Classification of Pinus patula, I? tecunum 

R oocarpa, R caribaea var. hondurensis, 

and Related nomic Entities 

A.E. Squillace and Jesse I? Perry, Jr. 

Abstract 

Stem xylem terpenes of 75 pine populations were studied to 

deterrnirre relationships among taxonomic entities. Typical 

Pir~xs patalc populations octurrirlg in areas north and west of 

Oaxaca, Mexico, had very high proportions of Q-phellandrene 

and low proportions of other constituents. Terpene 

compositions of populations of variety longipedz~ncltlata in 

northern Oaxaca were similar to that of the typical variety, 

while those of populations in southern Oaxaca resembled that 

of P. tecanunaanii. Typical P. lecunumanii from populations 

in Chiapas (Mexico), Guatemala, and southwestern Honduras 

contained high proportions of a-pinene, carene, limonene, and 

il-phellaadrene. Popda?inns in southern Mexico, Guatemala, 

Honduras, Nicaragua, and Belize that contaixied very high 

proportions of a-pinene and low proportions of other 

constituents were judged to be typical P. oocarpa. Other 

populations in Guatemala, Honduras, and Xicaragua tended 

to resemble both P. oocarpa and P. rlec?~nvmanii and were 

judged to be atypical P. oocarpa. Our results suggest that the 

two species hybridize at middle elevations, where they occur 

together. 0 ther researchers regard the atypical P. oocarpa 

populations as P. oocarpa, P. patala ssp. tecunumanii, or 

P. oocarpa var. ochoterenae. Most atypical P. oocarpa were 

more sirnilar to P. oocarpa than to P. fecanvmanii. They 

were definitely more sinlilar to P. oocarpa than to P. patgia 

and hence should not be referred to the latter taxonomically. 

P. caribaea var. hondvrensis trees differed from others mainly 

in that they contained high proportions of a-pinene and 

P-phellandrene and low proportions of other constituents. 

Hybridization with P. oocarpa occurs where the two species 

occur together at low elevations, 

Keywords: Monoterpenes, taxonomy. 

Introduction 

Ailthougl-1 diAFerences between Pinus patvla Schiede 

and Deppe and P. ooearpa Sclriede are usually clear, 

there is considerable controversy about the taxonomy 

and idelltities of the related entities P. pat~la var. 

io?tgzped.rkncalata Loock.. P. tecunamanzz j§ch\vd.) 

Eguiluz-Piedra and Perry (1983) [syn. P, patltka ssp. 

lecununzanii (Eguiluz and Perry) Styles], and P. 

oncarpa var. ochoterenae Martihez. The history of 

the problems involved has been discussed well by 

others including Styles ( 19'76 and 1985), Styles and 

Hughes (19831, Locklzart (1985, 1990b), Dvorak and 

Raymond (19911, and Perry (1991). In particular, 

tshere is disagreement about (I) the taxonomic status 

of P, oocarpa var. ochoferenae and P. patula var. 

longipeduncudata, (2) the extent of the range of P. 

tecunvmanzz (Styles and McCarter 1988), and (3) the 

extent of variation within entities. 

Here we analyze data on te-rpenes obtained from 

populations of the species in an attempt to shed light 

on various problems of identification and taxonomy. 

Data previously reported by others and our own 

previorisly unreported data are utilized. VCTe also 

report results of a study of natural hybridization 

between P. oocarpa and P. caribaea var. hondurensis 

(hereafter the varietal epithet will be omitted for the 

sake of brevity). These results explairr some of the 

variation among P. oocnrpa populatiorrs. 'ctTe also 

briefly summarize morphological data reported by 

others for the taxa discussed here. 

Materials and Methods 

Terpene compositions of 2,196 trees in 75 populatiolis 

(apps. I and 2) were studied. Thirty of the 

populations were sampled by the authors. The 

remainder: and sonre of those we sampled, wre 

sampled by others. Data from populations sampled 

by more than one author u3ere combined when the 

results were similar. CIiherl results were not similar, 

names were changed slightly and data kept separately. 

In all cases, oleoresin was obtained from stem xylem 

tissue about 1.5 m above ground level, Most of our 

samples were collected over several years prior to 

1988 and were analyzed by techniques described by 

Perry (1987). In these analyses, the sesquiterpene 

longifolene was not identified. I11 1988, six additional 

populations (rvios. 117, 18, 19, 27, 39, and 42) 

were sampled as part of a study of hybridization 
between P. oocarpa and P. caribaea. Proportions of 

longifolene were determined for these populations. 

In the 1988 sampling, composition of the turpentine 

was obtained by gas chromatography of a sample of 

the whole oleoresin dissolved in methyl tert-butyl 

ether (20 mg in 1 mL) using a 15-m (0.25-mn~ od) 

DX-1 fused silica column (J&WT Scientificll Folsom, 

CA), with a temperature program of 50 'C (15 rnin) 

followed by 2 OC/min at 110 OC to remove free resin 

acids from the column. 

In statistical analyses, we considered only the 

monoterpenes that frequently occurred in large 

proportions (a- pinene, P- pinene, carene, limonene, 

and P-phellandrene) and the sesquiterpene longifolene. 

Some authors reported large proportions of terpenes 

other than those listed above. Inclusion or exclusion 

of particular terpenes can change the relative 

proportion of each terpene appreciably. In order to 

minimize such effects, we renormalized data for all 

reports in which longifolene was assessed so that the 

sum of all constituents (the five monoterpenes plus 

longifolene) added to 100 percent. For those samples 

in which longifolene was not assessed, we renormalized 

the proportions to sum to 90 percent to provide an 

approximate allowance for the omission of longifolene 

(the proportion of this constituent averaged roughly 

10 percent). 

A preliminary examination of individual tree data for 

some populations showed that some individuals had 

relatively high proportions of both limonene and 

P-phellandrene, while others did not. We deternlined 

the percentage of trees having at least 10 percent 

limonene and at least 6 percent P-phellandrene in 

each population for which individual tree data were 

available. Such trees were characterized as "high-LP 

trees." 

