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Reu. MILS.Argentina Cienc. Nat., n.s. 3(1/: 1-12,2001 Buenos A~res.ISSN 1514-5158

A formal definition of the Para~no-Punanbiogeographic subre&on and its provinces, based mainly on ta%a

Juan J. MORRONE

Museo dc Zoologia, Facultad de Ciencias, UNAM, Apdo. Postal 70-399,04510 Mexico RE,Mhxico Abstract: The Paramo-Pundn subregion comprises the Andean hig-hlanck between western Venezueid and cen- tral western Argentina. It comprises five provinces: North Andean Paramo (high peaks of Vcnozuela, , Emador; and northern , above 3000 m), Coastal Peruvian Desert (narrow strip along the Pacific Ocean coast, from northern Paru to northern ),Puna (southernBolivia, northern Chile, northern Argentina, and southern Peru), Atmama (northern Chile), and Prcpuna (central western Argentina, fiom Jiijuy to northern Mendoza). The subregion and its provinces are defined and characterized by endemic twra, specially .

Key words: Paramo, Puna, South America, biogeography, regianaliarition,tracks

The Andean-Palagonian domain of Cabrera & RESULTS Willink (1973), comprising the Altoandean, Punan, Desert, Central Chilean, and Patagonian I'aramo-Punan subregion provinces, does not constitute a natural (Figs. 1,2) biogeographic unit. Recent studies (Fjeldsg, 1992; The Paramo-Punan subregion comprises the Morrone & Urtubey, 1997; Morrone et al., 1997; Andean highlands between western Venezuela Posadaset al., 1997) have shown that the Central and central western Argentina (Cabrera & Chilean and Patagonian provinces are not closely Willink, 1973; Morrone, 1994b, 1996, 1999). it related to the rest. Furthermore, the Paramo comprises five provinces (Fig. 1): North Andean exhibits historical relationships with the Paramo, Coastal Peruvian Desert, Puna, Atacama, remaining Andean-Patagonian provinces, in spite and Prepuna provinces. The track of the of being quite different from an ecological Macrostyphlus generic group (Coleoptera: viewpoint (Morrone & Urtuhey, 1997; Posadas et Curculionidae) (Fig. 21 is typical of the subregion. al., 1997). Based on these studies, I (Morrone, Synonyms.Andean district (Cabrera & Yepes, 1999) redefined the Paramo-Punan suhregion. My 1940: 16), Suhandean district (Cabrcra & Yepes, objective herein is to formalize this new 1940: 151, Altoandean province (Cabrera, 1951: biogeographic scheme for the Paramo-Punan 23, 1953: 107, 1957: 337, 1958: 200; Cahrera & subregion, defining it and its provinces, and Willink, 1973: 841, Andean domain (Ringuelet, providing the taxa endemic to them. In contrast 1961: 156), Subandean province (Fittkau, 1969: with previous schemes that emphasized 642), Central (Sick, 1969: 463), Andean taxa, I detail herein mainly animal taxa. province (Fittkau, 1969: 642; Carpintero, 1998: 1481, Quichua Altoandean district (Cabrera, 1971: MATERIAI, AND METHODS 30, 1976: 52), Punan centre (Muller, 1973: 921, Punan region (Rivas-Martinez & Tovar, 1983: Distributional data for this study were 5161, Punan subregion (Rivas-Martinez & Nava- obtained from the literature. Names and rro, 1994: map), Punan area (CoscarOn & classification of the bird and mammal taxa fbllow Coscar6n-Arias, 1995: 726), Central Andes Sibley & Monroe (19901 and Nowak (19911, biorcgion (Dinerstein et al., 1995: map), and respectively For the subregion and each province, Paramo-Punan suhregion (Morrone, 1999: 11). the distribution of an endemic tason is Vegetation. Shrub steppes, with small forests represented as an individual track (see Morrone ofPolylepis tomentella (Cabrera & Willink, 1973). & Crisci, 1995; Craw el al., 19991. Dominant plant include Acanthollppia 2 Reuzsta del Museo Argerzt~node CLenczas Naturales, n. s. 3 (I), 2001

