Marie Galante, where the Pleistocene record is well established. Finally, two distinctive periods stand out. A significant number of disappearances occurred during the Archaic and Ceramic periods. These include the disappearance of Buteo swainsoni, the giant form of Tyto or Patagioenas inornata from Basse Terre and Grande Terre, and the forest birds Turdus lherminieri and Cinclocerthia ruficauda from La Désirade and Marie Galante. Their disappearance could reflect very early human pressure on the environment, or an underestimating of their real age of disappearance for lack of historical documentation, in particular for the islands of La Désirade and Marie Galante.

177The second period sees a similar number of disappearances and corresponds to the Historical period. Evidence from the fossil record and historical sources documents the loss of nesting sites for Pterodroma hasitata and Puffinus lherminieri from the main islands of Guadeloupe, the disappearance of the Psittacidae from Basse Terre and Grande Terre, of the Rallidae from La Désirade and Marie Galante, and of Athene cunicularia. This loss is evident despite the fact that only a few sites can be reliably attributed to this period. With that said, direct radiocarbon dates obtained on Puffinus lherminieri and Athene cunicularia remains from Grande Terre provide secure evidence for their extirpation in historical times. Further direct dates on bones of locally disappeared taxa should make it possible to establish the role of this period more precisely in the recomposition of the islands’ bird communities.

178Compared to islands with reliably dated bird fossils such as Cuba, the LADs in the Lesser Antilles tend to show a similar time pattern, with most disappearances occurring during the Holocene (Orihuela et al., 2020) or change in the taxonomic diversity in the Late Quaternary (Zelenkov & Belichenko. 2022). Our results differ from those of Steadman and Franklin (2020) for the Bahamas where the same proportions of extinctions were observed in the late Pleistocene and early Holocene.

4.3 - Extinction factors

179By correlating environmental changes and LAD, it is possible to estimate that most of the documented disappearances or extirpations took place after humans arrived in the archipelago. The previous hypothesis of environmental modification at the end of the last glaciation as the primary cause underlying the disappearance of Caribbean species (Pregill & Olson, 1981) no longer seems tenable. Instead, the focus should be on the direct or indirect impact of human activity.

180Understanding the factors or causes of species extinction or extirpation is delicate. For the Caribbean this has been addressed by correlating an extinction event with the environmental changes based on LAD, for example (e.g., MacPhee & Flemming, 1999; Cooke et al., 2017).

181However, this approach has its limitations, in that species extinction proceeds through a succession of phenomena where a species is unable to maintain its numbers as a consequence of environmental variations, rendering relict populations sensitive to various environmental hazards (Simberlof, 1986; Lande, 1993). Assigning an extinction event to a particular cause therefore appears often pointless. An illustrative case is the extirpation of Pterodroma hasitata from Guadeloupe. The disappearance of this taxon has been reported as the result of the 1847 earthquake that caused the collapse of some slopes of the Soufrière massif where the species nested (e.g., Goetz et al., 2012). This is Noble’s (1916) record of an account provided by an old Soufriere resident that the petrel was no longer heard following this quake. The species was extensively overhunted from the first years of European presence in the Guadeloupe Islands (Breton, 1978, Du Tertre, 1667-1671), and catches remained considerable for decades (Froger, 1698; Longin, 1848; Lawrence, 1891), to the point that “the Negroes and the small inhabitants do not live on anything else during the season” (Labat, 1724, t. 1, part. 2, p. 110). Only the hunts reported at the end of the first part of the 19^th century showed a significant reduction in the number of catches (Révoil, 1877). Father Labat, who took part actively in these hunts at the end of the 18^th century, notes that “the difficulty of hunting these birds preserves the species, which would have been destroyed entirely many years ago, according to the bad custom of the French, if they did not withdraw to places that are not accessible to everyone” (Labat, 1724, t. 1, part. 2, p. 110). The collapse of the most inaccessible slopes of the Soufrière and the surrounding massifs therefore only affected a colony that was already reduced both numerically and spatially. The overhunting practiced in the previous decades contributed substantially to the extirpation of this bird.

182The combination of processes leading to the disappearance of a species from the Guadeloupe bird community is also illustrated by Athene cunicularia. Several ornithologists (Noble, 1916; Bond, 1936; Danforth, 1939) have suggested that the disappearance of this species from the Lesser Antilles was due to the mongoose, insofar as this ground-nesting bird was exposed to the introduction of a carnivorous predator and there was a correlation between the date when the last specimens were observed in Marie Galante and the introduction of the mammal on the island (in 1890 according to Lorvelec et al., 2021). Ober’s inability to obtain this bird in Guadeloupe despite a particular interest by the naturalist in acquiring specimens during his major bird collection in 1877, attests to its disappearance from both Grande and Basse Terre being earlier and, by the same token, to other mechanisms involved in the disappearance of this bird from the islands of Guadeloupe (Gala et al., 2020).