Relatio~lships among populations were deterri~i~led 

mainly by means of principal component analyses 

(Isebrands and Crow 1975) of population mean 

proportions of terpenes. The first of the 3 analyses 

involved only the 5 rnonoterpenes and all 75 

populations (P.C. Analysis 1). The second involved 

populations 1-48, in which both longifolene and the 

The use of trade or firm names in this publication is for 

reader illformation and does not imply endorsement by the 

U.S. Department of Agriculture of any product or service. 

five monoterpenes were assessed (P.C. Analysis 2). 

The third analysis involved the five rnonoterpenes of 

populations 49-75 (P.C. Analysis 3). 

We tentatively classified all populations into five 

major groups, mainly on the basis of major differences 

in terpene composition, but also partly on geographic 

location: 

Groups A-1 and A-2. Populations having very 

high P-phellandrene, in areas north of Oaxaca and 

northern Oaxaca, Mexico, respectively. 

Groups B-1, B-2, B-3, and B-4. Populations having 

high a-pinene, high carene or high limonene or 

both, and high P-phellandrene, in southern Oaxaca 

(Mexico), Chiapas (Mexico), Guatemala, and 

Honduras, respectively. 

Group C. Populations having both high a-pinene and 

high /?-phella.aadrene. 

Group D. Populations having very high a-pinene with 

low proportions of other terpenes. 

Groups E-1 and E-2. Populations having very high 

a-pinene and moderate proportions of carene or 

limonene or both, and populations having lower 

proportions of a-pinene and higher proportions of 

carene or limonene or both, respectively. 

Analyses of variance were conducted on the 

population means of terpenes. These compared all 

possible pairs of groups, mainly to determine the 

extent of interaction between groups and terpenes. As 

an example, the analysis comparing groups A-1 and 

A-2 (containing 4 and 3 populations, respectively) was 

as follows: 

Source of Degrees of 

variation freedom 

Groups 1 

Populations in groups 5 

Terpenes 4 

Groups x terpenes - 4 

Tot a1 14 

We also summarized data on morphological traits of 

needles and cones reported by others. These were 

used mainly to aid in judging results of terpene 

analyses. 
Results 3, the first three componerlts accounted for similarly 

large proportions of variation. 

Principal CompolneriC Analyses 

In PC Analysis 1, the first three prit~cipal components 

accounted for about 91 percent of the variatron 

in mean relative contents of terpenes in the 75 

populations. The first component was heavily 

weighted for a-pinene, 8-pinerre, 3-pfieilandrene, 

and liri~o~~ene in that order. The second and third 

compo~lents were most heavily weighted for carem 

and for lirnonene, respectively. In PC A~ialyses 2 and 

Ordiz-lation of the populations along pairs of principal 

cornponent,~ (eigenvectors) is shown in fighres 1-4. 

Figures 1 and 2 strongiy suggest that two rnitjor 

clusters are present. Group A and 73 populatiorls are 

on the left side of both figures, while poplllations of 

other groups tend to be on the right. Group A and B 

populations usually had relatively lower proportions of 

a-pinene and higher proportions of P-ghellandrene 

than had those on the right. 

Analysis 1 (1 -75) Groups 

A-1 and A-2 El 

B-1 and B-2 - 

B-3 and B4 II 

c A 

0 

E-1 @ 

E-2 @ 

Figure 1-Ordination of all 75 populations by principal 

components 1 and 2 of P.C. Analysis 1. 

Groups A-1 and A-2 fl-phellandrene (were LP trees). The four populatio~ls 

Terpenc compositions of the seven populations in located northwest of Oaxaca, Mexico (group A- 1). 

these groups were very similar. Each p~pulat~ion had were considered P. paldn by authors reporting 

an average of at least 75 percent B-phellandrene and terpene composition, and the three in nortliern 

olrly small amounts of other constituents (table 1). Oaxaca (group A-2) were considered P. paillla var. 

Very- few individuals had both liigll limoilene a~ld high longzpedracvlata (see app. 1). 
Analysis I (1 -75) 

B-1 and 8-2 - 

8-3 and B-4 Ill 

P.C. analyses (figs. 1, 2, and 4) showed that 

groups A-1 and A-2 are closely clustered and 

indistil~guishable on the basis of terpenes alone. 

TuIorphological data were available for only a few of 

the seven populatioris (table 2). Group A-l trees bad 

shorter peduncles and sxnalfer ratios of cone length to 

width than had group 12-2 trees. The Santa Maria 

Papalo population, which was considered P. pcatuta hy 

Ilvorak and Raymorld (1991), differed from P. pat~~ia 

populations only in Itlaving greater ratios of cone 

length to width, Gone shape in the Ixtlan popuiatisll 

was similar to cone shape for P. yalula, but trees 

of the Istlan populatior~ had longer peduncles than 

had P. pailzrla trees and were more typical of the 

i'ongzpedun czlata variety. 

Groups 

A-1 and A-2 El 

Figure 2-Ordination of dl 75 populations by principal 

components f and 3 of P.C. Analysis 1. 

Groups B-1, B-2, B-3, a11d B-4 

Trees in these groups had less P-pheiiandrene, more 

a-pinene, and more carene or limonene or both, than 

had trees in the A groups, and there were more LP 

trees ill groups B-1, B-2, B-3, and B-3 than in the A 

groups (table 1). Trees in groups B-1, B-2, B-3, and 

B-4 also tended to have shorter needles, Illore tleedIes 

per fascicle, and longer peduncles than had A-group 

trees (table 2). 

Terper-rcs and morpl~otogical traits of two of the B-l 

poputations (64 and 67) tended to resen~ble those of 

populations in group A-2. Terpenes of populations 

58 and 70 tended to be more like those irt offler B 

groups (table 1 and fig. 41, but populations 58 and 

70 had many more internal resin ducts than had 

populations in other B groups. 

Most of the remaining group B populations had high 

limonene and 3-phellandrene and rather sir~~ilar 

morphological traits. were considered by the 

authors to be B. -tecuntlt-tzanzz, especially ifmose 

irr Guatentala and TtTonduras. There were severat 

appreciable diRerences within and between these 

groups: 

(1) The Chiapas populations (k3-2) tended to 

have higher carene than others. and one of them 

(population 61) had no LP trees even though 

proportions of firnonene and P-phellandrene were 

substantial. 