leptostachya, and Trichocereuspasacana (Cabre- ra, 1971, 1976; Cabrera & Willink, 1973; Rivas- Martinez & Tovar, 1983; Fjeldss, 1992). Taxa. GNETQPRYTA. Gnetopsida. Ephedrales. Ephedraceae: Ephedra breana, E. multiflora, and E. rupestris (Hunziker, 1995). NIAGNQLIOPITYTA. Ma@oliopsida. Apiales. Apiaccae: Azorella colnpacta (Martinez, 1989). . :Barnadesia spp.,Lophopa- ppus, Perezia ciliosa, P. coerulescens, and P. purpurata (Katinas, 1995; Morrone & Urtubey, 1997). Rosales. Rosaceae: Polylepis (Simpson, 1979; Fjeldd, 19921. ANNELIDA. lilirudinea. Giossiphoniiformes. Giossiphoniidae: Adaeto- bdella cryptica and Mninbdella batracophila (Ringuelet,19851. ARTiiIROPQDA. Arachnids. Arancae. Ciubionidae: Trachelopachys bicolor (Platnick,1975). Pseudoscorpiones. Chthonnidae: Austrochtl~oniusboliuianus (Vitali-Di Castri, 1975). Solifugae. Ammotrechidae: Pseudocleobis alticola (Maury, 1983, 1998). Crustacea. Anomopoda. Chydoridae: Pleurozus caca (Paggi, 1998). Hexapoda. Coleoptera. Carabidae: Am- blygnathus giluipes peruanus, Notiobia aqui- lalorum, andN.peruuiana (Noonan,1981a, b; Ball & Maddison, 19871; Curculionidae: Acroriellus, Acrorius, Listroderes howdenae, L. punicola, Macrostyphlus generic group, Nacodius, Pan- deleleius baccatus, P. cornelli, P. smithsoni, Premnotrypes generic group, Rhigopsidius, and Trichocyphus (Kuschel, 1956; Lanteri, 1989; Morrone & LoiLono, 1992; Morrone, 1993,1994a- d, 1.997; Morronc & Ocampo, 1995; Anderson & Morrone, 1996; Howden, 1996; Ocampo & Morrone, 1996); Dytiscidae: Bunites (Bachmann & Tr6mouilles, 1982); Silphidae: Nic?ophorics scrulator (Peck & Anderson, 1985); Trogidae: Polynoncus peruanus (Scholtz, 19901. Diptera. Ephydridae: Mirnapsilopa (Lizarralde de Grosso, 1982); Simuliidae: Giganlodax cortesi species group, G. incapucara, Simulium albolinealum, S. blancasi species group, and S. horcoshuspi (Goscaron, 1991). Hemiptera. Belostomatidao: Belostoma dallasi and Lethocerus mazzai (Contartese & Bachmann, 1987); Corixidae: Ectemnostega stridulata (Bachmann, 19811; Fig. I. Provinces oftheParamo-Punan subregion. Miridae: Khinacloaaricana species group (Schuh & Schwartz, 1985). Hymenoptera. Pompilidae: Adirosles and Pompilocalus edmondii species hastulata, Adesmia horridiusciila, Atriplex group (Roig-Alsina, 1985, 1989); Sphecidae: micropt~ylla,Baccharis boliuie~zsi.s,B, incarum, Chloriunperuanum (Willink,1951). . Buddleia spp., Clzenopodium chilense, Chuqui- Pieridac: Infraphulia, Phulia, Piercolias, and raga atacamensis, Cortaderia speciosa, Ephedra Pierphuliu (Field & Herrera, 1977). Neuroptera. breana, Fabiana densa, Lophopappus spp., Hemerobiidae: IIeinerobius cmtralis (Monserrat, Nardophyllum armatum, Parastrephia lepi- 1996); Nemopteridae: Veurise (Stange & Williner, dophylla, P. pi~ylicaefolia,Polylepis tornentella, 1981). Orthoptera. Acrididae: I'onderacris Ribes spp., Solanum quadriparlilum, Slipa (Ronderos & Carbonell, 1971; Ronderos & Morrorre: The bzogeograph~cParamo-Punan subregzon 3

