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ISSN OOOI-6799 Acta Phytotax, Geobot. 45 (2):95-109 (1994)

Distribution of infraspecific taxa of Petunia axillaris

(Solanaceae) in Uruguay as revealed by discriminant analyses

TOSHIO ANDOi, SHINICHI IIDAi, HISASHI KOKUBUNi,

YOSHIHIRO UEDAi and EDUARDO MARCHESI2

iFkeculty 2Flicultad of Hbrticalture, Chiba U}iiversity, 648 Matsudo, Chiba 271; de Agronomia, Uhiversidad de la Republica, Garzon 780, Montevideo, U)uguay

Abstract. A survey of native habitats of Petunia species over 5 seasons revealed the co-

occurrence of two subspecies of P. axittaris sensu tato in Uruguay. Typical subsp. axiUaris and subsp. parodii were found in south eastern and north western departments, respectively, Using floral measurements (length of corolla tube, corolla limb, pedicel, and calyx lobes, and the length of corolla tube1length of corella limb ratio) of specimens collected in these de-

partments, we performed stepwise discriminant analyses. After substituting measurements of

174 specimens collected in the whole of Uruguay into the three discriminant functions thus

obtained, we determined them to bc either subsp, c[xittaris or subsp. parodii. Distribution of

the two subspecies thus determined by the discriminant functions was roughly delimited by

the Rio Negro. A considerable number of doubtfu1 specimens were found in the south west- ern dcpartmcnts, which are on opposite sides of the river de La Plata from the place where secondary intergradation betweell thc two subspecies had been suggested by Wijsman (1982) to have occuiTed,

Key words: discriminant function, disnibution, gardcn petunia, geographical taxa, Petunia urillaris, Solanaceae.

Received Plebruary 14, i994; accqpted September 30, l994

In 1834, the breeding of garden petunias was started by crossing Petu- nia axitlaris (Lamarck) Britton, Sterns & Poggenburg and P. integrijlotia (Hooker) Schinz & Thellung (Sink, 1984), which had been collected at Montevideo, Uruguay (Lamarck, 1793; Wijsman, 1982), and Rio Negro, Uruguay (Hooker, 1831), respectively. Although these species also grow in Argentina and Brazil (Fries, 1911), the parents of the first Petunia hybrid originated in Uruguay. To know the genetic background of garden petu- nias, detailed information about the species of Petunia native to Uruguay would be usefu1. During tours for five seasons to observe the habitat of Petunia species in Uruguay, we found the co-occurrence of two subspecies of P. axiUaris, that is, subsp. axillaris (short corolla tube with large corolla limb) and subsp. parodii (Steere) Cabrera (long corolla tube with small corolla limb). Subspecies parodii was initially found in Formosa Province in Argentina,

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96 Acta Phytotax. Geobot. Vol, 45

and had not been reported in Uruguay (Wijsman, 1982). In order to develop appropriate methods to distinguish subsp. axiUaris from subsp. parodii and to know the precise distribution of these subspe- cies of P. axillaris in Uruguay, we conducted discriminant analyses using floral characters measurable in dried specimens.

Materials and Methods

Specimens studied One hundred and thirty-four specimens of Petunia axiUaris sensu lato were collected during five visits to Uruguay (Nov. 15 to Nov. 29, 1988; Nov. 25 to Dec. 4, 1989; Nov. 8 to Nov. 25, 1990; Oct. 31 to Nov. 10, 1991; Dec. 9 to Dec. 19, 1992). Some of the specimens were donated to the MVFA (Laborat6rio de Botanica, Facultad de Agronomia, Monte- video, Uruguay) as indicated in Appendix 2. The distance that we traveled in Uruguay exceeded 20,OOO km. In addition, forty specimens of P. axilla- ris sensu tato collected otherwise and kept in the MVFA were also used for this study.

Characters for discriminant analysis Even though the condition of stamens, that is, didynamous (subsp. ax- illaris) or not (subsp. parodii), had been regarded as an important charac- ter to distinguish infraspecific taxa of this species (Steere, 1931), it was dif- ficult to correctly measure this feature in the fragile inner corolla organs without damaging the specimens. Therefore, characters related to stamen condition were not used in this study. For the study of herbarium specimens, Wijsman (1982) had recom- mended using the length of the corolla tube (from the lower end of the calyx to the point separating the tube and limb) to distinguish subspecies of P. axiUaris sensu tato from each other. In addition to this parameter, the length of the corolla limb (from the point separating the tube and limb to the tip of the limb), pedicel, and calyx lobes (from the base of the calyx lobe to the tip of the longest calyx lobe) were also measured in this study. The corolla tube length 1 corolla limb length ratio was also used since a similar ratio (corolla tube length 1 corol]a limb diameter) had been re- garded as a good feature fbr distinguishing live infraspeeific taxa of P. axil- laris sensu tato (Wijsman, 1983). The mean value of all floral measure- ments per specimen was used for subsequent analysis.

Discriminant analysis Stepwise discriminant anaiysis was performed by use of the SPSSX software program provided by the Computer Center of Chiba University. The criterion used for determining the selection of characters was the mini- mized Wilks' lambda. The lower limit for selecting or removing a character was based on a partial F-ratio, which was set at F l 1.000 to enter and F < 1.000 to remove. The minimum tolerance level was set at O.OOI. The re- maining conditions for the stepwise discriminant analysis were set to the default values as described in the User's guide of SPSSX (SPSS Inc.,

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1986). Using floral measurements of specimens collected in the south eastern (Canelones, Maldonado, and Montevideo; 33 specimens of typical subsp. axillaris) and north western departments (Artigas and Salto; 27 specimens of typical subsp. parodii), we performed stepwise discriminant analysis. By substituting measurements of the remaining specimens into discri- minant functions thus calculated, we determined all specimens to be either subsp. axillaris or subsp. parodii.

Results and Discussion

F7equency distribution of corolla tube length Frequency distribution of corolla tube length measured in all 174 specimens of P. axillaris sensu lato is shown in Fig. 1. We could not con- firm a clear separation of the frequency distribution into two groups (groups of subsp. axillaris and subsp. parodii) as had been shown by Wijs- man (1982). According to his figure, which is sirnilar to Fig. 1, he con- cluded that dried specimens with corolla tubes shorter than 46 mm and those exceeding 46 mm could be identified as subsp. axillaris and subsp. parodii, respectively.

71ypicat subsp. axillaris and subsp. parodii in Uruguay Observations of flowers of live plants in their native habitat revealed that all Petunia axiltaris sensu lato growing in the south eastern depart- ments (Canelones, Maldonado, and Montevideo) were typical subsp. axit- inris, as they had fbur strongly didynamous stamens (2 long, 2 medium, and 1 short), a short corolla tube, and a larger corolla limb. As mentioned earlier, Montevideo is the type locality of this subspecies.

30 :oE'agop6nosEsz

20

10

30 ao 50 6a 70

Length of corolla tube (mm)

Ft{/}. 1. Frequency of corolla tubc length in the specimens of Uruguayan Petunia axillan's used in this study.

