Rev. Mus. Argentina
Cienc. Nat.,
n.s.Z(li:
1-15,
2000
Duenos Area ISSN 1514-5158Biogeographic delimitation of the Subantarctic subregion
and its provinces
J u a n J. MORRONE
Museo de Zoologfa, Facultad de Ciencias, UNAM, Apdo. Postal 70-399, 04510 Mexico D.F., Mhxico. Abstract: The Subantarctic subregion, which belongs to the Andean region of the Austral kiiigdom, comprises the austral Andes from 37" south latitude to Cabo de Piornos, the archipelago oFsouthern Chile and Argentina, and the Malvinas, South Georgia, and Juan Ferndndez islands. The Subantarr::ie subregion comprises six provinces: Maulo (southern Ciiiie and Argentina, between 31-37" south latitude), Valdivian (southern Chile and Argentina, south of the Mauic province, reaching 47" south latitude), Magellanic forest (southern Chile from 47" south latitude to Cabo de Ilornos, and southern Argentina in small portions of westcrn SanLa Cruz and Tierra del Fuem), Magellanic moorland (southern Chile and Argentina, limited by the Marellanie forest urovinee
(Chilean islands of Masatir~ra o r Robinson Cmsoe, Masafuucra or Alcjandro ~ e k i r k , and Santa Clara, situated in the Pacific ocean, 600 k m west of Valpar.aiso, at 33' south latitude).
Key words: Subantarctic, biogeography, South America.
T h e S u b a n t a r c t i c s u b r e g i o n basically compriscs the austral Andes and several small i s l a n d s of s o u t h e r n A r g e n t i n a a n d Chile (Morrone, 1996d, 1999). Its reinarkable biota has attracted t h e attention of several authors. Some of them (Monrds, 1958; Kuschel, 1960; Cabrera
& Willink, 1973; Cabrera, 1976; Crisci et a/., 1991; M o r r o n e , 1996c, d ) e m p h a s i z e d t h e distinctiveness of the Subantarctic biota and its close relationship with t h e Australasian biota. C a b r e r a (1976) listed several p l a n t genera distributed in t h e Subantarctic subregion a s weil a s in Australia, New Zealand, and/ or New G n i n e a , n a m e l y , A b r o t a n e l l a , A r i s t o l e l i a , Astelia, Carpha, Donatia, Drapetes, Eucryphia, C a i m a r d i a , L a u r e l i a , L o m a t i a , L u z u r i a g a , Marsippospermum, ~Vothofagus, Oreobolus, Orites, Phyllacne, Pseudopanax, Rostlzowia, Schoenus, Selliera, Tetrachondra, and Veroni- ca.
Within South America, t h e Subantarctic subregion h a s been placed in the Andean region (Morrone, 1999), where i t is considered to be most closely related to the Central Chilean subregion (Morrone, 1994b; Morrone e t a l . , 1997; Troncoso
& Romero, 1998). In addition, this subregion shows also some biotic relationships with the P a r a n a subregion of t h e Neotropical region (Kuschel, 1960; C a b r c r a , 1976; Morrone &
Lopretto, 1994; Maury et al., 1996), which could evidence a former connection between these areas.
My objective i s t o f o r m a l i z e a new biogeographic scheme for t h e S u b a n t a r c t i c s u b r e g i o n , providing t h e t a x a e n d e m i c or characteristic to it and its provinces.
MATERIAL AND METHODS
Distributioilal d a t a for t h i s s t u d y were o b t a i n e d f r o m t h e l i t e r a t u r e . N a m e s a n d classification of the bird a n d mammal taxa follow Sibley & Monroe (1990) and Nowak (19911, r e s p e c t i v e l y . F o r t h e s u b r e g i o n a n d e a c h province, the distribution of a n endemic taxon is represented a s a n individual track (see Morrone
& Crisci, 1995; Craw et al., 1999).
RESULTS Subantarctic subregion (Figs. 1, 2)
2 Reursta del Museo Argentina d e CI
six provinces (Fig. 1). T h e t r a c k ofGermainiellus (Fig. 2 ) i s representative of t h e subregion.
Synonyms. Moist Andean zone (Shannon, 1927: 3), Chilean district (Cabrera &Yepes, 1940: 161, S u b a n t a r c t i c f o r e s t s (Soriano, 1950: 331, Subantarctic province (Cabrera, 1951: 23, 1953: 107, 1.958: 200, 1971: 37, 1976: 72; Cabrera &
Willink, 1973: 97; Morrone, 1994b: 191, 1996c: 106; Posadas e t al., 1997: 2; Carpintero, 1998: 148; Roig, 1998: 140),AustraIcordilleran domain ( R i n g u e l e t , 1 9 5 5 b : 841, S u b a n t a r c t i c Ibrests region (Hueck, 1957: 401, Araucanian subregion ( M o n r d s , 1 9 5 8 : 1 4 5 ; R i n g u e l e t , 1 9 6 1 : 1 5 6 ; Rapoport, 1968: 751, Austral-cordilleran domain ( R i n g u e l e t , 1 9 6 1 : 1601, A n d e a n - P a t a g o n i a n forests (Ragonese, 1966: 351, Chilean province (Fittkau, 1969: 6421, SouthernAndes (Sick, 1969: 4651, Subantarctic domain (Cahrera, 1971: 36; Cahrera & Willink, 1973: 96; Cabrera, 1976: 711, N o t h o f a g u s c e n t r e ( ~ M i i l l e r , 1 9 7 3 : 1551, P a t a g o n i a n province ( R i n g u e l e t , 1 9 7 5 : 1071, Andean Subantarctic region (Rivas-Martinez &
Tovar, 1983: Chilean subregion (Flint, 1.98% 11, V a l d i v i a n - M a g e l l a n i a n r e g i o n ( R i v a s - Martinez & Navarro, 1994: map), Subantarctic a r e a (Coscaron & Goscaron-Arias, 1995: 7261, A l t o a n d e a n province (Roig, 1 9 9 8 : 139), a n d Subantarctic subregion (Morrone, 1999: 13).