Cigliano, 1991); Tristiridae: Crites, Incacris, comidae:Abrocoma cinerea (Redford& Eisenberg, Paracrites, and Punacris (Cigliano, 1989). 1992); Caviidae: Cavia tschudii (Redford & VERTEBItATB. Actinopterygii. Cyprinodon- Eisenherg, 1992); Chinchillidae: Chinchilla tifbrmes. Cyprinodontidae: Orestini (Costa,1997). hrevicaudata (Redford & Eisenherg, 1992); Amphibia. Anura. Leptodactyiidae: Telmatobius Muridae: Abrothrix albiuenter, Ausliscomys spp. (Ringuelet, 1961;Muller, 1973; Lavilla, 1985). sublimis, Chinchillula sahamae, and Graonqvs Aves. Anseriformes. Anatidae: Anas armata garleppi (Redford & Eisenherg, 1992).Xenarthra. herlepschi, A. cyanoptera orinomus,A, flauirvstri.~ Dasypodidae: Chaetophioctus nationi (Redford& oxyptera, A. puna, and Chloephaga melanoptera Eisenberg, 1992). (Navas, 1977; Olrog, 1984). Apodiformes. Apodidac: Aeronautes andecolus and Cypseloides North Andean Paramo province rostchildi (Olrog, 1984). Ciconiiformes. hcci- (Fig. 3) pitridae: Buteo poecilochroirs (Muller, 1973); High peaks of' Venezuela, Colombia, Ecuador, Charadriidae: Charadrius alticola, Phegornis and northern Peru, above 3000 m. The track of m,itchelli, and Recuruir-ostra andina (Miiller, Gigontodaxceruicornis (liiptera:Simuliidae) (Fig. 1973); Falconidae: Phalcobaenus megalopterus 3) is typical of this province. (Olrog, 1984); Laridae: Larus ser~ranus(Olrog, Synonyms. Incasic province (Fittkau, 1969: 1984);Phoenicopteridae: Phoenicoplerus andinus 6421, Northern Andes (Sick, 1969: 4611, Paramo and P. jamesi (Miillcr, 1973; Olrog, 1984); province (Cahrera & Willink, 1973: 66; Morrone, Podicipcdidae: Podiceps taczanowskii (Muller, 1994b: 190, 1996: 108, 1999: 13; Rivas-Martinez 1973); Scolopacidae: Gallinago andina (Muller, & Navarro, 1994: map; Posadm et al., 1997: 2), 1973; Olrog, 1984); Threskiornitidae: Plegadis North Andean centre (Miilier, 1973: 45), Bogota ridgwayi (Muller, 1973; Olrog, 1984). Co- suhccntre (Miiller, 1973: 46), Northern Andean lumhiformes. Columhidae: Metriopelia aymam, province (Ringuclet, 1975: 107), Paranlo region 1!4. ceciliae, and M,morenoi (Miiller, 1973; Olrog, (Rivas-Martinez & Tovar, 1983: 516j, Norandean 1984). Gruiformes. Rallidae: Fulica cornuta, F area iCoscar6n & Coscariin-Arias, 1995: 7261,Cor- gigantea, and Gallinula chloropus garmani dillera de Merida Paramo ecoregion (Dinerstein (Muller, 1973; Olrog, 1984; Navas, 1991). et al., 1995: 107), Northern Andean Pararno l'asseriformes. Fringillidae: Diuca speculifera, ecoregion (Dinerstein et al., 1995: 1071, Santa Idiopsar brachyurz~s,Poospiza garleppi, Phrygilus Marta Paramo ecoregion (Dinerstein et al., 1995: atriceps, P. dorsalis, P. erythronotus, P.punerzsis, 107), and Cordillera Central Paramo ecoregion Sicalis lutea, and S. uropygialis (Mullcr, 1973); (Dinerstcin et al., 1995: 108). Furnariidae:Asthenes dorbignyi, A. maculicauda, Vegetation.Dflerent habitats are found inthis A.punensis,A, wyatti, Cinclodes atacamensis, and province, namely, grasslands, moorlands, hogs, river Schizoeca helleri (Muiler, 1973); Passeridae: valleys, and sc~vhhyvegetation. There are large Anthus fiir~catas(Mililer, 1973);Sturnidae:Mimus steppes of Festuca and Deyeuxia (Poaceaej, with dorsalis (Muller, 1973); Tyrannidae: Mu- typical "frailejones" (Asteraceae:Espeletiinaej, and scisaxicola juninensis and M. rufiuerlex (Multer, fbrests at the lowests parts (Cahrera&Willink, 1973; 1973). Piciformes. Picidae: Picumnus dor- Hammen, 1998; Sturm, 1990; Luteyn, 1999). bignyanus and Veniliornis/i.ontalis (Olrog, 1984). Dominant plant species are Calamagrostis spp., Psittaciformes. Psittacidae: Aratinga nitrata Espeletia spp., Espeletiopsis spp., Gentiana spp., (Olrog, 1984). Struthioniformes. Rheidae: Rhea Gunnera magellanica, Hypericum laricifolium, pennata tarapacensis (Muller, 1973). Tina- Miconia buxifolia, Paepalanthus spp., and Rubz~s miformes. Tinamidae: Nothoprocta ornata and spp. (Cahrera & Willink, 1.973; Hernandez et al., Tinamotispentlandii (Muller, 1.973;Olrog, 1984). 1992h). Restrepo & Duque (1992) analyzed the Trochiliformes. Trochilidae: Eriocnemis glau- vegetation oftho Llano de Paletara (Colombian cen- copoides and Microstilbon burmeiateri (Olrog, tral cordillera), distinguishing eight paramo 1984). Mammalia. Artiodactyla. Camelidae: vegetation types: "f~ailejonal"(with Espeletia Vicugna (Mullel-, 1973; Redford & Eisenherg, hartwegiana, Blechnum loxense, and Sphagnum. 1992); Cervidae: Hippocamelus (Muller, 1.973). suncto,josephense), "chuscal" (with Chusquea

. " " .. Pcrca, 1994); Mnstelidae: Conepatus chinga "pajonal" (witbCalamagrostis intermedia),thicket (Muller, 1973). Chiroptera. Molossidae: with Hypericurn lancioides and Blechnum loxense, Mormopterus kalinowskii (Redford & I

Taxa. FILICOPHYTA. Filicopsida. Mammalia. Rodcntia. Caviidae: Cauiaporcellus anulai~nae(Hernandez et al., 1992a); Mnridae:

