Diversity and conservation status of Aegla spp. (Anomura, Aeglidae): an update

Abstract

The speciose genus Aegla Leach, 1820 is the only valid extant genus of the anomuran family Aeglidae, bearing 83 known species. This diversity may be even greater since there is some evidence for cryptic speciation. The genus is endemic to southern South America, occurring in freshwater habitats. We assessed the conservation status of 82 species of Aegla and found almost 70% of them under some level of threat, which represents a concerning proportion. Major threats to the group include freshwater pollution with urban, agricultural and industrial effluents, habitat modification and fragmentation, riparian forest removal, among others. Conservation measures are required to mitigate the major threats to freshwater ecosystems along rivers where the group occurs.

Key words
Anomuran crabs; Decapoda; freshwater crabs; South American creeks

Introduction

Taxonomically, the family Aeglidae was formerly included within the anomuran superfamily Galatheoidea, but morphological and molecular evidence ( Martin and Abele, 1986; Tudge and Scheltinga, 2002; Pérez-Losada et al., 2002a; Ahyong and O’Meally, 2004) questioned this position. A reappraisal by McLaughlin et al. (2007) elevated Aeglidae to superfamily rank (Aegloidea).

Aegla Leach, 1820 is the only extant genus of the family Aeglidae and can be found in rivers, streams, and lakes of southern South America, occurring from the Rio Grande basin, on the border of São Paulo and Minas Gerais states, Brazil ( Bueno et al., 2007), to Duque de York Island, South River basin, Chile ( Oyanedel et al., 2011). This genus includes 83 described species ( Bond-Buckup et al., 2008; 2010a; 2010b; Santos et al., 2012; 2013; 2015; Moraes et al., 2016) and probably new species will be uncovered in the next years ( Fig. 1).

Figure 1
Cumulative number of know species of the genus Aegla Leach, 1820, since 1900.

The use of molecular techniques together with traditional systematics has helped to confirm several new species, as the number of morphological characters for taxonomic use is limited by the conservative morphotype of the group ( Bond-Buckup and Buckup, 1994; Pérez-Losada et al., 2004; Santos et al., 2009; 2010; 2012; 2013). Moreover, the existence of cryptic species cannot be ruled out since molecular markers and geometric morphometrics have already pointed to this possibility ( Bartholomei-Santos et al., 2011; Marchiori et al., 2014; 2015). This level of diversity is greater than that observed for crayfishes of the family Parastacidae ( Almerão et al., 2015) and brachyuran crabs of the genus Trichodactylus Latreille, 1828 ( Yeo et al., 2006), other crustacean groups inhabiting southern South American continental waters. Bond-Buckup et al. (2008) estimated that 36.5% of the 63 species described at that time were under threat of extinction. Pérez-Losada et al. (2009) assessed the conservation status of 66 aeglid species and found that 32% of them were threatened, that is, species falling within the International Union for Nature Conservation (IUCN) categories Vulnerable (VU), Endangered (EN), and Critically Endangered (CR). Many factors contribute to threat the group along its distribution in southern South America, such as decline in habitat quality, water contamination by extensive use of pesticides in agriculture or by urban sewage, silvicultural practices, construction of hydroelectric plants, among others ( Pérez-Losada et al., 2002b; Bond-Buckup et al., 2008; Baumart and Santos, 2010; Santos et al., 2012). The presence of some species in restrict areas, as in headwaters, several of them with a few records of occurrence, highlights even more the level of threat to the group’s conservation ( Magris et al., 2010).

In this study, we present data on the diversity of species of the genus Aegla, as well as updated information on the distribution of these species in different South American hydrographic basins. Moreover, based on data from several scientific collections and from years of work and experience of the authors, we evaluated the conservation status of the aeglid species and discussed the main threats to the genus diversity.