Vegetation. Forests w i t h dominant species belonging t o t h e A u s t r a l genera Nothofagus, D a c r y d i u m , S a x e g o t h a e a , A u s l r o c e d r u s , Pilgerodendron, a n d Fitzroya (Rothkiigel, 1916; Skottsberg, 1921, 1960; C a h r e r a , 1971, 1976; C a b r e r a & W i l l i n k , 1 9 7 3 ) . T h e r e a r e a l s o moorlands a n d other a r e a s lacking Nothofagus, e.g., t h e J u a n Fernandez, S o u t h Georgia, a n d Malvinas islands. Inclusion of t h e l a t t e r within t h e S u b a n t a r c t i c s u b r e g i o n , h o w e v e r , w a s postulated by Ringuelet (1955a) a n d supported by more recent cladistic biogeographic analyses (Morrone, 1993a; Morrone et a l . , 1994; Posadas et al., 1997).
Taxa.
FUNGI. Cyttariales. Cyttariaceae: Cyttaria darwinii, C, berteroi, C, hooheri, C, hariotii, C. espiriosae, C. johowii, and C. exigua (Elumphries et al., 1986; Crisciet al.. 1988: Sebers, 1991: Itoin-Juiient,~. Cyperaceae: Carpha, Oreobolw, andschoenus (Cabre- ra, 19763. Juncales. Juncaceae: Marsipposperinu,n and Rastkowia (Cahrera, 1976). Liliales Liiiaceae: Astelia, Lwuriaga, and Piiilesia (Cabrera, 1976). Restionales. Centrolepidaeeae: C a i ~ n a r d i a (Cabrera, 1976). Magnoliopsida. Apiales. Apiaceae: Azorella filamentosa,A. lycopodioides, and A. selago (Martinez,
enczus Naturales, n. s. 2 (I), 2000
1989); Araliaceae: Pseudopanax (Cahrera, 1976). Asterales. Asteraceae: Ahrotanella, Eriocaulon, Leucheria coerulescens, L , diernii, L . !nagnu, Macrachacnium,Perezia nutans, and Triptilion acizillae (Cabrera, 1976; Katinas et al., 1992; Katinas, 1995). Campanulales. Donatiaeeae: Doriatia (Cabrera, 1976). Fagales. Nothaiagaceae: Nothofagus (Cranwell, 1964; Fleming, 1964; Humpbries, 1981; Humphries & Parenti, 1.986; Humphrios et al., 1986; Grehan, 1991; Seherg, 1991; 1511 & Jordan, 1993; Linder & Crisp, 1995). Laurales. Gomorlecaccae: Go,norteca iCabrera, 1976); Monirniaeeae: Laurelia (Cabrera, 1976). Malvales. E1aeoeaqaceae:Aristotelia chilensis (Coode, 1984). Myrtalcs. Myrtaceae: Tepualia (Cahrera, 1976); Onagracoae: E p i l o b i u n ~ australe, E . niuale, E . puberulum, and Fuchsia inirgellanica (Solomon, 1982; Belly, 1989); Thymeleaci,ae i)ropetrs (Cabrera, 1976). Proteales. Proteaceae:/i'i~tii~~i/trium and Orites (Cabre- r a , 1976; Weston & C r x i p , 1987, 1994, 1996). Restionales. Restionaceat: (Ellsrnore, 1991). Rosales. Eucrmhiaeeae: Eucwphi(i (Ciibrera, 1976). Santalales.
Veronica (Cabrere, i976).
ANNELID^
Mirudinea. Glossiphoniiformes. Glossiphoniidae: Glossiphonia ~msrmhrina andflelobdella scutifera (Ringuelet, 1985). Hirudiniiormes. Americobdellidae: Arnrricobdella (Ringuelet, 1968); Semiscolecidac: Patagoniobdella (Ringuelet, 1985). ARTNROPODA. Arachnids. Araneae. Anyphaenidae: Acanthoceto cinereus, A, pi- chi, Aporatea, and Ferreria (Gerschman de Pikelin & Schiapelli, 1975; Ramirez, 1997); Dipluridae: Seotinoecus (Schiapelli & Gerschman de Pikelin, 1968); Mimetidae: Onolus (Platnick; 1993). Opiliones. Caddidae: Austropsopilio (Cokendolpher & Maury, 1990). Pseudoscorpionida. Chthoniidae: Austrochthonius chilensis (Vitali-Di Castri, 1975); Vachoniidae: Beierohisium (Qitali-Di Castri, 1970). Chilopada Geophilomorpha. Geophilidae: Schendyluides (Pereira et al., 1997; Pereira, 1998). Crustacea. Anomapoda. llyoeryptidae: Ilyoeryptus breuidentatus (Paggi, 1998). Capepoda. Canthocamptidae:Aitheyella crenulata (Menu Marque & Bosnia, 1986). Decapoda. Aegiidae: Ae& alaealufi and A. dertticulata (Morronc & Lopretto, 1994; Morrone, 1996b). Hexapoda. Coleoptera. Attelabidac: Minurus (Morrone & Posadas, 1998); Belidae: Atractuc/ius,Diconlylus, and 1Fichophtalmus(Kuschol,ubregion a n d ctsprouznces
3
Magelianic
forest
Fig. I. Provinces of the Subantarctic subregion.