~ ~ Thomasomys paramorurn (Muller, 1973); psida. Cyperales. Poaceae: Muhlenbergia cleefi Sciuridae: Sciurus granatensis variabilis (Laegaard, 1995). Magnoliopsida. Asterales. (Hernandez et a/., 1992a). Asteraceae: Coespeletia spicata, Espeletia, Espeletiopsis corymbosa, and Ruilopezia atro- Coastal Peruvian Desert province purpurea (Sturm, 1990). Capparales. Brassi- (Fig. 4) caceae: Draba arauquensis (Santana, 1995). Narrow strip along the Pacific Ocean coast, Haloragales. Gunneraceao: Gunnera antioquensis from northern Peru to northern Chile. The track and Q. cuucanu (Mora, 1984). Scrophuiariales. of Conepatus rex inca (Mammalia: Mustelidaej Scrophulariaceae: Aragoa (Fernandez, 1991). (Fig. 4) is typical of this province. Violales. Passiiloraceae: Passiflora truxillensis Synonyms. Desert province (Cabrera & (Escohar, 1988). ANNELIDA. Elirudinea. Wiilink, 1973: 89; ~Morrone,1999: 121, Coastal Hirudiniformes. Cylobdcllidae: Blanchardiella desert district (Cabrt:r;l & Willink, 1973: 90), fuhrmani (Ringuelet, 1974). AR1'NIEOPODA. Cardonales district ((:ahrera&Willink, 1973:91), Crustacea. Decapoda. Pseudothelphusidae: Andean Pacific cen?.r-e (Muller, 1973: loo), Strengerianini (Rodriguez & Campus, 1989). Peruvian subcentre [hfuller, 1973: 101), Guayas Iilexapoda. Coleoptera. Curculionidae: How- province (Ringuelet, 1.975: 107), PaciEic desert deniola nitidipennis, N.sulcipennis, Minetes, region (Rivas-Martinez & Tovar, 1983: 516), Ph.~llolhrozaristidis, andRupanius (Osella, 1980; Pacific Peruvian desert region (Rivas-Martinez & Voisin, 1986, 1991; Howden, 1992; Morrone, Navarro, 1994: map), Peruvian desert province 1.995);Staphylinidae:Polylobus belen,P. chingaza, (Rivas-Martinez & Navarro, 1994: map), Desert P. chisaca, and P. monserruteae, and Pseudopais area (Coscar6n & Coscar6n-Arias, 1995: 726),and wygodzinkyi (Herman, 1975; Kiimaszewski & ecoreson (Dinersteinet al., 1995: Sturm, 1991). Diptera. Agromyzidae: Liriomyza 111). bogotensis, L. cirriformis, L. marioecamilae, L. Vegetation. Scarce, there is only permanent menthauura, L. mosquerenais, L, santafecina, L. vegetation at the sides of the rivers and near the subachoquensis, Ophiornyia flocl~sa,0. pun- sea. Between 1500-3000 m altitude arborescent ctohalterata, and 0,sulcata (Sanahria deAri.valo, cacti are abundant, amongwhich grow shrubs and 1993a, b);Simuliidae: Gigontodax ceruicornis, G. herbs when it rains (Cabrera & Willink, 1973). paramurum and G. siberianus (Wygodzinky & Dominant plant species include Acacia Coscar6n, 1989). Hymenoptera. Pompilidae: macranllla, Ambrosia fruticosa, Caesalpinia Chirodamus paramicola (Roig-Alsina, 1985). spinosa, Diplostephium tacorense, Echinopsis Orthoptera. Acrididae: Bogotacris, Chibchacris, pachanoi, Inga feuillei, Kaegeneckia lanceolata, and Tifnotes(Ronderos & Cerdn, 1982; Ronderos Lemai,reocereus cartwrightianus, L. laetus, Neo- & Cigliano, 1991). VERTEBMFA. Amphibia. raimondia arequipensis, Paspalurn uaginatum, Anura. Bufonidae: Atelopus tamaense (Marca et Prosopis limensis, P.pallida, Salicornia ambigua, a/., 1989);Hylidae: Gastrotheca espeletia and G. Schinus areira, Tillandsia latifolia, T.purpurea, elenae (Duellman, 1989). . Elapidae: and T.straminea (Cabrera & Willink, 1973;Rivas- Martinez & Tovar, 1983). , " ' . Taxa. ARTHROPODA. Arachnida. (Hillis, 1985). Aves. Anseriformes. Anatidae: Araneae. Gnaphosidae:Echemoides aguilari and Anas cyanoptera borreroi and A.georgica niceforei E,penai (Platnick & Shadab, 1979). Hexapoda. (Hernandez et al., 1992a). Ciconiformes. Podi- Coleoptera. Carabidae:Notiobiamofletti (Noonan, cipedidae: Podiceps andinus (Ifernandez et a/., 1981a, b);Curculionidae: Galapaganus lacertosus 1992a). Coraciiformes. Momotidae: Momotus andG. squamosw (Lanteri, 1992);I-lydrophilidae: momota oliuaresii (Hernandez et a/., 1992a). Enochrus waterhousei (Ferndndez,1997). Diptera. Gruiformes.Ral1idae:Fulica americana colombia- Simuliidae: Simulium blancasi (Coscar6n & na (Hernandez et al., 1992a). Passeriformes. CoscarCin-Arias, 1995). Memiptera. Miridae: Certhiidae:Cainpylorhynchus griseus bicolor and Rhinacloa cajanzarca and R. peruana (Schuh & Cistothorus meridae (Muller, 1973; Hernandez et Schwartz, 1.985). VEXWEBRATA. Squamata. a/., 1992a); Tyrannidae: Muscisaxicola macu- Iguanidae: Tropidurwperuuianus (MiiUer,1973); lirostris niceforei (Hernandez et a/., 1992a). : heterolepis (Muller, 1973). Tinamiformes. Tinamidae: Crypturellus kerriae Aves. Passeriformes. Fringillidae: Atlapetes and C. saltuarius (HernAndez et al., 1992a). nationi andxenospingus concolor (Muller, 1973). Morrone The b~ogeographccParamo-Punan subreg~on 5

Figs. 2-7. Individual tracks. 2, Paramo-Punan subregion (Macrostyphlus generic group); 3, North Andean Paramo province (Gigantodaz ceruicornis); 4, Peruvian Coastal Desert province (Conepatus rex inca); 5, Puna province (Epilobiun pedicelare); 6, Atacama province (Telmatobius halli); 7, Prepuna province (Cyanoliseus palagonus andinus).

