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Rev. Mus. Argentino Cienc. Nat., n.s. 18(1): 9-30, 2016 ISSN 1514-5158 (impresa) ISSN 1853-0400 (en línea)

Recent changes in plant and animal distribution in the southern extreme of the Paranaense biogeographical province (northeastern Buenos Aires province, Argentina): Ecological responses to climate change?

Elián Leandro GUERRERO1 & Federico Lisandro AGNOLIN2, 3

1División Plantas Vasculares Dr. Ángel L. Cabrera, Herbario LP. Museo de La Plata. La Plata, Argentina. [email protected]; 2Laboratorio de Anatomía Comparada y Evolución de los Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”. Buenos Aires, Argentina; 3Departamento de Ciencias Naturales y Antropología, Fundación de Historia Natural “Félix de Azara”, Universidad Maimónides. Buenos Aires, Argentina.

Abstract: The Northern region of Buenos Aires province, in Argentina, suffered an uninterrupted increase in rainfall and temperature values during the last decades. The aim of the present contribution is to analyze the ef- fects of such climatic change in the distribution of local flora and fauna. The analysis resulted in the recognition of distributional shifts in a total of 115 species. These changes represented the progressive southern expansion of species typically present in subtropical areas. Because all these changes have a similar orientation and direction, and included a large number of plant and animals with different ecological requirements, it is here postulated that those geographical changes are probably a response to climatic alteration.

Key words: Buenos Aires; climate change; riparian forest; species distribution.

Resumen: Cambios recientes en la distribución geográfica de plantas y animales en el extremo sur de la provincia biogeográfica Paranaense (noreste de la provincia de Buenos Aires, Argentina): ¿Respuestas ecológicas al cambio climático? La región norte de la provincia de Buenos Aires, en la Argentina, sufrió un incremento ininterrumpido de los valores de precipitación y temperatura durante las últi- mas décadas. El objetivo de esta contribución es analizar los efectos de dicho cambio climático en la distribución de la flora y fauna locales. El análisis resultó en el reconocimiento de corrimientos en la distribución de un total de 115 especies. Estos cambios representaron una expansión progresiva hacia el sur de especies presentes típica- mente en áreas subtropicales. Como todos estos cambios tienen una orientación y dirección similares, e incluyen un gran número de plantas y animales con diferentes requerimientos ecológicos, se postula que estos cambios de distribución geográfica son probablemente una respuesta a la alteración climática.

Palabras clave: Buenos Aires; cambio climático; bosques en galería; distribución de especies. ______

INTRODUCTION of distribution alongside large rivers to the La Plata River latitude (Cabrera, 1971; Figure 2). Buenos Aires province is located in East- From an Ecoregional framework, Burkart et al Central Argentina. It is biogeographically in- (1999) named this complex “Delta and Paraná tegrated within the Pampean biogeographi- Islands Ecoregion”. Riparian rainforest’s plants cal province, of the Chacoan dominion in the belong to the Paranaense biogeographical prov- Chacoan sub-region (Cabrera, 1971; Morrone, ince (Cabrera & Dawson, 1944; Cabrera, 1971) 2014; Figure 1). Most of Buenos Aires prov- whereas coastal-forest plants and animals are ince is characterized by grasslands (locally widespread in Chaco and Paranaense provinces named “pampas”) and Celtis ehrenbergiana xe- (Ringuelet, 1955). Then, in northeastern Buenos rophytic woods (locally named “talares”) both Aires province, Paranaense flora and Subtropical typical of the Pampas biogeographical province fauna intergrades with Pampean biota, forming (Morrone, 2014). However, subtropical gallery an ecotone (the “subtropical-pampásico” ecotone rainforests and coastal forests from northeast- sensu Ringuelet, 1961, 1981). ern Argentina have their southernmost limit Climatic change is largely known to affect 10 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Fig. 1: Map of northeastern Argentina and bordering Fig. 2: Physical features of the study area and sites countries. A: Biogeographical units. In grey = mentioned in the text. Paraná River floodplain (Delta and Paraná Islands ecorregion of Burkart et al., 1999); Dotted = Pampa Biogeographical province sensu Morrone (2014). B: first, to show that geographical changes in spe- Political division. Argentinean provinces mentioned cies are not isolated and that there is a common in the main-text: MI = Misiones; FOR = Formosa; tendency to extend the southern limit of sub- CORR = Corrientes; S. FE = Santa Fe; CORD = tropical plant and animal species; and second: Córdoba; E.R. = Entre Ríos; B.A. = Buenos Aires. to examine if the observed shifts on animal and plant species distribution could be attributed to plant and animal distribution, having diverse climatic change. effects on biotic communities and ecosystems (McCarty, 2001; Walther et al., 2002; Parmesan, MATERIALS AND METHODS 2006; Walther, 2010). Climatological studies in Argentina show that during the 20th century, Geographical settings Buenos Aires province suffered an uninterrupted We analyzed the northeastern sector of increase in precipitation and temperature values Buenos Aires province, where Paranaense ri- (Berbery et al., 2006; Menéndez, 2006; Servicio parian forests have their southernmost patches Meteorológico Nacional, 2015; Barros et al., along the coast of the Lower Paraná and La Plata 2015). Having in mind such large-scale climat- rivers (Figure 2). We restricted our research to ic change, a poleward shift in the geographical this area because it was largely considered as the distribution of plant and animal communities is best known region in Argentina from a biological expected. viewpoint, and because it was the focus of study In spite of that, an integrative study about the of several researchers during the 19th and 20th impacts of recent climate change on Argentinean centuries. Consequently, large biological collec- ecosystems is still lacking. Studies about changes tions are available from this area since historical in species geographical distribution related to cli- times. This allows comparison between old and matic variation were mainly conducted in mam- recent collections. We did not include central and mals (e.g. Udrizar Sauthier et al., 2005; Fracassi southern Buenos Aires province, even though et al., 2010), medically important snakes (Nori some recent cases of shifts on animal species et al., 2013), insects (Farina, 2006; Medone et distributions were probably caused by climate al., 2015), and arachnids (Guerrero, 2014a). change (e.g. Zamorano & Scillato-Yané, 2008; Furthermore, there is a single phenological study Doumecq-Milieu et al., 2012). This is because of changes in arriving times of a migratory swal- those areas were poorly sampled by researchers low (Spescha et al., 2004). Nevertheless, each in historical times. one of these works focus on single species or on a From north to southeast, Pergamino, San small taxonomic group of animals. On the other Nicolás, Ramallo, San Pedro, Baradero, Zárate, hand, plant species range shifts deserved almost Campana, Pilar, Zelaya, Tigre and San Isidro no attention. The aim of this paper is twofold: districts, together with Martín García Island Guerrero & Agnolin: Recent changes in plant and animal distribution 11 encompasse the northern boundary of Buenos seums (Acosta, 2014). Therefore, we infer that Aires province, including Paraná’s Delta and knowledge of haverstmen in this area is relative- slope talares; from Buenos Aires City to the ly good. Furthermore, in this part of the prov- southeast are located the districts of Avellaneda, ince several harvestmen that distribute along Quilmes, Berazategui, Ensenada, La Plata, the Argentinean Mesopotamia (the geographical Berisso, Magdalena and Punta Indio; Punta Lara area encompassed between Paraná and Uruguay Natural Reserve is placed between Berazategui rivers) have their southern limit of geographical and Ensenada districts, and Parque Costero del distribution (Ringuelet, 1959; Acosta, 2002). On Sur Natural Reserve in Magdalena and Punta the contrary, because the Araneae are not well Indio districts. studied on a zoogeographical point of view, only a brief analysis of two well-studied genera is Taxon selection added. Analyzed taxa were chosen under two main Lepidoptera distribution. Lepidoptera species criteria. Firstly, we selected relatively well-stud- were shown to be very useful in climatic change ied species from the biogeographical point of view. studies, with many responses to that phenom- On the other hand, we analyzed if a comparison enon (Kocsis & Hufnagel, 2011). During the last between old and recent researches was possible. decade, butterfly communities in northeastern Consequently, we analyzed selected vascular Buenos Aires province were analyzed in detail plants, harvestmen (Arachnida, Opiliones), spi- in some sparse localities (Núñez Bustos, 2007, ders (Arachnida, Araneae), Rhopalocera butter- 2008, 2009; Núñez Bustos et al., 2013). However, flies (Insecta, Lepidoptera), and some vertebrates, well-sampled old lepidopteran collections were including birds (Aves), chelonians (Testudines), made by researchers by late 19th and early 20th and mammals (Mammalia). Present paper is centuries (Breyer, 1939b; Hayward, 1973; Canals, based on an exhaustive review of published liter- 2000). In this way, it is possible to made useful ature and our own research. Distributional areas comparisons between old collections and recent in each case were determined from bibliographi- observations. When Hayward’s comprehensive cal data such as catalogues and reviews. Below is catalogue (1973) is compared with recent works a brief review of the bibliography and collections (Canals, 2000; Núñez Bustos, 2007; 2008; 2009; we examined for each group: Núñez Bustos et al., 2013), a significant number Vascular plant distribution. The oldest well- of species were added recently to Buenos Aires referenced collections from La Plata River coast fauna. are those of John Tweedie (between 1825 and Bird distribution. The avifauna of Buenos 1832) and Carlos L. Spegazzini (in 1880 decade). Aires city and neighboring region was stud- Nikolai M. Alboff, Carlos Berg, Lucien Hauman, ied by the late 19th century by several authors Cristóbal Hicken, and many others collected a (d’Orbigny, 1847; Burmeister, 1868; Lynch big amount of specimens in the area by the late Arribálzaga, 1878; Holmberg, 1878, 1898; Sclater 19th and early 20th centuries. Collections and & Hudson, 1888; Narosky & Gallegos, 1992), and works on the provincial flora increased quickly, since then, this coastal zone has been considered and the monumental work “Flora de la provincia as the best known area of Argentina from the or- de Buenos Aires”, directed by Ángel L. Cabrera nithological point of view. Later works, like those (1963-1970) summarized the knowledge of plant of Olrog (1963, 1979) and Narosky & Di Giacomo species distribution in this province up to that (1993) allow good comparisons and recognition date. Therefore this document became the ba- of main distributional patterns. sis for any further study, including the present Chelonia distribution. Knowledge of the contribution. Additional data was achieved from distribution of reptiles and amphibians in the specialized works, especially about ferns (e.g. northeastern sector of Buenos Aires province is Capurro, 1961). still patchy and biased. Among them, it is pos- Arachnida distribution. We included in this sible that the best known group are the turtles, item the clades Opiliones and Araneae. Opiliones probably because they are easy to observe and in Argentina were studied in great detail by Raúl identify. Furthermore, as indicated by Crick & A. Ringuelet. Analyses of Ringuelet´s mono- Sparks (1999), the turtles are very sensitive and graph about Argentinean species of the clade climate-affected taxa, mainly because their nest- (1959) shows that northeastern Buenos Aires ling is climate-driven. Information on geographic province is the area with most previous col- distribution of chelonians was taken from the lection sites, probably due to its proximity to exhaustive papers by Freiberg (1977), Gallardo Buenos Aires and La Plata natural science mu- (1977), and Cabrera (1998). 12 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

