The Biogeography and Evolution of the Asteraceae
in the Atacama and Peruvian Deserts

Michael O. Dillon
Department of Botany
Center for Evolutionary and Environmental Biology
Field Museum of Natural History, Chicago, IL 60605-2496

and

Todd Vision
Department of Ecology & Evolutionary Biology
Princeton University
Princeton, New Jersey

ABSTRACT

INTRODUCTION

Occupying roughly 3500 kms or nearly one third the length of the western coast of South America, the Peruvian and Atacama Deserts are among the driest on earth and exert a profound effect upon the meager flora of this region (Rundel et al. 1990). The flora of this area is largely restricted to fog dependent, lomas formations and has been estimated at over 1100 species of flowering plants, gymnosperms, and ferns. Within the desert formation from northern Peru to central Chile, the family Asteraceae is represented by 53 genera and nearly 120 species. These taxa range from pantropical weeds to narrow endemics; however, the average number of species per genus is frequently quite small.

With some 16 species, the genus Senecio has the greatest number of species. This paper is a detailed accounting of the diversity patterns exhibited by the Asteraceae within the lomas formations, and provides discussions of the probably origins for the various genera.

Study Area

The greater part of the western coast of South America [5•00'-29•55'S] is occupied by the Atacama and Peruvian Deserts which form a continuous belt for more than 3500 km along the western escarpment of the Andes from northern Peru to northernmost Chile. The phytogeography and ecology of these deserts have been reviewed in detail (Rundel et al. 1991) and a floristic inventory of the entire area has been completed (Dillon, in prep.). While the desert is continuous from Peru to Chile, the topography, climate, and vegetation of each desert is distinct. This aridity is controlled by three climatic anomalies (Trewartha 1961, Johnson 1976). 1) An abrupt climatic transition both to the north and south results in a poorly developed steppe climate along the margins; 2) brief periods of heavy rainfall and relatively high temperatures associated with rare, but recurrent, El Niño events (Dillon 1985, Dillon and Rundel 1989, Quinn et al. 1987) occasionally affect parts of the desert, bringing wet tropical conditions; and 3) there is remarkable temperature homogeneity along the entire latitudinal extent of the deserts. This pattern of temperature stability results from the influence of cool, sea-surface temperatures associated with the south to north flow of the Humboldt (Peruvian) Current. Also important is the influence of strong atmospheric subsidence associated with a positionally stable, subtropical anticyclone (Trewartha 1961). The result is a mild, uniform coastal climate with the regular formation of thick stratus cloud banks below 1000 m during the winter months (Prohaska 1973).

Where coastal topography is low and flat, this stratus layer dissipates inward over broad areas with little biological impact, but where isolated mountains or steep coastal slopes intercept the clouds, a fog-zone develops with a stratus layer concentrated against the hillsides. These fogs, termed garúa in Peru and camanchaca in Chile, are the key to the extent and diversity of vegetation throughout the deserts of the western coast. This moisture allows the development of fog-zone plant communities termed lomas formations, literally "small hills", between sea-level and 1000 m. Other authors have referred to these communities as the "fertile belt" (Johnston 1929), "fog oases" (Ellenberg 1959) or "meadows on the desert" (Goodspeed 1961, Goodspeed and Stork 1955). In Peru and northern Chile, there exist over 60 discrete localities (Figure 1) which support vegetation, including Peruvian offshore islands such as Las Viejas, San Gallan, and San Lorenzo. While the actual desert covers nearly 200,000 sq. km., the actual area covered by vegetation, even during periods of maximum development, is probably less than 4000 sq. km.

Lomas formations occur as discrete communities or islands of vegetation separated by hyperarid habitat devoid of plant life. Since growth is dependent upon available moisture, climatic patterns determine plant distributions. Topography and substrate combine to influence patterns of moisture availability and area of suitable habitat. The ecological requirements and tolerances of individual species ultimately determines community composition and species endemism exceeds 40% (Rundel et al. 1991).

Current estimates (Dillon unpubl. data) for the total number of species represented within the Peruvian lomas formations are 72 families, 284 genera, and 557 species; and total numbers within the Atacama formation are 73 families, 196 genera, and 373 species. Of the total of nearly 1000 species, only 68 species (ca. 7%) are distributed within both Peruvian and Chilean formations. The families containing the greatest diversity from throughout the Atacama and Peruvian Deserts include, Asteraceae (57 genera, 147 spp.), Cactaceae (14 genera, 92 spp.), Nolanaceae (2 genera, 79 spp.), Leguminosae (26 genera, 71 spp.), Solanaceae (19 genera, 64 spp.), Malvaceae (9 genera, 56 spp.), Apiaceae (13 genera, 51 spp.), and Boraginaceae (7 genera, 51 spp.).

The Asteraceae, with 54 genera and 122 species, is the most diverse family within the Atacama and Peruvian deserts. All tribes common to western South America are represented within the lomas communities (Table 1). The most diverse tribe is the Heliantheae (16 genera, 25 spp.), followed by Mutisieae (10 genera, 21 spp.), Astereae (6 genera, 14 spp.), Inuleae s.l. (5 genera, 7 spp.), Eupatorieae (4 genera, 17 spp.), Lactuceae (3 genera, 4 spp.), Liabeae (2 genera, 3 spp.), Cynareae (1 genus, 5 spp.), and Anthemideae (1 genus, 2 spp.). For most genera, the species diversity is low, and only four genera are represented by five species or more, including Senecio (16 spp.), Ophyrosporus (9 spp.), Polyachyrus (6 spp.), and Centaurea (5 spp.)