Trochiliformes. Trochilidae: Thaumaslura cora Puna province (Miiller, 19733. Mammalia. Carnivora. Muste- (Fig. 5) Iidae: Conepatus rex inca (Miiller, 1973). Southern , northern Chile, northern Chiroptera. Furipteridae: Ainorphochilus Argentina, and southern Peru. The track of schnablii (Redford & Eisenherg, 1992); Epilobiumpedicelare (Myrtales: Onagraceae) ((Fig. Phyllostomidae: Sturnira bogotensis (Pacheco & 5)is typical of this province. Patterson, 1992). Kodentia. Muridae: Abrothrix Synonyms. Punan province (Cabrera, 1951: mollis and Phyllotis gerbillus (Muller, 1973; 23, 1953: 107, 1957: 337, 1958: 200, 1971: 32; Patton &Smith, 1992). Cabrera & Willink, 1973: 87; Cabrera, 1976: 59; 6 Reuzsta del Museo Argentzno de Cuzrzczas Naturales, n. s. 3 il),2001

Morrone, 1994h: 190, 1.996: 108; Posadas et al., Orestias (Parenti, 1981). Aves. Passcriformes. 1997: 21, Altoandean province (Cahrera, 1971: 30): Formicaridae: Grullaria andicolu (Fjelds$1992). High plateau region (Pefia, 1966: 4): Northern Mammaiia. Rodentia. Muridae: Ahrothrix Andean cordillera region (Pefia, 1966: 5), North orophilus (Patton & Smith, 1992). Andean centre (Miiller, 1973: 341, South Andean Cuyan province (Ringuelet, 1975: 107), Titicaca Atacama province province (Ringuelet, 1975: 107), Argentincan- (Pig. 6) Atacaman province (Rivas-Martinez & Navarro, Northern Chile, limiting to the south with the 1994: map), Bolivian province (Rivas-Martinez & Central Chilean subregion (Morrone, 2000). The Navarro, 1994: map), Peruvian province (Rivas- track of Telnzatobius halli (Anura: Lcptodac- Martinez & Navarro, 1994:map), Bolivim montane tylidae) (Fig. 6)is typical of this province. dry ibrests ecoregion (Dinersteinet al., 1995: 101), Synonyms. Northern coast region (Pciia, Central Andean Puna ecoregion (Dinersteinetal., 1966: 8),Northern desert region (Peiia, 1966: 71, 1995: 108), ecoregion Andean region (O'Brien, 1971.: 198), Atacaman (Dinerstein et al., 1995: 1081, Central Andean Dry region (O'Brien, 1971: 1991, Coquimban district, Puna ecoregion (Dinerstein et ai., 1995: 108), in part (Cabrcra & Willink, 1970: 91!, Andean Humid Punan province (Morrone, 1999: 121, Cen- Pacific centre (Miillcl-, 1973: loo), Chilean tral Punan province (Morrone,1999: 1.21, andArid subcentre (Miiller, 1973: 101), Atacaman area Punan province (Morrone, 1999: 12). (Artigas, 1975: map), Atacaman hiperdesert Vegetation.Montane grasslands, constituted province (Rivas-Martinez & Navarro, 1994: map), by different communities.of hunchgrasses, Desert area (CoscarOn & Coscar6n-Arias, 1995: montane grasslands, small shrubs, trees, and 726),Atacanlan desert ecoregion (Dinersteinet al., herbs (Cabrcra & Willink, 1973; Dinersteinel a1 ., 1995: 1111, and Atacaman province (Morrone, 1995). 1999: 12). Taxa. MAGNOLIOPmrA. Magnoliopsi- Vegetation.Generally scarce, although there da. Asterales. Asteraceae: Chuquiraga ataca- are rich communiiies, the lomas formations, mensis and C. kuscheli (Ezcurra et al., 1997). supported by the winter fogs formed over cool Myrtales. Onagraceae: Epilobium fragile, E. Pacific Ocean currents (Dinerstein et al., 1995). pedicellare, Fuchsia austromontana,F boliuianus, Taxa. IMAGNOLIOPNUTA. Magnoliopsi- F. cochabamba species group, F. simplicicaulis, and da Asterales. Asteraccae: Chuquiraga ulicina ii'. tincla species group, Berry, 1982; Solomon, (Ezcurra et al., 1997). ARTNROPODA. 1982). ARTHROPODA. Arachnida. Araneae. Arachnida. Araneae. Dipluridae: Chilehexops Clubionidae: Trachelopachys bidentatus, T. platnichi (Coyle, 1966); Nemesiidac: Acantho- rnachupicchu, and T. larnza (Platnick, 1975). gonatus alegre andFlamencup,sii (Golobofi; 1995). Scorpiones. Bothriuridae: Brachistosternus Solifugae. Ammotrechidae: Chileolracha (Maury, andinus (Maury, 19733. Crustacea. Decapoda. 1987, 1998). Hexapoda. Coleoptera. Carabidae: Aeglidae: Aegla humahuaca., A. intercolala, A. Notaphz~saricensis and Nothocys marcidas juju.yana, A. ringueleti, A. sunlorenzo, and A. (Jeannel, 1962); Curculionidae: Lislroderes septentrionalis (Bond-Buckup & Ruckup, 1994). robustior (Morrone, 1993). Diptera. Simuliidac: Hexapoda. Coleoptera. Carabidae: Notiobia Gigantodax cortesi, G. eremicus, G.ja2unchuspi, schnusei (Noonan, 1981a, b); Staphylinidae: G. praealtus, Simulium hectoruargusi, S,putre, Neobisnius senzirufus (Frank, 1981). Diptera. and S. tenuipes Wygodzinky & Coscarbn, 1989; Simuliidae: Gigantodux awa, G. chacabanzba, G. Coscarbn, 1991, 1998). Hemiptera. Miridae: patihuaycensis, and G. punapi (Wygodzinky & Polymerus atacamensis and Rhinacloa azapa CoscarOn, 1989). Hemiptera. Enicocephalidae: (Schuh & Schwartz, 1985; Carvalho & Carpinte- Enicocephalus taeuberi (Grimaldi et al., 1993); ro, 1986). Lepidoptera. : Hypsochila Miridae: Rhinacloa betanzos and R, juli (Schuh wagenknechti sulfurodice, Infraphulia ilyodes, & Schwartz, 1985); Nepidae: Curicta peruuiana and Pierphulia r. rosea (Field & Herrera, 1977). (Keffer, 1996). Hymenoptera. Pompilidae: Orthoptera. Tristiridae: Elasmoderini (Cigliano, Caliadurgus subandinus (Roig-Alsins, 1982). 1989). VERTEBRATA. Amphibia. Anura. Lepidoptera. Lycaenidae: Terra altilineatu Lcptodactylidae: Telnzatobius halli (Muller,1973). (Johnson, 1992); Slenomidae: Setiostonza Aves. Passeriforn~es.Rhinocryptidae: Pteropto- earobasis (Duckworth, 1971). Neuroptera. chos megupodius (Mullor, 1973); Sturnidae: Hemerobiidae: Hemerobius montsae (Monserrat, Mimus thenca (Miiller, 1973). Mammaiia. 1996). Orthoptera. Tristiridae: Atacamacridinae Rodentia. Ctenomyidae: Ctenomys robustus (Cigliano, 1989). VERTEBRATA. Actinopte- (Miiller, 1973; Contreras & Yafiez, 1995); rygii. Cyprinodontiformes. Cyprinodontidae: Octodontidae: Octodon degus (Miiller, 1973). Morrone: The biogeographic : Paramo-Pl~nunsubregion 7