(2) Like the B-2 populations, the Guatemala 

populations (B-3) had relatively high limonene and 

3-phellandrene, but their carene content was variable. 
c A 

Atypical P. oocarpa 0 0 

caribaea in Honduras and Nicaragua 

PC 1 Decreasing a-pinene and 8-phellandrene 

Figure 3-Ordination of populations 1 through 48 

by principal components 1 and 3 of P.C. Analysis 2. 

(3) Most of the Honduras populations (B-4) had less 

P-phellandrene, but they had appreciable percentages 

of LP trees. Content of carene was variable, and 

peduncles were longer than in other groups. 

(4) P.C. analyses (fig. 4) showed fairly distillet 

clustering of the B-2, 13-3, and 13-4 populations. 

Populations 62, 63, 68, and 75 tended to be outliers, 

but their rnorphologieal traits do not seen1 to be out 

of line with those of typical P. tec?tnumanzi. 

(5) Cone Iength/width ratios decreased and peduncle 

lengths increased, going from B-1 to B-4 populations 

(table 2). 

The authors judged that all of these populatio~ls 

(table I) were P. carzbaea (app. 1). The group as a 

whole differed from others in having high a-pinene, 

high @-phellandrene, and snlall amounts of ot'her 

consti tuerits. Mean proportion of longifolene was 

generally lower for group C populations than for 

others. Terpene compositior~ varied little among 

locations, although the three Belize populations had 

lower than average Iongifolene. 

P-phellandrene content of populations 38, 39, and 41 

was somewhat lower than average, possibly because 

some P. oocarpa x P. caribaea hybrids were present. 

On average, trees in populations 38 and 39 had 

fewer internal ducts than others, and this suggests 

Iiybridization (such data were not available for 

populatio~l 41). Note that populations 38, 39, and 41 

occur at relatively high elevations (app. 1) and are in 

a son~ewhat intermediate position between P. oocarpa 

and P. caribaea in figure 3. Presence of hybrids in 

population 38 (Santa Clara) was also suggested by 

Salazar (1983). 

Group D 

Most of these populations were considered P. oocarpa 

(app. I) and had similar terpene compositions. The 

group D populations had much higher proportions of 

a-pinene and lower proportions of P-phellandrene 

than had the A and I3 populations. Proportions of 

P-pinene averaged 0.9, 1.5, 4.4, 6.2, and 5.9 percent in 

the A, B, C, D? and E populations. 

Rilorphological data were scanty, but trees in most 

group D populations had more needles per fascicle, 

more resin canals (more of which were in the septa1 

position), and much lower cone lengthlwidt h ratios 

than had trees in the group A and B populations 

(table 2). 
Figure 4-Ordination of populations 49 through 75 by 

principal components 1 and 2 of P.C. Analysis 3. 

The two Belize populations differed from others in 

having lower than average proportions of longifolene, 

which is curious because Belize populations in group 

C also had lower proportions of longifolene than had 

others in group C. h~lorphological data were available 

only for population 19, and trees in that population. 

unlike most P. oocarpa, had no septal resin canals. 

Populations 17 and 18 had higher than average 

proportions of P-phellandrene. They were sainpled 

because appearance suggested that P. oocarpa x 

P. cnrzbaea hybrids were present. Roughly equal 

~lrimbers of suspected hybrids and typical P. oocarpa 

trees were selected for sampling. High content of 

P-phellandrene is believed to be indicative of natural 

hybridization, which will be discussed later. 

Group E 

These populatioxls (table I), like populations of P. 

oocarpa. are characterized by high prctport,io~ls of 

a-pinene and low p-phellandrene, but they differ 

fro111 populations of P. oocnrpa in having appreciable 

proportions of carene or lirilonerie or botli. Many of 

t-hese populatiorls were co~lsiderecf to be P. oocarpa 

or its variety ochoferenae, but several were co~lsidered 

P. tecztr1uma~2ii or P. patula spp. fecununzanzz 

moderate proportions of carene or limonene or 

both (group El) from those having relatively high 

proportions of those terpenes (group E-2). The forrlier 

also had higher proportions of a-pinene than had the 

latter, as would be expected because of constraint. 

Principal component analyses showed that group E- 1 

populations were relatively similar to each other and 

were close to the typical P. oocarpa populations, and 

that tlie group E-2 populations were more variable 

and more distinct from typical P. oocarpn (figs. 1, 

2, and 4). Also, both of the E groups were closer to 

P. oocarpa than to P. patula populations (figs. 1 

and 2). Population 31 (group E-1) is an outlier and 

is within the cluster of P. oocarpa populations (fig. 

3). Population 21 (group IF;-2) is within the cluster of 

P. -tecunumanii populations (figs. 1 and 2). 

Morphological differences between group E-1 and 

group E-2 populations were not appreciable. But 

these populations had fewer resin canals and fewer 

canals in the internal and septal positions than had 

group L) populations. Population 28 (La Laguniila) 

has very high carene but resembles typical P. oocarpa 

in most of its morphological traits. McCarter arid 

Birks (1985) considered it typical P. oocarpa. 
Azialy ses of Variazlcc 

Arialyscs of variarlcc often sho~ved. highly significarit 

diffkrences arilong relative terpene corltertts of the 

various groups, but groups had been forrlled partly 

on the hasis of terpcae compositiori. Ifowever, 

interact ions bcttveetl lerperies and groups also often 

difiretl appreciafjly and were indicative of tllc degree 

of sirlrilarity among grotips (table 3). 

l'tte data suggest relative sit~iilarity among grotips as 

follows, the distilrice between poilits reprcsenti~lg 

approximate averages of pertinent interactions: 

Tiltus, groups A-1 artci 12-2 are bery similar arid are 

rrlore silnilar to B-I tltail to other groups. Groilps B-2, 

B-3, and B-4 are also similar and are closer to 23-1 

tllart to others, artd so on. We think it significant that 

q~testiorlablc group E-2 falls bet ween group D (nlostly 

I" ooorar-]la) and the U-2, B-3, 13-4 cluster jniostlq P. 

teczlr2trntan~z). Also, groups E-l and E-2 arc. closer to D 

than to the A-1, A-2 cluster (ritostly P. pnttlfu). Tltese 

results agree well with the P.C. aitalyses. 