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On the other hand, all Petunia axillaris sensu lato growing in the north western departments (Artigas and Salto) were considered to be typical subsp. parodii because of 4 longer stamens ef equal length with 1 short sta- men in addition to a long corolla tube and smaller corolla limb as de- scribed by Steere (1931). This area is close to the type locality of this sub- species, that is, Formosa Province, Argentina. Even though the degrees of the didynamous condition were not re- corded in our field survey, stamens of P. axillaris sensu lato observed by us in the departments south of and north of the Rio Negro were strongly and weakly didynamous, respectively. Variations in stamen condition of live plants grown from seeds will be reported elsewhere. Except for stamen condition, differences in the floral characters men- tioned above were also confirmed in dried specimens (Table 1). According to Wijsman's (1982) criterion as mentioned earlier (critical length of corol- la tube = 46 mm), it is also reasonable to say that the south eastern and north western taxa of P. axillaris sensu lato are subsp. axillaris and subsp. parodii, respectively. As shown in Table 1, differences betwecn five characters in specimens from south eastern departments and those from north western departments were significant. In addition to the corolla tube length, therefore, all other characters measured in this study could be used as additional characters to discriminate these taxa from each other, even though Wijsman (1982) had stated that no character distinguishing subsp. axillaris from subsp. parodii had been found except for corolla tube length.

Discriminant analysis At first, the discriminant analysis was performed using all five charac- ters (length of corolla tube, corolla limb, calyx lobes and pedicel, and ratio of corolla tube length to corolla limb length). Pedicel length and corolla tube length ! corolla limb length ratio, however, were removed from the optimum subset for discriminating subspecies since their F values were less than 1.000. The discriminant function thus obtained was as follows: - Zi = O.2168763Xi + 0.4312565X2- O.2530165X3 9.321480 ･･･････････････function 1

TABLE 1. Comparison of floral characters rneasurable in dried specimens of Petunia axiUaris subsp. axitlatty from south eastern departments (Caneloncs, Maldonado & Montevideo) and subsp. paro- dii from north westem departments (Anigas alld Salto),

Length of Ratio

Corolla Corolla Calyxlobe Pedicel(mm) Tube1

tube (mm) limb (mm) (mm) limb

Subsp. axillaris 41.758,7 22.117.7 7.5lo.e 29.038.5 1.923.45

Subsp. parodii

All charactcrs wcre significantly different at u O.1 % level using the t-test.

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Xi=corolla tube length in mm X2=calyx lobe length in mm == X3 corolla limb length in mm The length of the corolla limb is sometimes difficult to measure de- pending on the condition of the specimen. If data of corolla limb length are not available, an alternative discriminant function, which was calcu- lated from the lengths of the corolla tube, calyx lobe, and pedicel, was obtained as follows: Z2= O.2147034Xi+ O.2841115X2- 13.03891 ･･･････････････function 2 Xi=corolla tube length in mm X2=calyx lobe length in mm In this case, pedicel length was again removed from the optimum subset of characters. If data from corolla tube length only were employed as in Wijsman's (1982) work, the fbllowing discriminant function was obtained: Z3 == O.2432504Xi- 11.99833 '･･････････････function 3 Xi=corolla tube length in mm This function indicated that the critical value between subsp. axitlaris and subsp. parodii js approximately 49 mm (more precisely 49.33 mm) in the length of corolla tube, which was 3 mm longer than the value fer Wijs- man's (1982) criterion. Briefly, a specimen of Petunia axillaris sensu lato can be determined as subsp. axillarts or subsp. parodii if discriminant scores, Zi, Z2 and Z3 values mentioned in the functions, are minuses or pluses (Appendix 1).

Statisticat features of discriminant fiznctions Statistical features evaluating these functions are listed in Table 2. Wilks' lambdas of the respective functions were significant, indicating the clear separation of the group means. The canonical correlation coefficients of the respective functions indicated a close association between the discri- minant score and the group, as shown in Table 3. In contrast, Box's M values and their significant level of the functions varied considerably depending on the function (Table 2). Their significant levels were too high to ensure the equality of the group covariance mat- rices except for function 3 (significant at a 5 % level). After substituting all measurements into the 3 functions obtained

TABLE 2. Statistical features of the 3 discriminant functiens obtained frem floral charucters in the in- fraspecific taxa of Petunia exiltaris in Uruguay,

Box's M Wilks'lambda Canonical Function (significant level %) correlatien cocfficient

Function 1 12.222 ( 7.3) O.100*O.170*O,187* O.949O.911O.902 Functien 2 5,940 (12,7) Function 3 5.802 ( 1,7)

* Significant at O.1% level.

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TABLE 3. Standardized canonical discriminant function ceefficiellts for the 3 discriminant fimctions obtained in this study.

Length of Function Corolla tube corolla limb Calyx lobe -O.778 Function 1 O.892e.8831.0oo O.508O.335

Function 2

Function 3

(Appendix 1), we calculated probabilities of discrimination error in the re- spective discriminant functions as follows: p = O.214 % (function 1), p == 1.473 % (function 2), and p = 1.991 % (function 3). Doubtfu1 ranges of the discriminant functions at a O.5 % level were -O.668 -O.131 -O.877 calculated as follows: < Z2 < (function 2) and < Z3 < O.149 (function 3). No doubtful range of function 1 at a O.5 % level existed. In function 3, the doubtful range is otherwise described as 45.7 mm to 49.9 mm for the length of the corolla tube at a O.5% level. Unfortunately, we hesitate to select the best one among the 3 discri- minant functions mentioned above for discriminating subspecies because each function had its own weak point. In such cases, it is preferable to col- lectively identify subspecies by using all 3 discriminant functions.

Distribution ofsubsp. axMaris and subsp. parodii in Ukuguay Localities of specimens used in this study are shown in Fig. 2. Petunia axillaris sensu lato actually grows throughout Uruguay except in the north eastern department of Rivera, even though an available distribution map of P. axiUaris sensu lato had excluded north western departments such as Artigas and Salto from its distribution area (Wijsman, 1982). It is found in great abundance in the southern departments along the Atlantic coast and along the river de La Plata. As reported by Aoki & Hattori (1991), corolla shape in species of Petunia is significantly modified by the ambient growing temperature. Strictly speaking, therefore, the morphology of the corolla should not be compared between specimens from different habitats. Even though such modification of the floral morphology by the environment could be in- volved, our herbarium study revealed the following points; As shown in Fig. 2, solid and open circles indicated the localities of the specimens that were regularly predicted by all discriminant functions as subsp. axillaris (90 specimens) and subsp. parodii (48 specimens), respec- tively. Triangles denote the location of doubtfu1 specimens (36 specimens) because they were determined as doubtful at a O.5 % level by any one of the discriminant functions, or were not regularly determined as either subsp. axillaris or subsp. parodii (Appendix 1). In Uruguay, subsp. axillaris and subsp. parodii are apparently south eastem and north western taxa, respectively, roughly separated by the Rio Negro, although considerable numbers of doubtful specimens were in- cluded (Fig. 2). This finding supports the opinion that these are geog-

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The Atla"rtc Ocean

FiG. 2. Specimen localities of Petunia zaxiltarts senstdato and thc prediction uccording to the discrimi- nant functions obtained in this study, Solid and open circles are determined Tegularly by all func- tions as subsp. axittaris or subsp. parodii, respectively. Triang]es represent doubtful determinations at aO.5 % level by any one function, or are those specimens not regularly determined to be subsp. axiilaris or subsp, parodii.

raphical subspecies of P. axillaris sensu lato (Cabrera, 1977, Wijsman, 1982). Two specimens from PaysandU (Ando & Shibata U265 & U267) were determined as subsp. axillaris by all functions, but our observation of fresh flowers in the natural habitat showed that they were subsp. parodii, having 4 stamens of equal length. The overall small size of the corolla, possibly caused by the environment in a given year, could be the reason for the erroneous determination (data not shown). It is interesting to note the distribution of doubtfu1 specimens, because they were conccntrated in the south western departments, such as in Col- gnia and San Jos6 (Fig. 2). Wijsman (1982) had suggested that secondary mtergradation between the geographical subspecies of P. axillaris sensu lato had occurred on the Argentina side of the Plata region (Argentina side of the river de La Plata, Buenos Aires, and the border of the Uru- guay river), which is on the opposite side of the river de La Plata from the department of Colonia. Even though further research will be necessary be-

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fore any conclusive statement can be made, the possibility that intergrada- tion between subsp. axillaris and subsp. parodii had also occurred in the south western departments of Uruguay, that is,on the Uruguayan side of the Plata region, can not be denied.