These representatives are a complex mixture of species from various origins and generally referred to the following broad categories: 1) wide-ranging pantropical or neotropical "weedy" species found within and outside of the lomas, 2) disjuncts from adjacent Andean source populations, either Peru or Chile, 3) lomas endemics (Peruvian and Chilean) with no extralomas populations, but widely distributed within the lomas, 4) narrow lomas endemics found in only one or a few lomas formations, 5) amphitropic disjuncts, primarily from arid and semi-arid regions of North America.
Patterns of Distribution
1) Wideranging Pantropical or Neotropical weeds.

2) disjuncts from adjacent Andes

2a. Peru
2b. Chile

3) Lomas endemics (genera and species)
Monotypic genera: Gypothamnium pinifolium Phil., Oxyphyllum ulicinium Phil. (both Chilean)
Endemic species:
Ambrosia , Baccharis, Centaurea, Erigeron, Encelia, Erechtites, Haplopappus, Helenium, Helogyne, heterospermum, Gypothamnium, Onoseris, Ophryosporus, Oxyphyllum, Perityle, Senecio.

4) Narrow endemics (one or a few lomas forations)
Ambrosia dentata
5) Amphitropical disjuncts:
A few North American species with amphitropic disjunct distributions are present in the lomas formations. For the most part, these are not the usual Mediterranean species commonly disjunct from California to central Chile (Constance 1963, Raven 1963, 1972). Rather, these species have their origins in the Sonoran or Mojave Deserts and their distributions are often restricted to the lomas formations. Several primarily North American genera have endemic species within the coastal deserts, including Ambrosia dentata (Cabr.) Dillon, Encelia canescens Lam., Viguiera weberbarueri S.F. Blake, and other primarily North American species represented within the Peruvian and/or Atacama Deserts include Malacothrix clevelandii A. Gray, Bahia ambrosioides Lag., Amblyopappus pusillus Hook. & Arn., and Perityle emoryi Torr.

Although many of the Atacama and Peruvian Desert species have obvious floristic affinities with the Andean Cordillera, the level of endemism in the isolated coastal vegetation is extremely high for some families, e.g., Bromeliaceae, Cactaceae, Malvaceae, Aizoaceae, Portulacaceae, Solanaceae, and Poaceae.
The greatest number of endemic genera are to be found in southern Peru between 15•-18•S latitude, and northern Atacama formations, specifically between 24•14'-26•21'S latitude.
While the arid environment is continuous between Chile and Peru, there appears to be a significant barrier to dispersal between 18• and 22•S latitude in extreme northern Chile. The lack of topographic barriers to provide fog moisture in the region between Arica and Antofagasta produces conditions with drought too severe for most plants to survive. Less than 7% of the combined Peruian and Chilean lomas flora, estimated at nearly 1000 species, occur on both sides of this region. Despite the floristic barrier in the region south of Arica, a number of species have become successfully established over the entire length of the Atacama and Peruvian Deserts, such as, Bidens triplinervia H.B.K., Encelia canescens Lam., Gamochaeta purpurea , Perityle emoryi, Pluchea chingoyo, Polyachyrus annuus, Vasquezia oppositifolia. A number of genera containing coastal perennials with species on each side of the barrier, including Ophryosporus, Senecio, Ageratina, Baccharis, Erigeron, and Gnaphalium.

Senecio Case Study



ASTERACEAE OF THE LOMAS FORMATIONS

Derived from LOMAFLOR, Jan 1995

1.1. Acmella alba (L'Her.) R.K. Jansen

General Distribution: Andean and coastal, northern Peru (0-1500 m) - Peru Depts. CA, LI, LL
Latitudinal Distribution: 7•54'S to 12•17'S
Lomas Distribution: [10] Manzano; Cerro Cabezon; Cerro Campana; Cerro Chiputur; Lupin; Lachay; Iguanil (Chancay); Pasamayo; Amancaes; Lurin.
Life Form: annual herbs

1.2. Acmella oleracea (L.) R. K. Jansen

General Distribution: Andean and coastal, northern Peru (0-500 m) - Peru Depts. AN, JU, LI, LL.
Latitudinal Distribution: 7•54'S to 12•52'S
Lomas Distribution: [7] Cerro Cabezon; Lupin; Lachay; Iguanil (Chancay); Ancon; Lurigancho (Lima); Antocongo; Amancaes; Quilmana
Life Form: annual herbs

2.1. Ageratina articulata (Hieron.) King & H. Robinson

General Distribution: Andean and eastern versant, rare in coastal northern Peru (500-3000 m) - Peru Depts. AM, CA, HU, LL
Latitudinal Distribution: 7•58'S
Lomas Distribution: [1] Cerro Campana
Life Form: shrubs

2.2. Ageratina azangaroensis (Schultz-Bip. ex Wedd.) King & H. Robinson

General Distribution: Widespread Andean, rare in coastal northern Peru (500-4500 m) - Peru Depts. AN, AR, AY, CA, CU, LA, LI, LL, PA, PU.
Latitudinal Distribution: 8•18'S
Lomas Distribution: [1] Cerro Negro
Life Form: shrubs

2.3. Ageratina remyanum (Phil.) King & H. Robinson

General Distribution: Chilean endemic
Latitudinal Distribution: 24•36'S
Lomas Distribution: [1] Miguel Diaz
Life Form: shrubs

