A m e r J of P o t a t o Res (2001) 78:197-207

197

Taxonomy and New Collections of V lld P o t a t o Species in Central and Southern Peru in 1999 Alberto R. Salas 1, David M. SpooneV*, Z6simo HuamS.n ~, Rafael Vinci Torres Maits Roel Hoekstra ~, Konrad Schtiler ~, and Robert J. Hijmans ~ 1International Potato Center (CIP), Apartado 1558, Lima 12, Peru. ~United States Department of Agriculture, Agricultural Research Service, Department of Horticulture, University of Wisconsin, 1575 Linden Drive, Madison, WI, 53706-1590, USA. ~Instituto Nacional de Investigaci6n Agraria (INIA), Estacidn Experimental Santa Ana, Huancayo, Peru. 4plant Research International, Centre for Genetic Resources The Netherlands (CGN), Droevendaalsesteeg 1, P.O. Box 16, 6700 AA Wagenmgen, The Netherlands SInstitut fiir Pflanzengenetik und Kulturpflanzenforschung (IPK) Gatersleben, Genbank Aussenstelle Grof~ Ltisewitz, (Institute of Plant Genetics and Crop Plant Research [IPK], Gatersleben, Genebank External Branch Grog Ltisewitz), Parkweg 1, D-18190 Grog Ltisewitz, Germany Corresponding author. Tel: 608-262-0159; Fax: 608-262-4743; email: [email protected].

p u b l i s h e d b y C. O c h o a in 1999. T h i s p a p e r r e p o r t s t h e col-

ABSTRACT

l e c t i o n a n d n e w s p e c i e s i d e n t i f i c a t i o n s o f t h e 1999 collecPeru contains about half of the described wild potato taxa,

and many of these

are not yet preserved

in

genebanks. This paper reports results of the second of a

tions, and germplasm conservation and survival of the 1998 and 1999 collections. In addition, chromosome c o u n t s a r e p r o v i d e d f o r 134 a c c e s s i o n s f r o m t h e 1998 a n d

s e r i e s o f f i v e p l a n n e d c o l l e c t i n g e x p e d i t i o n s t o P e r u . Col-

1999

l e c t i o n s w e r e m a d e in t h e c e n t r a l P e r u v i a n d e p a r t m e n t s

chomatophilum var. subnivale ( 2 n = 2x = 24), S. megis-

expeditions,

including

first

reports

for

S.

of Ancash, Huancavelica, La Libertad, and Lima, from

tacrolobum subsp, p u r p u r e u m ( 2 n = 2x = 24), and S. mul-

M a r c h 8 t o A p r i l 25, 1999. T h e y f o l l o w c o l l e c t i o n s in 1998

tiinterruptum var. multiinterruptum f. albiflorum (2n --

in t h e s o u t h e r n P e r u v i a n d e p a r t m e n t s o f A p u r i m a c , A r e -

2x = 2 4 ) ; w e a l s o r e p o r t t h e f i r s t t r i p l o i d c o u n t o f a n

q u i p a , Cusco, M o q u e g u a , P u n o , a n d T a c n a . We c o l l e c t e d

a c c e s s i o n o f S. immite.

101 g e r m p l a s m a c c e s s i o n s , i n c l u d i n g f i r s t g e r m p l a s m coll e c t i o n s o f t h e f o l l o w i n g 22 S o l a n u m t a x a : S o l a n u m

a m a y a n u m , S. a n a m a t o p h i l u m , S. a r a h u a y u m ( l o s t in g e r m p l a s m i n c r e a s e ) , S. augustii, S. bill-hookeri, S. can-

RESUMEN E1 P e r d t i e n e a p r o x i m a d a m e n t e l a m i t a d d e las t a x a

tense, S. chavinense, S. chomatophilum var. subnivale, S. c h r y s o f l o r u m , S. g r a c i l i f r o n s , S. h a p a l o s u m ,

S.

huarochiriense, S. h y p a c r a r t h r u m , S. jalcae, S. monilif o r m e , S. m u l t i i n t e r r u p t u m f. longipilosum, S. multiinterruptum var. m a c h a y t a m b i n u m , S. peloquinianum, S. r h o m b i l a n c e o l a t u m , S. s i m p l i c i s s i m u m , S. t a u l i s e n s e ( l o s t in g e r m p l a s m i n c r e a s e ) , and S. wittmackii. I n addition, new collections were made of the under-collected s p e c i e s S. hast~forme ( t h r e e c o l l e c t i o n s ) . T h e a b o v e t a x o n o m y is t h a t u s e d in p l a n n i n g o u r e x p e d i t i o n , t h a t w e compare to a new treatment

of Peruvian wild potatoes

Accepted for publication December 21, 2000. ADDITIONAL KEY WORDS: Solanum sect. Petota.

EXPLANATION OF ABBREVIATIONS: CIP, International Potato Center (Centro Interuacional de la Papa), and a herbarium code used by this organization for their herbarium in La Molina, Peru CGN, Centre for Genetic Resources, The Netherlands INIA, Instituto Nacional de Investigacion Agraria (National Institute for Agrarian Research), Peruvian Ministry of Agriculture IPK, Institut fitr Pflanzengenetik und Kulturpflanzenforschung, Gatersleben, Grog Ltisewitz, Germany MOL, Herbario Weberbauer, Departamento de Biologia, Secci6n Botfinica, Universidad Nacional Agraria, La Molina, Peru NRSP-6, National Research Support Program-6, Sturgeon Bay, Wisconsin, USA (the United States potato genebank, formerly called the Inter-Regional Potato Introduction Project, IR-1) PTIS, Herbarium of NRSP-6 WAG, National Herbarium Nederland, Wageningen University branch, Wageningen, The Netherlands

198

AMERICAN JOURNAL OF POTATO RESEARCH

de p a p a s i l v e s t r e d e s c r i t a , muchas de las c u a l e s ailn n o