This work could not have been accomplished wjthout the kind assistance of Mr. Masao Udagawa and Mr. Tomohiro Kamegawa, Montevideo, Uruguay, in surveying the habitats of Petunia.

References

Aoki, C. & K. Hattori. 1991, Attempt to classify Petania species on the basis of corolla shape. Jap. Journ. Breeding 41: 433-442. Cabrera, A. L. 1977. Novedades en las Solanaceas de Entre Rios. Hickenia 1(14): 79-81.

Fries, R. E. 1911. Die Arten der Gattung Petunia. Kungl. Svenska Vetenskapsakad. Handl, 46: 1-72,

Hooker, W. J, 1831, Saipigtossis integrijblia. Entirc lcavcd Saipigtossis. Curtis's Bot. Mag. 58: t.3113.

Lamarck, J, B, P. A. M, 1793, Nicotiana axitlaris, Tableau Encycl, Meth, Trois Regnes Nat. Bot, 2: 7, Sink, K, C, 1984. Taxonomy, In K, C, Sink (cd,), Petunia. Springer-Verlag. SPSS INC, (author unknown). 1986, Discriminant. In SPSSX User's Guide, 2nd edition. pp, 689-714, SPSS Inc, Steere, W. C. 1931, Petunia parodii, a new species of the subgenus Pseudenicotiana from Argcntina. Pap. Michigan Acad. Sci. Arts Let. 13: 213-215. Wijsman, J. R. 1982. 0n the inter-relationships of certain species of Petunia I. Taxonomic notes on the parental species of Petunia hybrida, Acta Bot. Neerl, 31: 477-490, . 1983. 0n the inter-relationships of certain species of Petunia II, Experimental data: Crosses between different taxa. Acta Bot. Neerl, 32: 97-107,

Appendix 1 -O.082,602,992.693.572.sc DiscTiminant scores of 174 specimens of Petunia Ando & Kokubun U3S 2,593.243.933.543,193.56O.493.143.523.013.472.96

axillatis sensu lato by 3 discriminant functions Ando & Iida U217

obtained in this study. Doubtful ranges at O,5 % Ando & Iida U218

level are underlined, Ando & Iidu U22e Ando & Iida U221

Discriminant scores Ando & Iida U223 Departments and Canelones speclmens -1.99-3.17-1.03-],62-2,23-2.06-1.26-2,51-1.32-1.58-1.75-1.42･-1,17-251-1.54-1.10-1.47-1.42 Func. IFunc. 2 Func, 3 M.B.Berro 446 M.B.Berro 6692

Artigas N.de Medina sln

N.G.Zorron 840 3,784.642,952,504.012,633.501.863.482191.763.111.091.372.111.452.44O.782.541.392.113,081.141.502231.382.69O.651.751.38 Ando & Kokubull Ul

N.G.Zorron 1025 Ando & Buto U175

Marchesi 10224 Ando & ]ida U242

Ando & Kokubun U27 Cerro Largo -1.67-1,81-1.45-1.67-1.66-1,54-o.os-o,"-2.20 Ando & Kokubun U29 Millot 57S

Ando & Kokubun U31 Lema et at. 7951

Ande & Kokubun U32 Lema et al. 7964 1.08-O.84-1,81 O,50-O.31-2.08

Ando & Kokubun U34 Ando & Kokubun U55

Ando & Kokubun U35 Ando & Shibata U258

Aiido & Kokubun U37

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-2.31-4.02-2.51-2,30-O.65-2.78-2.66-2.67-3.40--3.74-2.41-3,SO-3,47-4,44-2.20-3.17-2.38-1,69-1,47-2.02-O.41-2.27-1.78-1.54-2,e7-O.44 Colonia ArrMaga et al. 2069 -O,32-O.81-O.44-O.93-1,39-O.20 Puerto et al. 5311 Del Puerto et al. 9016 O.62-1.M O.29-1,26-O.79-1.62-1,89-O,76-o.ca Del Ando & Kokubun Uco Ando & Kokubum U61

Ando & Kokubun U91 O.67-1,13-2,10-O.65-IA2-3.10 Ando & Kokubun U63

A]do & Kokubun U92 Ando & Kokubun U65 -1.98-2.13-1.94-2.05-3.76-2.14-1.78-IM-2.88-1,49-1,80-･1,54-1,78-1,54-1,66-3.10-t.71-1.71-O.91-2.10-1,M-1.61 Ando & Kokubun U93 Ando & Kokubun U74

Ando & Watanabe U98 Aiido & Watanabe U14g

Ando & Watanabe U99 O.26-o.os Ando & Watanabe U149

-osc-O,Ol U151 Ando & Wutanabe UIOO Ando & Watanabe Ando & Watanabe UIOI O.58-O.66-L16-1,88-O.08 O,16 Ando & Watanabe U152 -O.24 Ando & Watanabe Ulee O.16-O.35-1,51 Ando & Watanabe Ul53 -O.72-1.73 Ando & Watanabe UI08 Ando & Buto U176

Ando & Buto U173 Ando & Buto U177

Ando & Buto U199 O.06 O.24 Aiido & Buto U178

Ando & Buto U200 O.96 1.37 O.53 Ando & IidaU244 Durazllo Ando & Iida U245

Zilianl et al, 10579 5.lg-1.91-1,99 4.l9 3.81 Montevideo -O,30 -O.20 -1.97-O,63-2,02-3,93-4.20-2.73-2.16-1.36-3.05-Lca-1,30-1.72-3.47-3.22-2.71-1,oo-1.46-1.58-1.30-1,78-2,39-3.48-3.73-3.oo-1.es-1,13-1.32 Ando & Kokubun U5 M,B,Berro 3Sll

-1.12 -1.13 Ando & Kokubun U21 M,B,Berro 6034 Ando & Buto U159 2.49 1,73 1,62 M,B.Berro 6693 Flores RosengurttB-2115

-O.93 -2.24 -2.63 360 Rosengurtt B-621 NrMaga Ando & Buto U167 4.17 2.35 2,11 izaguirre et al. 10728 -O.56 -1.38 -1,44 Ando & Iida U2as ? 1045 -3.29 -1,57 -1.3e Ando & Shibata U268 Ando & Kokubun U86 Rorida Ando & Buto UIS7 -121 -1,78-e.95-3.00-1.85-O,86-O.35 Gallinal et aL PE-5036 O.27 Paysandtl -2.14 -1.04 Ando & KQkubun U2 Ando & Watanabe U124 2,32 2,49 221 -2.60 -2.95 Ando & Kokubun U3 Ando & Watanabe U126 1.13 1.oo-122 1.28-2.03