Germainiellus, G e r n ~ a i n i u s , Hauersiella, a n d Sir~ophlueris (Kuschcl, 1951, 1987; Wood, 1986; Clark & Burke, 1988; Lanteri & O'Brien, 1990; Morrone, 1992b, e, 1993a, 1994a, d, 1995, 1996a; Morronc et al., 1994; Lanteri & Morrone, 1995; Morronc & Roig- Juiient, 1996); Dytisicidae: Lancetrs (Bachmann & Tr(?mouilles, 1981); Nemonychidae: Nannomacer (Kuschel, 1954, 1983; Morrone, 1996dl; Scarabcidae: Frichius (Zunino, 19841; Staphylinidae: Neqphu,~us (Thayel; 1989); Tcnebrionidae: Adelium spp. (Peiia, 18911. Dintera. Chiranomidae: Aphrateniinae, Diamesinae, and Podonomopsis-Rhrociilu,~ (Itoig- Juiient, 1994h); Ephydridao: Nutiocoenia pollinosa, Scatella neglectus, and S , sturdeear~us (Mathis, 1980); Mvcetoiihilidae: Echinupodiurn spp. (Duret, 1975, 1976); ~ y d i d a e : Mitrodetus ( ~ r t i ~ a s & ~ a p a v e r o , 1990); Sirnuliidae: Cnesiamima, Gigantadax antareticus, G. cannenae, G. fe,ni,ieus, G. ignieulus species group, G. margi,miis, G. rufkscens, G. r.ufidulu~, G. trifidus, and Parastrosi,nulium (Wygodzinky & Coscarbn, 1989; Coscar6n, 1991); Tipulidae: Elnoretta, Euualdiuiana, and Valdiuiana (Jong, 1989). Hemiptera. Corixidae: Sigara egbcrtae, S . trimaculata, and S . vuriloehe (Bachmann, 1981); Eriococcidae (Humphrios et ol., 1986; Seberg, 1991); Peloridiidae: Pantinia spp. and Peloridiurn spp. (Burckhardt & Agosti, 1991); lleduviidae: Microtornus gayi (Giacchi & Coscarbn, 1992). Hymenoptera. Diapriidae: Gladieauda (LoiBcono, 1988); Formicidae: Lasiophanes (Kusnezov, 1951); Pompilidae: Chirodamus agenius a n d Sphictustethus (Roig-Alsina, 1985, 1987); Tiphiidae: Cosila chilensis (Cenise, 1984). Lepidoptera. Cossidae: Chileco~nadia (Gentili, 1989; Schoorl, 19891; Ilepialidae: Andeabatis, Calada, Callipielus, Dalaca, and Parapielus (Nielsen & Itobinson, 1993); Micropterigidae: Heterobathmia spp. (Kristensen & Nielsen, 1979; Ilumphries et al., 1986); Noctuidae: Paraeuzoa (Jana-SBenz, 1989; Angulo, 1990); Paiaephatidae (Davis, 1986; Grehan, 1991); Pieridae:
Hypsochila argyrodile (Field & H e r r e r a , 19771. Neuroptera. Memerobiidae: Conchopterella, Gayomyia, Hemerobius chilensis, H. nehoi, H. stcnopterus, and Megalomus nigratus (Oswald, 1993; Monserrat, 1996, 1997). Odonata. Austropetaliidae: Rheopetalia rex (Carle, 1996); Cordu1idae:Rialla uillosa ( M u z h , 1995); Libellulidae: Sympetrunz uillosurn (Muzbn, 19951; Neapetalidae: Neopetalia punctata (Muzbn, 1995); I'etaluridae: Phenes r. raptor (Muzbn, 1995). Orthootera. Tristiridae: Tropidostethini (Cigliano, 198961. Plecaptera. ~ m p h i p n o i d a e (MU& & Bachmann, 1998); Eustheniidae: Neuroperla escheridingi and Neuroperlopsis patris (Bachmann, 19951; Notonemouridae: Neofulla, Neonemura, and Udamocercia (Bachmann, 19951. Trichoptera. Hydrobiosidae (Angrisano, 1995); Hydropsychidae: Srnicridea dreora, S . rrrircronata, S . penal, and S . pucara (Flir~L, 1989). VF,KTEBRATA. Amphibia. Anura. Leptodactylidae: B<rtrachyla leptopus (Barrio, 1967a); Rhinodermatidai?: Rhinoderma (Williams & Eeheverria, 19951. Aves. Anserifomes. Anatidae: A n m specularis, Chloephaga hybrida, C , poliocephula, C. rubidiceps, Tachyeres pataclionicus, and T . pteneres (Navas, 1977; Olrog, 1984; PeBa, 1992). Ciconiiformes. Accipitridae: Ruteo oenlralis (Sibley & Monroe, 1990); Faiconidae: Phalcobaenus albogularis (Olrog, 1984); Phalacrocoracidae: Plialacrocorax atriccps (Olrog,
(Olrog, 1984). Gruiformes. Rallidae: Gallinula Inelnnops (Navas, 1.991). Passerifomes. Fringillidae: Carduelis barbata (Olrog, 1984); Furnariidae: Apizrastura spinicauda, Cinelodes antarcticus, C. oustaleti, Pygarrhichus albogularis, a n d Syloiorhynchus des,nursii (Miiller, 1973; Olrog, 1984); Muscicapidae: Turdus falcklar~dii (Olrog, 19841; Rhynacyptidae: Pteroptochus tanzii (Olrog, 1984). Psittaeiformes. Psittucidae: Enicognalhw lkrrugineus and E. leptorhynchus (Olrog, 1984). Troehiliformes. Trochilidae: Sephanoides sephanioides (Olrog, 1984). Mammalia. Artiodactyla. Cervidae: Hippocornelus bisulei~s and Pudu puda (Redford & Eisenberg, 19921. Carnivora. ~clidae:>elisguigna (Redford &
ise en berg,
19921. Chirootera. Vespcrtilionidae: Myotis ehiloe~lsis (Redford
&
ise en berg,
1992). ~ i c r o b i o t h e r i a . Microbiothcriidae: Dromiciops australis (Redford &Eisenberg, 19921. Rodentia. Muridae: Abrothrir longipilis, Auliscomys micropus, Chelwnys maeronyx, Euneomys chinchilloides, Georrcs uuldiuianus, and
lrencrnys tarsalis Wedford & Eisenberg, 1992).