information was available during the preparation of the “Flora de la provincial de Buenos Aires” (Cabrera, 1963-1970). More recently, the exhaus- tive efforts of Flavio Moschione (1987, 1992) and Moschione & Klimaitis (1988) resulted in the ad- dition of 17 new records of native plant species to Punta Lara’s flora. Finally, Giudice et al. (2011) collected other three fern species not previously found in that locality. The works commented here were conducted in the same 6.6 hectares of the Punta Lara Natural Reserve. It is surprising that after 100 years of botanical research in this relatively small area, the increase in species rich- ness did not reach the asymptote. The progressive enrichment of the flora in Fig. 3: Southernmost Erythrina crista-galli L. forests this region, as noted by the above mentioned au- in Samborombón Bay. A-B: General view of the trees thors, is due to the conquest of plants from more along the river coast (winter condition). C: Teucrium vesicarium Mill., a typical herb of the La Plata River northern locations. The “bugre”, Lonchocarpus hygrophilous forests. nitidus, is a particular case. In Buenos Aires prov- ince it grows in the lower Paraná Delta, Martín Mammalia distribution. Small mammals, García Island, Avellaneda and Quilmes districts, especially rodent, have been found as valuable and has its southernmost populations in the sources indicative of climate regime (Cameron & Punta Lara Natural Reserve. This legume tree is Scheel, 2001). In Argentina, most studies regard- one of the most important components of Punta ing detailed distribution and taxonomy of small- Lara riparian rainforest (e.g. Cabrera & Dawson, mammals were carried out by the relatively re- 1944; Dascanio & Ricci, 1988) and Martín García cent efforts of Elio Massoia by the 1960 decade. Island forest (Arturi & Juárez, 1997). However, An important contribution is that of Galliari et it is curious that there are no collected specimen al. (1991), which stands as the main compilation from there or any another part of the province of mammalian records of the province up to that up to 1925 (Paraná Delta, 1925, collected by A. C. date. Since then, the studies on micromammal Scala, deposited in LP herbarium). Because the assemblages in Buenos Aires have been greatly species grows in an area that was visited several increased (e.g., Udrizar Sauthier et al., 2005; Teta times by botanists during the end of the 19th and et al., 2010, 2014; Pardiñas et al., 2010), but un- early 20th centuries, it is unlikely that this big fortunately species distribution in Buenos Aires ornamental plant was overlooked in the Punta province is still far from being well-understood. Lara forest. Its distributional shift may have oc- Therefore, we focus our work on middle mam- curred at the end of 19th century, and represents mals, which are well represented in public collec- an example of early distributional expansion. tions, they are usually well known by hunters, and are mentioned in 19th century explorers’ books. Changes in plant communities distribution Leaving aside individual species range ex- RESULTS tensions, an expansion of the entire Paranaense coastal woods is documented. When comparing Changes in vascular plants distribution the southern limit of this plant community as A total of 44 distribution changes in vascular observed by Vervoorst (1967) with today limit, plant species was confirmed (Table 1). we realize this wood advanced more than 25 Many novel species records are here made for kilometers to the south, from Salvador Grande the first time in the Punta Lara gallery rainforest. Point to the northern limit of Samborombón Bay First explorations to this Natural Reserve were (both sites in Parque Costero del Sur Natural made by Nicolás M. Alboff, Carlos L. Spegazzini Reserve, Punta Indio district). In this regard, and Lucien Hauman by the late 19th and early lineal patches of a coastal forests are present in 20th centuries. Later, Ángel L. Cabrera and col- a beach ridge between 35º 30,219’ S – 57º 9,795’ laborators studied extensively this rainforest be- W and 35º 30,447’ S – 57º 10,019’ W (Figures 2 tween 1926 and 1944 (Cabrera & Dawson, 1944). and 3). There, the characteristic tree Erythrina Plant collections in that place did not stop after crista-galli is associated with Acacia caven, that, and late to Cabrera & Dawson’s work, new Sesbania punicea, S. virgata, Senna corymbo- Guerrero & Agnolin: Recent changes in plant and animal distribution 13