-1,S7 -2,14 Ando & Watanabe UI09 Ando & ButoU197 O.36 -1,59 -1.36 Ando & Watanabe UllO Ando & Buto U198 2.13 1.45 O.55 -O,97 -O.61 Ando & Watanabe Ul13 Ando & Iida U237 3.el 2.54 2.23 Lavalleja Ando & Iida U238 3.S4-O,80-327 3,06-1.67-4.57 2,52-1,54-4,70 -2,67 -2,38 -2.03-2,51-1.05-2.15-3.00-L30-2.51--1.U5-2.03-1,78-1.IS-e.69-1.0S-1,17-1.95-1.30-1.54 Del Puerto et al. 7542 Ando & Shibata U265 -2.81 -2,78 Lombardo et at. 11918 Ando & Shibata U267 -104 -O.9S Ando & Kekubun U49 RioNegro -2.91 -2.69 Ando & Kokubun U51 RosengurttB-4079 5.261.S22.242.S21,oo3,501.881.883.12L163.571.961.452.45O.65 -3.71 -3.35 Ando & Kekubun U58 Ando & Bute Uloo

Ande & Kokubun U60 -1.48 -1.55 Ando & Buto U162 -2.89 -2.50 Ando & Kokubun U76 Aiido & Buto U163 -O.64 -1.09 Ando & Kokubun U79 Ando & Shibata U262 -2,45 -2.23 Ando & Kokubun U80 Rocha -2.11 -1.89 -2,06-2,56-4,34-2.51-3.16-2.50-225-2.50-3.25-O.70-2.06-2.05-1.an-3.27-1.67-2.03-2.64-L89-2.34-3.35-1.06-1.42-2.27-2.03-3,co-1.30-1,47-2.IS-1.66-2,"-2.88 Ande & Kokubun U83 RosengurttB-2459 -2.08 -1.S3 Ando & Watanabe U 141 Arrillaga 206 -1.72 -O.89 Ando & Watanabe U143 Rodriguez et al, 8140 -1,71 -1.29 Ando & Watanabe Ulas Ando & Kekubun U67 -2,08 -1.60 Ando & Watanabe U147 Ando & Kekubun U68 -Z.81 -2.03 Ando & Buto U204 Ande & Kokubun U69 -2.06 -1.le Ando & Buto U206 Ando & Kekubun U71 -1.8S -1.48 Ando & Iida U2S3 Ando & Kokubun U72 Maldonado Ando & Kokubun U73 -3.94 -2.55 -2.e3 M.C.Berro 7200 Ando & Buto U154 o.gl-1.17 -3.65 -3.95 -3.48 Arrillaga et al. 1599 Ando & Buto U156

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-4,26-2,85-2as-1,93-2.66-2.47-1,87-e.39-1.91-3.15-2.57-2,34-2.24-1,71-O.75-O,78--1.49-L19-1.53-O,15-2,51-1.Sl-O.64-O.76-1."-1,22-1,61-O.15 Ando & Buto U179 Del Puerto 1002 -O.02-O.05-1.37-O.16-O.57

Ando & Buto U180 Dcl Pllerto 2185 -O.93-O.81-020 Ando & Buto U181 J,Ren 10883 O.38 -O.92-O.08 Aiido & ButoU182 Del Puerto et al. 12691 O.63-O,20-O.91-1.66-O.91-1.28-122-1,96-2,Ol Ando & ButoU183 Ando & Kokubun U25 1.13-2.11-1.42-O.34-1,68-O,76-1,30-1.78 IA4-1.66-1.05-O,20 Ando & Buto Ulsc Ando & Kokubun U85

Ando & Buto U185 AndQ & Kokubun U87

Ando & Bute U186 Ando & Kokubun U88

-O,99-2,24-1,49-1.80-O.71 Ando & IidaU247 Ando & Kokubun U89 -1,05-O.27-O,81-2.07

-2.03-1.78-l.64 Ando & IidaU248 Ando & Watanabe U96 Ando & IidaU251 Ando & Watanabe U97

Ando & lida U252 Ando & Buto U174 Salto Soriano Millot 502 4.792.641.733.623.174.175.444.382.652.38O.822,791,721.313.352.664.025.4942Y2.031.64O,983.33L141.282.6e2.673,815.394,371.891.45O.82 M.B,Berro 2951 1,99 O.79 e.41 Del Puerto et al. 11474 Ando & Watanabe UI04 1,59 O.97 O.60 Ando & Kokubun U26 Ando & Buto U166 2.45-O.72-O.83-o,ca 2.01-･1.72-2,02-1.68 1.92-2.07--2,32-2,03 Ando & lida U224 dndo & Buto Ul 69 Ando & Iida U225 Ando & Buto U170 Ando & Iida U226 Ando & Buto U171 Ando & iida U228 Ando & Buto U201 O.80 1.65 1.58 Ando & Iida U230 Tacuarcmb6

Ando & IidaU232 Ando & Kokubun U19 2.88228O.953,933.433,03O.683.293,402.72O.312.79 Ando & Shibata U271 Ando & Kokubun U20 Ando & Shibata U272 Ando & Watanabe U121 San Jos6 2indo & BLto U195 -O,87-O.54-O,32-O.32 L.C.de Zolessi 318 O,13O,62O,12 Treinta y Tres -1,87-1,80-1.25-2.25-O.91-2,22 N.G.Zerroll1562 Ando & Buto Ul87 -O.47 -O.64 Del Puerto et al. 676 Ando & Buto U188

Appendix 2

7>lp}ical forms of Petunia axillaris subsp. axillaris [[he following specimens wcre determined to be Petunia axillaris (Lamarck) Britton, Sterns & Pog- gellburg subsp. axillaris according to all discrirninant functions obtained in this study. Dep. CANELONES: Independencia (8 Nov. 1899 M, B. Berro 446 MVFA); Las Hedras (13 Jan. 1913 M. B. Berro 6692 MVFA); Costas del Rio Sta. Lucia (29 Oct. 1933 IVieves de Medina sln MVFA); Route 5, O,3 krn E. of La Paz to Camino Uruguay, S34"45'48" W56013'14" (16 Nov. 1988 T. Ando & H. Kokubun Ul MVFA, Ando); Route 8, Pando, S34'43'35" W55a5747" (14 Nov. 1990 T. Ando & K. Buto UIZ5 MVFA, Ando); Route 5, Betw ¢ en Las Piedras and Progreso, S34042'44" W56013'03" (5 Nov, 1991 T. Anclo & S. Iida U242 MVFA. Ando), Dep. CERRO LARGO: Ruta 8, km 381, litosol sobre esquistos de Rincon de Alonso (26 Oct. 1961 MiUet 5Z5 MVFA); Sierra de Rios, 5 km antes de pico mas alto (23 Nov. 1968 Lema, Olano, Rodriguez & K)'aU 7951 MVFA); Route 7, 9.3 km N.E. oi Santa Clara de Olimar to Tupambae, S32as238" WS4"52'18" (12 Dcc. 1992 T. Ando & K. Shibata U258 MVFA, Ando). Dep. COLONIA; Colonia del Sacramento, 5 m, (Z3 Dec. 1988 71 Ando & H. Kokub"n U92 MVFA, Ando); Parque Forestal, 10 m (23 Dec. 1988 T, Ando & H. Kokubun USB MVFA, Ando); Juan Jackson (23 Nov. 1990 T. Ando & K. Bute UIZ3 MVFA, Ando). Dep. DURAZNO: Route 5, 12.3 krn N. of Rio Yi to Tacuaremb6, 280 m (18 Nov. 1988 ZAndo & H. Kokubun (121 MVFA, Ando). Dep. FLORES: Rio Yi entre Arroyos Matallzas y Carpinteria, Estancia de Quinteros (25 Nov. 1936 Rosengurtt B-621 MVFA); Route 3, 13.0 km S.E, of Greco to Trinidad, S33012'41" W57e05'5e", 93 m (4 Nov. 1991 Z Ando & S. fida U240 MVFA, Ando); Route 23, 14,1 km N. of Ismael Cortinas