Material and Methods

We assessed and updated the distribution of 82 species of Aegla. Based on the updated distribution data, we revised the conservation status for each species previously assessed until 2016, according to the rules established by the International Union for Conservation of Nature ( IUCN, 2012). There are five quantitative criteria (A - E) to be applied in order to evaluate if a taxon is under threat, and in which category the taxon should be included (EX - extinct; EW - extinct in wild; CR - critically endangered; EN - endangered; VU - vulnerable; NT - near threatened; LC - least concern; DD - deficient data; and NE - not evaluated; CR, EN and VU represent threat categories). The assessment employs sub-criteria to justify the assignment of a taxon to a certain category. Each species was evaluated in relation to its Extent Of Occurrence (EOO; sub-criterion B1) and Area Of Occupancy (AOO; sub-criterion B2), using data from literature ( Bond-Buckup and Buckup, 1994; Jara and Palacios, 1999; Jara et al., 2003; Galves et al., 2007; Santos et al., 2009; 2010; 2012; 2013; 2014; 2015; Bond-Buckup et al., 2010a; 2010b; César and Damborenea, 2010; Oyanedel et al., 2011; Rocha and Bueno, 2011; Boos et al., 2012; Satterlee et al., 2012; Giri and Collins, 2014; Moraes et al., 2016) and from sampling records in scientific collections (Universidade Federal do Rio Grande do Sul, Universidade Federal de Santa Maria, Pontifícia Universidade Católica do Rio Grande do Sul, Fundação Zoobotânica do Rio Grande do Sul, Universidade Regional do Alto Uruguai e das Missões - Campus Erechim, Universidad Austral de Chile). We determined the AOO for species with less than four records, using an area measure unit (each pixel corresponds to an area of 1 km²). In this case, AOO ranged from 1 to 3 km². For species with four or more records, we calculated the EOO. We assessed data for land-use classification for the whole occurrence area of Aeglidae from the Harmonized World Soil Database ( http://webarchive.iiasa.ac.at/Research/LUC/External-World-soil-database/HTML/), from the World Land Use - Land cover from Food and Agriculture Organization of the United Nations (FAO) ( http://www.fao.org/geonetwork/srv/en/metadata.show?id=12749&currTab=simple), as well as from SOS Mata Atlântica and Instituto Nacional de Pesquisas Espaciais, 2011 ( http://mapas.sosma.org.br).

Results

The list of species, with author(s) and year of description, as well as their distributions, are presented in Tab. 1. Records indicate the presence of aeglids in rivers, lakes and streams in catchments in Argentina (14 species), Bolivia (1 species), Brazil (52 species), Chile (22 species, considering Aegla intermedia Girard, 1855), Paraguay (1 species) and Uruguay (4 species), some species occurring in more than one country ( Figs. 2, 3).

Table 1
Known species of the genus Aegla Leach, 1820, distribution, previous and current conservation status assessments according to IUCN (2012) Red List categories (CR - critically endangered; EN - endangered; VU - vulnerable; NT - near threatened; LC - least concern; DD - deficient data; EX - extinct and NE - not evaluated). The numbers in parentheses, after the name and author of the species, indicate their location in the maps of figures 2 and 3.