Correlations between elevation arrd terpcne contents 

in prttati\re P, tecunumnnzi, atypical P. ooculpa, ad 

typical P. ooearp popilllatioris are given in table 4. The 

followi~ig points are of interest. 

1. The strong negative correlation (-0.67) bett\lcen 

elevatiort and a-pinene contcrit in P. tecuntinrnnti 

populations may il~dicate that there is aplprcciaI>le 

ilitrogression of P. ooeurpa (which has very higlt 

a-pine~ie) at low ele\rations and Little or ilo irrt rogression 

at higtl elevations. The same situation may be true for 

the strong positive correlation (0.55) between cle\iation 

and percent LP trees. The increasing trentl in LP trees 

may be due to introgression of P. ooen9-pn (which tias 

very few LP trees) into 1'. tecunzdmarzia stands at the 

lower elevations. 

2. AItl~ough carene content was very low in typical P. 

oorarp, level of this tcrpene was strongly correlated 

(0.56) with etevation. This correlation probably results 

from iilt rogression of P. teczr~lumanga jwfrich has 

appreciable carene coiite~tt) into high-eievation P. oocap-pa 

stands. 

3. The strong negative correfatiorl (-0.66) between 

elevatio~l and a-pinene content in atypical P. oucrsrp 

poptllations sl~ggests that at least some trees in these 

populations are P. tecununza~zaz x P. oocarp llyhrids. 

Atypical populatiorts at relatively high elevaliorrs probably 

receive genes from P. teeunumantz (which has low 

P-pinelie), while those at low elevations receive genes 

from P. ooccarp (wlrich has relatively high /3-pinene). 

?Villiam S. Dvorak2 contends--based 0x1 nrimerolls field 

observations-that tlre two species hybridize freqr~en t ly 

where they occur together at n~iddle elesations. 

Anlong the 23 A and B populations for whidr data 

on peduncle length were available, values of t ltis trait 

increased, going from northwest to southeast. Rarrett 

(1972) sllowed a similar treitd for P. patuln and its 

variety tongi~dunculntu. When latitude axid lorlgitude 

were ilideperzderlt variables and peduncle le~rgtlr the 

dependc~tt variable, the n~ultiple correlation coeficie~it 

was 0.00 and was Iliglrly significant. The trcncl, however, 

appeared to be "stepped9%-that is, peduncle lengths were 

relatively consistelit witlriri locations other tltart northern 

Oaxaca (table 5). Data for the E and D populations 

were scarce and did not show a trend. The rest~lts 

suggest that the R po~lrrlations are related to (and rnay 

have originated from) P. patula and that tlre It: and D 

populations are not related to P. ~~(ltula. 

Evidence of Hybridization. between 

P. oocaupa and I", cavibaea 

As we have mentioucd, the presence of high 

j3-phelland rene trees in low-elevation P. oocarpn 

populations was thoriglit to indicate that some 

populations corltained hybrids. Table 1 ancl data from 

Nikles (1966) and Burley and Green (1977), indicate 

that P. caribaea trees generally contain appreciable 

proport ions of P-phelland rene, while most P, oocnrpa 

trees have low P-phellandrene. High P-phellandrene in 

cortical oleoresin is dominalit over low in some pirre 

species (Squillace 1982). This may also be true in xylem 

oleoresin. But even if it is not, the progeny of crosses 

between the two species are likely to have higher average 

content of 8-phellandrene, Since P, caribtsets occurs at 

low elet~ations (10 to 700 m in this study), hybridization 

is rnost likely to occur in P. oocnrp stands at relatively 

low elevations and within the range of P. carzbcn. 

To estilnate the extent of hybridization, we compared 

proportions of high P-phellandrene trees in populations at 

Dvorak, W'ililiarn S. 1992. Letter dated March 4 to Anthony 

E. Squillace. On file with: A.E. Squillace, School of Forest 

Resources and Conservation, Gniversity of Florida, Gainesville, 

FL 32611. 
8'75 to 1550 rn with j>ruportians of high 3-phellancirene 

trees in ilopulations at 550 to 700 ID {table 6). (b:ilj 

papuiations for rvhicl.1 incfillidnai-tree data were atailable 

and that were within the range of P. ~(ir~baea were 

consider~rt. Only 3.8 percent of the trees in thgs eight 

high-elevation ~mpulat ions corit airled high proportions 

of d-pirellandserre, wtit~reas 16,.f percent sf trees in the 

-five low-elevation poplrlatiorls had high ~tropor"ions of 

3-lahellandrene. All of the 15 illdividual P car-rlia9;a trees 

had high P-plneltandrcnc. Thtrs, appreciable hyl>ritIization 

seerns to be occr~rring in the low-elevation P, oucarpa 

populations. Evidence of hybridization betweer1 6. 

uacarpcr and P. caribaecr also has beer1 reportcbri by 

Btlrley arrd Green (1979), Styles and others (l9"iL ,and 

firnandez cle la Iteguera anthoti~c~rs (1988a. f98Rb). 

r '( I ire apparent presence of hybrids in tire five low-elevdtiiln 

poprtlatio~~s (Nos. 16, 117, 18, 19, and 20) is also rcflectcci 

in the j~rincipal component analyses in figure 3. 'The 

low-elevdtion populations tend to occur betwcern other 

P. oocnrpn and P. cap-zbnca popufations. Note alw that 

three I). carzbaea populations (Nos. 38, 39, and 41) 

tend to be in somewhat il~terrnedistte positions lwtrvct~aa 

If. ccirltcsea and P. oocarpa in figure 3. As inight kit? 

expected, these three 1" crnl-zhaea populations were at 

relatively high elevations for the species (700, i;"ii. and 

500 m, respectively). Salazar (1983) found eviclencc. of 

lrybridization in the Santa Clara populatio~l. 