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to Trinidad, S33051'47" WS7"02'e2" (16 Dec. 1992 Z Ando & K, Shibata U268 Ando). Nov. 1988 Tl Ando & Dep. FLORIDA: Route 5, 1.1 krn S. of Rorida to Canelones, 140 m (16 CB H. Kokubun U2 MVFA, Ando); Route 5, La Cruz, 120 m (16 Nov. 1988 T Ando & H. Kokubun Ando); Route 7, Rio Santa Lucia, S34e12'49" W55044'34' (29 Nov. 1989 T. Ando & H. Watanabe UI09 Ande). P"erto & Marehesi Dep, LAVALLEJA: entre Casupa y Polanco, en cerrito (27 Oct, 1968 Del MVFA); rs42 MVFA); entre Minas y Pan de Azucar (21 Dec. 1972 Lombardo & Del Puerto IJ918 Route 8, 4.4 km N.E. of Arroyo Campancro Grande to Trcinta y Tres, 130 m (24 Nov. 1988 71 Ando & H. Kokubun U49 MVFA, Ando); Route 8, 4.2 km N. of Mariscaltt to Treinta y Tres, 150 m (24 Nov. 1988 T. Ando & H. Kokubun U5i MVFA, Ando); Route 8, 9.0 km S, of Hraraja to Minas, 9e km N, of Aigufi,170 m Nov. m (26 Nov. 1988 T. Ando & H. Kokubun [L58 Ando); Route 39, 10.6 (26 1988 T. Ando & H. Kok"b"n U60 Ando); Route 13, 8.9 km W. of Arroyo del Aiguti to Minas. 190 m W. of Arroye del (27 Nov. 1988 T. Ando & H. Kokuhun U76 MVFA, Ando); Route 13, 16.8 krn N.E. of Aiguti to ]dinas, 140 m (27 Nov. 1988 T. Ando & H. Kekubun U79 Ando); Route 8, g.5 km Arroyo del Pcnitente to Treinta y Tres, 160 m (27 Nov. 1988 T. Ando & H. Kok"bun U80 wwA, Ando); Route 60, Between Minas and Pan de Azucar, 40 K, 190 m (27 Nov. 1988 Z Ando & H. Kokubun U83 MVFA, Ando); 3.0 km N.E. of Minas to Arroyo del Perdido, 30 m (2 Dec. I989 T. Ando & H. Wl:tanabe U141 MVFA, Ando); 33.2 km N.E. of Minas to Arroyo del Perdido, 210 rn (2 Dec, 1989 T. Ando & H. Watanabe U146 Ando); 53.6 km N.E. of Minas to Arroyo del Perdido, 60 m "Penitcnte'" (2 Dee. 1989 T. Ando & H. vatatanabe Ul47 MVFA, Ando); Cttscadas de Agua del (Z4 Nov. 1990 T. Ando & K. Buto [J204 Ando); Gruta y Cerro Arequita, S34e17'23" WS5016'05" (24 Dec. 1990 71 Ando & K. Buto U206 Ando); Route 14, 15.1 km N.W. of Lascano to Jos6 P. Varcla, Rio Cebollati, S33036'22" W54020'21", 98 rn (7 Dec. 1991 T. Ando & S. fida U253 MVFA, Ando). Oct. Dep. MALDONADO: MaLdonado (21 Oct. 19I3 M. C. Berro Z200 MVFA); Ruta 12 (12 1963 Arritlaga, Izaguirre & Brescia 1599 MVFA); Punta Ballena (23 Apr, 1964 Arrillaga, Ifaguirre & Laguardia 2069 MVFA); Playa Hermosa (7 Oct. 1965 Del Puerto & Marchesi 5311 MVFA); Route le9, 11.5 km S.E. of Aigua to Rochu, Arroyo Saucc, 140 rn (26 Nov. 19S8 T. Ando & H. Kokub"n U61 Ando); Route 109, 19.8 km S.E. of Aigua to Rocha, 330 m (26 Noy. 1988 T. Ando & H. Koku- bun U63 MVFA, Ando); Route 13, 15.7 km N.E. of Aigua to Lascallo, Arroyo Sarandi Grande, 40 m of Minas to Maldonado, 250 (27 Nov. 1988 T. Ando & H. Kvkubun U74 Ando); Route 12, 21.5 km S. Minas to Maldonado, m (3 Dec. 1989 T. Ando & H. Wdtanahe [ll48 Ando); Route 12, 28,6 km S, of tOO m (3 Dec. 1989 T. Ando & H. VVatanabe Ul49 MVFA, Ando); Routc 10, 2.4 km W. of Maldona- do (3 Dec. 1989 T. Ando & H. Watanabe [ll51 MVFA, Ando); Punta Negra (3 Dec. 1989 T. Ando & H. IVatanabe Ul52 Ando); Route 10, Arroyo Solis Grande (3 Dcc. 1989 T. Ando & H. Wdtanahe U153 MVFA, Ando); Routc 9, 1.9 krn E. of Arroyo Pan de Azucar to San Carlos, S34046'50" WS5013'11" (14 Noy. I990 71 Ando & K. Buto UJ 76 Ando); Route le, 7.4 km E. of La BaTra de Mal- denado to La Paloma (15 Nov. 1990 T. Ando & K. B"to CU 77 MVFA, Ando); Route 10, 6.3 km E. oi km E. of L. Jos6 Ignacio to La Puloma (15 Nov. 1990 T. Ando & K. Buto tU78 Ando); Route 9, 28.5 Solis to San Carlos, Arroyo Sauce, 121 K, S34"44'23" W55004'46", 102 m (6 Nov. 1991 T. Ando & S. Iida U244 MVFA, Ando); Route 9, 4,6 km W. of Arroyo Garzon to San Carlos, S34n41'37" W54e34'17Z 38 m (6 Nov. 1991 T. Ando & S, fida U245 MVFA, Ando). Dep. MONTEVIDEO: Cerro (10 Nov. 1906 M, B. Berro 3511 MVFA); Malvin (1 Mar. 1911 M. B. Berro 6e34 MVFA); Punta Carretas (21 Nov. 1912 M. B. Berro 6693 MVFA); Pajas Blancas, en 360 costa maritima (26 Sept. 1937 Rosengttrtt B-2115 MVFA); Parque Lecoq (30 Scpt. 1956 Arrillaga MVFA); Ccrro (13 Oct. 1971 fzagzairre, Laguardia & Marchesi 10728 MVFA); costa Ramirez (.? I045 MVFA); Route 1, O.3 km E. of Rio Santa Lucfa to Montevideo, 5 m (21 Dcc. 1988 T. Ando & H. Kekubun U86 MVFA, Ando); Cerro, S34053'17" W56015'35" (10 Nov. 1990 71 Ando & K. Buto U157 MVFA, Ando). Dcp. PAYSANDO: Casa Blanca, S32023'48" WS8009'43" (15 Dec. 1992 T. Ando & K. Shibata U265 Ando); Route 90, 49.7 km E. of Esperanza to Guich6n, S32023'2e" W57a26'05" (15 Dec. 1992 T. Ando & K. Shibata U267 MVFA, Ando), Dep. ROCHA: Arroyo La Pantallosa (5 Feb. 1938 Rosengurtt B-2459 MVFA); Sta. Teresa (Jan. 1956 Arrittaga 206 MVFA); Parque Sta. Tcresa (23 Dec. 1968 Rodriguex, Coll & Biagioni 8140