Vol. 78

paper reports collections in the departments of Ancash, Huan-

est~n conservadas en bancos gen6ticos. Este articulo

cavelica, La Libertad, and Lima, from March 8 to April 25, 1999.

c o n s i g n a los r e s u l t a d o s de la s e g u n d a de u n a s e r i e de

This is the second of a series of five planned collecting expedi-

c i n c o e x p e d i c i o n e s p l a n i f i c a d a s de r e c o l e c c i 6 n e n el

tions to Peru, and follows collections made in 1998 in the south-

Peril. Se colect5 e n los d e p a r t a m e n t o s de Ancash, Huan-

ern Peruvian departments of Apurlmac, Arequipa, Cusco,

cavelica, La L i b e r t a d y Lima, e n t r e el 8 de m a r z o y el 25

Moquegua, Puno, and Tacna. This paper details the collection

de a b r i l de 1999. F u e r o n l a c o n t i n u a c i 6 n de c o l e c c i o n e s

and taxonomy of the 1999 collections, germplasm survival of the

r e a l i z a d a s e n 1998 e n el s u r del Peril, e n los d e p a r t a -

1998 collections, and chromosome counts of 1998 and 1999 col-

m e n t o s de Apurimac, Arequipa, Cusco, Moquegua, P u n o y

lections.

T a c n a . R e c o l e c t a m o s 101 a c c e s i o n e s de g e r m o p l a s m a ,

We planned and conducted our 1998 and 1999 collections

i n c l u y e n d o las p r i m e r a s colecciones de g e r m o p l a s m a de

prior to a monograph of the Peruvian wild potatoes (Ochoa

las siguientes 22 t a x a de S o l a n u m : S o l a n u m a m a y a n u m ,

1999). These prior data (mainly Hawkes 1990, see Spooner et

S. anamatophilum,

( p e r d i d a al i n c r e -

al. 1999) documented 36 wild potato taxa in the targeted cen-

m e n t a r g e r m o p l a s m a ) , S. augustii, S. b i l l - h o o k e r i , S.

tral Peruvian departments of Ancash, Huancavelica, La Liber-

c a n t e n s e , S. c h a v i n e n s e , S. c h o m a t o p h i l u m var. s u b n i -

tad, and Lima. A total of 24 of these taxa had no germplasm

S. a r a h u a y u m

vale, S. c h r y s o f l o r u m , S. gracil~frons, S. h a p a l o s u m , S.

collections in any genebank worldwide (Spooner et al. 1999).

h u a r o c h i r i e n s e , S. h y p a c r a r t h r u m , S . j a l c a e , S. monil-

The goal of this expedition was to collect these 24 taxa and

i f o r m e , S. m u l t i i n t e r r u p t u m f . l o n g i p i l o s u m , S. m u l t i i n -

other rare taxa.

t e r r u p t u m var. m a c h a y t a m b i n u m , S. p e l o q u i n i a n u m , S. r h o m b i l a n c e o l a t u m , S. s i m p l i c i s s i m u m , S. t a u l i s e n s e

MATERIALS AND METHODS

( p e r d i d a al i n c r e m e n t a r g e r m o p l a s m a ) y S. w i t t m a c k i i . A d e m ~ , colectamos u n a especie poca colectada: S. h a s -

The report of the 1998 expedition (Spooner et al. 1999) con-

t i f o r m e ( t r e s c o l e c c i o n e s ) . La t a x o n o m i a a q u l u s a d a es

tains details not repeated here regarding permission to collect,

l a d e l p l a n e a m i e n t o la e x p e d i c i 6 n q u e f u e c o m p a r a d a

ecogeographic data, a list and map of Peruvian type localities of

c o n el n u e v o t r a t a m i e n t o de papas s i l v e s t r e s p e r u a n a s

wild potatoes for all of Peru, and further details on taxonomy

p u b l i c a d o p o r C a r l o s O c h o a e n 1999. E s t e a r t i c u l o

that relates to this report. All methods reported there were fol-

i n f o r m a s o b r e la colecciSn y la i d e n t i f i c a c i 6 n de n u e v a s

lowed on the 1999 expedition, with the exception of the targeted

especies de las c o l e c c i o n e s de 1999 y de la c o n s e r v a c i 6 n

collection areas in central Peru, rather than southern Peru.

y s u p e r v i v e n c i a del g e r m o p l a s m a de las c o l e c c i o n e s de

Herbarium vouchers were deposited at the herbaria of CIP,

1998 y 1999. Asimismo, se p r o p o r c i o n a el c o n t e o de cro-

MOL, PTIS, and WAG (herbarium codes MOL and WAG follow

m o s o m a s de 134 a c c e s i o n e s de las e x p e d i c i o n e s de 1998

Holmgren et al. [1990]; PTIS will apprear in the next edition; CIP

de S . m e g i s -

is not listed there). An unpublished report of this expedition that

t a c r o l o b u m subsp, p u r p u r e u m ( 2 n = 2x = 24), S. m u l t i -

includes many details not reported here is deposited at CIP,

i n t e r r u p t u m var. m u l t i i n t e r r u p t u m f. albO"lorum ( 2 n =

CGN, INIA, IPK, NRSP-6, and other genebanks.

y 1999, i n c l u i d o el p r i m e r r e p o r t e

2 x = 24) y S. v e l a r d e i ( 2 n = 2x =24); t a m b i ~ n infor-

In addition, we obtained somatic chromosome counts of

m a m o s s o b r e el p r i m e r c o n t e o de triploide de u n a acce-

germplasm of the 1998 and 1999 collections, grown in green-

siSn de S. i m m i t e .

houses at the CIP station in Huancayo, Peru. Mitotic counts were obtained from root tips by the acetocarmine squash tech-

INTRODUCTION Wild and cultivated potato species have proven value in

nique (Smith 1974). Ochoa (1999) was not available for trip planning until after both the 1998 and 1999 expeditions were completed. We

breeding programs for disease resistance, environmental toler-

changed the identifications of some of our 1998 collections

ances, and other agronomic traits of interest (Ross 1986;

based on taxonomic changes in Ochoa (1999). We also made

Hawkes 1990; Spooner and Bamberg 1994; Ochoa 1999; Jansky

changes based on observation of more mature plants of 1998

2000). Peru contains about one-half of the described wild potato

and 1999 collections at the CIP station in Huancayo, Peru, in

taxa, and many of these are not yet preserved in genebanks. This

March 2000 (Tables 1, 2).