Species	Country: Hydrographic basin (main rivers)	Previous assessment (Reference)	Current Status: Criterion
Aegla abtao Schmitt, 1942 (1)	Chile: Toltén River basin (Colico, Allipen, Toltén, Donguil, Pedregoso rivers, Caburgua, Colico lakes); Valdivia-Cruces River basin (Cruces, Pirén rivers); Valdivia-Calle Calle River basin (Huanehue river); Queule River basin (Queule River); Bueno River basin (Curileufu, Ralitrán, Huilma, Pilmaiquen, Negro, Rahue rivers, Mihue Lake); Maullín River basin (Maullín River, Llanquihue, Todos Los Santos lakes); Chamiza River basin (Chamiza River), Puelo River basin (Puelo River), Chiloé Island (San Juan, Huicha, Butalcura rivers, Huillinco Lake).	LC (Pérez-Losada et al., 2009)	LC
Aegla affinis Schmitt, 1942 (2)	Argentina: Colorado River basin (Colorado, Barrancas, Chico rivers); Desaguadero River basin (Atuel, Tunuyan, Chico, Barrancas, Malargüe rivers, La Matancilla, El Salto, Claro creeks). Chile: Maule River basin (Maule Lake, Maule River).	LC (Pérez-Losada et al., 2009)	CR: B1ab(iii)+2ab(iii)
Aegla alacalufi Jara & López, 1981 (3)	Chile: Reloncaví River basin (Reloncaví Fiord); Petrohué-Puelo River basin (Cochamó River); Huequi River basin (Huequi River); Chiloé Island (Puchagrán, Chadmo rivers, Huilinque, Tarahuín lakes); Palena River basin (Palena River); Yelcho River basin (El Amarillo River, Yelcho Lake); Madre di Dios Island; Duke of York Island (North, South, Pollux, Thompson rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla araucaniensis Jara, 1980 (4)	Chile: Valdivia River basin (Leufucade, Calle Calle rivers, Challahuin, Chesque, Coihueco, Quilquil, Quilen, Puquiñe, Huillilelfun, Puente Negro, Ñancul creeks, Riñihue lake); Bueno River basin (Curileufu, Puquitre, Coinco, Lumaco, Sagllue, Pichimaule rivers, Cañal creeks); Petrohué River basin (Cayutue River, Todos los Santos Lake); Maullin River basin (Lahuen Ñadi, Machete creeks); Chilóe Island (Caulín, Huicha, Gamboa creeks, Tarahuin Lake).	LC (Pérez-Losada et al., 2009)	LC
Aegla bahamondei Jara, 1982 (5)	Chile: Bío Bío River basin (Rafael River); Lebú-Paicavi River basin (Caramávida-Tucapel, Pingueral, Caramávida, Cayucupil, Butamalal rivers).	VU: D2 (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla brevipalma Bond-Buckup & Santos 2012 (6)	Brazil: Uruguay River basin (Matador River).	CR: B2ab(iii) (Santos et al., 2012)	CR: B2ab(iii, iv)
Aegla camargoi Buckup & Rossi, 1977 (7)	Brazil: Uruguay River basin (Apuaê-Inhandava, Pelotas rivers).	EN: B1ab(iii,iv) (Brasil, 2014)	EN: B1ab(iii, iv)
Aegla carinata Bond-Buckup & Gonçalves, 2014 (8)	Uruguay: Uruguay River basin (Cuñapiru Creek).	NE	CR: B2ab(iii, iv)
Aegla castro Schmitt, 1942 (9)	Brazil: Paraná River basin (Upper Paranapanema,Tibagi, Ivai Rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla cavernicola Türkay, 1972 (10)	Brazil: Southeastern Atlantic system (Ribeira do Iguape River basin, Areias Grot and Areias de Baixo Cave).	CR: B2ab(iii,v) (Maia et al., 2013)	CR: B2ab(iii, v)
Aegla cholchol Jara & Palacios, 1999 (11)	Chile: Imperial River basin (Pichilumaco, Pichi-Cautín, Traiguén, Colpí, Quillen, Chol Chol, Boroa, Cautin, Quepe rivers); Toltén River basin (Donguil River).	VU: A2ae (Pérez-Losada et al., 2009)	VU: B1ab(iii)+2ab(iii)
Aegla concepcionensis Schmitt, 1942 (12)	Chile: Bío Bío River basin (Manantiales, Arabian Stadium creeks); Andalién River basin (unnamed creek at Villa Vergara, Nonguén River, Pineda Lake); Itata River basin (creek in Cerro Cayumanqui).	CR: A2ae+B1ab(i,iii,v) (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla denticulata denticulata Nicolet, 1849 (13)	Chile: Bío Bío River basin (Malleco River); Budi River basin (Budi Lake); Toltén River basin (Mahuindanche, Donguil, Huiscapi, Curileufu rivers); Lingue River basin (Lingue River); Valdivia River basin (unnamed creek at Villa Calafquen, Calafquen Lake); Bueno River basin (Bueno, Negro rivers); Maullín River basin (Llanquihue Lake); Chiloé Island (Caulín, Huicha creeks, creeks affluent to Huilllinco Lake, Tarahuin Lake).	LC (Pérez-Losada et al., 2009)	LC
Aegla denticulata lacustris Jara, 1989 (14)	Chile: Bueno River basin (Rupanco Lake).	NT (Pérez-Losada et al., 2009)	CR: B1ab(iii)+2ab(iii)
Aegla expansa Jara, 1992 (15)	Chile: Bío Bío River basin (Hualqui River).	EX (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla franca Schmitt, 1942 (16)	Brazil: Paraná River basin (Grande River).	VU (Bueno et al., 2007)	CR: B2ab(iii)