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106 Acta Phytotax. Geobot, Vol, 45

MVFA); Route 15, 10.1 km S.E. of Rocha to La Paloma, 40 m (Nov. 1988 T. Ando & H. Kokubun U67 Ando); La Paloma, 40 m (26 Nov. 1988 T. Ando & H. Kokubun U68 MVFA, Ai}do); Route 15, 8.6 km N. of Rocha to Velazquez, 50 m (27 Nov. 1988 Z Ando & Hl Kokub"n U69 MVFA, Ando); Route 15, 32.8 krn N. of Rocha to Velazquez, 60 m (27 Nov, 1988 T. Ando & H. Kbkubun U71 MVFA, Ando); Route 13. 7.6 km N.W. of Velazquez to Aiguti, S33059'04" W54020'20", 4e rn (27 Nov. 1988 T. Ando & II. Kokubun U72 Ando); Ro-te 13, 5.S km N.E. of Arroyo del Alferez to Lascano, 40 m (27 Nov. 1988 T. Ando & H. Kokubun UZ3 Ando); Route 13, 35.6 km N.W. of Castillos to Velazquez, S34ne3'05" W54e02'iK}" (S Nov. 199e T. Ando & K. Buto U156 MVFA, Ando); Route 9, O.4 km,N.E. of Arroyo de Rocha to Rocha, S34030'58" W54"20'44" (15 Nov. 1990 T, Ando & K, Buto Cll7P Ando); Route 10, 7.0 krn S.VV. of Agutts Dulces to La Patoina (15 Nov. 1990 T. Ando & K. Buto tU80 Ando); Route 9, Fortaleza y Parque Nacional dc Santa Teresa, S33058'30" W53']33'30" (15 Nov. 1990 T. Ando & K. Buto Ul83 Ando); Route 15, 9.0 km N. of Rocha to Velazquez (16 Nov. 1990 T. Ando & K, Buto U184 Ando); Route 15, 6.1 km S.W. of Lascano to Aiguti, S33043'04" W54015'Ol" (16 Nov. 1990 71 Ando & K. Buto tU85 Ando); Fortaleza y Parque Nacional dc Santa Teresa, Playa La Mosa, S33a58'32" W53C31'47", 70 m (7 Nov. 1991 T. Ando & S. fida CJ248 Ando); Route 13, 4,O km E, of Arroyo de la India Muerta to Castillos, S34003'41" W54"12'02", 45 m (7 Nov. 1991 T. Ando & S. Jida CL251 Ando); Route IS, 12.6 km N. of Velazquez to Lascttno, S330S7'10" W54019'27", 102 m (7 Nov. 1991 Tl Ando & S, tida U252 MVFA, Ando). Dep. SAN JOSE: Route 1, O.5 km W. of ATToyo Pavon to Colonia del Sacramento, 20 m (10 Dec. 1988 Z Ando & H, Kokubun U85 MVFA, Ando); Route 1, 3,O km W. of Rio Santa Lucia to Colonia del Sacramento (21 Dec. 1988 Z Ando & H. Kok"bun U87 MVFA, Alldo); Route 1, O.S km E. of Arroyo Cufre to Montevideo, 10 m (23 Dec, 1988 Z Ando & H. Kokubuti U89 Ando); Routc 23, 5.9 km S.E, of Guaycuni to San Jos6 de Mayo, S34e06'34" W56059'45" (13 Nov. 1990 TL Ando & K. Buto U174 MVFA, Ando). Dep, SORIANO: Route 14, 11.3 km W, of Arroyo Grande to Cerro Vera (13 Nov. 1990 T. Ando & K. Buto [ll69 Ando); 17.8 km S. of La Tabla to Palmitas (t3 Nov. 1990 T. Ando & K. Buto U170 MVFA, Ando); Route 2, 42.0 km S.E. of Palmitas to Cardona (13 Nov. 1990 Z Ande & K. B"to Vi71 MVFA, Ando). Dep. TREINTA Y TRES: Route 17, 11.6 km W, of (iral. Enrique Martinez to Treinta y Tres (16 Nov. 1990 T, Ando & K. Buto Ul87 Ando); Route 8, 31.1 km N. ef Treinta y Tres to Melo (16 Nov. 1990 T, Ando & K. Buto U188 Ando).

7)Lpicatfornzs of Petuniu axillaris s"bsp. parodii The following specimens weTe determined to be Petunia axillaris subsp. parodii (Stccre) Cabrera according to all discriminant functions obtained in this study. Dep, ARTIGAS: Estancia Timbauba, Arroyo Tres Cruccs, cerca de la barra (Feb. 1955 Nbemi Garcia Zorron 840 MVFA); Puntas Cuaro, Paso Campamento (Feb. 1955 IVbemi Garcia Zorron 1(125 MVFA); Tomas Gomensoro (16 Jan. 1971 Marchesi 1(]224 MVFA); 16.3 krn S.E. of Bella Union to Tomas Gomensore (22 Nov. 1988 T. Ando & H. Kokubun U27 MVFA, Ando); Route 30, 16,1 km E, of Tomas GomensoTo to Anigas (22 Nov. 1988 71 Ando & hr. Kokubun LC29 Ando); Route 30, 21.1 km E. of Tomas GoTnensoro to Artigas (22 Nov, 1988 T. Ando & H. Kokubun U31 Ando); Route 30, Arroyo Cuaro Grande (22 Nov. 1988 Z Ande & H. Kokubun tL32 MVFA, Ando); Route 30, 5.4 krn E. of Javier de Viana to Artigas (22 Nov, 1988 T. Ando & H. Kokubun CL34 Ando); Routc 30, 5,3 krn S. of Artigas to Tranqucras, S30025'40" W56e27'29" (22 Nov. 1988 T. Ando & H. Kokubun as5 Ando); Route 30, 11.0 km S. of Cda. de Brum to Tranqueras, 60 m (22 Nov. 1988 T. Ando & H, Kbk"bun U37 Ando); Route 3U, 25.0 km N.W. of Sarandi dcl Arapey to Artigas, S30048'15" W56015'57", 246 m (2 Nov. 1991 T Ando & S. Iida U217 MVFA, Ando); Route 30, 37.0 krn N.W. of Sarandi del Arapey to Artigas, S30042'30" W56'19'18", 176 m (2 Nov. 1991 T. Ando & S. Iida U218 MVFA, Ando); Route 4, Arroyo de los Chullchos, S30034'50" W56037'12", 155 m (3 Nov. 1991 T Ando & S. Iida V220 MVFA, Ando); Route 4, Arroyo Cuaro Chico, S30"42'40" W56042'41", 178 m (3 Nov. 1991 T. Ando & S. lida U221 Ando); Route 4, Arroyo Cuaro Grande, S30"46'52" W56046'37", 135 m (3 Nov. 1991 T. ilnde & S. Iida tL223 MVFA, Ando). Dep. COLONIA: Punta Gorda, S33"54'57" W58e24'49" (20 Nov. 1990 Z Ando & K. Buto U200

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107 December 1994 ANDO et al. : Distribution of Petunia axiUarts