2001

SALAS, et al.: P E R U POTATO COLLECTION

TABLE 1--Prior ge~nplasm collections at NRSP-6 and new germplasm

199

RESULTS AND DISCUSSION

collections made on the 1999 expedition in central Peru. The taxonomy of this list differs f r o m that in the report of the 1998 expedition (Table 2 of Spooner et al. 1999) by the incor-

Taxonomic Overview The target species and germplasm holdings prior to

poration of taxonomic changes of Ochoa (1999), by the dde-

the 1999 e x p e d i t i o n for t h e central Peruvian depart-

tion ofS. limense, shown by Ochoa (1962, pg. 297) to be a

ments of Ancash, Huancavelica, La Libertad, and Lima

synonym of S. chacoense, and by the listing of taxa in all

are presented in Table 1. This is a subset of Table 2 of

departments in which they were collected in 1999.

S p o o n e r et al. (1999) for t h e s e four departments, and with taxonomic changes by Ochoa (1999). As detailed in

Department

Ancash

Taxon

Prior germplasm collections from Peru ( all departments) at NRSP-6

S. albicans (Ochoa) Ochoa 19 S. anamatophilum Ochoa 0 S. ancophilum Correll 1 0 S. augustii Ochoa 0 S. • Ochoa2 4 S. chomatophilum Bitter var. chomatophilum 18 S. chomatophilum var. subnivale Ochoa 0 S. dolichocremastrum Bitter3 4 S. medians Bitter, ploidy and variety undetermined 4 9 S. multiinterruptum var. multiinter~tptum 5 0 S. orophilum Correll 1 S. pdoquinianum Ochoa 0 S. sogarandinum Ochoa 2 Huancavelica S. amayanum Ochoa 0 S. bill-hookeri Ochoa 0 S. brevicaule Juz. 6 168 S. gracilifivns Bitter 0 S. huancavelicae Ochoa 7 0 La Libertad S. albicans (Ochoa) Ochoa see Ancash S. • Ochoa2 see Ancash S. chiquidenum Ochoa var. chiquidenum 8 5 S. chomatophilum var. chomatophilum see Ancash S. hastiforme Correll 1 S. immite Dunal See Lima S. ja/cae Ochoa var. ja/cae 9 0 S. mochiquense Ochoa 10 4 S. multiinterruptum var. machaytambinum Ochoa 0 S. sogarandinum Ochoa 2 S. taulisense Ochoa 0 S. yamobambense Ochoa 0 Ulma S. • Ochoa 11 0 S. brevicaule Juz. 6 168 S. cantense Ochoa 0 S. chancayense Ochoa 10 2 S. ehomatophilum var. chomatophilum See Ancash S. huarochiriense Ochoa 0

New germplasm collections from each department in 1999 2 1 4 1 1 1

Table 1, these involve four classes of changes: (1) the n e w recognition of one formerly synonymized taxon, (2) the placement of four formerly recognized taxa in synonymy, (3) the description of two n e w taxa, and (4) n e w hypotheses for hybrid origins for three taxa. Our taxonomic observations are presented below.

S e r i e s Piurana Solanum series Piurana is one of 19 tuber-bearing series recognized by Hawkes (1990), who included 15

2 (1 lost) 3

s p e c i e s , d i s t r i b u t e d f r o m s o u t h e r n C o l o m b i a and

1

(with 11 species). This series contains some of the mor-

E c u a d o r ( w h e r e four s p e c i e s o c c u r ) to central Peru phologically most diverse species in sect. Petota, but the

9 (2 lost) 1 2 1 1 1 1 1 1 3 1

morphological definition of the series and its taxonomic limits remain controversial. The m o s t distinctive features of set. Piurana, used by all major taxonomists of sect. Petota (Correll 1962; Hawkes 1990; Ochoa 1999) are the globose to ovoid fruits and coriaceous glossy leaves. The p r o b l e m is that t h e s e traits vary so m u c h in species placed in other series that it is difficult to clearly decide w h a t to include in ser. Piurana. All the above a u t h o r s have e x p r e s s e d d o u b t as to the limits of the

2 3 3 1 3 0 1 3 1 (lost) 0 1 (lost) 1 (lost) 4 0 1 1

series, but perhaps the clearest statement of this difficulty was by Correll (1962: 139): "This series, more than any of the others, may be considered a catchall. Paradoxically, its component species are held together not so m u c h by their similarity as by their differences." O c h o a ' s (1999) t r e a t m e n t o f t h e series c l e a r l y reflected these differences of opinion. Relative to the treatment of the 11 Peruvian species of ser. P / u r a n a of H a w k e s (1990), O c h o a (1999) i n c l u d e d t w o s p e c i e s H a w k e s placed in set. Tuberosa (S. chiquidenum, S. h u m e c t o p h i l u m ) , and p l a c e d s p e c i e s that H a w k e s included in ser. Piurana in two other series (S. • galdosii in ser. Cuneoalata and S. jaleae in ser. Ingifolia). In addition, Ochoa placed S. pascoense in synonymy

200

AMERICAN JOURNAL OF POTATO RESEARCH

TABLE 1 - - C o n t i n u e d .