2.4. Ageratina sternbergiana (DC.) King & H. Robinson

General Distribution: Widespread Andean, rare in coastal northern Peru (500-4500 m). - Peru Depts. AM, AN, AP, AR, AY, CA, CU, HU, HV, JU, LI, LL, PA, PU
Latitudinal Distribution: 7•58'S
Lomas Distribution: [1] Cerro Campana
Life Form: shrubs

3. Amblyopappus pusillus Hook. & Arn.

General Distribution: Amphitropic disjunct (NA origin), rare in northern Chile (x-x m).
Latitudinal Distribution: 22•05'S to 26•07'S
Lomas Distribution: [3] Tocopilla; Paposo; Pan de Azucar
Life Form: annual herbs

4.1. Ambrosia artemisioides Meyen & Walp.

General Distribution: Widespread Andean, rare in coastal southern Peru (500-4000 m) - Peru Depts.: AR, MO, TA
Latitudinal Distribution: 17•50'S
Lomas Distribution: [1] Tacna
Life Form: shrubs

4.2. Ambrosia dentata (Cabrera) M.O. Dillon

General Distribution: Peruvian coastal endemic (0-500 m) - Peru Depts.: AR
Latitudinal Distribution: 15•22'S
Lomas Distribution: [1] Jahuay
Life Form: shrubs

5.1. Baccharis juncea (Lehm.) Desf.

General Distribution: Non-endemic
Latitudinal Distribution: 23•37'S
Lomas Distribution: [2] Salar del Carmen; Quebrada San Ramon
Life Form: perennial herbs

5.2. Baccharis salicifolia (Ruiz & Pav.) Pers.
(syn.: B. glutinosa Pers.)

General Distribution: Widespread throughout the Neotropics, rare in northern Peru & Chile (0-3500 m)
Latitudinal Distribution: 7•54'S to 25•22'S
Lomas Distribution: [5] Peru: Cerro Cabezon; Cerro Campana; Cerro Negro; Lomas. Chile: Quebrada San Ramon.
Life Form: shrubs

5.3. B. linear-leaved morph cf. kingii Cuatrecasas 1982 :166.

General Distribution: (?Endemic morph?)
Latitudinal Distribution: 7•54'S to 8•19'S
Lomas Distribution: [5] Cerro Cabezon; Cerro Campana; Cerro Chiputur; Cerro Negro; Viru;
Life Form: shrubs

5.4. Baccharis taltalensis I.M. Johnst.

General Distribution: Northern Chilean endemic
Latitudinal Distribution: 24•36'S to 25•03'S
Lomas Distribution: [2] Miguel Diaz; Paposo
Life Form: shrubs

6. Bahia ambrosioides Lag.

General Distribution: Amphitropic disjunction (NA origin?)
Latitudinal Distribution: 22•05'S to 26•09'S
Lomas Distribution: [7] Tocopilla; La Chimba; Caleta Botija; Miguel Diaz; Paposo; Taltal-Hueso Parado; Pan de Azucar;
Life Form: shrubs

7. Bidens triplinervia H.B.K

General Distribution: Non-endemic (Peru & Chile)
Latitudinal Distribution: 11•21'S to 24•56'S
Lomas Distribution: [4] Lachay; Caleta Botija; Aguada Panul; Quebrada El Rincon
Life Form: perennial herbs

8.1. Centaurea atacamensis (Reiche) Johnst.

General Distribution: Chilean endemic
Latitudinal Distribution: 22•05'S to 26•09'S
Lomas Distribution: [6] Paposo; Quebrada San Ramon; Pan de Azucar; Tocopilla; Quebrada Cascabeles; Quebrada de la Cortadera
Life Form: perennial herbs

8.2. Centaurea cachinalensis Phil.

General Distribution: Chilean endemic
Latitudinal Distribution: 28•47'S
Lomas Distribution: [0]
Life Form: perennial herbs

8.3. Centaurea chilensis H. & A.

General Distribution: Chilean endemic
Latitudinal Distribution: 27•53'S to 28•04'S
Lomas Distribution: [1] Copiapo-Carrizal;
Life Form: perennial herbs

8.4. Centaurea floccosa H. & A.

General Distribution: Non-endemic
Latitudinal Distribution: 29•42'S
Lomas Distribution: [0]
Life Form:

8.5. Centaurea melitensis L.

General Distribution: Non-endemic; widely distributed in SA
Latitudinal Distribution: 24•56'S
Lomas Distribution: [1] Quebrada El Rincon;
Life Form: annual herbs

9.1. Chaetanthera glabrata (DC.) Meigen

General Distribution: Non-endemic
Latitudinal Distribution: 25•01'S to 35•25'S
Lomas Distribution: [7] Paposo; Palo Varado; Taltal-Hueso Parado; Quebrada de la Cortadera; Pan de Azucar; Caleta Obispito; Copiapo-Carrizal
Life Form: annual herbs

9.2. Chaetanthera linearis Poepp.

General Distribution:
Latitudinal Distribution: 25•03'S to 30•20'S
Lomas Distribution: [4] Paposo; Taltal-Hueso Parado; Quebrada de la Cortadera; Caleta Obispito
Life Form: annual herbs