Fig. 4: Some sedimentary features of Punta Indio ridges (Parque Costero del Sur). Redrawn from Guerrero (2014a). A: General view of a stratigraphic cut in Erythrina crista-galli forests at Punta Indio. B: Detail of the same stratigraphic cut. Note the plastic buoy taken from the lower part of the strata. C: Recently formed ridge. Arrows indicate plastic waste embedded in the geoform. sa, Celtis ehrenbergiana, Elymus scabrifolius, distributional change was proposed as a climate- Cynodon dactylon, Hibiscus striatus, Hydrocotyle driven phenomenon that produced first geomor- bonariensis, Teucrium vesicarium, Eclipta pros- phological modifications, then physiognomic trata, Tillandsia aëranthos, Passiflora caerulea, changes in the flora, and finaly harvestmen colo- Ipomoea cairica and Araujia sericifera. This asso- nization (Guerrero, 2014a). ciation conforms a Paranaense woodland of very Among spiders, we recognize distribution- recent origin, showing that the forest is actually al changes in Nephila clavipes and Neotrops extending its distribution to southern latitudes. pombero. The first one is a good example of a It is also worth mentioning that several hu- recent southward expansion. This subtropical mid forests within the Parque Costero del Sur spider reached Corrientes province as its south- Natural Reserve may have appeared within the ern limit up to the 2000s, but to present time oc- area during the 20th century, as inferred from curs on almost every humid coastal wood from sedimentary evidence (Guerrero, 2014a). In fact, Paraná Delta to Punta Indio district (Guerrero there is a fine sand stratum of about one meter et al., 2012). Dispersion of Nephila clavipes was thick carrying human solid waste under the soil related to a flood event that swept some spiders where the forest grow up, and in some places a downstream to north Buenos Aires. Neotrops thin mud stratum with plastic fragments under- pombero is a common species in northeastern lies the previous one (Figure 4). On this basis, Argentina and Paraguay (Grismado & Ramírez, it was concluded that the forests obviously grew 2013). Its sudden appearance in Costanera Sur up after these strata have settled down, and the Ecological Reserve in 2011 was attributed to a presence of plastics within the sediment indi- recent dispersal (Grismado & Ramírez, 2013; cates they were deposited during the 20th century Zapata & Grismado, 2015). Many other spider (Guerrero, 2014a). species may have reached the study area in the last decades, but unfortunately lot of research is Changes in Arachnida distribution needed (Zapata & Grismado, 2015). For example, Table 2 summarizes taxa with recent exten- Micrathena furva (Keyserling 1892) was recorded sions on their geographical distributional area. by Levi (1985) up to Punta Lara Natural Reserve, A total of 6 harvestmen (Opiliones) species but nowadays it extends southwards to Parque have increased their distributional range from Costero del Sur Natural Reserve (Arachnological north to south along the La Plata River fluvial collection of La Plata Museum, La Plata, Buenos system: Discocyrtus prospicuus, Discocyrtus Aires, Argentina: MLP_Ar 19407). testudineus, Gryne orensis, Hernandaria scabri- cula, Metalibitia argentina and Varinodulia insu- Changes in lepidopteran distribution laris. The southward expansion of Mesopotamian Southward expansions were detected in but- harvestmen species is strikingly coincident with terfly species from different localities at Buenos the increase of rainfall and temperature in Aires province. 33 taxa were found as having im- Buenos Aires province (Guerrero, 2014a). This portant distributional changes (Table 3). 14 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Morpho epistropheus argentinus, considered actual specimens and was considered as dubious the national butterfly due to its beauty, is a big (Richard & De La Fuente, 1992; Cei, 1993; Alcalde and conspicuous species on its adult and larval et al., 2012). Consequently, Williams (1991) did stages. Farina (2006) followed the increase of its not include the species within Buenos Aires her- range extension across the eastern part of the petofauna, a criterion followed by many authors. province between 1985 and 2003. Up to 1939 the In the last decade this turtle was found in sev- species was known from a few localities of north- eral localities of northern Buenos Aires province: ern Buenos Aires province, its southern limit San Isidro and Campana districts, and Buenos being at Punta Lara (Breyer, 1939a). To early Aires city (Pereira & Haene, 2003; Chebez, 2000s it expanded its range to Punta Piedras 2009; Agnolin et al., 2014), possibly Punta Lara (Punta Indio district) and General Madariaga (Saibene et al., 2012), and in Magdalena, which district (Klimaitis, 2000). Finally, a few years constitutes its current southernmost locality later it was detected in the southern General (Alcalde et al., 2012). Thus, we consider these re- Pueyrredón district (Farina, 2006). As its favor- cords corroborate the hypothesis of a geographi- ite host plant, Scutia buxifolia (Rhamnaceae), cal expansion in this species. exists in the xerophitic woods or “talares” almost Hydromedusa tectifera was recently found since the beginnings of the 20th century (Parodi, in Sierra de La Ventana, 495 kilometers south 1940), Farina (2006) concluded that the range of its previously known geographic distribution expansion of this butterfly could be connected (Di Pietro et al., 2012). We did not include this with the climatic change. Also, Hypanartia bella, record here because it is unclear if it is due to a Doxocopa laurentia laurentia, Phoebis neocypris, recent natural expansion or human introduction. and P. sennae marcelina showed the same pat- However, the new record here reported at Punta tern of southward extension through the “ta- Indio district (Arroyo Salvador Grande, near the lares” (Farina, 2006). bridge of Route 11, 7 January 2009) may be an Another remarkable case is Adelpha syma, evidence of a natural southward expansion of the of which the original distribution encompasses species following La Plata River coast. southeastern Brazil, Uruguay, Paraguay and northeastern Argentina, was not reported Changes in bird distribution in Buenos Aires province prior to the 2000s Several authors considered that some bird (Hayward, 1973; Willmott, 2003). Klimaitis species inhabiting the La Plata River coast ex- (2000) argued the species arrived to the area due panded their southern distribution in recent to passive raft dispersal downstream the Paraná times (Haene, 2006; Chimento et al., 2011; Godoy or Uruguay rivers or by active dispersal along the et al., 2012; Agnolin & Rivero, 2014). Based on Atlantic coast. Nowadays it is a common butter- available data, at least 24 different species of fly in several sites of northeastern Buenos Aires birds can be considered as being more widely dis- province (Núñez Bustos et al., 2013). tributed than previously known (Table 5). Finally, some species increased their abun- All the species included in the table were dance during the last decades. For example, scarce or absent from Buenos Aires province Núñez Bustos et al. (2013) mention that Badecla but they are frequent nowadays and were re- argentinensis, which had scarce past records in ported from many localities. Among them, most Buenos Aires, became much more frequent in re- taxa are related to woodland habitats, both cent decades. aquatic (Anhinga anhinga, Tigrisoma lineatum, Aramides cajanea) and terrestrial (Penelope ob- Changes in chelonian distribution scura, Columba picazuro, Columbina talpacoti, As a result of comparing old works with recent Tapera naevia, Crotophaga ani, Leucochloris papers and observations, we found that three tur- albicollis, Picumnus cirratus, Melanerpes tle species changed their geographical distribu- candidus, M. cactorum, Phacellodomus ru- tion and expanded its geonemy (Table 4). ber, Thamnophilus caerulescens, Hemitriccus The case of the Red eared slider Trachemys margaritaceiventer, Euscarthmus meloryphus, dorbignyi merits some explanatory com- Basileuterus leucoblepharus, Saltatricula mul- ments. This species is frequent in northeastern ticolor, Saltator aurantiirostris, Cyanocompsa Argentina, north to Entre Ríos province (Barco brisonii), whereas a minor component includes & Larriera, 1991; Richard & De La Fuente, taxa of more opened areas and generalist hab- 1992; Cabrera, 1998). Freiberg (1977; probably its (Elaenia spectabilis, Phylloscartes ventralis, based on Marelli, 1924) mentioned its presence Sporophila hypochroma, Sporophila cinna- in Buenos Aires, but this report was not based on momea) (Nores et al., 2005). Guerrero & Agnolin: Recent changes in plant and animal distribution 15