Vol. 78

A c h l o r o p l a s t DNA r e s t r i c t i o n site p h y l o g e n e t i c study of sect. Petota a n d immediate outgroup relatives in

Department

Taxon

Prior germplasm collections from Peru ( all departments) at NRSP-6

S. hypacrarthrum Bitter 0 S. immite Dunal 4 S. medians Bitter var. autumnale 3 0 S. medians Bitter var. medians 3 9 S. medians Bitter var. undetermined3 0 S. multiinterruptum Bitter var. multiinterruptum 5 9 S. multiinteT'ruptum var. multiinter~uptum f. albiflorum 5 0 S. neoweberbaueri Wittm. 10,12 0 S. simplicissimum Ochoa 0 S. soga7undinum Ochoa see Ancash S. wittmackii Bitter 0

New germplasm collections from each department in 1999 5 (1 lost) 2

s e c t . E t u b e r o s u m ( B u k a s o v a n d K a m e r a z ) A. C h i l d ( S p o o n e r a n d Castillo 1997) defined four clades. One of t h e s e four clades i n c l u d e d all s e v e n e x a m i n e d species o f ser. P i u r a n a a n d s e v e n o t h e r s p e c i e s p l a c e d b y H a w k e s in ser. C o n i c i b a c c a t a , Megistacroloba, Y u n g a s e n s i a , a n d T u b e r o s a . A n o t h e r c h l o r o p l a s t DNA r e s t r i c t i o n site s t u d y b y Castillo a n d S p o o n e r (1997)

8 3 2

While s o m e of t h e s e s p e c i e s a s s i g n e d to o t h e r s e r i e s

6

m a t c h o u r "gestalt" i m p r e s s i o n s of ser. P i u r a n a (S.

a d d i t i o n a l l y a d d e d S. c h o m a t o p h i l u m

to t h i s clade.

a n d r e a n u m , S. c h a n c a y e n s e , S. c h o m a t o p h i l u m , S. 0 0 2 1 7 (1 lost)

iOchoa (1999) recognized S. rhomboideilanceolatum var. ancophilum Correll as S. ancophilum (Correll) Ochoa. Ochoa (1999) provided an invalid spelling change from his original spelling of this species in 1952 to S. rhombilanceolatum. 2Ochoa (1999) designated S. • (type locality in Cajamarca) as a hybrid between S. anamatophilum and S. peloquinianum. 3Ochoa (1999) placed S. chavinense as a synonym ofS. dolichocremastrum. 4Ochoa (1999) recognized two varieties of S. medians: var. autumnale (diploid) and var. medians (triploid), and synonymized S. weberbaue~g (type from department of Tacna) in the former variety. Distinction of these varieties rests on morphological and ploidy characters and we here rely on ploidy determinations for identifications. ~Ochoa (1999) made S. chrysoflorum (from Lima), S. moniliforme (from Ancash), and S. multiinterruptum f. Iongipilosum (from Ancash) synonyms of S. multiinterruptum var. multiinterruptum. He also recognized the new taxon S. multiinterrupture var. multiinterruptum f. albiflorum Ochoa. 6Ochoa (1999) placed S. hapalosum (from Lima Department) in synonymy with S. bukasovii Juz. We temporarily treat the 168 collections of S. brevicaule here in a very broad and likely artificial sense to include S. bukasovii and many other species. Solarium brevicaule may later be treated as the two species S. bukasovii and S. brevicaule (see text). 7Ochoa (1999) described this new species. sOchoa (1999) placed S. chiquidenum var. caehicadense as a synonym of S. chiquidenum var. chiquidenum. ~~ species grow in the coastal desert lomas and are best collected in June to September. ~Ochoa (1999) recognized S.jalcae var. pubescens (Correll), creating the autonym var. jalcae. ~'Ochoa (1999) designated S. • as a hybrid between S. medians Bitter and S. wittmackii Bitter. '2Ochoa (1999) designated S. • as a hybrid between S. medians and S. chancayense.

m o c h i q u e n s e , S. i m m i t e ) , o t h e r s a p p e a r misplaced (S. h u a n c a b a m b e n s e , S. s o g a r a n d i n u m , S. tundalomense). Two possible explanations for these problems are misidentifications of g e r m p l a s m u s e d in t h e s e studies or "chloroplast capture". The latter p h e n o m e n o n , c o m m o n in plants, is c a u s e d b y hybridization b e t w e e n m e m b e r s of different species differing b y c h l o r o p l a s t types, foll o w e d b y b a c k c r o s s i n g to the p a t e r n a l parent, resulting in d i s c o r d a n c e b e t w e e n "species trees" a n d " m a t e r n a l gene trees" (Wendel a n d Doyle 1998). E x a m i n a t i o n w i t h biparentally inherited m o l e c u l a r m a r k e r s will b e n e e d e d to resolve the true phylogenetic relationships of ser. P i u rana. O u r c o l l e c t i o n s in 1999 g a t h e r e d g e r m p l a s m o f species useful for c o n t i n u e d morphological a n d molecular resolution of the limits of set. P i u r a n a . Relative to t h e c o n c e p t of ser. P i u r a n a b y O c h o a (1999), w e coll e c t e d S. cantense, S. e h i q u i d e n u m , and S. h y p a e r a r t h r u m . Relative to o u r a g r e e m e n t w i t h a n e x p a n d e d c o n c e p t of the series b y the c h l o r o p l a s t DNA studies of S p o o n e r a n d Castillo (1997) a n d Castillo a n d S p o o n e r (1997), w e additionally collected S. c h o m a t o p h i l u m and S. i m m i t e . These latter t w o species have r o u n d to ovoid fruits a n d glossy leaves p r e s e n t in o t h e r m e m b e r s of t h e series, a n d o u r b e s t g u e s s o f r e l a t i o n s h i p s b a s e d o n t h e s e c h a r a c t e r s (gestalt a p p r o a c h ) a n d the cpDNA data p l a c e s t h e m in ser. P i u r a n a . We additionally s u s p e c t t h a t the following species we collected to b e m e m b e r s of

w i t h S. c h o m a t o p h i l u m . In s u m m a r y , ser. P i u r a n a i n c l u d e s

t h e series: S. blanco-galdosii, S. h u a r o c h i r i e n s e , S. jalcae, S.