Maule
province(Figs. 1 J 3 )
S o u t h e r n Chile a n d Areentina. between
-
34-37' south latitude. T h e t r a c k of ~ h a e t a n t h e r a serrata (Fig. - 3 ) is representative of this province.
4 Reuzsta del Museo Argentcno d e Cc
Willink, 1973: 99; Cabrera, 1976: 731, Valdivian region (O'Brien, 1971: 2031, Maule district (Ca- b r e r a & W i l l i n k , 1 9 7 3 : 9 8 ) , V a l d i v i a n a r e a (Artigas, 1975: map), Valdivian province (Rivas- Martinez & Navarro, 1994: map), Maule (Roig- J u i i e n t , 1 9 9 4 b : 1 8 1 ) , a n d M a u l e p r o v i n c e (Morrone, 1999: 14).
V e g e t a t i o n . Transitional temperate forests, w i t h s o m e biotic e l e m e n t s from t h e C e n t r a l Chilean subregion. Dominant piant species a r e Acaenu p i n n a t i f i d a , Alstroemeria a u r a n t i u c u , Aristotelia chilensis,Boccharisconcaoa, Berberis
buxifolia, Boquila trifoliata, Chusquea couleu, C o r t a d e r i a p i l o s u , C r y p t o c a r i a m a m m o s a , Escallonia uirgata, L a p a g e r i a rusea. Laureliu s e m p e r u i c e n s , M y r t u s l u m a , N o t h o f a g i t s alexandri,
N
domlieyi,N
leoni,N.
obliqua,N.
pltmilio, Pernettya m u c r o n a l a , Perszu lingue, P o d o c a r p u s n n b i g e n a , R h a p h i d h a ~ n n z u s spinosum, andRibes magellanica (Cabrera, 1971, 1976; Cabrera & Willink, 1973). There are small a r e a s with forests ofAraucaria a r a u c a n a (Peiia, 1966; Cabrera & Willink, 1973).Taxa.
CONIFEROPHWA. Coniferopsida.Coniferales. Araucarinceae: Araucaria araucana (Ca- brera, 1976; Cabrera & Willink, 1973; Covas, 1995b). MAGNOLIOPHYTA. Magnoliopsida. Asterales. Asteraceae: Chaelanti~era linearis var taltalensis, C. serrata, and ZFiptilio~z achilleue (Morrono ct al., 1997). ARTIIROPODA. A r a c h n i d a . Araneac. Anyphaenidac: Aca,~thueeto ladormida (Ramirez, 1997); Dipluridae: Seuti~zoecus cinereopilnsus (Schiapelli & Gerschman de Pikelin, 1968); Gnaphosidae: Apodrns.sode,s mercedes, A. P I L C O ~ , and
Eciiernoides rnalleco (Platnick, 1983; Flatnick & Sliiidab, 1983); Migidae: Malleco,nigas (Goloboi'i & Platnick, 1987); Nemesiidae: Aeanti~ogo~zotus brunnew, A. hualpen, A. mulchen, A. nai~uelbuta, A. rucintu, and A. tolhuaca (Goloboff, 1995). Opiiianes. Triaenonychidae: Arar~canobunus (Mufioz, 1973). Pseudoscorpionida. Chthonidae: Sathruchtlcoriiiis pefhuri (Vitali-Di Castri, 1974). Cmstacea. Dccapoda. Aeglidae: Aegla baiiamondei, A. erpansa, and A. speetabilis (Jara, 1992; Bond-Buckup & Buekup, 1994). Hexapoda. Coieoptera. Anthribidae: Dinoee,~trus signatipes (Morrone & Roig-Juiient, 1995); Beiidae: Atractuciius a r g u s , Callirhynchi,ius rxyuisitus, Dicordylus balteatus, and Oxycrrrspedus (Kuschcl, 1959; IUgueta, 1986; Morrone & Raig-Jufient, 1995; Morrone & Posadas, 1998); Bi~prestidac: Mastogenius sulcicollis, Mendizabalia g. geierrnair~i, Pterobothris c . corrosus, and l'rigorioge~ziunt angulosurn ruginosurn (Bellamy & Moore, 1990; Roig-Juficnt, 1994b); Carabidae: Barjpusparalellr~s, Cero~lossus c. chile,csis, C. e. temucensis, C.darwini lnagrllanicus,C. ualdiuiar subnitens, Cnemelobus gerjnair~i, Cyliridern ci~ilunsis, andNotliobroscus (Roig-Juiient & Ciechino, 1989; Roig- Juiient, 1990, 1994~1, b; Roig-Jufient & Ball, 1995); Caridae: Caeno~ninurus (Kuschel, 1992; Morronc &
rnccas Nalurales, n. s. 2
il),
2000Roig-Juiient, 1995; Morrono & Posadas, 1998); C1eridac:Eurymetopum frigidus,E.gayi, andE.uittula (Solervieens, 1886); Curcuiionidae: Argorhinus albolineatus, A. bouiei, A. nitens, A. suturnlis, Antl~onumus aruucanus, A. cl~ilicoln, Araucarietus, Amucarius chilensis, A. major, A. medivs, A. minor, Caluertius, C y l y d r o r i ~ i ~ ~ v s iriaeyuatus, Dosyden~a