Aegla franciscana Buckup & Rossi, 1977 (17)	Brazil: Uruguay River basin (Apuaê-Inhandava, Pelotas rivers); South Atlantic system (Caí, Sinos, Taquari-Tainhas, Tramandaí, Mampituba rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla georginae Santos & Jara, 2013 (18)	Brazil: Uruguay River basin (Perau Creek).	EN: B1ab(iii) (Santos et al., 2013)	CR: B2ab (iii)
Aegla grisella Bond-Buckup & Buckup, 1994 (19)	Brazil: South Atlantic system (Upper Jacuí, Taquari-Tainhas, Ijuí, Passo Fundo rivers).	VU: B1ab(iii,iv) (Brasil, 2014)	VU: B1ab(iii, iv)
Aegla hueicollensis Jara & Palacios, 1999 (20)	Chile: Valdivia River basin (Futa, Chaihuín, Colún, Hueicolla rivers).	VU: B1ab(iii,iv) (Pérez-Losada et al., 2009)	NT
Aegla humahuaca Schmitt, 1942 (21)	Argentina: Paraná River basin (Bermejo, Dulce, Juramento, Grande rivers).	LC (Pérez-Losada et al., 2009)	VU: B1ab(iii, iv)
Aegla inconspicua Bond-Buckup & Buckup, 1994 (22)	Brazil: South Atlantic system (Guaíba, Caí, Taquari-Antas, Sinos, Gravataí, Tramandaí rivers).	VU: B1ab(iii,iv) (Brasil, 2014)	VU: B1ab(iii, iv)
Aegla inermis Bond-Buckup & Buckup, 1994 (23)	Brazil: South Atlantic system (Sinos, Caí, Tramandaí rivers).	EN: B1ab (iii, iv) (Brasil, 2014)	CR: B1ab(iii, iv)
Aegla intercalata Bond-Buckup & Buckup, 1994 (24)	Argentina: Mar Chiquita system (Las Lajas River); Dulce River basin (Los Sojas River tributaries); Valle Central River basin (Las Trancas River).	LC (Pérez-Losada et al., 2009)	VU: B1ab(iii, iv)
Aegla intermedia Girard, 1855	Chile: Maipu River basin (Maipu River; record not confirmed after description).	NE	NE
Aegla itacolomiensis Bond-Buckup & Buckup, 1994 (25)	Brazil: South Atlantic system (Gravataí, Sinos rivers).	VU: B1ab(iii,iv) (Brasil, 2014)	VU: B1ab(iii, iv)
Aegla japi Moraes, Tavares & Bueno, 2016 (80)	Brazil: Paraná River basin (Tietê River).	VU: B2aD2 (Moraes et al., 2016)	VU: B2aD2
Aegla jaragua Moraes, Tavares & Bueno, 2016 (81)	Brazil: Paraná River basin (Tietê River).	CR: A4eB2a (Moraes et al., 2016)	VU: B2aD2
Aegla jarai Bond-Buckup & Buckup, 1994 (26)	Brazil: Uruguay River basin (Canoas, Pelotas rivers); South Atlantic system (Itajaí-Açu River).	LC (Pérez-Losada et al., 2009)	LC
Aegla jujuyana Schmitt, 1942 (27)	Argentina: Paraná River basin (Chico, Grande, Huasamayo rivers, Paco Creek).	LC (Pérez-Losada et al., 2009)	LC
Aegla jundiai Moraes, Tavares & Bueno, 2016 (82)	Brazil: Paraná River basin (Tietê River).	VU: B2aD2 (Moraes et al., 2016)	VU: B2aD2
Aegla laevis (Latreille, 1818) (28)	Chile: Maipo River basin (Maipo River); Maule River basin (Maule, Putagán rivers).	CR: A2ae (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla lancinhas Bond-Buckup & Buckup, 2015 (30)	Brazil: Southeastern Atlantic system (Upper Ribeira do Iguape River sub basin, Lancinhas Grot).	EN: B1B2ab(iii) (Santos et al., 2015)	EN: B2ab(iii)
Aegla lata Bond-Buckup & Buckup, 1994 (31)	Brazil: Paraná River basin (Tibagi River, Apertados Creek).	EX (Pérez-Losada et al., 2009)	CR: B1ab (i, iii, iv)
Aegla leachi Bond-Buckup & Santos 2012 (32)	Brazil: Uruguay River basin (Galafre River, Marombas River tributaries, Passo Fundo Creek).	VU: B1ab(iii) (Santos et al., 2012)	VU: B1ab(iii, iv)
Aegla leptochela Bond-Buckup & Buckup, 1994 (33)	Brazil: Southeastern Atlantic system (Ribeira do Iguape River sub basin, Paiva’s Grot).	CR: B2ab (iii,v) (Maia et al., 2013)	CR: B2ab (iii, v)
Aegla leptodactyla Buckup & Rossi, 1977 (34)	Brazil: Uruguay River basin (Silveira River); South Atlantic system (Taquari-Antas River).	EN: B1ab(iii,iv) (Brasil, 2014)	EN: B1ab(iii, iv)
Aegla ligulata Bond-Buckup & Buckup, 1994 (35)	Brazil: South Atlantic system (Taquari-Tainhas, Tramandaí rivers).	EN: B1ab(iii,iv) (Brasil, 2014)	EN: B1ab(iii, iv)
Aegla longirostri Bond-Buckup & Buckup, 1994 (36)	Brazil: Uruguay River basin (Ibicuí River); South Atlantic system (Vacacaí-Mirim, Jacuí, Pardo, Taquari-Tainhas, Antas, Caí, Sinos rivers).	LC (Pérez-Losada et al., 2009)	DD
Aegla loyolai Bond-Buckup & Santos 2015 (37)	Brazil: Paraná River basin (Pajanduvas River).	EN: B1B2ab(iii) (Santos et al., 2015)	EN: B2ab(iii)
Aegla ludwigi Santos & Jara, 2013 (38)	Brazil: Uruguay River basin (Cambará Creek, branch of Potiribu River).	CR: B1ab(iii) (Santos et al., 2013)	CR: B2ab (iii)
Aegla manni Jara, 1980 (39)	Chile: Valdivia River basin (Futa River, Buenaventura, Joaquines creeks); small Coastal basins (Millalafquén, Huiko creeks).	VU: D2 (Pérez-Losada et al., 2009)	VU: B1ab(iii)+2ab(iii)
Aegla manuniflata Bond-Buckup & Santos, 2009 (40)	Brazil: Uruguay River Basin (Ibicuí-Mirim, Toropi rivers, Taquara, Itaimbé creeks).	EN: B1ab (iii, iv) (Brasil 2014)	EN: B1ab(iii, iv)