s o m e very distinctive species, a n d various a u t h o r s h a v e m a d e

p e l o q u i n i a n u m , and S. s i m p l i c i s s i m u m . We will include t h e m

t h e i r b e s t j u d g e m e n t as to t h e species to include in it b a s e d o n

in future studies of ser. Piu~una. We h a d difficulty distinguishing

individual a s s e s s m e n t s of c h a r a c t e r states t h a t are n o t clearly

c o l l e c t i o n s 7 3 2 0 and 7325 as S. c h o m a t o p h i l u m or S. j a l c a e

defined (a "gestalt" approach).

a n d o u r identifications of these m a y change. Additional p l a n n e d

2001

SALAS, et al.: P E R U P O T A T O C O L L E C T I O N

TABLE 2 - - - S u m m a r y o f 1 9 9 8 a n d 1999 w i l d p o t a t o ( S o l a n u m sect. P e t o t a ) a n d close o u t g r o u p species (S. l y c o p e r s i c o i d e s , S. s u a v e o l e n s ) g e r m p l a s m collections, c h r o m o s o m e counts, a n d t h e i r s u r v i v a l a t CIP.

Coll. Map no locality of the 1999 collections (Fig. 1)~ 7201 7202 7203 7204 7205 7206 7207 7208 7209 7210 7211 7212 7213 7214 7215 7216 7217 7218 7219 7220 7221 7222 7223 7224 7225 7226 7227 7228 7229 7230 7231 7232 7233 7234 7235 7236 7237

Somatic Germplasm chromosome status 3 number (2n) Species2

Solanum tuberosum subsp. andigena S. acaule S. suaveole~s S. yungasense S. limbaniense (type locality) S. megistacrolobum subsp, toralapanum S. raphanifolium S. raphanifolium S. marinasense (type locality for S. cuzcoense) S. raphanifolium S. lignicaule S. brevicaule S. brevicaule S. tarapatanum (type locality) S. t a r a p a t a n u m 4 S. tarapatanum S. urubambae (type locality) S. laxissimum S. bre~icaule S. piUahuatense (type locality) S. bre~icaule S. lignicaule S. tuberosum subsp. andigena S. lignicaule (type locality) S. u r u b a m b a e 5 (type locality ofS. multiflorum) S. u r u b a m b a e 5 (type locality of S. multiflorum) S. santolallae (type locality of S. santolallae f. velutinum) S. santolallae (type locality of S. santolallae f. velutinum) S. buesi i (type locality) S. buesii S. urubambae 6 (type locality of S. viUuspetalum) S. buesii S. buesii S. incasicum (type locality, see text) S. buesii S. sawyeri? (type locality, see text) S. santolallae

48 48 36 24

CIP CIP, USA CIP, USA CIP CIP

24 24 24

CIP, USA CIP, USA CIP

24

CIP, USA Lost CIP, USA CIP, USA CIP CIP CIP CIP CIP CIP CIP, USA

24 24 24 24 24 24 24 24 24 24 24 24

CIP CIP, USA CIP, USA

48 24

CIP, USA CIP

24

CIP CIP, USA

7238 7239 7240 7241 7242 7243 7244 7245 7246 7247 7248 7249 7250 7251 7252 7253 7254

7255 7256 7257 7300 7301 7302 7303 7304 7305

34 34 35 35 35 29

7306

30

7307 7308 7309 7310

30 32 31 26

7311 7312

28 28

7313 7314 7315 7316 7317 7318 7319 7320 7321

27 27 4 3 3 6 6 6 7

CIP, USA 24 24

CIP, USA CIP, USA Lost Lost CIP, USA Lost

24 24 48 24

CIP, USA CIP, USA CIP CIP

7322 7323 7324 7325 7326 7327 7328 7329 7330 733 7332

S. sawyeri? (type locality, see text) 48 24 S. chiUonanum (type locality) 24 S. velardei (type locality) S. marinasense 4 24 8. marinasense 24 24 S. acroscopicum (type locality) 24 S. lycopersicoides (type locality) S. acaule 48 S. tacnaense 24 S. acroscopicum 7 24 S. acroscopicum 7 24 S. acroscopicum 7 24 S. sandemannii (type locality) 24 S. t a c n a e n s e 4 24 S. sandemannii 24 S. acaule 48 S. megistacrolobum subsp. megistacrolobum f. purpureum (type locality) 24 S. Iongiusculus (type locality) S. m a r i n a s e n s e 7 24 S. aymaraesense (type locality) 24 S. medians var. autumnale 24 S. medians var. autumnale 24 S. medians var. autumnale 24 S. medians var. autumnale 24 S. medians 36 S. b~vvicaule (type locality of S. hapalosum) 24 S. gracilifrons (at or near type locality) 24 S. bill-hookeri (type locality) 24 S. amayanum (type locality) 24 S. huancavelicae (type locality) 24 S. medians var. autumnale (type locality) 24 S. huarochiriense (type locality) 24 S. multiinterruptum var. multiinterruptum 24 S. medians 36 S. immite 36 S. immite 24 S. ja/cae var. ja/cae (type locality) 24 S. ja/cae var. ja/cae 24 S. albicans 72 S. albicans 72 S. jalcae var. jalcae 24 S. chiquidenum var. chiquidenum (type locality of S. chiquidenum var. cachicadense) 24 S. chomatophilum var. chomatophilum24 S. chomatophilum var. chomatophilum24 S. hastiforme 24 S. ja/cae var. ja/cae 24 S. sogarandinum 24 S. chomatophilum var. chomatophilum 24 S. sogarandinum 24 S. hastiforme 24 S. albicans 72 S. chiquidenum var. chiquidenum 24 S. sogarandinum 24

201

CIP CIP, USA CIP, USA CIP CIP CIP, USA CIP, USA CIP, USA CIP CIP, USA CIP, USA CIP, USA CIP, USA CIP CIP, USA CIP, USA

CIP, USA Lost CIP, USA CIP, USA CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP

CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP CIP

202

AMERICAN JOURNAL OF POTATO RESEARCH

TABLE 2 - - C o n t i n u e d . Coll. Map no locality of the 1999 collections (Fig. 1) 1 7333