a i i n ~ ~ c e l l a , E i ~ i ~ ~ g i u ~ , E u ~ ~ 1 ~ ~ tesselati~s, G e n i o ~ ~ ~ e ~ n n u s chilie,isis; Hybreoleptops aureosignatw, Lan~iarhinus horridus, Listroderes breuirostris,Megialorr~etides spp., Megalov~elis andigena, Nothofagi,ioides, Ornuides validus, Planus, Polydrusus roseus, Rhyepl~erres b.ayi, R . lateralis, Tartarisus perforatipennrs, and T. subfnscintus (Kuschei, 1952, 1966; May, 1967; Riihm, 1987; Clark & Burkc, 1988; Morrone, 199Za, 199Ga, e; Morrone & Roig-Juiienl. 1995); Endornychidae: Chileobius cekalouici aiiri (:, notatus (Pakaluk & Slipinski, 1990); Megalapodidae: Palopitagoides (Kuschcl & May, 1996); Ncnionychidae: Mecomaeer, Na,o~ornoei.r gcrmaini, Ithynclcitornacer, and lthyriciiito~naterinus (Kusi:hel, 1954, 1959, 1983; Morrone & Roig-Juiient, 1995). Ilyrnenoptera. Brudynobaenidae: Bradynobaenus australis (Genisc, 1086); Formicidae: Pogonomynnez uerrniculatus (Kusneaov, 1978). Lepidoptera. Choreutidae: Nyr (I-Ieppncr, 1982); Cossidae: Andesiana, Austrocossus, Sel~ausisea,S~~rcossus, andl%ilariglaw (C-entili, 1989; Schoorl, 1989); Palaepbatidao: Metapiiatus cirrhus,M. siriuatus,Palaephatus latus, andP, leueao.utus (Davis, 1986). Odonuta. Austrupetaliidae: Ophiuprtalia araucanu and 0. pudu (Carlc, 1996). Orthoptera. Tristiridae: Elysiacris angusticollls (Cigliano, 1989a; Roig-Juiient, 1994b). Trichoptera. Hydropsyehidae: Smicridea eorr~plicatissima,S. redunca,S. tregala,and S. turgida (Flint, 1989). VERTEBIMTA. Amphibin. Anura. Leptodactylidae: Telrnatobufo bullucki (Cei, 1973). Mammalia. Rodentia. Gtenomyidae: Ctenomys maulir~ud (Redford & Eisenberg, 1992); Octodontidae: Aeonaemys sagei and Oelodun bridgesi (Redford & Eisenberg, 199#.
Valdivian p r o v i n c e (Figs. 1 , 4 )
Southern Chile a n d Argentina, s o u t h o f t h e Maule province, reaching 47' south latitude. The track of Crirzodendron hookerianum (Fig. 4) is representative of this province.
S y n o n y m s . Valdivian forest (Soriano, 1950: 33; Kuschel, 1960:541;Roig-Jufient, 1994b: 1811, Subantarctic forests subregion (Hueck, 1957: 40), hlountain zone (Kuschel, 1960: 54.9, Valdivian region (O'Rrien, 1971: 2031, Valdivian forest region (Pefia, 1966: 141, Valdivian cordillera r c g i o n (Pefia, 1966: 151, Caducifolius forest district (Cabrera, 1971: 37. 1976: 73; Cabrera &
Morrone The Subantarctic subreg~on a n d zts prouznces
5
6 Reursta del Museo A r g e n t ~ n o de Czenczas Naturales, n. s. 2 (l), 2000
Marcinez & Navarro, 1994: m a p ; Morrone, 1999: 141, and Valdivian temperate fbrests ecoregion (Dinerstein et al., 1995: 101).
V e g e k a t i o n . Temperate forests (Cabrera &
W i l l i n k , 1973). D o m i n a n t plant species are A n e m o n e m u l t i f i d a , A u s t r o c e d r u s c h i l e n s i s , Berberis spp., Blechnum chilense, Chrysanthe- m u m leucanthemum, Chusquea couleu, Colletia s p i n o s a , C o r i a r i a r u s c i f o l i a , D a s y p h y l l u m d i a c a n t h o i d e s , D i o s t e a j u n c e a , E u c r y p h i a c o r d i f o l i a , F a b i a n a i m b r i c a t a , F r a g a r i a chiloensis,Fitzroya cupressoides, Geuuina auella- n u , L a u r e l i a p h i l i p p i a n a , L o m a t i a h i r s u t a , Maytenus boaria,Mutisia decurrens,M. spinosa, Myrceugenella apiculala, Nothofagus antarctica, N . hetuloides,
N.
domheyi,N.procera,N obliqua,N .
procera,N.
pumilio, Podocurpus nubigenus, Scirpus californicus,Senecio microcephalus, and Weimannia trichosperma (Soriano, 1950; Cahre- ra, 1971, 1976; Cabrera & W i l l i n k , 1973).T a x a . C O N I F E R O P f I Y T A . C o n i f e r o p s i d a . Coniferales. Cuprcssaceae: Austroeedrus and Fitzroya (Cabrera, 1976; Covas, 1 9 9 5 ~ ) ; Podoearpaeeae: Podocarpus nubigenus andSaregothaea (Covas, 1995a). NLAGNOLIOPHYTA. Magnoliopsida. Asteralcs. Asteraceae: Chaetantizera brael~ylepis and C. elegar~s var. rlegairs (Morrone et al., 1997). Maivales. Elaeocarpaceae: Criiiodendron hookeriariurn (Coode, 1987). Myrtales. Onagraceae: Epilobium obscurum (Solomon, 1982). Santalales. Misodendraceae: Misodendrum anguEatum,M, braehystachyu,n, andM. gayanurn (Zavaro et al., 1997). M O L L U S C A .