MVFA, Ando), 10579 Dep. DURAZNO: Arroyo Chi!eno Grandc (21 Mar. 1971 Ziliani, Biagioni & Calero MVFA); Route 4, 14.5 km S.E. of Baygorria to Carlos Reyles (11 Nov. 1990 Tl Ando & K. Buto U159 MVFA, AiidO)･ Ando). Dep. FLORESt Gruta del Palacio (13 Nov. 1990 T. Ando & K. Buto tit67 193K Nov. 1989 Dep. PAYSANDU: Route 26, Between Tambores and Rio Queguay Grande, (30 T. Ando & H. Wdtanabe U124 MVFA, Ando)l Route 26, 9.5 km W. of Arroyo de los Corrales to 26, O.2 km W. of Paysanda (30 Nov. 1989 T. Ando & H. Watanabe U126 MVFA, Ando); Route 3, 2.0 Arroyo Buricayupi to Paysandti (18 Nov. 1990 T. Ando & K, Buto U198 MVFA, Ando); Route km S. of Rio Dayman to PaysandU, Arroyo los Chanchos, S31M28'34" W57054'07", 32 m (4 Nov. 1991 T, Ando & S. Iida U237 Ando); Route 3. Termas de GuaviyU, Arroyo Guaviyti, S31'SO'34" W57e53'02", 39 m (4 Nov. 1991 71 Ando & S. Iida U238 MVFA, Ando). Patos, Estan- Dep. RIO NEGRO: ori11as del Ilio Uruguay, Rincon dc las Galtinas, Arroye de los 4, 10.1 km N.W. of to cia de Goenaga (22 Oct. 1942 Rosengurtt B-4079 MVFA); Route Baygonia 1990 T. Ando & Merccdes (11 Nov. 1990 T. Ando & K. Bute U160 Ando)l Nuevo Berlin (12 Nov, K, Buto U162 Ando); S. of Nueve Berlin, S32059'IS" WS8"03'06" (12 Nov. 1990 71 Ando & K. Buto U163 Ando); Las Cafias, S33alO'13" WS8"21'18" (14 Dec. 1992 11 Ando & K. Shibata U262 MVFA, Ando). Dcp. SALTO: 4 km de bifurcacion Esta. Ana (C. Termas) hacia B. Brum (19 Oct. 1961 Miltot 502 MVFA); entre Termas del Arapey y Ruta 3 (13 Oct. 1972 Del Puerto & Marchesi 11474 MVFA); Routc 3, Rio Arapey Grande, S30D58'40" W57041'40" (21 Nov. 1988 Tl Ando & H. Kokubun U26 MVFA, Ando); Route 4, Rio Arapey Chico, S31002'17" W56"53'52" (3 Nov. 1991 T. Ando & S. Iida U224 Ando); Between Lluveras and Termas de Arapey, S30058'43" W57026'30", 154 m (3 Nov. 1991 T. Ando & S. Iida U225 MVFA, Ando); Between Lluveras and Termas de Arapey, Rio Arapey Chico, S30"57'06" W57"27'20", 154 m (3 Nov. 1991 T. Ando & S. fida U!l26 Ando); Betweep Santa Ana and Termas de Arapey, S30054'39" W57031'07", 154 m (3 Nov, 1991 T. Ando & S. Iida CL228 Ando); Ter- mas del Arapey, Rio Arapey Grandc, S30"56'15" W57032'Ol", 9g m (3 Nov. 1991 T. Ando & S. fida U230 moA, Ando); Route 3, Between Zanja Honda and Salto, S31015'37" W57046'24", 116 m (3 Nov. 1991 Z Ando & S. Iida U232 MVFA, Ando); Route 31, 6.1 km W. of Rincon Valentin to Salto, S31018'41" W57"15'04" (17 Dec. 1992 T. Ando & K. Shibata U271 MVITA, Ando); Route 31, 11.7 km E. of Rincon Valentin to Tacuaremb6, S31"lg'45' W57U05'35" (17 Dec. 1992 T, Ando & K. Shibata U272 MVFA, Ando). Dep. SORIANO: Vcra (22 Dee. 1902 M. B. Berro 2951 MVFA); Routc 21, 29.6 km N.E. of Do- 4.4 km W. lores to Mercedes (27 Nov. 1989 T. Ando & H. Wlttanabe UI04 MVFA, Ando); Route 14, of CeTro Vera to Mercedes (12 Dec. 1990 T. Ando & K. Buto Ul66 MVFA, Ando); Playa Agraciada, S33"48'50" WS8"25'39" (20 Nov. 1990 Z Ando & K. B"to V201 MVFA, A]do). Dep. TACUAREMBC): Route 5, 1.2 km N.E, of Curtina to Tacuaremb6, 290 m (18 Nov. 1988 T. Ando & H. Kokubun UI9 MVFA, Ando); Route 5, 12,2 krn N. of Rio Negro to Tacuaremb6, 280 m Eden to C18 Nov, 1988 T. Ando & H. Kokubun U20 MVFA, Ando); Route 26, 4.5 km W. of VaLle km W. of Rio Paysandti (30 Nov. 1989 T, Ando & H. Walanabe U121 moA, Ando); Route 26, 26.1 Queguay Grande to Paysandti (18 Nov. 1990 T. Ando & K. Buto LU95 Ando).

Doubij'ut forms of Petunia axillaris s"bsp. axilLaris and subsp. parodii The fo11owing specimens were doubtfully determined, or were not regularly determined to be subsp. axillaris or parodii according to one of the thrcc discriminant functions obtained in this study. Dep, ARTIGAS: Route 30, 24.0 km S, of Cdtt. de Brum to Tranqueras, 60 m (22 Nov. 1988 Z Ando & H. Kokubun tL38 Ando). Dep. CERRO LARGO: Sierra de Rios (5 km antes de pico mas alto) (23 Nov. 1968 Lema, Ola- no, Rodriguez & Krall 7964 MVFA); Routc 8, 2,5 km N. of Rio Tacuari to Melo, 2oo m (25 Nov. 198S 71 Ando & H, Kokubun U55 MVFA, Ando). Dep. COLONIA: Ruta 21 y Arroyo Viboras (27 Oct. 1969 Del Puerto & Marchesi 90J6 MVFA); Route 1, O.9 km W. of La Paz to Colonia del Sacramento, 10 m (23 Dec. 19S8 T. Ando & H. Koku- bun U90 MVFA, Ando); Route 1, O.2 krn W. of Arroyo Sauce to Colonia del Sacramcnto, 5 m (23 Dec. 198S T. Ando & H. Kokubun U91 Ando); Route 1, 20,4 km W. of Arroyo Sauce to Colonia del

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108 Acta Phytotax ． Geobot ． Vol．45

Sacramento 26 Nov．工989 T ．Ando ＆ H ． （ Watanabe σ 98 Alldo）； Co 且onia dcl Sacramento（26 Nov ． 1989T ． Ando ＆ H ． iVatanabe U99 MVFA ，A 皿do）； Route 21，2 ．1 km W ． of A 酊 oyo Miguelete to Car− melo （27 NGv ．1989 T． Ando ＆ H ． Watanabe ひ 100 Ando ）i Carmelo （27 Nov．1989 T ．．4ndo ＆ H ． Watanahe UIOI MVFA ， Ando ）； Carme］o （27 Nov ．1989 累 緬 ＆ 〃 ． Watanaba UIO2 MVFA ° 厂” 厂” ， Ando ； Route 23 35 km ．of te Juan Jackson ） ， S lsmaelConinas ， S53 5934 W57 ℃6 17 （28 Nov ．1989 τ Ando ＆ H ， W 廨 欄 わε U！lO8 MVFA Ando ， ）；85 km S．of Conchillas（20 Nov ．1990，T．Ando 【fiκ， β uto UJ 99 Ando ）．