Aegla marginata Bond-Buckup & Buckup, 1994 (41)	Brazil: Paraná River basin (Paranapanema, Upper Iguaçu rivers); Southeastern Atlantic system (Ribeira do Iguape River, Paranaguá Bay).	LC (Pérez-Losada et al., 2009)	LC
Aegla meloi Bond-Buckup & Santos 2015 (42)	Brazil: Paraná River basin (tributary of Iguaçu River).	CR: B2ab(iii) (Santos et al., 2015)	CR: B2ab (iii)
Aegla microphthalma Bond-Buckup & Buckup, 1994 (43)	Brazil: Southeastern Atlantic system (Ribeira do Iguape River sub-basin, Santana Cave).	CR A4e+B2ab(iii,v) (Maia et al., 2013)	CR: A4e+B2ab(iii,v)
Aegla muelleri Bond-Buckup & Buckup, 2010 (44)	Brazil: South Atlantic system (Passa Quatro River, Espingarda Creek).	NE	VU: B2ab(iii)
Aegla neuquensis Schmitt, 1942 (45)	Chile: Simpson River basin (Simpson, Pollux rivers). Argentina: Neuquén River basin (Neuquén River); Negro River basin (Negro, Limay, Aluminé, Collón Curá rivers, Nahuel Huapi, Tromen, Aluminé, Huechlafquen, Lolog lakes); Chubut River basin (Chubut, Mayo, Chico, Tecka, Senguerr rivers, Fontana Lake).	LC (Pérez-Losada et al., 2009)	LC
Aegla oblata Bond-Buckup & Santos 2012 (46)	Brazil: Uruguay River basin (Caronas, Lava-Tudo, Périco Redondo rivers, Engenho Velho Creek).	VU: B1ab(iii) (Santos et al., 2012)	EN: B1ab(iii, iv)
Aegla obstipa Bond-Buckup & Buckup, 1994 (47)	Brazil: South Atlantic system (Guaíba Lake, Lower Jacuí, Ratos, Camaquã rivers).	EN: B1ab(iii) + 2ab(iii, iv) (Brasil, 2014)	EN: B2ab(iii, iv)
Aegla occidentalis Jara, Pérez-Losada & Crandall, 2003 (48)	Chile: Paicaví River basin (Caramávida, Tucapel rivers, Lanalhue Lake); Lleu Lleu River basin (Lleu Lleu Lake).	LC (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla odebrechtii Müller, 1876 (49)	Brazil: Uruguay River Basin (Irani, Peixe, Canoas, Pelotas rivers); South Atlantic system (Itajaí-Açu River).	LC (Pérez-Losada et al., 2009)	LC
Aegla papudo Schmitt, 1942 (50)	Chile: Choapa River basin (Choapa, Illapel rivers); Ligua River basin (Ligua River, Papudo Creek); Catapilco River basin (Catapilco River); Aconcagua River basin (Aconcagua River); Marga Marga River basin (Marga Marga River); Maipo River basin (Mapocho River).	CR: A2ae (Pérez-Losada et al., 2009)	EN: A2ae
Aegla parana Schmitt, 1942 (51)	Brazil: Paraná River basin (Upper and Lower Iguaçu, Timbó, Canoinhas rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla parva Bond-Buckup & Buckup, 1994 (52)	Brazil: Paraná Basin (Upper, Middle and Lower Iguaçu River); South Atlantic system (Itajaí-Açu, Cubatão do Sul, Cedro rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla paulensis Schmitt, 1942 (53)	Brazil: Southeastern Atlantic system (Cubatão River).	VU: B2aD2 (Moraes et al., 2016)	VU: B2aD2
Aegla perobae Hebling & Rodrigues, 1977 (54)	Brazil: Paraná River basin (Peroba Grot, Tietê-Piracicaba River).	VU: B2ab(iii) (Pérez-Losada et al., 2009)	CR: B2ab (iii)
Aegla pewenchae Jara, 1994 (55)	Chile: Rapel River basin (Claro, Cachapoal rivers, Chimbarongo Creek); Mataquito River basin (Mataquito River); Maule River basin (Maule, Lircay, Longaví, Putagán rivers); Itata River basin (Cato, Ñuble, Chillán rivers); Bío Bío River basin (Bío Bío, Laja rivers); Imperial River basin (Traiguén, Quino rivers); Toltén River basin (Donguil River).	LC (Pérez-Losada et al., 2009)	LC
Aegla plana Buckup & Rossi, 1977 (56)	Brazil: South Atlantic system (Caí, Taquari-Tainhas rivers).	EN: B1ab(iii,iv) (Brasil, 2014)	EN: B1ab(iii, iv)
Aegla platensis Schmitt, 1942 (57)	Argentina: La Plata River basin (La Plata River, Garupá Creek); Mar Chiquita system (Dulce River, Tipas, Loro, Singuil rivers); Uruguay River basin (Itacaruaré River, Santa Maria Creek); Martín García Island. Brazil: Uruguay River basin (Parizinho, Passarinhos, Chapecó, Guaraim, Moinho, Ibicuí, Ijuí, Passo Fundo, Piratini, Quarai, Santa Maria, Turvo-Santa Rosa-Santo Cristo, Várzea, Jacutinga rivers); South Atlantic system (Lower Jacuí, Caí, Sinos, Camaquã, Gravataí, Mirim-São Gonçalo, Sinos, Butuí-Icamaquã rivers, Guaíba Lake). Paraguay: Paraguay River basin (Jejuí River). Uruguay: La Plata River Basin (La Plata River, Miguelete Creek); Uruguay River Basin (Uruguay, Negro, Tacuarembó, Quaraí revers); South Atlantic system (Cebollatí River).	LC (Pérez-Losada et al., 2009)	DD
Aegla pomerana Bond-Buckup & Buckup, 2010 (58)	Brazil: South Atlantic system (Itajaí-Açu, Itapocu rivers).	NE	VU: B1ab(iii, iv)
Aegla prado Schmitt, 1942 (59)	Brazil: South Atlantic system (Mirim Lagoon, Jaguarão River). Uruguay: Uruguay River basin (Negro River); La Plata River basin (Miguelete, Malvin creeks).	LC (Pérez-Losada et al., 2009)	LC