(Platnick & Forster, 1987); Piemcsiidae: Acanthogonatus coufusus, A . franki, and A .
subcalpeianus (Goloboff, 1995). Opiliones. Triaenonychidae: Diasia (Maury, 1987). Crustacea. Decapoda. Aeglidae: Aegla araucaniensis, A . riolimaya~~a,A. d. dentieulata,A. d. lacustris,A. manni, and A. rostrata (Band-Buckup & Buekup, 1994; Morrone & Lopretto, 1994). Hexapoda. Coleoptera. Anthrihidae: Dinocentrus tuberculosus and Sistellorhynei~us postieallis (Morrane & Roig-Juiient, 1995); Brentidae:Apionpacizymerurn (Morrone & Raig- Juiient, 1995); Carabidae: Ceroglossus ualdiuiae chiloensis, C. d . darwini, C. sprciosus, Creobius euduuxy, Cylinders gortnazi, Nothanilll~s luisae, Notaphus stenoderus, Thalassobius testaceus, 7kechisibus obtusiuseulus, and T oerobates (Jeannei, 1962; Roig-Juiient, 1994b); Cleridae: Euryrnetopum proteus (Roig-Juiient, 1994b; Solervicens, 1986);
Curculionidae: Aegorhinus fascicularis, A, kuscheli,A. ?naer;tus, Anthonornus berberidis, Berberidicola cren!ilatus, Cnernocorlus ualdivianus, Epaetius, Euealus fasciolatus,E. unicolor,Falklandius chilensis, P.peckorurn,Gayus elegans,Germainiellus attenuatus, G.pur~ctiuentris,IIeter~nragdalis heteronyz,Listroderes
obrieni, Myelobius biuculatus, M. faseiolatus, N e o p s i l o r h i ~ ~ u s , Nothofagius, Nothofagobius, Nototactus, Pachytrogus crassirostris, Philippius, Puranius fasciculiger, Rhyephenes clathratus, Tartarisus griseus, and Wittmerius (Elgueta, 1985; Clark & Burke, 1988; Marrone, 1990,1993a, b, 1996a, e; Morrone &Anderson, 1995; Morrone & Roig-Juiient, 1995; Morrone rt al., 1997); Nemonyehidae: Nannornacer wittrneri (Kuschel, 1959; Morrone & Roig- Juiient, 1995); Scarabeidae: Frickius variolosus (Iiowden, 1982); Staphylinidae: Pseudopsis adustipennis (Herman, 1975). Diptera. Simuliidae: Gtgantodax igniculus (Wygodzinky & Coscarbn, 1989); Tipulidae: Yaldiviaria edwarsina and Y . shannonL~a (Jong, 1989). Hymenoptera. Formicidae: Anthichyhonidris bidentatus (Roig-Juiient, 1994b): Ichneumonidae: N o t o p h r i ~ d o s (Porter, 1993). Lepidoptera. Palaephatidae. Apophatus, Metaphatus ichnius, Palaephatus a1biti.i-ruinus, P. arnplisaccus, P. fusciterrninus, P. luteolus, P nielseni, P. spinosus, P. striatus, Sesornrnata albirnaculata, S . leuroptera, and S . paraplatysaris (Davis, 1986; Roig-Juient, 1994b). Neuroptera. Hemerobiidao: Megalomus dernocratieus and M. stangei (Monserrat, 1997). Odonata. Austropetaliidae: Ophiopetalia diar~a (Carle, 1996). Orthoptera. Acrididae:Naiiuelia nnthrucina (Ronderos & T u r k , 1989). Trichoptera. Hydrobiosidae: Australobius ( S c h m i d , 1989). V E R T E B R A T A . A m p h i b i a . Anura. Leptodactylidae: Batrachyla antartandica, Hylorirta syluatica, and Telmatobufb australis (Barrio, 1967a. b; Cei, 1973). Mammalia. Paucitubercuiata. Caenolestidae: Rhyncholestes raphanurus (Redford & Eiseherg, 1992; Contreras & Yafiez, 1995). Rodentia. Muridae: Abrothrix sanborr~i (Redford & Eisenberg, 1.992).
M a g e l l a n i c
forest
p r o v i n c e (Figs. 1,4)Southern Chile f r o m 47' south latitude to Cabo de Hornos, andsouthcrn Argentina i n small portions o f western S a n t a Cruz and Tierra del Fuego. T h e track ofGigantodru briophyi (Fig. 4) i s representative of t h i s province.
Morrone: The Subantarctic subregion a n d its provinces 7
V e g e t a t i o n . M o r e x e r i c f o r e s t s , w i t h a b u n d a n t Nothofagus betuloides t r e e s , a n d evergreen s w a m p forests (Cabrera & Willink, 1973; Dinerstein et al., 1995). Dominant plant species include Drimys winteri, Embolhrium coccineum, Maytenus magellanicus, Nothofagus a n t a r c t i c a , N . b e t u l o i d e s , a n d N . p u m i l i o (Rothkiigel, 1916; Kuschel, 1960; Tuhkanen et a l . , 1990).
T a x a . CONIFEROPHYTA. Coniferapsida. Coniferales. Podocarpaceae: Lepidotham~zus fonkii (Covas, 1995a). MAGNOLIOPHUTA. Magnolia- psida. Myrtales. Onagraceae: Epilobium conjiigena (Solomon, 1982). ARTHROPODA. Arachnids. Araneae. Dipluridae: Seotinoecw fasciatus (Schiapelli & Gerschman de Pikelin, 1968). Nexapoda. Coleoptera. Carabidae: Antaretonornus eomplanatus, Cascellius aeneoniger, C. grauesii, Notaphiellus cchaiouici, Notaphus knscheli, Notholopha otrum, Peryphus rufoplagiatus, Pseudurnigadops oualis, Dechisibusantarcticus, and% hornesis (Jeamel, 1962; Roig-Juiient, 1994b, 199513); Curculionidae: Antarctobius gerrnaini, A. hyadesii, A. lacunosus, Cylydrorhinus ursinus, C. uittatus, Falklundinpsis, Germainiellus Eoeuirostris, and G , lugens (Morrone, 1992b, 1993%; Morrone & Anderson, 1995; ~Morrone & Roig-Juiient, 1995); Perimylopidae: Parahelops yuadricollis (Roig-Juiient, 199410). Legidoptera. Noetuidae: Paraeuzoa perdita (Jana-SBenz, 1989; An- gulo, 1990). Neuroptera. IIemerobiidae: Megalomus australis (Monserrat, 1997). VERTEBRATA. Mammalia. Rodentia. Muridae: Abrothrix lanosus (Redford & Eisenberg, 1992).