5

7334

14

7335 7336 7337 7338 7339 7340 7341

14 10 10 9 8 8 8

7342 7343 7344 7345 7346

10 9 11 11 11

7347

11

7348 7349 7350

12 12 12

7351 7352

12 16

7353

17

7354

17

7355 7356

13 19

7357

19

7358

19

7359

18

7360

18

7361 7362 7363 7364 7365 7366 7367

22 22 21 21 21 21 23

Somatic Germplasm chromosome status ~ number (2n) Species2

S. multiinterruptum var. machaytambinum (type locality) S. multiinterruptum var. multiinterruptum S. sogarandinum S. peloquinianum S. peloquinianum (type locality) S. albicans S. • S. hastiforme S. chomatophilum van chomatophilum S. taulisense (type locality) S. • S. ancophilum S. ancophilum (type locality) S. chomatophilum var. subnivale (type locality) S. chomatophilum var. subnivale (type locality) S. ancophilum S. ancophilum S. chomatophilum var. chomatophilum S. dolichocremastrum S. multiinterruptum var. multiinterruptum S. dolichocremastrum (type locality of S. chavinense) S. dolichocremastrum (type locality ofS. chavinense) S. oropizilum (type locality) S. multiinterruptum van multiinterruptum S. multiinterruptum van multiinterruptum (type locality of S. multiinterruptum f. longipilosum) S. multiinterruptum var. multiinterruptum S. multiinterruptum var. multiinterruptum (type locality of S. moniliforme) S. multiinterruptum van multiinterruptum S. medians var. autumnale S. medians var. autumnale S. wittmackii S. simplicissimum S. hypacrarthrum (type locality) S. hypacrarthrum S. hypacrarthrum

24

CIP

24 24 24 72 24 24

CIP CIP CIP CIP CIP CIP CIP

24 24 24 24

CIP Lost CIP CIP CIP

7368 7369 7370 7371 7372 7373 7374

23 23 23 23 22 21 21

7475

21

7376

21

7377 7378 7379 7380 7381 7382 7383 7384 7385

20 20 20 20 25 25 25 25 25

7386 7387

24 24

7388

24

7389 7390 7391 7392 7393 7394 7395 7396 7397 7398

33 19 19 23 23 23 23 23 16 14

7399 7400

15 15

Lost 24 24 24

CIP CIP CIP

24 24

CIP CIP

24

CIP

24

CIP

24 24

CIP CIP

24

CIP

24

CIP

24

CIP

24

CIP

24 24 24 24

CIP CIP CIP CIP CIP CIP CIP CIP

24 24 24

S. cantense 24 S. medians 36 S. cantense 24 S. medians S. cantense (type locality) 24 S. wittmackii 24 S. multiinterruptum var. multiinterruptum (type locality) 24 S. multiinterruptum var. multiinterruptum 24 S. multiinterruptum var. multiinterruptum f. albiflorum (type locality of S. chrysoflorum) 24 S. immite 24 S. simplicissimum 24 S. medians S. wittmackii 24 S. wittmackii S. cantense 24 S. medians van autumnale 24 S. hypacrarthrum S. multiinterruptum var. multiinterruptum f. albiflorum (type locality) 24 S. sogarandinum 36 S. multiinterruptum var. multiinterruptum 24 S. chomatophilum var. chomatophilum 24 S. brevicaule S. medians S. anamatophilum (type locality) 24 S. wittmackii 24 S. wittmackii 24 S. hypacvarthrum S. wittmackii 24 S. arahuayum (type locality) S. augustii (type locality) 24 S. multiinterruptum var. multiinterruptum 24 S. albicans 72 S. multiinterruptum var. multiinterruptum

Vol. 78

CIP CIP CIP CIP CIP CIP CIP CIP

CIP CIP CIP CIP CIP Lost CIP CIP Lost

CIP CIP CIP CIP Lost CIP CIP CIP CIP CIP CIP Lost CIP CIP CIP Lost

~See Spooner et al. (1999, Table 3) for map localities of the 1998 collections. 2Species names of the 1998 collections in bold are changed from those listed in Spooner et al. (1999, Table 3), see footnotes here and text. 3CIP, USA = Germplasm at CIP and also transferred to USDA quarantine for disease screening, with later transfer to NRSP-6 for a seed increase, CIP = Germplasm still all at CIP, lost = germplasm died in attempts to increase it at CIP. r accessions were collected when young and these more accurate identifications wilt be possible with flowering material observed at CIP. ~Ochoa (1999) placed S. multiflorum (this accession collected at the type locality of this name) in synonymy with S. urubambae. 6Ochoa (1999) placed S. viUnspetalum (this accession collected at the type locality of this name) in synonymy with S. urubambae. rPrevionsly listed as Solanum unidentified species.

2001

SALAS, et al.: PERU POTATO COLLECTION

203

collections in northern Peru will allow even more members of

a single highly polymorphic taxon would be convenient, but con-

this series to be collected.

trary to the natural (monophyletic; e.g., Baum and Donoghue

It is possible that the series may be even further expanded

1995) classification we are pursuing in sect. Petota.

by including a character not used by any taxonomist to date. All

A third problem is the apparent expansion of the complex

species placed by Hawkes (1990) and Ochoa (1999) in ser. P/u-

by additional species we observed in the field in 1998 and 1999

r a n a possess tubers in a monilifonn arrangement (arranged like

and data of Ochoa (1999). These additional species include S.

beads on a string, see for example Figures 37, 113, 173, 178 in

aymaraesense Ochoa, S. bill-hookeri Ochoa, S. huancavdicae

Ochoa [1999]), unlike the more typical arrangement of single

Ochoa, S. orophilum Ochoa, S. puchupuchense Ochoa, S. sax-

tubers placed at the end of stolons. This character, like many in

atilis Ochoa, S. sawyeri Ochoa, S. tapojense Ochoa, and S.

sect. Petota, is difficult to assess because some tubers are con-

velardei Ochoa.

stricted in the middle, and it is not clear if they represent transitions to this trait.