Aegla renana Bond-Buckup & Santos, 2010 (60)	Brazil: South Atlantic system (Caí River).	CR: B1ab(iii, iv) (Brasil, 2014)	CR: B2ab(iii, iv)
Aegla ringueleti Bond-Buckup & Buckup, 1994 (61)	Argentina: Paraná River basin (Salado River).	LC (Pérez-Losada et al., 2009)	CR: B2ab(iii)
Aegla riolimayana Schmitt, 1942 (62)	Argentina: Negro River basin (Limay, Aluminé, Chimehuin rivers, Moquehue, Huechlafquen, Aluminé lakes, Jones Creek); Valdivia River basin (Lácar Lake). Chile: Valdivia River basin (Cruces, Leufucade, Antilhue, Reyehueico, San Pedro-Calle Calle rivers); Bueno River basin (Lake Puyehue); Chilóe Island (Huincha, Butalcura rivers, Huillinco Lake).	NT (Pérez-Losada et al., 2009)	LC
Aegla rosanae Campos Jr., 1998 (78)	Brazil: Southeastern Atlantic system (Paraíba do Sul River).	CR: B2abiii (Moraes et al., 2016)	CR: B2abiii
Aegla rossiana Bond-Buckup & Buckup, 1994 (63)	Brazil: South Atlantic system (Araranguá, Tramandaí rivers).	LC (Pérez-Losada et al., 2009)	EN: B1ab(iii, iv)
Aegla rostrata Jara, 1977 (64)	Chile: Toltén River basin (Caburga, Colico, Villarica lakes); Valdivia River basin (Huanehue, San Pedro, Cau Cau rivers, Calafquén, Neltume, Panguipulli, Riñihue lakes).	LC (Pérez-Losada et al., 2009)	LC
Aegla saltensis Bond-Buckup & Jara, 2010 (65)	Argentina: Paraná River basin (Pasaje, Juramento rivers).	NE	VU: B2ab(iii, iv)
Aegla sanlorenzo Schmitt, 1942 (66)	Argentina: Paraná River basin (San Lorenzo River, Los Berros Creek).	LC (Pérez-Losada et al., 2009)	EN: B2ab(iii)
Aegla scamosa Ringuelet, 1948 (67)	Argentina: Colorado River basin (Mendoza, San Juan rivers, Uspallata, Villa, El Infiernillo, Água Negra creeks).	LC (Pérez-Losada et al., 2009)	DD
Aegla schmitti Hobbs III, 1979 (68)	Brazil: Paraná River basin (Paranapanema, Upper Iguaçu, Paraná, Tibagi, Ivai, Timbó rivers); Southeastern Atlantic system (Ribeira do Iguape River).	LC (Pérez-Losada et al., 2009)	LC
Aegla septentrionalis Bond-Buckup & Buckup, 1994 (69)	Bolivia: Paraná River basin (Salo, Sella rivers). Argentina: Paraná River basin (Bermejo, Pilcomayo, Arenales rivers).	LC (Pérez-Losada et al., 2009)	EN: B1ab(iii, iv)
Aegla serrana Buckup & Rossi, 1977 (70)	Brazil: Uruguay River basin (Peixe, Santa Rita rivers); South Atlantic system (Caí, Taquari-Tainhas, Sinos, Tramandaí rivers).	VU: B1ab(iii,iv) (Brasil, 2014)	VU: B1ab(iii,iv)
Aegla singularis Ringuelet, 1948 (71)	Argentina: Uruguay River basin (Pindapoy Creek). Brazil: Uruguay River basin (Apuaê-Inhandava, Turvo, Ijui, Passo Fundo, Várzea, Jacutinga, Peixe rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla spectabilis Jara, 1986 (72)	Chile: Imperial River basin (Chol Chol, Perquenco, Quepe, Cautín rivers); Toltén River basin (Donguil River).	CR: A2ae+B1ab(i,iii,v) (Pérez-Losada et al., 2009)	VU: B1ab(iii)+2ab(iii)
Aegla spinipalma Bond-Buckup & Buckup, 1994 (73)	Brazil: South Atlantic system (Jacuí, Taquari-Tainhas, Sinos, Upper and Lower Jacuí rivers); Uruguay River basin (Ijuí River).	VU: B1ab(iii, iv) (Brasil, 2014)	VU: B1ab(iii, iv)
Aegla spinosa Bond-Buckup & Buckup, 1994 (74)	Brazil: Uruguay River basin (Apuaê-Inhandava, Jacutinga, Canoas, Pelotas rivers).	LC (Pérez-Losada et al., 2009)	LC
Aegla strinatii Türkay, 1972 (75)	Brazil: Southeastern Atlantic system (Ribeira do Iguape River, Diabo Cave, Tapagem Grot).	LC (Pérez-Losada et al., 2009)	EN: B2ab(iii)
Aegla talcahuano Schmitt, 1942 (29)	Chile: Cachapoal River basin (Antivero River); Tinguiririca River basin (Chimbarongo River); Maule River basin (Maule, Lircay, Putagán, Liguay, Perquilauquén rivers).	VU: A2ae (Pérez-Losada et al., 2009)	EN: B1ab(iii)+2ab(iii)
Aegla uruguayana Schmitt, 1942 (76)	Argentina: Paraná River basin (Areco, Dulce, Tercero, Paraná, Baradero rivers); Uruguay River basin (Yeruá, Molino creeks); Martín García Island; La Plata River basin (Cepeda, Manantiales creeks). Brazil: Uruguay River basin (Ibicuí, Quaraí, Santa Maria rivers); South Atlantic system (Guaíba, Lower Jacuí, Negro, Mirim, São Gonçalo, Icamaquã rivers). Uruguay: Uruguay River basin (Uruguay River); La Plata River basin (Rosario River, Las Vacas, San Carlos creeks); South Atlantic system (Cebollatí River).	LC (Pérez-Losada et al., 2009)	LC
Aegla vanini Moraes, Tavares & Bueno, 2016 (79)	Brazil: Southeastern Atlantic system (Claro River).	VU: B2aD2 (Moraes et al., 2016)	VU: B2aD2
Aegla violacea Bond-Buckup & Buckup, 1994 (77)	Brazil: South Atlantic system (Lower Jacuí, Lake Guaíba tributaries).	EN: B1ab(iii, iv) + 2ab (iii, iv) (Brasil, 2014)	EN: B1ab(iii,iv)