Magellanic moorland province (Figs. I, 5)
Southern Chile a n d Argentina, limited by the Maeellanic forest nrovince to t h e east.
-.
The ~~ ~ track of Notocascellius hyadesii (Fig. 5) is representa- tive of this province.Synonyms. Magellanic moorland (Godley, 1960: 467), Magellanic moorland zone (Kuschel, 1960: 5441, Southern Pacific region (Pefia, 1966: 16), Magellanic region (O'Brien, 1971: 204), Aus- t r a l P a c i f i c z o n e ( C e k a l o v i c , 1 9 7 4 : 3011, P a t a g o n i a n i c e zone (Cekalovic, 1974: 3031, M a g e l l a n i c a r e a ( A r t i g a s , 1 9 7 5 : m a p ) , Anstroandean province (Rivas-Martinez & Na-
varro, 1994: map), Fuegian province (Kivas- Martinez & Navarro, 1994: map), Eastern forest (Roig-Jufient, 1994b: 1821, Subpolar Nothofagus forests ecoregion (Dinerstein et al., 1995: 101), a n d Magellanic moorland province (Morrone, 1999: 14).
Vegetation. Moorlands (Cabrora & Willink, 1973). Dominant plant species include Astelia pumila, Donatia fascicularis, Drimys winteri, G a i m a r d i a a u s t r a l i s , ilymenophyllnm spp.,
Nolhofagus betuloides,
N.
pumilio, Oreobolus o b t u s a n g u l u s , S c h o e n u s a n t a r c t i c u s , a n d S p h a g n u m m a g e l l a n i c u m ( K u s c h e l , 1 9 6 0 ; Tuhkanen et al., 1990).Taxa. ARTNROPOUA. Nexapoda. Coleoptera. Brcntidae: Apion fuegienum (Cekalavic, 1974); Carabidac: Ceroglossus s . suturalis, Feroniomorpha lucida, Metius malachitichiis, Mimodrornius nigmtestaceus, and Notocascellius i~yadesii (Xoig- Juiient, 1994b, 199Sb); Caridae: Chilecar (Kuschel, 1992; Marronc & Roig-Juiient, 1995); Curculionidae: Aegorhinus vitul~s,Antaretobius rugirostris, A. yefael, Berbe~idir.02~ anratus, and 3iilurus (Cekalovie, 1974; Morrone, 1992b; Morrone &Anderson, 1995; Morrone & Roig-Juiient, 1995). Diptera. Chironomidae: Paroclilus pilusus (Roig-Juiient, 199413); Scatopsidae: Diarnphidicus chilensrs,(ilmorim, 1990); Simuliidae: Cigar~todax antarcticus. (>. brophyi, and G. rufidulus (Roig-Juiient, 1994b). YERTEBRBTA. Mammalia. Rodentia. Muridae: Abrothrk hershhovitzi and A. marhFumi (Redford & Eisenberg, 1992).
Malvinas islands province
Argentinean archipelago of t h e Malvinas or F a l k l a n d islands a n d South Georgia island, situated i n the south Atlantic Ocean, about 550 km from Tiorra del Fuego.
Synonyms. Insular province (Cabrera, 1951: 23, 1953: 107, 1958: 200, 1971: 40; Cabrera &
W i l l i n k , 1973: 1 0 2 ; C a b r e r a , 1976: 78), Patagonian grasslands ecoregion (Dinerstein et al., 1995: 1081, Cortaderal Malvinian district (Roig, 1998: 1.421, T u n d r a or Georgias district (Roig, 1998: 1421, a n d Mavinas islands province (Morrone, 199% 14).
Vegetation. Grasslands, steppes, and tundra (Cabrera & Willink, 1973; Dinersteinet al., 1995). D o i n i n a n t p l a n t s p e c i e s a r c A b r o t a n e l l a ernarginata, Acaena adscendens,A. magellanica, A. microcephala, Astellia p u m i l a , Azorella lycopodioides, A. selago, Baccharis magellanica, Blechnzcm tabulare, Bolax gummifera, Caltha appendiculata, Carex trifida, Cortaderia pilosa, Deschumpsia flexuosa, Empetr-urn r u b r u m , Festuca erecta, G a i m a r d i a a u s t r a l i s , J u n c u s sheuzerioides, Pernettya pumila, P o a annua, P . antarctica,
P.
flabellata, Pratia repens, Senecio candicans, S. littoralis, S . uaginata, Uncinia smithii, andveronica elliptica (Skottsberg, 1921; Davies, 1939; Moore, 1968; Cabrera, 1971,1976; Cabrera & Willink, 1973).8 Rrvzsla del MweoA?.gent!no rlr Czenczas Naturales, n. s. 2
ill,
2000(Cazzaniga, 1995). ANNELIDA. Hirudinea. Bryaceae: Bryurn fer,~a,~dezianurn (Mahii, 1995); Gloesiphoniifonnes. Glossiphoniidae: Adaetobdella Cryphaeaceae: Cyptodon cmssineruis (Mahu, 1995); rr~alvir~er~sis (Ringueiet, 1985; Lo~lretto, 1995a). Dicranaeeae: Ca,npyloous aberra,is. C. blindioides,
1992). C r u s t a c e a . Amphipoda. Gamrnaridae: Felklandella and Prae,fklklanddla (Stock & Platvoet, 1991; I,opretto, 1995b); 1lyallelidae:Hyalrlla ,ieo,zuma (Stock & P1atxroct, 1991). Anostraca. Branchinoctidao: Bmrr.chineeta gain' (Cohen, 1998). EIexapoda. Coleoptera. Byrrhidae: Ci~alcosphaeriurn soloz and C. erzderleirii (Voisin, 1987); Carabidae: Pseudomigadops darwini, and Y falklandicus (Jeannel, 1938, 1962; Roig-Juilent, 1994bj; Curculionidae: Ar~tarctobii~s abditus, A. bidentatus, A. falhla,~dieus, Caneortibtus bia,igulotus, C , uretai, Cylydrort~inus le,n,ziseatics, Fulkla,idius goliath, F. huscheli, F , tuirbificati~s, Gerrnainiellus salebrusus, Lartteriella, Maluinius. Puranius cl~amplioni, P. exsculpticollis, and P. seaber (Robinson, 1984; Voisin, 1987; Morrone, 1992b, c, 1993a, 1994~; Marrane &Anderson, 1995; Mmrone & Roig-Juiient, 1995:; Leiodidae: Falhucholei~a falkland~ca and Falko,ze~nadiis spherzisei (.Robinson,
1984; Voisin, 1987); Perimylopidac: Darwiiiella arnnrodies, Parah~lops . . fhlkla,~dicus. P. havemi. and P . qi~odrieollis (Vaisiri, 1087; Florcs, 1998); Pythidae: Poopizylax falhlandica (Voisin, 1087). Diptera. Ephydridae: Seatella nvglrcta and 9 . stiirdra,ra (Lizarralde de Grosso, 1998). Orthoptera. Gryllacrididae: Pari~denus falhla,idicus (Robinson, 1984). VERTEBRATA. Avos.Anserifonncs. Anatidae: Anas g. georgiea, Ctiloept~aga pieta leucoptern, and Tuehyeres brachypterus (Navas, 1977; Olrog, 1984). Ciconiiformes. Frrlconidae: Phalcobaei~us aiistratis (Olrog, 1984). Mammalin. Carnivora. Canidae: Dusicyon australis (Nowak. 1991).