10. Chionopappus benthamii Blake

General Distribution: Non-endemic; disjunct to adjacent Andes
Latitudinal Distribution: 7•58'S to 8•19'S
Lomas Distribution: [4] Cerro Campana; Cerro Prieto; Cerro Chiputur; Viru;
Life Form: perennial herbs

11. Chuquiraga ulicina (H. & A.) H. & A.

General Distribution: Non-endemic
Latitudinal Distribution: 24•33'S to 31•39'S
Lomas Distribution: [3] Caleta Botija; Cerro Perales; Pan de Azucar;
Life Form: shrubs

12.1. Conyza

General Distribution:
Latitudinal Distribution: 15•48'S to 17•45'S
Lomas Distribution: [4] Atiquipa; Mollendo; Mejia; Ilo;
Life Form: annual herbs

12.2. Erigeron paposanum Phil.

General Distribution: Chilean endemic
Latitudinal Distribution: 24•32'S to 25•07'S
Lomas Distribution: [5] Caleta Botija; Miguel Diaz; Quebrada El Rincon; Paposo; Quebrada Matancilla
Life Form: perennial herbs

13.1. Cotula australis (Sieb.x Spr.) Hook f

General Distribution: Non-endemic; introduced autodisjunct (Australian origin)
Latitudinal Distribution: 11•21'S to 17•45'S
Lomas Distribution: [5] Lachay; Iguanil (Chancay); Amancaes; Mejia; Ilo;
Life Form: annual herbs

13.2. Cotula coronopifolia L.

General Distribution: Non-endemic; introduced autodisjunct (Australian origin).
Latitudinal Distribution: 25•01'S to 25•13'S
Lomas Distribution: [2] Paposo; Quebrada Bandurrias
Life Form: perennial herbs

14. Encelia canescens Lam.

General Distribution: Endemic (Chile & Peru)
Latitudinal Distribution: 7•58'S to 35•03'S
Lomas Distribution: [15] Cerro Campana; Cerro Chiputur; Viru; Amancaes; Atocongo; Jahuay; Atiquipa; Atico; Camana; Mejia; Tacna; Quebrada El Rincon; Paposo; Chanaral-Barquito; Caleta Obispito;
Life Form: shrubs

15. Erechtites leptantha (Phil.) Cabrera

General Distribution: Chilean endemic
Latitudinal Distribution: 25•26'S; 23•33'S to 27•04'S
Lomas Distribution: [3] La Chimba; Caldera; Quebrada de la Cortadera
Life Form: annual herbs

16. Erigeron leptorhizon DC.

General Distribution: Peruvian endemic
Latitudinal Distribution: 7•54'S to 12•08'S
Lomas Distribution: [9] Cerro Cabezon; Cerro Campana; Cerro Chiputur; Cerro Negro; Lachay; Iguanil (Chancay); Pasamayo; Amancaes; Atocongo
Life Form: perennial herbs

17. Facelis retusa (Lam.) Sch.Bip.

General Distribution: Non-endemic
Latitudinal Distribution: 25•24'S
Lomas Distribution: [1] Taltal-Hueso Parado;
Life Form: annual herbs

18.1. Galinsoga caligensis Canne

General Distribution: Peruvian endemic?
Latitudinal Distribution: 11•21'S to 12•17'S
Lomas Distribution: [3] Lachay; Atocongo; Lurin;
Life Form: annual herbs

18.2. Galinsoga parviflora Cav.

General Distribution: Non-endemic
Latitudinal Distribution: 11•21'S to 12•52'S
Lomas Distribution: [3] Lachay; Lurigancho (Lima); Quilmana;
Life Form: annual herbs
18.3. Galinsoga quadriradiata R. & P.

General Distribution: Non-endemic
Latitudinal Distribution: 12•00'S to 12•08'S
Lomas Distribution: [2] Lurigancho (Lima); Atocongo;
Life Form: annual herbs

19.1. Gamochaeta purpurea (L.) Cabr.

General Distribution: Non-endemic
Latitudinal Distribution: 11•21'S to 30•41'S
Lomas Distribution: [11] Caleta Botija; Paposo; Lachay; Iguanil (Chancay); Amancaes; Quilmana; Camana; Mollendo; Mejia; Ilo; Miguel Diaz;
Life Form: annual herbs

19.2. Gamochaeta sphacelata (H.B.K.) Cabr.

General Distribution: Non-endemic
Latitudinal Distribution: 22•05'S to 25•24'S
Lomas Distribution: [4] Tocopilla; Miguel Diaz; Paposo; Taltal-Hueso Parado;
Life Form: annual herbs

20. Grindelia glutinosa (Cav.) Dun.

General Distribution: Non-endemic
Latitudinal Distribution: 15•48'S to 17•45'S
Lomas Distribution: [6] Loma de Okopa; Atiquipa; Chaparra; Mollendo; Mejia; Ilo;
Life Form: shrubs

21.1. Gutierrezia espinosae Acevedo

General Distribution:
Latitudinal Distribution: 23•33'S
Lomas Distribution: [1] La Chimba;
Life Form:

21.2. Gutierrezia gayana (Remy) Reich

General Distribution: Non-endemic
Latitudinal Distribution: 29•08'S
Lomas Distribution: [0]
Life Form:

21.3. Gutierrezia resinosa (H. & A.) Blake

General Distribution: Non-endemic
Latitudinal Distribution: 30•20'S to 30•41'S
Lomas Distribution: [0]
Life Form:

21.4. Gutierrezia taltalensis Phil.

General Distribution: Chilean Endemic
Latitudinal Distribution: 24•36'S to 29•50'S
Lomas Distribution: [4] Miguel Diaz; Taltal-Hueso Parado; Cerro Perales; Quebrada de la Cortadera;
Life Form: shrubs

22. Gypothamnium pinifolium Phil.

General Distribution: Chilean endemic
Latitudinal Distribution: 24•43'S to 26•41'S
Lomas Distribution: [11] Punta Plata; Paposo; Quebrada Cascabeles; Quebrada San Ramon; Taltal-Hueso Parado; Cerro Perales; Quebrada de la Cortadera; Quebrada El Nueve; Esmeralda; Chanaral-Barquito; Caleta Obispito
Life Form: shrubs

23.1. Haplopappus deserticola Phil.

General Distribution: Chilean endemic
Latitudinal Distribution: 24•32'S to 30•41'S
Lomas Distribution: [5] Caleta Botija; Paposo; Cerro Perales; Quebrada de la Cortadera; Quebrada El Nueve;
Life Form: shrubs

23.2. Haplopappus rosulatus Hall

General Distribution:
Latitudinal Distribution: 25•03'S
Lomas Distribution: [1] Paposo
Life Form: shrubs

24.1. Helenium aromaticum (Hook.) Bailey

General Distribution:
Latitudinal Distribution: 24•33'S to 31•39'S
Lomas Distribution: [3] Caleta Botija; Tortolas-Chepica; Esmeralda
Life Form:

24.2. Helenium atacamense Cabrera

General Distribution:
Latitudinal Distribution: 25•26'S
Lomas Distribution: [1] Quebrada de la Cortadera;
Life Form: annual herbs

24.3. Helenium litoralis (Phil.)
General Distribution:
Latitudinal Distribution: 25•01'S
Lomas Distribution: [1] Paposo;
Life Form:

Helenuim

General Distribution:
Latitudinal Distribution: 25•01'S
Lomas Distribution: [1] Paposo;
Life Form:

25. Helogyne hutchinsonii K. & R.

General Distribution: Peruvian endemic
Latitudinal Distribution: 16•14'S
Lomas Distribution: [1] Atico
Life Form: shrubs

26.1. Heterosperma diversifolia H.B.K.

General Distribution: Non-endemic
Latitudinal Distribution: 11•21'S
Lomas Distribution: [1] Lachay;
Life Form: perennial herbs

26.2. Heterosperma ferreyrii H. Robinson

General Distribution: Peruvian endemic
Latitudinal Distribution: 15•48'S to 17•50'S
Lomas Distribution: [2] Atiquipa; Tacna
Life Form: perennial herbs

27. Hypochoeris grandidentata (Phil.) Reiche

General Distribution:
Latitudinal Distribution: 24•36'S to 32•04'S
Lomas Distribution: [6] Miguel Diaz; Paposo; Taltal-Hueso Parado; Cerro Perales; Chanaral-Barquito; Caleta Obispito
Life Form: perennial herbs

28. Jungia axillaris (DC.) Spreng.

General Distribution:
Latitudinal Distribution: 10•33'S
Lomas Distribution: [1] Lupin
Life Form: perennial herbs

Vouchers: Cerrate 803, 812; Ferreyra 764A, 3922, 8061, 8372, 13790

29. Lactuca

General Distribution:
Latitudinal Distribution: 25•07'S to 25•26'S
Lomas Distribution: [2] Quebrada Matancilla; Quebrada de la Cortadera
Life Form:

30.1. Leucheria cerberoana Remy
General Distribution:
Latitudinal Distribution: 28•08'S to 33•09'S
Lomas Distribution: [0]
Life Form: annual herbs

30.2. Leucheria cumingii H. & A.

General Distribution:
Latitudinal Distribution: 25•03'S to 26•18'S
Lomas Distribution: [5] Paposo; Quebrada Matancilla; Quebrada Cascabeles; Taltal-Hueso Parado; Pan de Azucar
Life Form: annual herbs

30.3. Leucheria modesta (Phil.)

General Distribution:
Latitudinal Distribution: 22•05'S to 25•01'S
Lomas Distribution: [2] Tocopilla; Paposo
Life Form: annual herbs

31.1. Malacothrix clevelandii A. Gray

General Distribution: Western North American annual, autodisjunct in SA
Latitudinal Distribution: 26•33'S to 29•57'S
Lomas Distribution: Tocopilla
Life Form: annual herbs

31.2. Malacothrix coulteri A. Gray

General Distribution: Western North American annual, autodisjunct in SA
Latitudinal Distribution: 29•50'S
Lomas Distribution: Coquimbo
Life Form: annual herbs

32. Mutisia acuminata

General Distribution: Non-endemic
Latitudinal Distribution:
Lomas Distribution: Lima: Lomas de Asia, Grant 7487 (GH, NY, US).
Life Form: shrubs

33.1. Onoseris humboldtiana Ferreyra

General Distribution: Peruvian endemic
Latitudinal Distribution: 15•13'S
Lomas Distribution: [0] ?
Life Form: annual herbs

33.2. Onoseris odorata (D. Don) H. & A.

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 17•07'S
Lomas Distribution: [6] Cerro Cabezon; Cerro Chiputur; Viru; Atico; Camana; Mejia;
Life Form: annual herbs