Discussion and Conclusions 

Styles (1985) reported that po~>~tlatioi~s usually considereti 

to be 1" pte~la var. longzpdt~nculatn and P. fectrnrimanzt 

are extensions of P, pattila. Partly because of this 

apparent clinal trend, he declared the former two entitics 

to be P. pat~lla ssp. teeunumanii. However, he arrtl 

other authors (Birks and Barnes 1985; lllcCarter and 

Birks 1985; Styles aricI McCarter 1988) also desigl~ateci a, 

number of populations-pre'vio~~sly considered P. oocclr-pa 

or its variety ocfiotc.rcnae-as belongiirg to the mtcw 

sttbspecies (see app. I j. Most often mentioned are the 

populatio~ls showirlg rapid growth in pruvenancc: tests, 

such as Yucul, San Rafael, and Las Carnelias in Nicarngt~a 

and hft. Pine Ridge irt Belize. Our results strorlgly 

suggest that these four populations (plus other atj pica1 

P. mcarpca populations) are more related to f. oocat-pa 

than to P. pntula. 

Rap~d growt-h rnay not be a reliable criterion far judging 

taxonorriic status. The volume growth at ages 6 to 9 

years of 24 populations designated as either P. ooeargxl 

or subspecies teeunumanii in international provenance 

trials (Rirks and Barnes 1990) was found to be 

significantly correlated with mean annual rainfall of the 

provenance origin. The four populations mentioned above 

were located in areas of high rainfail. The trend was 

aiaparerr t also for P. oocas-pa populatisrrs in Guate~rlala, 

where populations in ilte rtortireast reccilrcii iriglr rairnfaii 

and exhibited rapid growth. The relatioriship agrees ttirli 

Salrrillac-c's (1966) rr-port stlggestirlg that. lialrlrai srrlccriu~a 

favors trees with inkiercrlt rapici. grorvt li rreorr: in iiis~ah of 

favorable climate than iri areas cf rrlifavlorable ~Iirriate. It 

m,zb. be that the poptilarions dcsignatcci as anc.irmbers of 

the new subspecies arc. thc r:rore rapid growers anrcrng 

P. oocnrpa or var. ochot~rttziie popnlathonrs as a rebult 

of natural selection arici nrr rrot geneticallj rel~tcci to I". 

lccunu~~anz~. 

Our rclstllts agree witll Lockhart'< reports ( 19913a, 1990b) 

ir~clicatirrg that populatitirrs csnsidrred to he merlrbers 

of the stabspecies arc Iligl~ly variable and not similar 

to P. pwtbla. Our result- also agree fairlj tv~4II with 

Dvorak and Rayrnortrt" ~rcpork (1991) on rnorpi~ologicai 

traits. That report siiggvired that many of the 

high-elc.rration papulati~~as (greater tisan ItirlO 111) lit 

Chiapa%, Guatemala, anti E1o:rcllrsras are P. !~c~rrrtrrriartzz 

ratlirr than P. ooca.iya var. otrholerr~~ac: j+the probable. 

absence of la. teczdnatr~inrtrt from Onxaca; and (3) rfre 

teradearc y for ~lmildrl k4 among popuiatiorrs witIlirl species 

that arc3 ~Iosely assciciat ecl geograp'lricnlly. 

In spite of the clinai trerad irr peduncle Lengths rtoleri 

earlier, there is some ctridcnce of disconrtirsuity between 

the Oaxaca pcrtpulatiorls and those to the south and 

east. 7 ltat is, most of flie group B poprrlations in 

Gliliapas, Guatemala, and IIonrlrlras are s~iffrcietrlly alike 

and clustered to be coalsiciered n separate species (1'. 

leeernuntnnzi) from varicly /angtr~edunct~lcrtct in Oaxaca 

and Cllinpas. Outliers occur, as mcntioncd earlier, but 

these 1x1;~~. be the result sf hybridization or otlrcr factors 

[Perry 1991). 

The relationships and taxollomy of the popridatioris we 

considered typical and atypical P. oocnvcr, arc- also 

controversial. Our resrrl ts suggest that the highly 

atypical populations are more related to P. oocarpn 

than to P. patula. They may co:ltain P. oocnrpa x P. 

lecunurnnnii hybrids and possibly also some P. ooeal-pcs 

var. oclzotewncse. The Belire populations diffijr solnewhat 

from typical P. ooctarptt both in terpene composition and 

in some morphological traits. But the5 lack high careae 

and timonene, unlike the populations we termed highly 

atypical P. oocarp or the oclaote~nae variety, and this is 

at variance with Hunt's (2962) designation of the Belize 

populations as oehote~nae. More study oft this is needed. 

A stlmmary of prominent differences between species 

and our opinions as to the taxonomic status of each 

population are given in table 7. Although the Belize 

populations of P. oocarpa are shown separately from 

those in other areas, we do not consider them separate 
taxonomic entities. The same is true for the Belize 

populations of P. carzhea. The populations we 

considered atypical P. oocarpu certainly need further 

study. A comprehensive study of morphological traits, 

especially location of resin canals, would be helpful in 

judging the taxonomic status of each population. 

Although this study was fairly successful in determining 

degrees of relationship among populations, we should keep 

in mind that a number of investigators who employed 

somewhat different techniques collected and analyzed the 

terpene samples. Further sampling of both cortical and 

stem xylem oleoresin by a single team would be desirable. 

Literature Cited 

Barret t , W.H.G. 1972. Variation de caracteres ~norfologicos 

en poblaciones naturales de Pinvs patvla Sclrledrt . et 

Cham. en Mexico. IDIA Suplemento Forestal. 7: 9-35. 

Barrett, W.H.G.; Golfari, Lamberto, 1962. Description 

de dos nuevas variedades del "Pino del caribe" (Pi~zus 

cariiraea hlorelet ) . Caribbean Forester. 23(2) : 59-71. 

Birks, J.S.; Barnes, R.D. 1985. Multivariate analysis 

of data from international provenance trials of Pinus 

oocarpa/Pinus paiula subspecies teczlnumanii. 

Commonwealth Forest Review. 64(4): 367-374. 