Juan Fernandez islands province
Chilean i s l a n d s of M a s a t i e r r a (= llobinson Crusoe), Masafuera (= Alejalidro Selkirk), a n d S a n t a C l a r a , s i t u a t e d i n thc Pacific Ocean, 600
km
west of Valparaiso, a t 33" south latitude. Synonyms. Juan F e r n a n d c z province (Ca- brera & Willink, 1973: 103), Fcrnandezian region ( T a k h t a i a n , 1986: 252). a n d J u a n F e r n 8 n d e z. . . (Mahii, 1995); Fissidentaceae: Fissides crassieuspes and I: feriiandrzianurn (Milhd, 1995); Hookeriaceae: Ao.ophyllum ter~icineroe (Mahd, 19951, Distichophy- llurn subelimbatum (Mahli, 1995); Neckeracone: Neckera roturtdata, Pilzrratella rnicrusticta, and
Tharnraobryium caroli (Mrihri, 1995); Orthotricaceae: Ulota fernandeziann (Mahli, 1995); Ptychomitriaceae: P t y c i ~ o r n i t r i u n ~ fer,iandezianurn (Mahii, 1995); Racopilaceae: Racopilum fer,randezinnu,n (Mahli, 1995); Thuidaceae: TiLuidii~m Jnasafurrae (Mahii, 1995). PILICOPllYTA. F i i i c o a s i d a . Fiiicales. Dicksoniaceae: Dichsoriia Li'rturiar~a (Murillo, 1988). MAGNOLTOPNYTA. Liliopsida. Arecales. Arecaeeae: Juaniaaustmlis it:abrera &Willink, 1973). Mamoliopsida. Asterales. Asteraccae: Centauroden- d r o n , Dendruseris, Phaenicoseris spp., Rea, Rhetinodeiidron, Robinsor~ia, and Yunquea (Cabrera & Willink, 1073; Stuessy et al., 1984,1990; Sanders et
e l . , 1987; Crawford r t al., 1992). Campanulales. Campanulaceae: Wahle,abergia fernandideziana coinpiex (Lammers, 1996). Caryophyllales. Chenopodiaccae: Cizenopodium snnctaeclarae (Stuessy rt al., 1990). Euphorbiales. Euphorbiaceae: Dysopsis hirsuta (Stuessy rt al., 1984). Magnoliales. 1,actoridaceae (Stuessy et al., 1984). Myrtales. Myrtaceae: Ugni selkirizii (Stuessyet al., 1984).Verbcnales.Verbenaceae:
Ri~apizitarnnus oe,~ustus (Rodriguez et al., 1983). Violales. F1aeourliaceae:Azara serrata (Stuessy et at., 1984). AttTHROPODk Wexapoda. Archaeognatha. Ki~sciielochilis (Wygodzinsky, 1.967; Camousseight, 1995a). Coleoptera. Carabidae: Tracicysarus (Straneo & Jeannel, i953); Curculionidae: Ariolethrus,
Juanorhinini,Pacicystylw, andStrorigylopterus ouatus (Kuschei, 1952; Wibmer & O'Brien, 1988). Diptcra. Simuliidae: Gigantodax kuscheli (Wygodzinky & Coscardn, 1989). Neuroptera, Hemerobiidae: Co,~chopterella izuscheli and C , maculata (Oswaid, 1993). Thysanoptcra. Thripidne: Physothrips shottsber,qi (Prado & Muiioz, 1995). Thysanura. Lepismatidac: Isolepisma annectens (Camousseight, 1995hj.
islands province (Rivas-Martinez & Navarro,
1994: map; Morrone, 1999: 14). ACKNOWLEDGMENTS
Vegetation.
F o r e s t s , s h r u b l a n d s , a n dmasslands (Cabrera . & willink. ,9731. ~ ~
1
a m grateful to ~ iPaula
~Posadas a n d Sergio ~ ~ t, ~ ~- ~~-
p i a n t s p e c i e s include
A~~~~
fernandeziana, Roig-Juiient for critically c o m m e n t i n g on acoprosAa
hookeri, ~ i kfemandeziana, ~ ~ ~ ipreliminary draft of t h e manuscript. ~ Drimysconfirtifalia,Dysopsis hirsuta, Empetrumr u b r u m , Escallonin callcotliae, i j h g a r a nluyu, BIBLIOGlZAPHY Juania australis, Myrceuge~zia schulzei, Pernettyo
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