We include below a brief account of some re- ent though very scarce in Punta Lara. Narosky markable species that, according to current re- & Di Giacomo (1993) reported a large amount of cords, habits, and visibility, are known to have localities at northern and eastern Buenos Aires expanded its geographical distribution south- province, and Pugnali and Chamorro (2008) and wards. Zelaya & Pérez (2008) indicated that this species The Darter Anhinga anhinga inhabits fresh- currently is common in Buenos Aires city. Also water environments surrounded by wooded Chimento et al. (2012) reported new records for areas of northeastern Argentina (Olrog, 1963; the Cardinal and said that it probably expanded Narosky & Di Giacomo, 1993). Narosky (1969) its geographical range to the south of the prov- was the first author reporting its presence in ince in recent times. Likewise, the Saltator Buenos Aires; its nestling was later corroborated Saltator similis was first found in the northern by López Lanús & Roda (1987). Since then, it extreme of Buenos Aires province by Fernández was reported from several localities of the prov- (1991). Since then, it was found in other locali- ince and Buenos Aires city. Its southernmost ties of the province, including Punta Lara and record is from the Punta Lara Natural Reserve Atalaya (Bodrati et al., 2001b; Mérida & Bodrati, (Narosky & Di Giacomo, 1993; Babarskas et al., 2006; Bodrati & Sierra, 2009), and Buenos Aires 2003; Pugnali & Chamorro, 2006; Pagano et al., city (Pugnali & Chamorro, 2008). 2012). Previous to the 1980s, Phacellodomus ruber The Guan Penelope obscura is common presence in Buenos Aires province was inferred in wood areas and forests from northeastern from a single skin coming from Avellaneda dis- Argentina (Olrog, 1963; Narosky & Di Giacomo, trict (Narosky &Di Giacomo, 1993). This spe- 1993). Césari & Alonso (1975) cited the species for cies is related to wooded areas and forests from the first time in the northern tip of Buenos Aires northern Argentina north to Santa Fe province province, and since then it was found in several (Olrog, 1963, 1979). However, since 1989 the localities, including the Paraná Delta (Narosky species has become more frequent in different & Di Giacomo, 1993), Otamendi (Babarskas et localities at northeastern Buenos Aires province al., 2003), Punta Indio (F.L. Agnolin pers. obs.), (Chebez & Haene, 1991; Narosky & Di Giacomo, Buenos Aires city (Eguía, 2012), and more recent- 1993; Bodrati et al., 2001a). Bodrati et al. (2001a) ly, maped in Punta Rasa (Narosky & Yzurieta, considerably expanded the knowledge of the spe- 2010). cies in the province, and indicated that it is rela- The Woodpecker Melanerpes cactorum tively frequent in Vuelta de Obligado (San Pedro was first reported for the province in the local- district), where it nests. Similarly, in the early ity of Ramallo (Maugeri & Montenegro, 2002). 20th century Thamnophilus caerulescens had Recently, Chimento et al. (2009) reported for only two records in northeastern Buenos Aires the first time that this species nests in La Plata province (Sclater, 1890; Holmberg, 1898), and city, 250 kilometers to the south of its previously was considered a rare species in the province by known distribution. More recently, the species Narosky & Di Giacomo (1993). More recently, was repeteadly reported for Buenos Aires and it was frequently reported in the studied area, La Plata cities (Chimento et al., 2011; Borsellino, with records from Zárate, San Isidro, Campana, 2012), and Vuelta de Obligado, San Pedro district San Pedro, Baradero and Llavallol districts, and (Chimento et al., 2011). The presence and nestling Buenos Aires city (Narosky, 1983; Barbetti et al., of the species on its southern geographical limit 1985; Babarskas et al., 2003; Bodrati et al., 2001a; was considered by Chimento et al. (2011) as prob- Pugnali & Chamorro, 2008; Lucero et al., 2011). ably driven by climatic changes in the region. Furthermore, in Baradero and Llavallol districts The Green-backed Becard Pachyramphus the species is relatively frequent (Bodrati et al., viridis was observed for the first time in Buenos 2001a). Aires province in 1989 (Narosky & Di Giacomo, Several other species that were considered 1993). Since then, the species expanded its distri- rare, scarce or hypothetical for Buenos Aires bution to several localities of the area (Krapovickas have been increasing its occurrence in the region. et al., 1992; Narosky & Di Giacomo, 1993; Grilli For example, up to late 1980s the Woodcreeper & Areta, 2002; Bodrati, 2001; Bodrati et al., 2006; Lepidocolaptes angustirostris was considered a Areta in Pagano & Mérida, 2009). Similarly, the very rare species at Punta Lara Natural Reserve Cardinal Paroaria capitata was not observed by (Klimaitis & Moschione, 1987), whereas more ancient authors in the province (e.g., Holmberg, recently it is considered “very common” and fre- 1898; Burmeister, 1868; Pereyra, 1938). Klimaitis quent in all environments (Pagano et al., 2012). & Moschione (1987) stated the species was pres- The species is also very common in Buenos 16 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Aires city (F. L. Agnolin pers. obs.), and has ex- In the same line, the new records of the mi- panded its southern and western distribution in cromammals Cryptonanus chacoensis, Deltamys several counties of the province (Haene, 2006; kempi and Oligoryzomys nigripes in more south- Chimento et al., 2012). The same may be said for ern localities than previously known (Udrizar Myiodynastes maculatus, which was a rare resi- Sauthier et al., 2005) may be due to an expan- dent on the northern tip of the province, with sion in their range. Likewise, Holochilus chac- only a few records (Narosky & Di Giacomo, 1993). arius was first recorded in Buenos Aires in 2004, Recent overviews indicate that the species is com- where it was never found previously (Voglino et mon at several localities, including La Plata city al., 2004; Courtalon et al., 2013). Nevertheless (F. L. Agnolin pers. obs.), Llavallol (Lucero et al., these four species were not considered in our 2011), Avellaneda and Quilmes districts (Godoy study because the extension of their southern et al., 2012), and Punta Lara Natural Reserve distributional limits could also be attributed to (Pagano et al., 2012). A similar pattern may be lack of historical collections. recognized for the Trushes Turdus rufiventris and T. amaurochalinus. The fomer was consid- DISCUSSION ered scarce by d’Orbigny (1847) and Withington (1888) in Buenos Aires neighborhood. However, This contribution indicates important chang- since the 1960s the species has been increasing es on the geographical distribution of 115 spe- its abundance in the province (Narosky, 2012). cies belonging to seven major groups of plants Nowadays, this Thrush is probably one of the and animals. Plants, butterflies and birds were most frequent birds in several localities, includ- the groups with the larger number of southward ing Buenos Aires city (Narosky, 2012; Narosky et expansions, with 44, 33, and 24 species respec- al., 1996; Haene, 2006) and expanded its distribu- tively, whereas arachnids, mammals and turtles tion towards the south in Buenos Aires province. were represented by 8, 3 and 3 respectively. In Similarly, T. amaurochalinus was an uncom- addition, an increase in abundance was detected mon species in the province, where it was fre- in a butterfly and some bird taxa. It is worth quent only in some streams of the Paraná Delta noting that several of these species have diver- (Pereyra, 1938). Daguerre (1922) indicated it was gent ecological habits. The plants that have ex- a very rare species at Rosas (Las Flores district). panded their distribution encompass all known It is today distributed along a large area in the kinds of growth represented in riparian forests province, including its southernmost limit, and it and marshes from northeastern Argentina and is very frequent along the coastal line, where this surrounding countries, with the single exception species increased its abundance at least since the of parasitic plants (Table 7). This fact suggests 1990s (Narosky & Di Giacomo, 1993; Roesler, there is no ecological bias in recently geographi- 2001; Maugeri, 2002; Haene, 2006). cally-expanded species. The same may be said for birds, in which insectivorous, frugivorous and pi- Changes in mammals distribution scivorous species have been expanded southward Three mammals species are known to have (De La Peña, 2010). changed their distributional limits in the last de- Almost all taxa expanding their southern cades (Table 6). limits of distribution are inhabitants of South- The occurrence in the last decades of some American subtropical forests and surrounding mid- to large mammals, including Dasypus wetlands, and show biogeographical affinities novemcinctus, Cerdocyon thous, and Procyon with Chaco and Paranaense biogeographical cancrivorus, in the north of Buenos Aires prov- provinces. There are no reports of species from ince was interpreted by some authors as a result the Pampa or Patagonia biogeographical prov- of more humid and temperate conditions in the inces that had increased their distributional area area (Zamorano & Scillato-Yané, 2008; Fracassi from south to north in the last decades. On the et al., 2010). The latter species was reported contrary, some Pampean taxa of Patagonian or by Fracassi et al., (2010) from several localities Central Argentinean affinities disappeared from of the province, including the Reserva Natural several localities and reduced their geographic Otamendi. This protected area was prospected in distribution in the last centuries, mainly due high detail during decades in search of mammals to human disturbance, but also probably due to (Heinonen de Fortabat & Chebez, 1997; Pereira et climatic change (Agnolin & Lucero, 2014; Teta al., 2003). Since these researches were unable to et al., 2014; Agnolin & Rivero, 2014). In this find P. cancrivorus, the arrival of the species in the regard, some taxa, like the rodent Ctenomys ta- area was probably recent (Fracassi et al., 2010). larum and the birds Sturnella loyca and Geositta Guerrero & Agnolin: Recent changes in plant and animal distribution 17 cunicularia, among others, moved towards more and carry lot of individuals and propagules of arid and cold regions (Agnolin & Lucero, 2014; diverse animal and plant species (Achaval et al., Agnolin & Rivero, 2014). 1979; Guerrero et al., 2012; Katinas et al., 2013). With the aim to correlate these changes with This phenomenon has taken place at least since the increase in some climatic values during the the colonial period (see e.g. Robertson, 1916), but lasts centuries, it is necessary to rule out other in most cases, when tropical species arrived to factors that may produce distributional shifts in some species. A main change that occurred Buenos Aires, they failed to colonize new envi- in the area of study during late 19th and the 20th ronments and died due to severe winter tempera- centuries is the raise of intensive agriculture, tures (Spegazzini, 1905; see Hudson commentar- which is clearly not a positive factor for native ies from Narosky & Gallegos, 1992). Otherwise, species (Rapoport, 1996). As stated by Hickling the local increase in temperature and humidity, et al. (2006), the use of landscape negatively af- and changes on hydrological parameters since fects species lacking dispersal abilities to spread the 1970s, may have enhanced and allowed the over these strongly modified and fragmented possibilities of raft dispersal and effective coloni- habitats. zation for many taxa. Human introduction of several exotic tree species in the Pampa grasslands may enable some bird taxa increase their southern limits of CONCLUSIONS geographic distributions (Chimento et al., 2012). Nevertheless, the area here studied had native First, we summarized, analyzed, and dis- woods, like the “talares” (Fig. 2), long before the cussed the distributional changes of plants and first European explorations (Delucchi & Charra, animals detected in recent times in northeast- 2012). ern part of Buenos Aires province, a well-known Probably, colonization of the La Plata River transitional area between forested galleries of coast by the riparian subtropical rainforest is a the La Plata Fluvial System and Pampas grass- climate-driven event (Schnack, 2000). In fact, it lands. One hundred and fifteen species belong- was hypothesized that it occurred when the Little ing to different major groups extended their Ice Age ended by the mid 19th century (Deschamps et al., 2002; Deschamps & Tonni, 2007; Guerrero, geographical ranges towards southerly locations 2014b). Whenever the time of appearance of in northeastern Buenos Aires province by late riparian forest in the studied area, it is prob- 20th and early 21th centuries. Geographical ex- able that this habitat richness steady increased tension was detected in species of each reviewed since the end of 19th century, as noticed early by taxonomical group. The changes of geographical William H. Hudson (Narosky & Gallegos, 1992) distribution limits observed in a large diversity and Carlos L. Spegazzini (1905). Paraphrasing of plant and animal taxa of ecologically diver- the last author, northeastern Buenos Aires prov- gent groups appear to be coeval with some well- ince flora changed from xerophyte to hygrophyte known climatic changes. Buenos Aires riparian between 1881 and 1905. Although more evidence forests as a whole show an incessant increment is wanting, many species of plants and animals probably arrived from northern locations in in species richness and a southward geographi- that period (Guerrero, 2014b). Among the evi- cal expansion that probably represent ecological dence listed in this paper, the tree Lonchocarpus responses to climatic changes. Although we sum- nitidus could be a good example of those shifts. marized distributional changes in well-sampled Most geographical changes listed in the pres- groups, it is highly probable that a greater part ent contribution may have taken place since of the flora and fauna is being also affected by 1970 to the date, when an important increase in the same pattern. temperature and rainfall has been registered in This paper is the first comprehensive analy- northern Buenos Aires province (Berbery et al., sis demonstrating the possibility of an ecologi- 2006; Servicio Meteorológico Nacional, 2015). cal response of a large number of animals and Furthermore, Menéndez (2006) recognized a raise in the discharge of the La Plata Fluvial plants to recent climatic change in Argentina. It System over the last four decades. It is well-doc- is clear that more studies are urgently needed, umented that in almost every raise in this fluvial since most other aspects of the ecological and system level, large masses of floating vegetation evolutionary responses to climate change remain (locally called “camalotales”) are dragged down largely unknown. 18 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Table 1: Selected plant species showing distributional changes. LP: La Plata Museum Herbarium, La Plata, Buenos Aires, Argentina; SI: Darwinion Herbarium, San Isidro, Buenos Aires, Argentina. PCSNR: Parque Costero del Sur Natural Reserve. PLNR: Punta Lara Natural Reserve.