Birks, J.S.; Barnes, R.D. 1990. Provenance ~ariat~ion 

in Pinzls caribaea, P. oocarpa, and P. patula ssp. 

tecunumanii. Tropical Forestry Papers 21. 40 pp. 

Burley, J.; Green, C.L. 1977. Variation of gum turpentine 

between provenances of Pinus ca~ibaea hlorelet and P. 

oocarpa Schiede in Central America. In: EEC symposium 

on forest tree biochemistry, Brussels; 73-108. 

Burley, J.: Green, C.L. 1979. Relationships of terpenes 

between exotic and natural populations of Pinus caribaea 

Morelet and P. oocarpa Schiede. In: Rudin, D., ed. 

Proceedings of a conference on biochemical genetics of 

forest trees; Umea, Sweden; 118- 135. 

Coppen, J.J.W.; Robinson, J.M.; Mullin, L.J. 1988. 

Composition of xylem oleoresin from five Mexican and 

Central American Pinus species growing in Zimbabwe. 

Phytochernistry. 27: 1731-1734. 

Coyne, J.F.; Critchfield, W.B. 1974. Iderttity and terpene 

composition of Honduran pines attacked by the bark 

beetle Dendroctonzls frontalis (Scolytidae). Turrialba. 24: 

327-331. 

Dvorak, W.S.; Raymond, R.H. 1991. The taxonomic 

status of ciosely related closed cone pines in Mexico and 

Central America. New Forests. 4: 291-307. 

Eguiluz-Piedra, T. 1984. Geographic variation in needles, 

cones, and seeds of Pi~us tecunumanii in CuatemaIa. 

Silvae Genetica. 33: 72-79. 

Eguiluz-Piedra, T. 1986. Taxonomic relationships of Pinus 

tecun?tmanii from Guatemala. Cornonwealth Forest 

Review. 65: 303-313. 

Eguiluz-Piedra, T.; Perry, J.P., Jr. 1983. Pinl~s 

tecltnumanii: una especie nueva de Guatemala. Ciencia 

Forestal (Mexico). 41 (8): 3-22. 

Fernandez de la Reguera, P.A.; Burley, J.; Marriatt, 

F.H.C. 198th. Putative hybridization between P. 

caribaea Morelet and P. oocarpa Schiede: a canonical 

approach. Silvae Genetica. 37: 88-93. 
firnandez r31e la Rrgr~cbra, P.A.; Marriotl, F.H.G.: 

BEE rley, 3. 1988L. Xlultiplc-set canor~ical analysis: arr 

applicatiort to fr>rr+try gerrrtics. Biornetrics. 4% 4: 875-880 

Want, D.R. 1962. Sor~te nvteb orr the pines cif Britislr 

f lCinduras. Ernpire Forest Review. 51: 13 4-1-15, 

Isebrands, 3.6.; Crow, Thomas R. 1975. 5r1trudnctiua1 to 

usrs and interpret ation of principal corirportent analysis 

in forest biolog). Ccn. Tech. Rrp. NC-17. St, Paul, XIS: 

U.S. Department of iigricut tilre, Forest Service, Xc,rth 

G'er~tral Forest Exp~rintent Station t 3 pp. 

IloR, P.M., Jr.: Mirsv, N.T. 1953, Compn4tior1 of gtrrn 

t~irpentines of pints. IG. .& report on Pinus oornt-pa 

arlct P. pse?idostrol.us var. oarncann horn Chinpas and 

P. cnoperi from Durango. Journal of tlae ii~-n~t-nican 

Pharmaceutical Association, Science Edit ion. -1 2: 46-49. 

Iloff, P.M., Jr.; Mirov, N,T. 1953. Ciir-tt~positiorr of gum 

turperttines of pines. 22. rZ report on Prnus rrtdzs and 

P. harfwegit frorn hlexico and P. ~nsslnrns from the 

Philippines. Journal of the American Ptrarmiaceutical 

Association. Scierrce Edition. 43: 742-745. 

Lockhart, Linda A. 1985. Investigatioaa of tropical pine 

rc.iin terpenes. Oxford, UIC: Oxford University. 206 pp, 

M.S. thesis. 

Lockhart, Linda A. 1990a. The intensive study of tropical 

pine gene resources. In: Hat temer, EI.H.; Finesdli, S*, 

eds. Biochemical markers in the population genetics of 

forest trees. The Hague, Netherlands: SPB Academic 

Publishing: 113-1 19. 

Lockhart, Linda A. 1990b. Chemotaxonomic relationships 

within the Central American closed-cone pines. Silvae 

Genetica. 39: 173-184. 

McCarter, P.S.; Birks, J.S. 1985. Pinvs patvia subspecies 

t ecu numanii : the application of numerical techniques to 

some problerns of its taxonomy. Commonwealth Forest 

Review. 64(2): 117-132. 

Mirov, N.T. 1961. Composition of gum turpentines of pines. 

Tech. Bull. 1239. kvashington, DC: U.S. Department of 

Agriculture. 158 pp. 

Nikles, Donald G. 1966. Comparative variability and 

relationship of Caribbean pine (Pinus carzbaen Rlnr.) 

and slash pine (Pinxs elliottii Engelm.). Raleigh, KC. 

Raleigh: North C-arolina State University, School of 

Forest Resources. 201 pp. Ph.D. dissertation. 

Perry, J.P. 1987. A new species of Pinvs from h4esico and 

Central America. Journal of the Arnold Arboretum. 

68: 447-459. 

Salazar, R. 1983. Gerlrkic variatiorr in rreedfes (if 6'27. ris 

carrlinea var. R O~JU reiLsts RCirr. et Golf. from x~ai~~rai 

stands. Silvae Genletica. 32: 52-59, 

Sncdecor, George W, 1956. Statistical i~zetIroiis. 5111 ed. 