Prepuna province (Flores & Roig-JuAent, 1997). Diptera. Culicidae: (Fig. 7) Culex cuyanus and C. riaJanus (Duret, 1968); Central western Argentina, from Jujuy to Simuliidae: Gigantodm cilicinus (Wygodzinky & northern Mendoza. The track of Cyanoliseus Coscar6n, 1989). VERTEBRATA. Amphibia. patagonus a~zdinus(Aves: Psittacidae) (Fig. 7)is Anura. Buronidae: Bu/a s, spinulosus (Gallardo, typical of this province. 1966); Hylidac: IIyla pulchella andina and Synonyms. Subandean district (Cabrcra & Melanophrynisci~sstelzneri spegazzinii (Galiardo, Yepes, 1940: 15), Prepunan province (Cabrera, 1966); Leptodactylidae: Odontophrynus occiden- 1951: 23,1953: 107,1958: 200,1971: 21; Cabrcra talis,Pleurodernacinerea, andP. nebulosa (Gallar- & Willink, 1973: 76; Cabn?ra,1976: 34; Morrone, do, 1966). Squamata. Iguanidae: Cupriguanus 1999: 12), Subandean province (Fittkau, 1969: scapulatus (Cei, 1973). Aves. Passeriformes. S42), Altoandean province (Cabrcra, 1971: 30; Furuariidae: Asthenes sieir~bachi(Olrog, 1984). Cabrera & Willink, 1973: 841, Cuyan Altoandean Psittaciformes. Psittacidae: Cyanoliseus pata- district (Cabrera, 1971: 31, 1976: 551, and gonus andinus (Darrieu, 1980). Southern Andean Steppe ecoregion (Dinersteinet al., 1995: 108). Vegetation. Shrub steppes with cacti (Cabre- r.a, 1971, 1976; Cabrera & Willink, 1973). I am gratcrul to Gustavo Flores and Sergo Dominant plant species arc Ahromeitiella Roig-Juiient for their useful comments on the breuifolia, Acacia uisco, Adesmia inflexa, manuscr~pt Raccharis salicifolia, Caesalpinia trichocarpa, Cassia crassirustris, Cercidiurn andicolo, Det~terocohniastrobilifera, Goci~natiaglutinosa, Plasopis ferox, Puya dickioides, Schinus areira, Acos~a,I,. E. & E. A. Maury 1998. Scorpionos. in: Tillandsia gilliensi, Trichocereus pasacana, T. Morrano, J. J. & S. Coscar6n (eds.), Aiodiversidad terscheckii, and Zucagnia puncluta (Cabrera, de artrdpodos argcntinos: Una peri.pectiua 1971,1976; Cabrera & Willink, 1973). biotazondmica, Ediciones Sur, I,a Plata, pp. 545-559. Taxa. MAGNOLIOPW-YTA. Magnoliopsi- Anderson, R. S. & J. J. Morrone. 1996. A new of microphtalrnic Rhytirrhinini from Andean leaf litter da. Apiales. Apiaceae: Azorella cryptantha (Coleoolera: Curculionidae). Enl. Scand. 27: 259- (Martinez, 1989). Asterales. Asteraceae: Chu- 278. quiraga echegarayi, C. erinacea subsp. hystrix, C. Artigas, J. N. 1975. Introducci6n a1 estudio par compu- ruscifolia, Dolic1ilasiurn, Proustiu cunei/alia var. tacidri de las iroas zoogeogrificas de Chile conti- cuneifolia, and P. cuneifolia var. mollis (Katinas, ncntal basndo en la dislrihucidn de 903 cspecies de 1995; Ezcurra ef al., 1997). MOLLUSCA. animales terrestres. Gayana, mise. 4: 1-25. Gastropoda. Basommatophora. Chilinidae: Hachmann, A. 0. 1981. Insecta. Hemiptera. Corixidae. Chilina mendozana (Castellanos & Gaillard, In: Ringuelet, R. A. (dir), Fauna de Agua Dulce de la ltephhlica Argentina, kcic, Huenos Aircs, 35(2): 1981).ANNELIDA. Hirudinea. Hirudinifonnes. 9 "mn l.',,". Cylobdeliidae: Orchibdella diaguita (Ringuelet, Bachmann, A. 0. & E. R. TrC.mauillcs. 1982. El gknero 1985). ARTNROPODA. Arachnids. Scorpiones. Aunites Spangler (Coleaptera, Ilytiscidae), nuevo Bothriuridae: Bolhrinrus olaen, Brachistasternus .~nara la Areentina. I'hvsis (Buenos Aires) B. borellii, B. pentheri, B, montanus, Timogenes 40(99):109-i'10. haplochirz~s,and T. sumatranus (Maury & Roig- Hall. G. E. & 1). It. Maddison. 1987. Classification and Alsina, 1977; Roig-Alsina, 1977; Roig-AIsina & evolutionary aspects of the species of the genus Maury, 1981,1984;Maury, 1982;Acosta & Maury, Amb!ygnalhu,s Dejean, with description of Plaiyn~etopsis,new genus, and notes about selected 1998). Soiifugae. Ammotrechidae: Pseudocleobis species of Seloiophorus Ilejean (Coleoptera: arzdinus and P. puelche (Maury, 1983, 1998). carabidae: Harpalini). Trans. Amer: Enlomo!. Sac. Crustacea. Amphipoda. Bogidiel1idae:Bogidiella 113: 189-307. talarnaavensis (1,onretto. 1995): In~olfiellidae: Berrv. E E. 1982. The svstemalics and evolution of . " 4, Ingolfiella uspallatie (Lopretto, 1995-1 Decapoda. Fuchsia scct. Fuchsia (Onagraceae).Ann.Missouri Aeglidae: Aegla intercalata, A, ringueleti, and A. Bot. Gard. 69(1): 1-198. scarnosa (Bond-Buckup & Buckup, 1994).Isopoda. Hand-Huckup, G. & L. Buckup. 1994. A familia Aeglidae Protojaniridae: CuyoJanira (Lopretto, 1995). (Cruslacea, Decapoda, Anomura). Ary. Zool., Srio Stygocarididae:Parustygocaris Pauio 32(4): 159-346. Syncarida. (Grosso Cabrera, A. & J. Yepes. 1940. Mamiferos sud-arnrrica- & Peralta, 1997). Hexapoda. Coleoptera. 1~0s'uida, cosli~rnbresy desci-ipcidn~.Historin Na- Carabidae: Barypus calchaquensis (Roig-JuAent, tural Ediar, Huenos hires. 1992); Staphylinidae: Neobisrzius ornni~-ufus - 1951. Territories Iitogeogrificos de la Repkblica (Frank, 1981); Tenebrionidae: Enlomoderes zupay Argentina. Bol. Soc. Argent. Bot. 4(1-2): 21-65. 8 Reulsta del Museo Argentzno de C~enczasNaturales, n s 3 iIi,2001