33.3. Onoseris sp. nov. vel O. odorata

General Distribution:
Latitudinal Distribution: 15•22'S to 16•55'S
Lomas Distribution: [4] Jahuay; Atiquipa; Capac; Mollendo;
Life Form: annual herbs

34.1. Ophryosporus floribundus (DC.) K. & R.

General Distribution: Non-endemic
Latitudinal Distribution: 12•01'S to 26•00'S
Lomas Distribution: [5] La Chimba; Cerro Perales; Esmeralda; Amancaes; Atocongo;
Life Form: shrubs

34.2. Ophryosporus galioides (DC.) K. & R.

General Distribution: Non-endemic
Latitudinal Distribution: 7•58'S
Lomas Distribution: [1] Cerro Campana;
Life Form: shrubs

34.3. Ophryosporus hartwegii (B.L. Rob.) K. & R.

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 9•28'S
Lomas Distribution: [2] Casma; Cerro Cabezon;
Life Form: shrubs

34.4. Ophryosporus hoppii (B.L. Rob.) K. & R.

General Distribution: Peruvian endemic
Latitudinal Distribution: 16•55'S
Lomas Distribution: [1] Mollendo;
Life Form: shrubs

34.5. Ophryosporus johnstonii B.L. Rob.

General Distribution: Chilean endemic
Latitudinal Distribution: 24•33'S to 24•45'S
Lomas Distribution: [2] Caleta Botija; Aguada Cardon;
Life Form: shrubs

35.6. Ophryosporus paradoxus (H. & A.) B. & H.

General Distribution:
Latitudinal Distribution: 29•59'S
Lomas Distribution: [0]
Life Form: shrubs

35.7. Ophryosporus peruvianus (Gmel.) K. & R.

General Distribution:
Latitudinal Distribution: 7•54'S to 11•21'S
Lomas Distribution: [5] Cerro Cabezon; Cerro Campana; Cerro Chiputur; Viru; Lachay;
Life Form: shrubs

35.8. Ophryosporus pinifolius (Phil.) K. & R.

General Distribution: Chilean endemic
Latitudinal Distribution: 22•33'S to 29•45'S
Lomas Distribution: [1] Cobija;
Life Form: shrubs
35.9. Ophryosporus pubescens (J.E. Sm.) K. & R.

General Distribution:
Latitudinal Distribution: 15•33'S to 15•50'S
Lomas Distribution: [1] Chaparra;
Life Form: shrubs

35.10. Ophryosporus triangularis Meyen

General Distribution:
Latitudinal Distribution: 22•05'S to 29•57'S
Lomas Distribution: [9] Tocopilla; Cobija; La Chimba; Aguada Panul; Quebrada El Rincon; Paposo; Quebrada Matancilla; Quebrada San Ramon; Quebrada de la Cortadera;
Life Form: shrubs

36. Oxyphyllum ulicinum Phil.

General Distribution: Chilean endemic
Latitudinal Distribution: 25•01'S to 26•09'S
Lomas Distribution: [7] Paposo; Quebrada Matancilla; Quebrada Cascabeles; Quebrada San Ramon; Taltal-Hueso Parado; Esmeralda; Pan de Azucar
Life Form: shrubs

37.1. Perityle discoidea (Phil.) Johnst.

General Distribution: Chilean endemic
Latitudinal Distribution: 22•05'S to 23•33'S

Lomas Distribution: [2] La Chimba; Tocopilla
Life Form: annual herbs

37.2. Perityle emoryi Torr.

General Distribution: Non-endemic; amphitropic autodisjunct (NA origin)
Latitudinal Distribution: 17•50'S to 29•59'S
Lomas Distribution: [21] Tacna; Tocopilla; Cobija; La Chimba; Salar del Carmen; Antofagasta; Caleta Botija; Paposo; Quebrad Matancilla; Palo Varado; Taltal-Hueso Parado; Quebrada de la Cortadera; Esmeralda; Pan de Azucar; Chanaral-Barquito; Caleta Obispito; Caldera;Baja Inglesa; Monte Amargo; Copiapo; Copiapo-Carrizal;
Life Form: annual herbs

38.1. Philoglossa peruviana DC.

General Distribution: Non-endemic
Latitudinal Distribution: 7•58'S to 17•50'S
Lomas Distribution: [12] Cerro Campana; Lupin; Lachay; Iguanil (Chancay); Ancon; Amancaes; Atocongo; Lurin; Quilmana; Atiquipa; Chaparra; Tacna
Life Form: perennial herbs

38.2. Philoglossa purpureodisca H. Rob.

General Distribution: Peruvian endemic?
Latitudinal Distribution: 7•54'S to 8•19'S
Lomas Distribution: [5] Cerro Cabezon; Cerro Campana; Cerro Cabras; Cerro Chiputur; Viru
Life Form: annual herbs

39. Pleocarphus revolutus D. Don

General Distribution:
Latitudinal Distribution: 29•49'S
Lomas Distribution: [0]
Life Form:

40. Pluchea chingoyo (H.B.K.) DC.

General Distribution: Non-endemic
Latitudinal Distribution: 9•04'S to 18•20'S
Lomas Distribution: [2] Cerro Chimbote; Mejia; Chile?
Life Form: shrubs