Name Southern limit Source Present southern Source known in the 20th limit century Abutilon pauciflorum A. St.- Northern Buenos Cabrera (1963- Quilmes and Guerrero 186 (LP); Hil. Aires slopes 1970) PLNR Moschione­ & Klimaitis (1988) Adiantopsis chlorophylla (Sw.) Paraná Delta Capurro (1961) PLNR Moschione (1987) Fée Aloysia gratissima (Gillies & Northern Buenos Cabrera (1963- PLNR Moschione & Klimaitis Hook.) Tronc. Aires slopes 1970) (1988) Araujia angustifolia (Hook. & Paraná Delta Cabrera (1963- PLNR and Berisso Moschione & Klimaitis Arn.) Steud. 1970) district (1988); Guerrero 471 (LP) Blepharocalyx salicifolius Berisso district Cabrera (1963- PCSNR Guerrero 19 (LP) (Kunth) O. Berg 1970) Canavalia bonariensis Lindl. Berisso district Cabrera (1963- PCSNR Galup (2009) 1970) Clytostoma callistegioides Punta Lara Cabrera (1963- PCSNR Galup (2009) (Cham.) Baill. Natural Reserve 1970) Combretum fruticosum Entre Ríos pro- Burkart (1969- Martín García Lahitte & Hurrell (Loefl.) Stuntz vince 1987) Island (1994) Dioscorea sinuata Vell. Punta Lara Cabrera (1963- Magdalena district Guerrero 381 (LP) Natural Reserve 1970) Dolichandra cynanchoides Entre Ríos pro- Burkart (1969- Costanera Sur Cabanillas et al. s.n. Cham. vince 1987) Ecological Reserve (LP) Doryopteris concolor (Langsd. Northern Buenos Capurro (1961); PLNR Moschione (1987) & Fisch.) Kuhn Aires slopes and Cabrera (1963- Martín García 1970) Island Doryopteris pedata (L.) Fèe Corrientes pro- De la Sota (1977) Martín García Ramos Giacosa et al. vince Island (2004) Doryopteris pentagona Pic. Corrientes pro- Zuloaga et al. PLNR Giudice et al. (2011) Serm. vince (2008) Enterolobium contortisiliquum Paraná Delta Cabrera (1963- Avellaneda and Guerrero et al. (2012) (Vell.) Morong 1970) Quilmes districts Gomesa bifolia (Sims) M.W. Paraná Delta and Cabrera (1963- PLNR and PCSNR Tonni s.n. (LP); Cellini Chase & N.H. Williams Buenos Aires City 1970); Correa et et al. (2009) al. (2009) Heteropterys glabra Hook. & Paraná Delta and Cabrera (1963- PLNR Moschione & Klimaitis Arn. Buenos Aires City 1970) (1988) Lonchocarpus nitidus (Vogel.) Entre Ríos pro- Lorentz (1878) PLNR Cabrera & Dawson Benth. vince (1944) Mimosa pigra L. var. pigra Paraná Delta Cabrera (1963- Avellaneda and Guerrero et al. (2012) 1970) Quilmes districts Mutisia coccinea A. St.-Hil. Paraná Delta Cabrera (1963- PLNR and Moschione & Klimaitis 1970) Magdalena district (1988); Galup (2009) Myrceugenia glaucescens Berisso district Cabrera (1963- PCSNR Galup (2009) (Cambess.) D. Legrand & 1970) Kausel Osmunda regalis L. Paraná Delta Capurro (1961) PLNR Moschione (1987) Guerrero & Agnolin: Recent changes in plant and animal distribution 19