A~nes, IA: Iowa State ITnir~en.sity Press 53 1r pp 

Squillace, A.E. 1966. Racial r;artaticpr~ iri s?asir pine ns 

abhcted by clirnnt lc. fiii ti,r \ 1-ti.s I\;ap YE-21 ."iIreville, 

XC: U.S. Depardta~r,nd of Ag11cu1t iire, Forest Scrvrrc, 

Saarrtheastern Forest Expel isrneilt Station. IO pp 

Styles, B,T, 1976. Studies of variation ier Centr-ni .$rnericn~~; 

pir~i.s, I. Tire Idcr~tiiy of Pzn-ris cocarpa var. oi.i~oie?cnr~n 

?vlnrtinez. Silr;ae (Tienclica 25: 109-1 17~ 

S~YICF, B.T. 1985. Tins* idtr~tity of 5dlwerdtfrgc.r-'4 C'clntrai 

Arrxerlcan pine, 111: Forest gcr;cxkics researc-4) inaforrrlntionl 

[report]. 13. Rorrre, Italy: Fi4113: 47-Zl. 

Styles, B.T.; Hughes, @.E, 298.3. Studies of variation in 

Central American pines. 3. Yotes or1 tile taxc~rlorriy and 

~~omnenclature of the pines and related gymnoqpermn 

in IIorzduras and ncljacent Latin American repirhlics. 

Bre~tesia. 21 : 269-291. 

Styles, B.T.; McGartcs, P.S, 1988. The botany, ecology, 

distribution, and conservation oh Pzrcus patltin ssp. 

tecunamaniz' in the Republie of tlonduras. Oxford, UK: 

Oxford Forestry Institute, Department of Plant Scie~rres. 

U~liversity of Oxford. 34 pp. 

Styles, B.T.; Stead, J.W.; Rolph, K.J. 1982. 

Studies of variation in Central American pines. 2. 

Putative hybridization between Pinxs caribaea var. 

hondzlrensis and P. nocarpa. Turrialba. 32: 229-242. 

Perry, Jesse P., Jr. 1991. The pines of Mexico and Central 

America. Portland, OR: Timber Press, Inc. 231 pp. 
Table I- rhtive content (percent) of t , artd percent high-LP trees 

Mean percent content Percent highPopulation nu LP trees 

Hi& &phelM 

Group ~-1~ (Vera Cruz, Puebla, and Hidalgo, Mexico) 

49 ~aculti~p~ 6 2 6 0 76 -- 0 

50 El Chico 9 1 1 3 7 6 -- 0 

51 Xoxocatla 8 1 1 5 77 8 0 

52 Huauchinan& 8 1 I 1 80 10 0 

Mean 8 1 2 2 77 9 0 

Group A-2 (northern Oaxaca, Mexico) 

55 Santa M. Papalo 7 2 6 0 7 5 -- 0 

53 Llano d. Flores 7 1 0 4 82 6 7 

54 Ixtlan 

Mean 

High a-pinene, high cxrene or limnene or both, and high f3-phelldrene 

Group B-1 (southern Oaxaca, Mexico) 

58 Tlrrcuache 23 2 43 8 14 -- k 6 

64 Las Trancas 17 4 9 5 54 -- 0 

67 El Manzanal 13 2 14 2 58 -- 0 

70 Juquila 25 2 13 6 43 11 21 

Mean 20 2 20 5 42 11 -7 

Group B-2 (Chiapas, Mexico) 

56 San Jose 12 2 49 8 19 -- -- 

57 Las Piedrecitas 17 2 28 22 16 15 24 

59 Rancho Nuevo 22 1 19 11 3 7 -- 17 

60 Napite & Teoptsca 25 2 2 4 15 24 - - 12 

61 Camino-Chanal 26 1 15 17 31 -- -- 0 

Mean 20 2 2 6 15 24 15 f3 

High *pine, high carene or liraonene or both, arid high #3--phelWrene 

Group B-3 (Guatemala) 

65 Pachoc 8 8 8 36 45 11 92 

66 La Soledad 17 2 1 3 7 32 10 68 

68 San Vicente 12 3 20 18 38 -- 23 

71 San Jose Pinula 12 1 6 38 3 3 -- 52 

73 San Lorenzo 3 3 0 0 3 3 21 13 -- 74 San Jeronimo 38 1 12 24 16 10 36 

Mean 20 1 6 31 31 11 

Group B-4 (Honduras) 

62 Montana Sumpul 30 1 30 21 5 13 21 

63 La Paz 37 2 4 42 5 10 20 

69 Las Trancas 13 1 13 4 1 2 2 -- 33 

72 Guaj iqui ro 2 9 1 6 44 9 10 30 

75 Celaque 17 3 6 18 43 13 11 

Mean 25 2 12 33 17 12 2 3 

See footnotes at end of table. 
Table 1- relative cmtmt (gescmt) of te 

Percent highPopulation LP trees 

Group C (Honduras) 

39 Los Limones 67 6 1 1 18 7 0 

40 Miravelles 62 5 3 3 21 5 -- 

41 Culmi 78 3 1 1 8 8 -- 

43 Guanaja Island 62 6 4 1 2 2 6 -- 

Mean 67 5 2 2 17 6 0 

Group C (Nicaragua) 

38 Santa Clara 69 4 4 1 16 6 -- 45 Alamicmba 71 4 1 1 18 5 -- 47 Karawala 59 3 9 0 20 9 -- 48 Lagma d. Pinar 57 3 7 1 24 7 -- 

Mean 64 4 4, 1 20 7 - - 

High ct-pinme and ephellandrene 

Group C (Belize) 

42 Mt . Pine Ridge 66 6 2 1 24 1 0 

44 Los Lomitas 72 5 2 1 18 3 -- 46 Melinda 66 4 2 1 2 6 2 -- 

Mean 68 5 2 1 23 2 0 

Hi& epinene 

Group D (Mexico) 

2 Dos Aguas 87 1 2 1 2 7 0 

3 Abosola 80 5 3 1 1 10 0 

Mean 84 3 2 1 2 8 0 

Group D (Guatemala) 

23 Unknown 74 10 7 1 1 8 -- 

1 Pueblo Viejo 78 2 4 1 0 14 -- 

4 La Cwnbre 86 4 1 0 0 9 -- 14 Unknown 88 3 0 1 0 8 -- 20 Conacaste 86 1 0 1 2 11 0 

Mean 82 4 2 1 1 10 fi 

5 San Juan 

7 Sigua tepeque 

8 Zamorano 

10 El Corozo 

11 Guaimaca 

12 Villa Santa 

15 Pimientilla 

17 Ocotillo 

18 V.d. Lepaguare 68 5 

Mean w 

Group D (Honduras) 