183The difficulty of understanding the influence of different factors in the disappearance of a species is further compounded by the fact that this influence can be indirect, for example, when it is expressed by a succession of disappearances within the same food web. The disappearance of the giant form of Tyto is a good example. The ecological hypothesis for the extinction of this bird is the joint disappearance of the predator, the owl, and its prey, the local form of endemic rodent, the Antillomys rayi (Gala et al., 2022). Recent work with this rodent species places its complete disappearance or at least its rarefaction in the first years of European colonization or the Contact period, as a consequence of the introduction of the black rat that potentially carried a pathogen (Goedert et al., 2020). In this case, the introduction of pathogens accompanying the appearance of Old World mammals may have led indirectly to the disappearance of this giant owl.

184One approach to identifying the potential causes of extinctions is to list the interacting factors that may have favored them globally, rather than looking for a particular cause. Based on a study of the extinctions of Pacific Island birds (Steadman, 2006), Steadman and Franklin (2020) listed seventeen factors that may have favored or, conversely, minimized the disappearance of native birds of the Bahamas (tab. 4). Three categories are identified: abiotic factors, indigenous biological factors, and cultural factors. In the case of the Bahamas, two abiotic factors (the small size of the islands and a flat, low topography), three native biological factors (a depauperate floral and faunal diversity and the absence of indigenous terrestrial mammals), and one cultural factor (an extensive island-wide settlement pattern) were found to favor extinctions. On the other hand, one abiotic factor (impoverished soils limiting the cultivation of the entire territory) may have offset the effects of human-induced environmental modifications, allowing avian populations to persist.

185The data collected during our work highlights the specificity of the islands of Guadeloupe with respect to the Bahamian model.

186The size of the islands, at least in that it allows the maintenance of a physiographic complexity and areas preserved from human impact, may account for the disappearance of birds from Marie Galante and La Désirade, while these same species only experienced a reduction in their habitats on Grande Terre, where they persist in the preserved forested areas of the island. This is the case with Cinclocerthia ruficauda and Turdus lherminieri, both present today in the swampy forests in the west of the island and repopulating the central sectors of the island in the process of reforestation since the 19^th century (Levesque et al., 2020).

187The deleterious role of the flat, low-lying topography of the islands is less clear. A significant part of the surface of La Désirade is formed by the slopes bordering the central plateau of the island. This relief is unfavorable to human settlement but does not seem to have limited the extinction of species, whether the birds identified in this work or terrestrial mammals and herpetofauna, as pointed out by Boudadi-Maligne et al. (2015). In this initial investigation of the disappearance of vertebrates from the island, the authors identified the cultivation and deforestation of the flat parts of the island, combined with the deforestation or grazing of sectors of secondary agricultural interest, leading to general erosion of soils inducing a collapse of the food web (Boudadi-Maligne et al., 2015). According to this interpretation, the scarcity of calcareous soils, which limits their ability to recover after erosion, appears to be a key factor promoting rather than limiting extinctions. In addition, free grazing of goat herds and the production of coal were important in historical times, eliminating sloped areas as refuge for native species. The exploitation of the entire surface of the small islands, combined with poor soils, therefore appear as primary factors contributing to the fragility of island ecosystems.

188Poor soil quality and the steep topography of the islands may, however, delay the disappearance of native avifauna. The swampy sectors, which were not suitable for historical exploitation, remain much the same on the main islands of Guadeloupe today. On the volcanic island of Basse Terre, the very steep slopes of the mountain range combined with regular, substantial rainfall that prevails on the highest reliefs preserve mountain forest. These two forests, swampy and mountain, today support the core population of species that have disappeared from the rest of Grande Terre, including Melanerpes herminieri, Cinclocerthia ruficauda and Turdus lherminieri. In the swampy area, soils are difficult to exploit, while in mountainous zones, the combination of steep relief and climate act as a brake on the disappearance of forest birds.

189An arid climate accompanied by a marked dry season is considered to be a factor leading to fragility, as it favors the burning of forests, whether for cultivation and logging for the production of charcoal or simply to favor hunting. The burning of savannas was, in fact, already practiced by indigenous Caribbean populations as a method of hunting to collect “half roasted” game (de Rosny, 1886). Such practices, whether they took place during the Amerindian or Historical period, may have favored the disappearance of species occupying drier habitats. The disappearance of such species is documented in the fossil record of the islands of Guadeloupe by Mimus gundlachii and Athene cunicularia.

190The only non-flying mammal species that may have populated the Guadeloupe Islands before the arrival of human populations is the Antillomys rayi (Brace et al., 2015). Its presence before the first Human occupation of the Guadeloupe Bank (ca. 3,000 BCE) remains subject to debate, however, while its presence on Marie Galante is only attested from the Amerindian times, with direct dating occurrence testifying its presence between 9^th and 14^th century AD (Goedert et al., 2020). Nevertheless, how this native rodent may have exerted pressure on avian populations remains unknown. The historical mammalian introductions (rat, cat, mongoose) on these islands may have favored the disappearance of species, at least indirectly, as in the case of the giant Tyto, by inducing the disappearance of the native rodent prey species. However, the effect of mammalian introductions remains difficult to assess, as shown by the disappearance of Athene cunicularia, which predated the introduction of the mongoose in Grande Terre and also disappeared from La Désirade where this carnivorous mammal was never introduced (Lorvelec et al., 2021).