Name Southern limit Source Present southern Source known in the 20th limit century Oxalis hispidula Zucc. Paraná Delta Cabrera (1963- PLNR Moschione & Klimaitis 1970) (1988) Passiflora morifolia Mast. in Córdoba, Chaco Deginani (2001) Berazategui dis- Cellini 78 (SI) Mart. and Corrientes trict provinces Philodendron tweedieanum Entre Ríos pro- Hurrell & PCSNR Hurrell & Delucchi Schott vince Delucchi (2008) (2008) Phyllanthus sellowianus Punta Lara Cabrera (1963- PCSNR Galup (2009) (Klotzsch.) Müll. Arg. Natural Reserve 1970) Pleopeltis minima (Bory) J. Paraná Delta Capurro (1961); PLNR and La Giudice et al. (2011); Prado & R.Y. Hirai Cabrera (1963- plata district Guerrero 173 ,519 (LP) 1970) Pleopeltis pleopeltifolia Corrientes pro- Zuloaga et al. La Plata and Guerrero 172 (LP); (Raddi) Alston vince (2008) Magdalena district Cellini 22 (SI) Pouteria salicifolia (Spreng.) Berisso district Cabrera (1963- PCSNR Galup (2009) Radlk. 1970) Psilotum nudum (L.) P. Beauv. Corrientes pro- De la Sota (1977) PLNR Moschione (1992) vince Rumohra adiantiformis (G. Martín García Capurro (1961) PLNR Moschione (1987) Forst.) Ching Island Sicyos polyacathos Cogn. Entre Ríos pro- Burkart (1969- Avellaneda and Guerrero et al. (2012) vince 1987) Quilmes districts Solanum sarrachoides Sendtn. Paraná Delta Cabrera (1963- PLNR Moschione & Klimaitis 1970) (1988) Spartina spartinae (Trin.) Santa Fe province Lewis & Pergamino district Lewis & Pire (2006) Merr. Collantes (1975) Tarenaya hassleriana Paraná Delta Cabrera (1963- PLNR Moschione & Klimaitis (Chodat) Iltis 1970) (1988) Terminalia australis Cambess. Berisso district Cabrera (1963- Magdalena district Guerrero 30, 518 (LP) 1970) and PCSNR Tessaria integrifolia Ruiz & Paraná Delta and Hicken (1910); Berisso district Guerrero 83 (LP) Pav. Buenos Aires City Cabrera (1963- 1970) Thelypteris abbiatti C.F. Reed Ensenada district Ponce (1987) Magdalena district Guerrero 382 (LP)

Thelypteris decurtata subsp. Ensenada district Capurro (1961); PCSNR Galup (2009) platense (Weath.) de la Sota Ponce (1987) Thelypteris hispidula (Decne.) Entre Ríos pro- Ponce (1987) Ramallo district Ramos Giacosa et al. Reed vince and PLNR (2004); Giudice et al. (2011) Thelypteris rivularioides (Fee) Entre Ríos pro- Ponce (1987) PLNR Moschione (1987) Abbiatti vince Thelypteris totta (Thunb.) Paraná Delta Capurro (1961); PLNR Ponce (1987); Schelpe Cabrera (1963- Moschione (1987) 1970) Tillandsia myosura Griseb. Colonia Zuloaga et al. PLNR Cellini et al. (2012) ex Baker Department, (2008) Uruguay Vicia macrograminea Burkart Escobar district Cabrera (1963- PLNR Moschione & Klimaitis 1970) (1988) Vigna adenantha (G. Mey.) Paraná Delta and Cabrera (1963- PLNR and Berisso Moschione & Klimaitis Maréchal, Mascherpa & Buenos Aires City 1970) district (1988); Buet 233 (LP) Stainier 20 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Table 2: Selected arachnid species showing distributional changes. PCSNR: Parque Costero del Sur Natural Reserve. PLNR: Punta Lara Natural Reserve.