Dep ．DURAZNO ： Route 5 0．5 ， km N ． of Du 丁azno to Tacuaremb6 ， Rio Yi，190 m （17 Nov ．1988 T ・Ando ＆ H ・Kokubun Us MVFA ， Ando ）． Dep ． FLORIDA ： Cerro Colorado San Pedro del Timotc ， ，SierradelOlivo（20ct ．1942 GaUiual，

Aragone Berga〃’ Campal （fi Rosengunt PE −5036 Route 7 15 8 km N E of Reboledo ， ， MVFA ）； ， ． ．． to

じ rro orado C Co 且 （29 Nov ．1989 T ． Ando ＆ H ． Watanabe ひ 1ヱO MVFA Ando Route 7 A π oyo dcl ° ’” ／ ” ， ）； ， Potrero 〔S33 46 42 W55D28 36 29 Nov ．1989 T．．4ndo ＆ σ MVFA Ando ）（ H ，Watanabe 1Z3 ， ）． Dep ， LAVALLEJA ： 16．8 km N ．E ． of Minas to Arroyo del Perdido，140 m （2 Dcc ．1989 T ．．4ndo ＆ 〃 ．Watanabe σ143 And （，）， ． Dep ． MALDONADO ： Route 109 30 ． km m ， 8 S．E ，of Aigu旨 to Rocha，340 （26 Nov ．1988 T． Ando （隻H ． Kokub “ n U65 Ando ）． Dcp ． PAYSANDU ： Route 26，05 km W ． of Arroyo de los CoTrales to Paysand 頓 18 Nov ．1990 τ：Ando 6とK ， B “ to U197 Ando ）． ° ’” ° ’” Dep ． ROCHA ： Route 13，27，9 km N ．W ．of Cast且los to Velazquez， S34 3 26 W53 58 04 （8 Nov ， ° ’ ” 1990 踊 η 40 ＆ κ ．Buto U154 MVFA Ando Route 9 0．8 E ． of Castillos to Chuy S34 11 41 ” ’” ， ）； ， ， W53 49 50 15 Nov ．1990 T ．Ando ° ” ° ’” （ （k K ． B “to σ 181 Ando ）； Laguna Negra ， S34 0げ 48 W53 3556 （15 Nov ．工990 T ． Ando ＆ K ． Buto 〔〃 82 Ando ）；4．9 km N ． of Cebollati tG Vergara 16 Nov ．1990 T． Ando 〔 ° ’” じ ’ ＆ K ．B 吻 U186 MVFA ， Ando ；Fortaleza Parq 皿 e Nadonal de Santa Teresa S335903 W53 33 3ぴ ） y ， ， 70m 7 Nov （ ．1991 T ． Ando ＆ S．lida U247 MVFA ， Ando ）． Dep ． SAN JOSE ： Arazatf （240ct ．1954五． Coレ8〃o de Zole∬ i 318 MVFA ）；Rincon del Pino （17

Nov ．1956 N ．Gar （：ia Zorron 1562 MVFA − ）； Kiyfi（21 Nov ．1961 Det Puerto ＆ Millot676 MVFA ）；Ara zati 27 Dec ．1961 Det PuertO （ 1002 MVFA ）； Kiy亘 （30 Scpt．1962 Del Puerto 2185 MVFA ）；Arenas cerca BarTa Santa Lucia （3 Nov ．1971 ノ、 Ren 10883 MVFA ）； Arazati （30 Jan．1976 Det Puerto ＆ ° ’ ” Marchesi 12691 MVFA ； Route 11，〔〕．6 km W ， of Rio Santa Lucfa to San Jos6 de Mayo S34 26 52 ° ’” ） ， W56 24 17 10 m 20 Nov ．1988 T． Ando ＆ H ． Kokubun U25 MVFA 2 O km S W of Kiya ， （ ，Ando ）； ． ． ． ， 5m （21 Dec．1988 T． Ando ＆ H ． Kokubun U88 MVFA ， Ando ）； Route 1， ．9 km W ． of Rio Santa Lucia to Colo血 a del Sac丁amento 26 Nov ＆ Watanabe こJ96 MVFA Alldo Route （ ．1989T．Ando H ． ， ）； 1 Between Eeilda Pau 皿ierand Alroyo ， Cufr6 （26 Nov．1989 T ．／lndo ＆ H ． Watanabe U97 Ando）． ＝ Abbreviatlons： MVFA Laborat6rio de Bot6nica， Facultad de Agronomia ， Casilla 1238， Montevideo ≡ ， Uruguay ；Ando private cellections of Toshlo Ando ，

摘 要

1 1 1 1 ・ 一 ・ ・ ・ 安藤敏 夫 飯 田新 国分 尚 上 田 善弘 Eduardo Marchesi2 ：標 本 の 判別 関数 に よ る

Petunin axillaris （ナ ス 科）亜種 の ウ ル グア イ 内分 布

ペ チ ュ ニ ア の は 1834 の P ．axiliaris と P ． integrifoliaの に る 品種 改 良 ， 年 交 雑 始 ま 。 少 な く と も当初 は ど ち ら も ウ ル グ ア イ 産 の も の が 使 わ れ た こ とか ら，品種 の 遺伝 的背 景 を解析 す る に は ウ ル グ ア イ の の を る こ と は で ， 産 両 種 実 態 解析 す 有意 義 あ ろ う。 本 報 で は ，まず ウ ル グ

ア イ 内 に P exillaris の 亜 の を い だ の べ ． 2 種 存在 見 し， そ 分布状 況 を調 た。5 季 に渡 る実 地 踏

査 の 結 果 ウ ル グ ア イ南 の び の は ， 東部 Caneloncs， Maldonado 及 Montevideo 3 郡 に ，花冠 が大 き く，花筒 が 短 く，2 強 雄ず い を もつ 亜 種 axiltaris が ，北西 部 の Artigas， Salto両 郡 に

は が さ が い つ 々 い ，花 冠 小 く， 花 筒 長 く， 等長 雄ず を も 亜 idiparodiiが 各 分布 し て る こ とが

分 か っ た 。 こ れ ら の か ら 計 5 郡 産 60 標 本 ，花筒 長 ， 花 冠 縁 部 長，花 茎長 ，が く裂 片 長 及 び

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December 1994 ANDO et al．： Distribution of Petunia axillaris 109

花筒 長 〆花 冠縁部 長 の 5 形質 を計測 し， ス テ ッ プ ワ イズ 判 別分析 に よ り両亜 種 の 判別 関数を ’ ル ら の 174 地 の 本 を判 定 得た 。 こ れ に他郡 産標 本 の 値 を代 入 して ，ウ グ ア イ全 域 か 言1 点分 標 した 、，ウ ル グ ア イ中央部 を流 れ る ネ グ ロ 川 を境 と して ，南東 部 に 亜 種 axillaris が ，北 西部 に 亜 種 parodii が分 布 して い た が ，境 界付近 の 南 西 部 に は 判 定 を保 留す べ きもの が 認 め ら 札 1 そ こ で 形態 が 遷 移 し て い る可 能性 が 示 唆 され た 。（〒 271 千 葉県松 戸市 松戸 648 千 葉 大 学 2Facultad 780 Montevideo 園芸学 部 ； de Agronomia， Univerg．idad de laRcpublica ， Garzon ， ， Uruguay ）

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