- 1953. Esquema. fitogeografico de la Repiihlica Ar- Oxford Biogeography Series 11, Oxford University gentina. Rev. Mus. La Plata (N 8.1, Rot. 8(33): 87- Press, New York. 168. Darrieu, C. A. 1980.Lasrazas geogrifiias deCyanoliseus - 1957. Lavegetacibn de la Puna argentina.Reu. In". patagonus (Aves Psittacidnej. Neotropica 26(76): Agric. ll(4): 317-412. 207-216. - 1958. Fitoeeomafia en la Areentina. Sunla Geoer" Dincrstein, E. D. M. Olson, D. J. Graham, A. L. Webster, 3: 101-20; " S. A. Primm, M. P Bookbinder & G. Ledec. 1995. - 1971. Fitogeografia de la Rcpiiblica Argentina. Rol. Unn eualuacidn del estado de conservacidn de las Soc. Armnt. Rot. 14!1-2): 1-42. ecoregiones te~reslr-esde America Latina y el Cari- - 1976. Rcgiones SitogeogrAficas argentinas. 11': be. World Bank, Washington, D.C., 135 pp. Kugler, W F (ed.1, Enciclopedia Argentina de Agri- Duckworth, W D. 1971. Neotropical Microtepidoptera cultura y Jardinoriii, 11, ACME, Buenos Aires, pp. XX: Revision of the gcnus Seliostoma (Lrpidoptera: 1-85. Stenomidae). Smilhsorz. Contrib. Zool. 106: 1-45. Cabrera, A. L. &A. Willink. 1973.Riogeografia dr Am6- Duellman, W E. 1989. Lista anotada y clave de los sa-