191A relatively impoverished fauna may also have contributed to the fragility of bird populations, as the absence or early disappearance of large terrestrial mammals led human populations to exert predation pressure on birds such as Procellariidae or Columbidae.

192The morphological traits and behavior of certain species certainly played an important role in their disappearance. This is the case of Pterodroma hasitata, reputed for the quality of their meat (Du Tertre, 1667-1671; Labat, 1724) and so attractive that salted fish were sent from the neighboring island of Dominica to supplement the local catch (Atwood, 1791). The ease of capture of ground-nesting species, such as petrels and shearwaters, may also have contributed to the collapse of populations. In the case of Puffinus lherminieri, however, this aspect must be qualified, insofar as, although this bird appears to have been hunted regularly by Amerindian populations, colonies nevertheless persisted throughout this period. The excavation and study of the Grotte Blanchard 2 nesting site suggests a reoccupation by birds of the southern coast of Marie Galante following the abandonment of this area by Amerindian populations (Mallye et al., 2018). Dates obtained on Puffinus lherminieri bones from the Amerindian cave of Morne Rita, which was both a nesting site and occupied by Amerindians, places the last occupation of this cave by birds between the 14^th and 19^th centuries. This would support the idea of a dynamic relationship between human collection pressure and the locations of colonies. In fact, the introduction of carnivorous mammals to Marie Galante appears to be the most parsimonious explanation for the abandonment of the area of by nesting bird colonies (Mallye et al., 2018).

193The pigeon (Patagioenas spp.) was also valued for its meat, as Father Labat reports, to the point that this bird remained the main game species hunted on the island throughout the historical period (Noble, 1916). The same was true of the Psittacidae, already consumed by Amerindian populations (Monpilliers, 1654) and which remained a choice food in the first centuries of the Colonial period, before they disappeared (Labat, 1724; Le Hirbec, 1890). These same birds, raised in Caribbean villages (de Rochefort, 1681), also attracted the attention of humans for the quality of their feathers, as did the dried hummingbirds offered as presents “to ladies who make earrings out of them” (Breton, 1665).

194A dataset including several islands sharing the same avifauna makes it possible to identify the factors that contributed to the disappearances of bird species in the Guadeloupe Islands. These factors appear to differ from those held to explain the recomposition of avian fauna in the Bahamas. The combination of three factors seems to be the main driving force behind disappearances in the islands of Guadeloupe: the small size of the islands, the absence of steep cliffs and an island-wide human settlement pattern and hence influence on the small islands, either directly (hunting, forest exploitation) or indirectly (cultivation and soil erosion, grazing, introduction of non-flying mammals). Secondary factors also played a role, namely an impoverished fauna, the absence or near absence of terrestrial mammals and specific ecological traits exposing certain taxa to human impact.

5 - Conclusion

195The archaeological and paleontological records are an important source of data for documenting the past avian biodiversity of the Guadeloupe Islands. Until a few years ago, most taxonomic determinations were at the supra-specific level (genus, family, and order) and were to be assessed in the context in which they were produced: small reference collections that are not representative of all current biodiversity, lack of determination keys for families of biogeographic interest (e.g., Columbidae and Mimidae). Our study has benefited from a large reference collection, new osteological descriptions and quantitative data for the reliable identification of avian remains, and a larger number of taxa have been identified in the 5 islands of the Guadeloupe archipelago (55 taxa). The number of fossil taxa is the largest of the Lesser Antilles (7 sites in Antigua have yielded 24 taxa and 3 sites in Barbuda 30 taxa) but it represents only a fraction of the current Guadeloupe avian community. The fossil record thus seems to provide only a partial representation of avian diversity, which we estimate to represent at best about half of the overall Guadeloupe breeding bird community.

196Among the identified taxa many disappearance events (e.g., extinction, local disappearance or extirpation) are documented and affected primarily forest birds (e.g., Mimidae, Turdidae and Psittacidae). The causes of biodiversity loss in the avian community are complex and the example of the Burrowing Owl shows the difficulty of answering these questions without a reliable chronological framework (direct dating of bones) or a combination of several types of data (ecological considerations, historical sources).

197The correlation between environmental changes and the dating of the last occurrences shows that most of the avian disappearances and extinctions examined in our research were recent and took place during the last millennia in the Guadeloupe Islands. The human presence in the archipelago is strongly connected with these extinction and extirpation events and birds have shared the same fate as many terrestrial vertebrates (e.g., snakes and lizards), condemned to disappear by direct and indirect human impacts.