Name Southern limit Source Present southern Source known in the 20th limit century Discocyrtus prospicuus Berisso district Ringuelet (1959) PCSNR and the Acosta & Guerrero (Holmberg, 1876) Atlantic seacoast of (2011) the province Discocyrtus testudineus Paraná Delta Ringuelet (1959) Quilmes and Guerrero et al. (Holmberg, 1876) Berazategui districts (2012); Guerrero (2014a) Gryne orensis (Sørensen, Santa Fe province Ringuelet (1959) San Pedro and San Guerrero (2012); 1879) Nicolás districts Acosta & Vergara (2013) Hernandaria scabricula Berisso district Ringuelet (1959) PCSNR Guerrero (2014a) Sørensen, 1884 Metalibitia argentina San Nicolás dis- Ringuelet (1959) PLNR Guerrero (2011) (Sørensen, 1884) trict Varinodulia insularis Canals, Punta Lara Ringuelet (1959) Berisso district Guerrero (2014a) 1935 Natural Reserve Nephila clavipes (Linnaeus, Corrientes pro- Levi (1980) PCSNR Guerrero et al. 1767) vince (2012) Neotrops pombero Grismado Corrientes pro- Grismado & Costanera Sur Grismado & & Ramírez, 2013 vince Ramírez (2013) Ecological Reserve Ramírez (2013)

Table 3: Selected butterfly species showing distributional changes. PCSNR: Parque Costero del Sur Natural Reserve. PLNR: Punta Lara Natural Reserve. Name Southern limit Source Present southern limit Source known in the 20th century Adelpha syma (Godart,1884) Misiones province Hayward (1973) PLNR Núñez Bustos et al. (2013) Adelpha zea (Hewitson, 1850) Misiones province Hayward (1973) Paraná Delta and Canals (2000); Costanera Sur Núñez Bustos Ecological Reserve (2008) Anarthria jatrophae jatrophae Entre Ríos and Hayward (1973) PLNR Núñez Bustos et (Linnaeus, 1763) Santa Fe provinces al. (2013) Antigonus liborius areta Misiones province Hayward (1973) PLNR Núñez Bustos et Evans, 1953 al. (2013) Cymaenes gisca Evans, 1955 Northeaster Hayward (1973) PLNR Núñez Bustos et Argentina al. (2013) Cymaenes tripunctata Entre Ríos pro- Hayward (1973) Berisso district Canals (2000) (Latreille, 1823) vince Diaethria candrena candrena Berisso district Klimaitis (2000) PCSNR Núñez Bustos (Godart, 1824) (2009) Doxocopa kallina (Godart, Corrientes pro- Hayward (1973) PLNR Núñez Bustos et 1824) vince al. (2013) Eantis thraso (Hübner, 1807) Corrientes and Hayward (1973) PLNR Núñez Bustos et Santa Fe provinces al. (2013) Emesis lupina melancholica South Brasil and Núñez Bustos PLNR Núñez Bustos et Stichel, 1910 Uruguay (2007) al. (2013) Guerrero & Agnolin: Recent changes in plant and animal distribution 21

Name Southern limit Source Present southern limit Source known in the 20th century Enantia lina psamathe Paraná Delta and Hayward (1973); PLNR Núñez Bustos et (Fabricius, 1793) Martín García Canals (2000) al. (2013) Island Hamadryas februa februa Martín García Canals (2000) PLNR Núñez Bustos et (Linnaeus, 1823) Island al. (2013) Heraclides astyalus (Godart, Entre Ríos and Hayward (1973) PLNR Núñez Bustos et 1819) Santa Fe provinces al. (2013) Hermeuptychia hermes Entre Ríos pro- Hayward (1973) PLNR Núñez Bustos et (Fabricius, 1775) vince al. (2013) Hypanartia bella (Fabricius, Córdoba and Hayward (1973) PCSNR and Atlantic Núñez Bustos 1793) Corrientes pro- seacoast (2009); Farina vinces (2006) Lycorea ilione ilione (Cramer, Corrientes pro- Hayward (1973) PLNR Núñez Bustos et 1775) vince al. (2013) Marpesia chiron marius Corrientes pro- Hayward (1973) Costanera Sur Núñez Bustos (Cramer, 1779) vince Ecological Reserve (2008) Mechanitis lysimnia lysimnia Corrientes pro- Hayward (1973) PCSNR Núñez Bustos (Fabricius, 1793) vince (2009) Ministrymon una (Hewitson, Misiones and Hayward (1973) Paraná Delta and Núñez Bustos 1873) Formosa provinces Martín García Island (2007) Monca telata penda (Evans, Santa Fe province Hayward (1973) Martín García Island Núñez Bustos 1955) (2007) Morpho epistrophus argentin- PLNR Breyer (1939a) PCSNR and Atlantic Núñez Bustos us Frühstorfer, 1907 seacoast (2009); Farina (2006) Ortilia ithra (Kirby, 1900) Entre Ríos and Hayward (1973) PCSNR Núñez Bustos Santa Fe provinces (2009) Paryphtimoides poltys Santa Fe province Hayward (1973) PLNR Núñez Bustos et (Prittwitz, 1865) al. (2013) Paryphtimoides zeredatha Corrientes and Hayward (1973) Martín García Island Núñez Bustos (Butler, 1869) Santa Fe provinces (2007) Phoebis neocypris neocypris Campana district Canals (2000) PCSNR and Atlantic Núñez Bustos (Hübner, 1823) and Martín García seacoast (2009); Farina Island (2006) Phoebis sennae marcelina PLNR Canals (2000) PCSNR and Atlantic Núñez Bustos (Cramer, 1777) seacoast (2009); Farina (2006) Polites vivex catalina (Plötz, Entre Ríos pro- Hayward (1973) PLNR Núñez Bustos et 1886) vince al. (2013) Priamides anchisiades capys Entre Ríos and Hayward (1973) Berisso district Canals (2000) (Hübner, 1809) Santa Fe provinces Pyristia nise tenella Córdoba province Hayward (1973) PLNR Núñez bustos et (Boisduval, 1836) al., (2013) Rekoa malina (Hewitson, Misiones province Hayward (1973) Magdalena district Canals (2000) 1869) Siproeta epaphus trayja Entre Ríos and Hayward (1973) PLNR Núñez Bustos et (Hübner, 1823) Santa Fe provinces al. (2013) Strymon canitus (H.H.Druce, Corrientes pro- Hayward (1973) PLNR Canals (2000) 1907) vince Urbanus zagorus (Plötz, 1880) Corrientes and Hayward (1973) Costanera Sur Núñez Bustos Córdoba provinces Ecological Reserve (2008) 22 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Table 4: Selected turtle species showing distributional changes. PCSNR: Parque Costero del Sur Natural Reserve.

Name Southern limit known Source Present southern Source in the 20th century limit Hydromedusa tectifera Magdalena district Cabrera (1998) PCSNR E. L. Guerrero, Cope, 1869 pers. obs. Phrynops hilarii (Duméril Zelaya and Zárate Williams (1991); Cabrera Magdalena dis- Derocco et al. & Bibron, 1835) districts (1998) trict (2005) Trachemys dorbignyi Entre Ríos province Barco & Larriera (1991); Magdalena dis- Alcalde et al. (Duméril & Bibron, 1835) Richard & De La Fuente trict (2012) (1992); Cabrera (1998)

Table 5: Selected bird species showing distributional changes. PCSNR: Parque Costero del Sur Natural Reserve. PLNR: Punta Lara Natural Reserve.