rica Latir~a.Monomafia " 13. Serie de Bioloria.-, OEA. pos marsupiales (Anura: I-iylidae: Gastrolheee) de Washington D.C. Colombia. Caldasia 16(76j: 106-111. Carpintero, D. L. 1998. Miridae. In: Morrone, J. J. & S. Duret, .l. F! 1968. Dos ospecics nuevas do Culex fCulex1 CoscarOn (eds.),Biodiuersidod de artrhpodos argen- de la Argentina (Dipteiit, Cuiicidae). IJhysis (Bue- iirios: Unaperspecliua biolmonrimica. Ediciones Sur, nos Aires) 27(763: 323-328. La Plata, pp. 144.150. Escohar, L. K. 1988. Passifloraceae: Passiflora, Camaiho. J. C. M. & D. I,. Carointero. 1986. Mirideos suhgeneros Tacsonia, Rati~ea,Manicate y Distephana. In: Pinto, P & G. Lozano (eds.1, Flora do Colombia, monow no. 10, Universidad Nacional de Colombia, Smtafi. do Bogota, pp. 1-138. Castellanos, Z. A. do & M. C. Gaillard. 1981. Molluscs: Ezcurra, C, A. Ruggicro & 3. V Crisci. 1.997. Phylogeny Gasteropoda: Chiiinidae. In: Ringuelet, R. A. (ed.), of Chuquiraga sect. Acanthophyllae (Asteraceae- Fauna de Agun Dulce de la Repiibliea Argentina. Barnadesioideaej, and the evolution of its leaf Feeic, Buenos Aires, 15(41: 19-51. morphology in relation to climate. Syst. Rot. 22(1): Cei, J. M. 1970. Comentarios sobre algunos generos de 151-163. iruanidos: Diololaen~us.Leiosourus. .. Aoeronristis . v Ferndndez. J. L. 1991. Nueva especie y comentarios Cupriguanus. Physis (Buenos hires) C, 32(85):269- morfolbgicos y fitogoogr6ficos en el genera Aragou 276. H. B. K. (Scrophulariaceae). Caldasia 16(78j: 301- Cieliano. M. M. 1989. Itevisibn sistematieade la familia 310. Tristiridae (Orthoptera, Acridoidea).Bol. Soe. Biol. FcmAndez, L. A. 1991. Nuevos aportes para el conoci- Cor~cepcidn60: 51-110. miento del ginero Enochrus Thornson (Coleoptera: Contarrteae. A. M. & A. 0. Bachmann. 1987. Distribu- IIydrophilidaej. Physis (Buenas Aires) B, 53(124- ci6n geogrstica de las Belostomatidae argentinas 1253: 21-29. (Heteioptora).Pt~ysisiBuenoshires) B, 45(1081: 25- Field, W D. & J. Herrera. 1977. The pierid of 27. tho genera Hypsochila Wreta, Phulia I-Ierrich- Contreras, L. C. & J. L. YBfiez. 1995. Marniferos. Irb: SchSfer, Infraphulia Fiold, Pirwhulia Field, arid Sirnonetti, J. A,, M. T. K. Arroyo, A. E. Spotorno & Piereulias Staudinger Smithson. Cor~trib.2001. 232: E. Lozada (eds.), Diversidad biol6gica de Chile, 1-64. Canicyt, Santiago de Chile, pp. 336-349. Fittkau, E. J. 1969. The fauna af South America. In: Coscarbn, S. 1991. Insecta Diptera. Simu1iidao.In: Cas- Fittkau, E. J. J, lilies, H. Klingo, G. H. Schwahe & tellanos, 2. A. de (ed.),fiuna de Agua Dulce de la H. Sioli (ods.),Biogeography and ecology in South Repiiblica Argentina. Fecic, Buenas Airos, 38(21: 1- America. 2. Junk. Tire Hague.- en. 624-650. 384. FjeldsA, J. 1992. Biogeograpl~icpatterns and evolution - 1998. Simuliidae. In: Morrone, J. J & S. 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