Yet a forth problem is the apparent breakdown of the formerly convenient geographical division of the two elements of the S. brevicaule complex at the Peruvian and Bolivian border.

Complex

Ochoa (1999) cited specimens ofS. sparsipilum (relatively wide

Perhaps the most difficult taxonomic problem in sect.

leaves and few interjected leaves) from the southern Peruvian

Petota concerns the species status and interrelationships of the

departments of Cusco and Puno. Two of our collections from

Solanum brevicaule

more than 30 taxa of the Solanum brevicaule complex. Spooner

Cusco that we provisionally label as S. brevicaule match the S.

et al. (1999) provided details of this complex that extends from

sparsipilum morphotype (7213, 7221). However, the morpho-

central Peru to northern Argentina. In short, members of this

logical and molecular studies of the S. brevicaule complex men-

complex show extensive variation, but apparently without any

tioned above placed this species into the southern element of

"species-specific" characters useful for the construction of keys

the complex. Our examination of photos in Ochoa (1999) sug-

and descriptions to clearly define them. Combined data from

gest to us that S. sawyeri (Cusco) also is a match for the S. spar-

morphology (van den Berg et al. 1998) and two molecular mark-

s i p i l u m morphotype (compare Figures 379, 389 of Ochoa

ers appropriate for investigation of closely related taxa (single-

[1999]).

to low-copy nuclear Restriction Fragment Length Polymor-

As practical taxonomists responsible for the stable and

phisms, RFLPs; and Random Amplified Polymorphic DNAs,

repeatable identifications of these economically important and

RAPDs) were all concordant in not being able to clearly define

widely used genetic resources, we are faced with a problem

these species (Miller and Spooner 1999). At best, only two highly

without an apparent easy solution. To address this problem we

polymorphic groups could be recognized, one from Peru, and

are conducting a replicated morphological study of the complex

another from Bolivia and Argentina.

in central Peru (Huancayo, elevation 3200 m, latitude 12~ lon-

The recognition of these two components of the complex

gitude 75~

This location more closely approaches that of cli-

(S. bukasovii as the Peruvian element, S. brevicaule as the Boli-

mate, elevation, and daylength than that of the first

vian and Argentinean element), however, is fraught with prob-

morphological study in the northern United States (Hancock,

lems. First, the morphological data can be used to define these

Wisconsin, elevation 328 m, latitude 44~

two "species" only with many characters that overlap greatly in

these morphological data likewise fail to clearly define taxa, it

longitude 90~

If

range, and then only with computer-assisted multivariate tech-

would further argue for placement of species in synonymy in

niques. Essentially, the characters that define them are imprac-

this group. However, it is yet unclear to us how this will be done

tical to use when germplasm collections from throughout the

considering our inability to clearly key out even the two larger

range are planted together and identifications are attempted

"species", one from Peru and the other from Bolivia and

without knowledge of place of collection, prior identifications,

Argentina. Until such studies are complete, we maintain identi-

or ploidy determinations.

fications of all members of the complex under the clearly artifi-

A second problem is that the two molecular markers

cially broad concept of S. brevicaule.

(RAPDs, RFLPs) suggested that the S. brevicaule complex was not a natural taxon, but rather each of the two elements formed

Solanum ineasicum

separate clades with other species that themselves were mor-

One of the important new collections of the 1998 expedi-

phologically distinct. To lump S. bukasovii and S. brevicaule as

tion was S. incasicum (collection 7234), known from a single

204

AMERICAN JOURNAL OF POTATO RESEARCH

Vol. 78

FIGURE 1 Route of the 1999 expedition to central Peru, and generalized sites for wild p o t a t o e s corresponding to "New Collections in 1999".

SALAS, eta/.: PERU POTATO COIJ,ECTION

2001

locality in Cusco Department. Our field collection of mature flowering and fruiting plants at the type locality was similar to

205

Germplasm Access and Plans f o r Future Collecting

one of the morphotypes of S. raphanifolium as illustrated in

Peruvian materials collected in 1999 and used in this study

Ochoa (1962). This morphological similarity, and the fact that

are not available for international distribution at the moment.

collection 7234 was found at the northern end of the range of

Peru is in the process of finalizing its germplasm access regula-

S. raphanifolium, and its growth in similar habitats as that

tion in conformity with the Convention on Biological Diversity

species led us to suspect that the two species were the same

and the Andean Pact Genetic Resources Decision 391. When the

(Spooner et al. 1999). Plants grown from seed of the original

regulation is officially in place, Peru will make the materials

collection lack the typical morphology of S. raphanifolium

available under the conditions of the official regulation in accord

and S. incasicum likely is a distinct species. Distinctive chloro-

with international agreements. However, any scientist wishing

pl as t DNA r e s t r i c t i o n site m a r k e r s for S. r a p h a n i f o l i u m

to work with the materials in Peru can easily do so. Figure 2 out-

(Spooner et al. 1991; Spooner and Castillo 1997) will allow us

lines the ten of the 24 departments in which we collected wild

to better determine the species limits and relationships of S.

potatoes in 1998 and 1999 and goals for future potato collecting

incasicum.

in the nine remaining departments with wild potatoes, planned

Solarium sawyeri Two important collections in 1998 were possibly S. sawyeri

(7236, 7238), known from a single site in Cusco Department.

after the regulation is in effect.

ACKNOWLEDGMENTS

The type locality was not exact (several kilometers west of

We thank Ned Garvey and Karen Williams of the USDA,

Vestapata), and we found that the correct spelling of this town

Agricultural Research Service, National Germplasm Resources

was Pistipat~- Our collections 0.4 and 1.7 km east of this town

Laboratory; CIP; IPK; STET Holland B.V.; and CGN for collecting

were close matches for the type, and we identified 7236 and

funds; and Stephen Eberhardt of the USDA, Agricultural

7238 as this species. However, the chromosome counts of these

Research Service, National Seed Storage Laboratory, for funds

two collections are both 2n = 48, while Ochoa (1999) provided a

to increase germplasm at CIP for the germplasm collected as

count ofS. sawyeri as 2n = 24.

vegetative material.