Figure 2
Distribution of Aegla Leach, 1820 in Argentina, Bolivia, Chile, and Paraguay (see Tab. 1 for the corresponding species number). In areas with less species diversity, we used a larger font size to facilitate the localization in the map. However, where diversity is high, to avoid overlapping the numbers, we reduced the font size, and if necessary we used a point with a line indicating the respective number.

Figure 3
Distribution of Aegla Leach, 1820 in Brazil and Uruguay (see Tab. 1 for the corresponding species number). In areas with less species diversity, we used a larger font size to facilitate the localization in the map. However, where diversity is high, to avoid overlapping the numbers, we reduced the font size, and if necessary we used a point with a line indicating the respective number.

Based on our assessment, 17 species were classified as critically endangered (CR), 21 as endangered (EN), 19 as vulnerable (VU), one as near threatened (NT), 21 as least concern (LC), three had deficient data (DD), and one was not evaluated (NE). Current and previous conservation status and the criteria used to assign each species to an IUCN category are also shown in Tab. 1.

Discussion

There are currently 83 known species of the genus Aegla, although the existence of A. intermedia is questioned. This species was described based upon specimens collected by the “U. S. N. Astronomical Expedition”, in tributaries of Maipu River, near to Santiago, Chile. Despite the effort of various researchers, the species has never been found again in nature and its type-series has disappeared ( Bond-Buckup and Buckup, 1994). Hence, the conservation status of this species was not evaluated.

Using updated data from the species distribution, we found 57 species of Aegla under threat or almost 70% of the 82 species evaluated. This proportion represents a significant increase in relation to previous assessments for the genus Aegla, which estimated nearly half of the present value ( Bond-Buckup et al., 2008; Pérez-Losada et al., 2009). Considering the most recent conservation status assessment for each species, from the 32 species previously assessed as LC, only 20 kept the same status; nine species changed the category to some level of threat and three have deficient data. Twenty-nine species kept the same threat category, but seven previously threatened species raised the threat level. Only one species changed from a threat category to NT status ( Tab. 1).