Name Southern limit Source Present southern Source known in the 20th limit century Anhinga anhinga Entre Ríos and Olrog (1963) Berisso district Narosky (1969); (Linnaeus, 1766) Santa Fe provinces López Lanús & Roda (1987) Aramides cajanea Statius- Martín García Dabbene (1917); PCSNR Pagano & Merida Muller, 1776 Island Pereyra (1938) (2009) Basileuterus leucoblepha- Entre Ríos and Olrog (1963, 1979) Quilmes district Montaldo et al. rus (Vieillot, 1817) Santa Fe provinces and PLNR (1987); Godoy et al. (2012) Columbina talpacoti North Buenos Hartert & Venturi PLNR Bodrati et al. (2006); Temminck, 1810 Aires, i.e. San (1909); Pereyra Pagano et al. (2012) Pedro and Ramallo (1938); Narosky districts (1983); Moschione­ et al. (1992); Narosky & Di Giacomo (1993); De La Peña (2013) Crotophaga ani Avellaneda dis- Hartert & Venturi PCSNR Pagano & Merida (Linnaeus, 1758) tricts (1909); Olrog (1979) (2009) Cyanocompsa brisso- Santa Fe province Holmberg (1898); PCSNR Pagano & Mérida, nii (Lichtenstein, 1823) and Paraná Delta Olrog (1963, 1979) (2009) Elaenia spectabilis Northern Santa Fe Olrog (1963, 1979) PCSNR Pagano & Merida Pelzeln, 1868 province (2009) Euscarthmus meloryphus Entre Ríos pro- Olrog (1979) PLNR Povedano & Wied, 1831 vince Moschione (1989); Pagano et al. (2012) Hemitriccus mar- Northern Entre Olrog (1979) PCSNR Pagano & Merida garitaceiventer Ríos and Santa Fe (2009) (d’Orbigny & Lafresnaye, provinces 1837) Leucochloris albicollis Pilar districts Pereyra (1965) PCSNR and Narosky (1983); (Vieillot, 1818) Atlantic seacoast Montaldo (1984); Narosky & Di Giaco­ mo (1993); Pagano & Merida (2009) Melanerpes cactorum Santa Fe and De La Peña (2006, La Plata district Chimento et al. (2009, (d’Orbigny, 1840) Entre Ríos pro- 2013) 2011); Borsellino vinces (2012) Guerrero & Agnolin: Recent changes in plant and animal distribution 23

Name Southern limit Source Present southern Source known in the 20th limit century Melanerpes candidus North Buenos Hudson (1870); Lynch PCSNR Pagano & Merida (Otto, 1796) Aires province Arribálzaga (1878) (2009) Pachyramphus viridis Santa Fe and Olrog (1979) Magdalena dis- Pagano & Merida (Vieillot, 1816) Corrientes pro- trict (2009) vinces Paroaria capitata Scarce or absent Holmberg (1898); PCSNR and Narosky & Di Giaco­ (d’Orbigny & Lafresnaye, in Buenos Aires Burmeister (1868); Atlantic seacoast mo (1993); Pagano 1837) province at the Hudson (1870); & Merida (2009); beginnings of the Pereyra (1938) Chimento et al. (2012 20th century Penelope obscura Corrientes and Olrog (1963) PCSNR and La F.L. Agnolin pers. Temminck, 1815 Santa Fe provinces Costa district obs.; Narosky & Yzurieta (2010) Phacellodomus ruber Santa Fe province. Olrog (1963, 1979) La Plata district Chebez & Haene (Vieillot, 1817) Accidental in (1991); Narosky & Avellaneda district Di Giacomo (1993); (see text) Bodrati et al. (2005); Pagano et al. (2012) Phylloscartes ventralis Entre Ríos pro- Olrog (1979) PLNR Klimaitis & (Temminck, 1824) vince Moschione (1987); Pagano et al. (2012) Picumnus cirratus Entre Ríos pro- Olrog (1963) Buenos Aires city Narosky & Di Temminck, 1825 vince Giacomo (1993) Saltator similis d’Orbigny Entre Ríos and Olrog (1979) Magdalena dis- Bodrati et al. (2001b) & Lafresnaye, 1837 Santa Fe provinces trict Sporophila cinnamomea Entre Ríos pro- Narosky (1973); Olrog Buenos Aires City Bodrati et al. (1997a); (Lafresnaye, 1839) vince (1979) Pugnali & Chamorro (2006) Sporophila hypochroma Entre Ríos pro- Olrog (1979); Wege & Buenos Aires City Bodrati et al. (1997b); Todd, 1915 vince Long (1995) Pugnali & Chamorro (2006) Tapera naevia (Linnaeus, Paraná Delta Holmberg (1898) PCSNR Pagano & Merida 1766) (2009) Thamnophilus caerules- North Buenos Sclater (1890); Llavallol district Lucero et al. (2011) cens (Vieillot, 1816) Aires province Holmberg 1898 Tigrisoma lineatum Avellaneda, Zotta & Da Fonseca PCSNR Pagano & Merida (Boddaert, 1783) Campana and (1936); Pereyra (2009) Escobar districts (1938); Eisenmann (1965)

Table 6: Selected mammal species showing distributional changes.

Name Southern limit Source Present southern Source known in the 20th limit century Cerdocyon thous (Linnaeus, Entre Ríos pro- Díaz & Lucherini Paraná Delta Fracassi et al. 1766) vince (2006) (Buenos Aires (2010) province part) Dasypus novemcinctus Entre Ríos pro- Abba & Vizcaíno Campana and Fracassi et al. Linnaeus, 1758 vince (2008) Zárate districts (2010) Procyon cancrivorus (Cuvier, Entre Ríos pro- Díaz & Lucherini Campana and Fracassi et al. 1798) vince (2006) Escobar districts (2010) 24 Revista del Museo Argentino de Ciencias Naturales, n. s. 18(1), 2016

Table 7: Plant species that expanded their distribution to northern Buenos Aires province and their growth habits. Name Growth habit Name Growth habit Abutilon pauciflorum Bush Passiflora morifolia Climber Adiantopsis chlorophylla Herb Philodendron tweedieanum Herb Aloysia gratissima Bush Phyllanthus sellowianus Bush Araujia angustifolia Climber Pleopeltis minima Epiphyte Blepharocalyx salicifolius Tree Pleopeltis pleopeltifolia Epiphyte Canavalia bonariensis Climber Pouteria salicifolia Tree Clytostoma callistegioides Climber Psilotum nudum Herb Combretum fruticosum Climber Rumohra adiantiformis Herb Dioscorea sinuata Climber Sicyos poliacanthos Climber Dolichandra cynanchoides Climber Solanum sarrachoides Herb Doryopteris concolor Herb Spartina spartinae Herb Doryopteris pedata Herb Tarenaya hassleriana Bush Doryopteris pentagona Herb Terminalia australis Tree Enterolobium contortisili- Tree Tessaria integrifolia Tree quum Gomesa bifolia Epiphyte Thelypteris abbiatti Herb Heteropterys glabra Bush Thelypteris decurtata subsp. pla- Herb tense Lonchocarpus nitidus Tree Thelypteris hispidula Herb Mimosa pigra var. pigra Bush or small Thelypteris rivularioides Herb tree Mutisia coccinea Climber Thelypteris totta Herb Myrceugenia glaucescens Bush or small Tillandsia myosura Epiphyte tree Osmunda regalis Herb Vicia macrograminea Climber Oxalis hispidula Herb Vigna adenantha Climber

ACKNOWLEDGEMENTS BIBLIOGRAPHY

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Recibido: 11-X-2015 Aceptado: 10-V-2016