41.1. Polyachyrus annuus Johnst.

General Distribution: Endemic (Peru & Chile)
Latitudinal Distribution: 16•14'S to 26•57'S
Lomas Distribution: [18] Tocopilla; La Chimba; Caleta Botija; Quebrada Medano; Paposo; Quebrada Matancilla; Quebrada San Ramon; Cerro Perales; Quebrada Los Zanjones; Tortolas-Chepica; Caleta Obispito; Atico; Camana; Mollendo; Mejia; Ilo; Tacna; Iquique
Life Form: annual herbs

41.2. Polyachyrus cinereus Ricardi & Weldt

General Distribution: Chilean endemic
Latitudinal Distribution: 23•33'S to 26•09'S
Lomas Distribution: [6] La Chimba; Paposo; Quebrada de la Cortadera; Tortolas-Chepica; Esmeralda; Pan de Azucar
Life Form: shrubs

41.3. Polyachyrus fuscus (Meyen) Walpers

General Distribution: Chilean endemic
Latitudinal Distribution: 22•05'S to 32•14'S; 23•33'S to 26•08'S
Lomas Distribution: [10] Tocopilla; La Chimba; Paposo; Cerro Perales; Quebrada de la Cortadera; Esmeralda; Pan de Azucar; Chanaral-Barquito; Caleta Obispito; Caldera; [2] La Chimba; Pan de Azucar
Life Form: perennial herbs

41.4. Polyachyrus gayi Remy

General Distribution:
Latitudinal Distribution: 29•57'S
Lomas Distribution: [0]
Life Form:

41.5. Polyachyrus mollendoensis Johnst.

General Distribution: Peruvian endemic
Latitudinal Distribution: 16•53'S
Lomas Distribution: [0]
Life Form: annual herbs

41.6. Polyachyrus poeppigii (Kunz ex Less.)Less.

General Distribution: Non-endemic
Latitudinal Distribution: 25•03'S to 28•28'S
Lomas Distribution: [5] Cerro Perales; Pan de Azucar; Caleta Obispito; Copiapo-Carrizal; Paposo
Life Form: perennial herbs

42.1. Proustia pungens
General Distribution:
Latitudinal Distribution:
Lomas Distribution: [1] Atiquipa
Life Form: shrubs

42.2. Proustia tipia (Phil.) Fabis

General Distribution:
Latitudinal Distribution: 24•48'S to 25•25'S
Lomas Distribution: [4] Quebrada Medano; Paposo; Taltal-Hueso Parado; Cerro Perales
Life Form: shrubs

43.1. Pseudognaphalium dombeyanum (DC.) Anderb.

General Distribution: Non-endemic
Latitudinal Distribution: 11•21'S to 29•08'S
Lomas Distribution: [5] Quilmana; Paposo; Lachay; Atiquipa; Mollendo;
Life Form: perennial herbs

43.2. Pseudognaphalium heterotrichum (Phil.) Anderb.

General Distribution: Non-endemic
Latitudinal Distribution: 24•56'S
Lomas Distribution: [1] Quebrada El Rincon;
Life Form: perennial herbs

44.1. Senecio abadianus DC.

General Distribution:
Latitudinal Distribution: 11•21'S to 31•55'S
Lomas Distribution: [12] La Chimba; Quebrada El Rincon; Paposo; Quebrada Cascabeles; Quebrada San Ramon; Cerro Perales; Quebrada de la Cortadera; Pan de Azucar; Caleta Obispito; Lachay; Iguanil (Chancay); Atocongo;
Life Form: perennial herbs

44.2. Senecio acarinus Cabr.

General Distribution:
Latitudinal Distribution: 15•22'S
Lomas Distribution: [1] Jahuay;
Life Form: shrubs

44.3. Senecio adenotrichius DC

General Distribution:
Latitudinal Distribution: 30•41'S
Lomas Distribution: [0]
Life Form:

44.4. Senecio almeidae Phil.

General Distribution:
Latitudinal Distribution: 25•01'S to 28•57'S; 25•26'S
Lomas Distribution: [3] Paposo; Palo Varado; Cerro Perales; Quebrada de la Cortadera;
Life Form: succulent-leaved shrubs

44.5. Senecio arnaldii Cabr.

General Distribution:
Latitudinal Distribution: 8•10'S to 8•19'S
Lomas Distribution: [2] Cerro Chiputur; Viru
Life Form: perennial herbs

44.6. Senecio bahioides Hook. & Arn.

General Distribution:
Latitudinal Distribution: 28•08'S to 29•56'S
Lomas Distribution: [0]
Life Form: shrubs

44.7. Senecio calciola Meyen & Walp.

General Distribution: Peruian coastal endemic.
Latitudinal Distribution: 16•55'S
Lomas Distribution: [1] Mollendo
Life Form: shrubs

44.8. Senecio cachinalensis Phil.

General Distribution:
Latitudinal Distribution: 24•56'S to 28•28'S
Lomas Distribution: [9] Quebrada El Rincon; Paposo; Quebrada Cascabeles; Taltal-Hueso Parado; Quebrada de la Cortadera; Esmeralda; Pan de Azucar; Caleta Obispito; Copiapo-Carrizal
Life Form: shrubs

44.9. Senecio lomincola Cabrera

General Distribution:
Latitudinal Distribution: 9•28'S to 11•21'S
Lomas Distribution: [2] Casma; Lachay
Life Form: shrubs

44.10. Senecio myriophyllus Phil.

General Distribution:
Latitudinal Distribution: 26•07'S to 26•09'S
Lomas Distribution: [1] Pan de Azucar
Life Form: shrubs