Eighteen new species were described in the last decade (see Tab. 1). It is noteworthy that all the recently described species are under some level of threat ( Santos et al., 2009; 2010; 2012; 2013; 2014; 2015; Bond-Buckup et al., 2010a; 2010b; Moraes et al., 2016). Two species previously considered extinct in the wild, Aegla expansa Jara, 1992 ( Pérez-Losada et al., 2002a) and Aegla lata Bond-Buckup & Buckup, 1994 ( Pérez-Losada et al., 2009), have been found again ( Galves et al., 2007; Ministerio del Medio Ambiente, Chile, 2013) and are currently categorized as endangered and critically endangered, respectively.

The high rate of endemism presented by many aeglid species, in association with habitat loss/fragmentation or even climatic events, has led many populations to decline ( Maia et al., 2013; Bueno et al., 2014). Forty years ago, many species were found in several streams and rivers within a basin, but currently, most of them are restricted to low order creeks ( Bond-Buckup and Buckup, 1994).

Threats to conservation of aeglids and other freshwater organisms in southern South America derive from different sources: removal of riparian forest, causing siltation ( Magris et al., 2010); habitat modification, fragmentation and destruction ( Cumberlidge et al., 2009; Reid et al., 2013); freshwater contamination with agricultural pesticide ( Magris et al., 2010; Negro et al., 2015; Stehle and Schulz, 2016); construction of dams that alter the flow and sometimes the temperature of the water as well ( Olsson, 2015); urban and agricultural activities which modify the physical and chemical characteristics of water bodies and freshwater biota composition ( Milesi et al., 2008; Hepp and Santos, 2009; Hepp et al., 2010; Magris et al., 2010); invasive species ( Palaoro et al., 2013; Loureiro et al., 2015), among others.

Threats to the family Aeglidae, specifically, have been identified and listed, as the use of biocides in wine and fruit farms in Chile, silviculture of exotic species, large-scale cultivation of potatoes and apples with extensive use of pesticides, and hog raising activities along southern rivers in Brazil ( Bond-Buckup et al., 2008), beyond severe deforestation in Paraguay ( Satterlee et al., 2012). Irrigated rice crops are responsible for the entrance of great amount of pesticides into the hydric systems ( Bhuiyan and Castañeda, 1995), what may constitute a threat to aeglids in southern Brazil, Uruguay, Paraguay and northwestern Argentina. These activities, although doubtless important, negatively impact the aquatic environment, and it is imperative that they respect the principle of sustainability ( Santos et al., 2012). In Argentina, the exotic golden mussel Limnoperna fortunae Dunker, 1857 was reported to settle on Aegla platensis Schmitt, 1942 affecting its populations ( Darrigran and Damborenea, 2006). In Bolivia, anthropic activity and indiscriminate extraction by local people affect populations of Aegla septentrionalis Bond-Buckup & Buckup, 1994 ( Flores, 2010), the only aeglid species occurring in that country.

It should also be taken into account that many aeglid species present a narrow distribution, and two species with a relatively wide distribution, Aegla longirostriBond-Buckup & Buckup, 1994 and Aegla platensis, may represent a complex of cryptic species as supported by genetic and morphogeometric evidence ( Bartholomei-Santos et al., 2011; Marchiori et al., 2014; 2015). If this is true, so the distribution area of each cryptic species will be narrower than the “grouped” species, which can result in different conservation status for each cryptic species, the reason for which data were considered deficient for these two current recognized species.

Our assessment of the conservation status of all the currently known species of Aeglidae is worrisome, with 70% of the species under some level of threat and 20% critically endangered. Measures to protect the aeglid fauna are urgent, such as the frequent monitoring of water quality in systems where the threatened species occur, following-up the species to obtain information on possible population fluctuation or decline, protecting and restoring habitats, and information spreading among the population on the importance of water courses for conserving the native fauna, as well as encouraging agricultural practices that do not harm the water quality.

Aeglids live preferentially in clean waters ( Bond-Buckup and Buckup, 1994), presenting a high demand for oxygen ( Dalosto and Santos, 2011), and thus making these crustaceans potential good indicators of water quality. Policies for continental water quality control will help to protect not only the aeglids but also the limnetic fauna as a whole.

Acknowledgements

We are grateful to Dr. Nilton Hebling, the first author's "scientific grandfather", for his wise teachings and wonderful legacy. We would like to thank CNPq for the productivity grant for SS (311142/2014-1).

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• 1
  This article is part of the special series offered by the Brazilian Crustacean Society in honor to Nilton José Hebling in recognition of his dedication and contributions to the development of carcinology in Brazil.

• 2
  Guest Editors: Maria Lúcia Negreiros-Fransozo and Adilson Fransozo

• Zoobank:

  http://zoobank.org/urn:lsid:zoobank.org:pub:A7252DAD-A10F-4110-94F0-890E2707F5EF

Publication Dates

• Publication in this collection
  2017

History

• Received
  10 Mar 2016
• Accepted
  01 Oct 2016