Our collections of these two accessions in disturbed areas

We thank Manuel Arca Bielick and Mario Rodrlguez of INIA

in a fruit plantation, and their similarity to cultivated potatoes,

for obtaining permission to collect in Peru; Luisa Huaccho (CIP)

led us to suspect that S. sawyeri was an escaped cultigen. Ochoa

for assistance in map production.

(1999) indicated that S. sawyeri was very similar to S. sparsip-

ilum, a species we place in the S. brevicaule complex. Solanum sparsipilum is so similar morphologically to the cultigen S. tuberosum that it commonly has been advanced as its close relative or parent (e.g., Hawkes 1956; Cribb and Hawkes 1986). We will need more collections at the type locality of S. sawyeri to determine the identity of collections 7236 and 7238 and the species status of S. sawyeri.

New Chromosome Counts C h r o m o s o m e counts are first r e p o r t e d here for S.

c h o m a t o p h i l u m var. subnivale (2n = 2x = 24), S. megistacrolobum subsp, purpureum (2n = 2x = 24), and S. multiinterruptum vat. multiinterruptum f. albifloram (2n = 2x = 24); we also report the first triploid count of an accession of S.

i m m i t e (7314). All other c h r o m o s o m e counts match those reported for these species before (Hawkes 1990; Bamberg et al. 1996; Ochoa 1999; Spooner and Hijmans in press).

LITERATURE CITED Bamberg, J.B., M.W.Martin, J.J. Schartner, and D.M. Spooner. 1996. Catalog of potato germplasm--1996. Potato Introduction Station, NRSP-6, Sturgeon Bay, Wisconsin. Baum, D.A~,and M.J. Donoghue. 1995. Choosing among alternative "phylogenetic" species concepts. Syst Bot 20:560-573. CastiUo, R.O., and D.M. Spooner. 1997. Phylogenetic relationships of wild potatoes, Solanum series Conicibaccata (sect. Petota). Syst Bot 22:45-83. Correll, D.S. 1962. The potato and its wild relatives. Contrib Texas Res Found 3:1-606. Cribb, P.J., and J.G. Hawkes. 1986. Experimental evidence for the origin of Solanum tuberosum subsp, andigena. In: D'Arcy,W.G. (ed.), Solanaceae: Biologyand Systematics. Columbia UniversityPress, New York, pp. 383-404. Hawkes, J.G. 1956. Taxonomic studies on the tuber-bearing Solanums. I. Solanum tuberosum and the tetraploid species complex. Proceed Linnean Soc London 166:97-144. Hawkes, J.G. 1990. The Potato: Evolution, Biodiversity and Genetic Resources. Belhaven Press, Oxford, UI~

206

AMERICAN JOURNAL OF POTATO RESEARCH

FIGURE 2 D e p a r t m e n t s in which wild p o t a t o e s were collected in 1998 a n d 1999, and planned f u t u r e d e p a r t m e n t s to collect.

Vol. 78

2001

SALAS, e t a / . : P E R U P O T A T O C O L L E C T I O N

Holmgren, P.K., N.H. Holmgren, and L.C. Barrett. 1990. Index herbariorum, Part I: The herbaria of the world. Ed. 8. Regnum Veg 120:1-693. Jansky, S.H. 2000. Breeding for disease resistance in potato. Plant Breed Rev 19:69-155. Miller, J.T., and D.M. Spooner. 1999. Collapse of species boundaries in the Solanum brevicaule complex (Solanaceae: S. sect. Petota): Molecular data. Plant Syst Evo1214:103-130. Ochoa, C.M. 1962. Los Solanum tuberfferos silvestres del Peril (Tuberarium subsecc. Hyperbasarthrum). Privately published. Ochoa, C.M. 1999. Las papas de Sudamerica= Peru (page 1). Centro International de La Papa (CIP), Lima, Peru. Ross, H. 1986. Potato breeding - - Problems and perspectives. Adv Plant Breed Suppl 13. Paul Parey, Verlag, Berlin, Germany. Smith, B.W. 1974. Cytological evidence. In: Vascular Plant Systematics, Harper and Row, New York, pp. 237-258. Spooner, D.M., and J.B. Bamberg. 1994. Potato genetic resources: Sources of resistance and systematics. Am Potato J 71:325-337. Spooner, D.M., and R. Castillo. 1997. Reexamination of series relationships of South American wild potatoes (Solanaceae: Solanum sect. Petota): Evidence from chloroplast DNA restriction site variation. Am J Bot 84:671-685.

207

Spooner, D.M., and R.J. Hijmans. In press. Potato systematics and germplasm collecting, 1989-2000. Am J Potato Res. Spooner, D.M., A. Salas-L, Z. Huam~_n, and R.J. Hijmans. 1999. Wild potato collecting expedition to southern Peru (Departments of Apurimac, Arequipa, Cusco, Moquegua, Puno, Tacna) in 1998: Taxonomy and genetic resources. Am J Potato Res 76:103-119. Spooner, D.M., K.J. Sytsma, and J.F. Smith. 1991. A molecular reexamination of diploid hybrid speciation of Solanum raphanifolium. Evolution 45:757-764. Van den Berg, R.G., J.T. Miller, M.L. Ugarte, J.P. Kardolus, J. Villand, J. Nienhuis, and D.M. Spooner. 1998. Collapse of morphological species in the wild potato S o l a n u m brevicaule complex (Solanaceae: sect. Petota). Am J Bot 85:92-109. Wendel, J.F., and J.J. Doyle. 1998. Phylogenetic incongruence: window into genome history and molecular evolution. In: Soltis, D.E., P. Soltis, and J.J. Doyle (eds.), Molecular Systematics of Plants II: DNA sequencing. Kluwer Academic Publishers, Boston. pp. 265297.