44.11. Senecio okopanus Cabr.

General Distribution:
Latitudinal Distribution: 15•48'S to 15•50'S
Lomas Distribution: [2] Atiquipa; Chaparra
Life Form: perennial herbs

44.12. Senecio paposanus Phil.

General Distribution:
Latitudinal Distribution: 24•32'S to 25•03'S
Lomas Distribution: [3] Caleta Botija; Quebrada El Rincon; Paposo
Life Form: shrubs

44.13. Senecio philippicus Regel & Koern.

General Distribution:
Latitudinal Distribution: 25•26'S
Lomas Distribution: [1] Quebrada de la Cortadera;
Life Form: shrubs

44.14. Senecio pseudalmeidae Cabr.

General Distribution:
Latitudinal Distribution: 25•23'S
Lomas Distribution: [1] Quebrada San Ramon;
Life Form:

44.15. Senecio smithianus Cabr.

General Distribution:
Latitudinal Distribution: 15•48'S
Lomas Distribution: [1] Atiquipa
Life Form: perennial herbs

44.16. Senecio troncosii Phil.

General Distribution:
Latitudinal Distribution: 25•07'S to 26•22'S
Lomas Distribution: [2] Quebrada Matancilla; Chanaral-Barquito
Life Form: shrubs

44.17. Senecio truxillensis Cabrera

General Distribution:
Latitudinal Distribution: 7•54'S to 7•59'S
Lomas Distribution: [3] Cerro Cabezon; Cerro Campana; Cerro Prieto.
Life Form: perennial herbs

44.18. Senecio vulgaris L.

General Distribution: Cosmopolitan weed
Latitudinal Distribution: 7•54'S to 25•25'S
Lomas Distribution: [2] Cerro Cabezon; Cerro Perales;
Life Form: annual herbs

45.1. Siegesbeckia flosculosa L'Her.

General Distribution:
Latitudinal Distribution: 8•10'S to 12•52'S
Lomas Distribution: [5] Cerro Chiputur; Viru; Lachay; Iguanil (Chancay); Quilmana
Life Form: perennial herbs

45.2. Siegesbeckia orientalis L.

General Distribution:
Latitudinal Distribution: 12•52'S
Lomas Distribution: [1] Quilmana
Life Form:

46. Simsia dombeyana DC.

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 8•10'S
Lomas Distribution: [2] Cerro Cabezon; Cerro Chiputur
Life Form: annual herbs
47.1. Sonchus oleraceus L.

General Distribution: Non-endemic
Latitudinal Distribution: 15•48'S to 25•01'S

Lomas Distribution: [6] Atiquipa; Camana; Paposo; Chala; Ilo; Tacna.
Life Form: annual herbs

47.2. Sonchus tenerrimus L.

General Distribution: Endemic?
Latitudinal Distribution: 22•05'S
Lomas Distribution: [1] Tocopilla
Life Form: annual herbs

48.1. Stevia hyssopifolia Phil.

General Distribution: Non-endemic
Latitudinal Distribution: 24•56'S to 25•26'S
Lomas Distribution: [4] Quebrada El Rincon; Quebrada Matancilla; Taltal-Hueso Parado; Quebrada de la Cortadera
Life Form: perennial herbs

48.2. Stevia melissaefolia (Lam.) Sch.-Bip.

General Distribution: Non-endemic
Latitudinal Distribution: 11•21'S to 15•48'S
Lomas Distribution: [5] Lachay; Iguanil (Chancay); Amancaes; Atocongo; Atiquipa; Juhuay-San Juan
Life Form: perennial herbs

49. Tessaria absinthioides (H. & A.) DC.

General Distribution: Non-endemic
Latitudinal Distribution: 25•33'S
Lomas Distribution: [1] Tortolas-Chepica;
Life Form:

50. Trixis paradoxa Cass.

General Distribution: Non-endemic?
Latitudinal Distribution: 7•54'S to 17•00'S
Lomas Distribution: [8] Atiquipa; Cachendo; Cerro Cabezon; Cerro Cabras; Cerro Chiputur; Viru; Atocongo; Chaparra
Life Form: shrubs

51. Vasquezia oppositifolia (Lag.) Blake

General Distribution: Non-endemic
Latitudinal Distribution: 7•58'S to 25•03'S
Lomas Distribution: [16] Cerro Campana; Cerro Chiputur; Viru; Lupin; Pativilca (Pacar); Lachay; Lurigancho (Lima); Amancaes; Atocongo; Atiquipa; Chaparra; Atico; Camana; Mollendo; Ilo; Paposo
Life Form: annual herbs

52. Verbesina saubinetioides Blake

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 8•19'S
Lomas Distribution: [5] Cerro Cabezon; Cerro Campana; Cerro Cabras; Cerro Chiputur; Viru
Life Form: shrubs

53.1. Viguiera pazensis Rusby

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 17•07'S
Lomas Distribution: [3] Cerro Cabezon; Mejia; Atiquipa;
Life Form: perennial herbs

53.2. Viguiera weberbaueri Blake

General Distribution: Peruvian endemic
Latitudinal Distribution: 16•55'S
Lomas Distribution: [1] Mollendo;
Life Form: shrubs

54. Wedelia latifolia DC.

General Distribution: Non-endemic
Latitudinal Distribution: 7•54'S to 8•19'S
Lomas Distribution: [5] Cerro Cabezon; Cerro Campana; Cerro Cabras; CerroChiputur; Viru
Life Form: shrubs