6 0 1 

0 1 2 

5 1 1 

1 1 0 

1 1 2 

4 1 1 

3 1 5 

0 1 loq 

2 1 12~ 

2 1 4 

High abpinene 

Group D (Nicaragua) 

6 Cusmapa 83 7 1 0 0 9 -- 9 Dipilto 7 9 4 4 1 0 12 -- 13 Las Camelias 83 7 1 1 0 8 -- Mean ?32 6 2 1 0 10 -- 

See footnotes at end of table. 
Percent highPspuPa t ion LP trees 

Group B (Belize) 

16 SIP, Pine Ridge 81 6 3 1 4 5 0 

19 Mt. Pine Ridge 86 6 2 1 2 3 

Mean 

kloderatdy hi& or li or both 

Group E-1 (Mexico) 

22 Rancho Muevo 70 0 0 23 0 7 -- 

Group E-l (Guatemala) 

24 Huehuetenango 72 3 12 1 0 12 0 

30 Bucaral 69 4 15 1 0 12 -- 

Group E-1 (Honduras) 

31 Joccin 68 10 11 1 1 10 0 

25 Zambrano 64 2 14 17 0 4 0 

26 Siguatepeque 67 3 6 9 5 20 4 

27 Cusuco 67 5 8 10 2 7 4 

37 Culmi 68 7 17 1 1 7 0 

Hean 69 4 10 13 1 9 1 

High care or liaronene or both 

Group E-2 (Mexico) 

21 San Cristobal 26 0 5 43 3 23 - - 

Group E-2 (Guatemala) 

28 La Lagunilla 23 2 60 0 0 

High carene or limmene or both 

Group E-2 (Honduras) 

32 San Francisco 51 5 24 6 1 11 

35 Villa Santa 40 7 41 3 1 8 

36 San Esteban 48 4 27 9 2 10 

Group E-2 (Nicaragua) 

29 San Rafael 35 6 48 1 1 10 0 

33 Yucul 18 14 57 0 3 8 

34 Las Hangas 

0 

50 8 24 1 5 12 -- 

Mean 36 6 36 8 2 12 d 

a Indicates contents were not assessed or that individual tree data necessary for determining 

percent high-LP trees were not available. 

b See text for mre detailed definitions of groups. 

Includes Pinal de holes, Queretaro. 

d Includes District 0 Federal. 

Includes Zacapoaxtla, Puebla. 

f Includes Chempil. 

The relatively high means here are likely due to hybridization with P. caribaea (see text). - 
Table 2 Logid traits of 

Needles / Needle Resin Location of resin canals Cone Cone length Peduncle 

Group fasicle length canals Internal Medial External Septa1 length +. width lqth 

Number 

3.2 

3.4 

4.3 

4.1 

- Percent - - - - - - 

91 0 1 

86 0 0 

54 1 0 

96 0 0 

87 2 2 

91 0 1 

12 0 0 

40 9 30 

94 0 0 

95 0 0 

" Based on data from Dvorak and Raymond (1991), McCarter and Birks (1985), Eguiluz-Piedra (1984), 

CAMCORE Cooperative (unpubl , ) , Eguiluz-Piedra and Perry (1983), Salazar (1983), Hunt (1962), and 

Perry (1991). 
for groups x terpene interaction (s,,), obtained 

of tespe concmtratio~~ inall pssible pirs 

b Group Population 

** = sipificmt at the 0,01 level. 

* = significant at the 0.05 level. 

a 'GT = where no is a type of average between the 

no numbers of populations in the two groups involved (Snedecor 1956). 

b See text for definitions. 
h 

% 

lrl 

? 

0, 

n 

\D 

9 

0, 

d m 

0, 

n 

P- 4 

.i 

w 

n * 

? 

c3, 

n 

v) 

9 

9 

w 

r. 

F 

7 w 
JCem j # ij '-Ci 0 0 

"EE 

%32 3. r-t+lrcd 

vl LC 3-4 g8sd 

a 

f-i 

5 

"iJ 

8 
Table 6--Frepuency distributions of percent Pphellmdrene in relatively high vs. low 

elevation Pinus oocarpa populations within the range of - P. caribaea and in - 

P. caribaea 

populations 

Class eleva t ionb elevat ionc - P. caribaead 

- - - - - - - - Number of trees - - - - - - - - 

Total 289 129 15 

Chi-square test of independence: 

High ePev, Low eleve Total 

High Fphellandrene 10 22 31 

Low Pphellanbrene 279 108 387 

Relatively low 

pphellmdrene 

Relatively high 

&phellandrene 

A* Significant at the 0.01 level. 

" Including atypical populations. 

b Population numbers 20, 21, 25, 27, 31, 32, 35, and 36. 

Population numbers 16, 17, 18, 19, and 20. 

d Population numbers 39 and 42. 
Table 7-Sumn~ary of major differences noted between taxonomic entities 

Avg . Avg . btion Avg . needles needle Resin ---.-.. ---- of resin Avg. cone @uncle ?YP 

Taxonomic entity a-pin. \faside length ds s lmgthlwidth leqth PVPU 

- - - - -- 

- - - - - - - - percmt - - - - - - - - Number - tM Nmber Percent - Ratio KN -- ------ 

P. patula var. - long i peduncuh~a 10 2 6 3 69 8 3.6 200 2.1 1&-82-&0 1.71 7 53, 54, 64, 67 58, 7Oc 

P. oocarpa, - moderately atypicald 68 3 10 9 1 8 3.8 184 24 6-54-44 1.04 21 22, 24, 25, 26, 27 

3, 37 

7 8 7 31210 4.6 198 3.4 24-34-10-33 .96 23 I through 15, 17, 18 

20, 23, 31 - 

P. car i baea - and Nicaragua 64 4 4 1 20 6 3.4 226 2.7 83-17-04 1.42 15 38, 39, 40, 43, 45, 47, 48 41 

Internal, medial, external, and sepcal, respectively. 

Population 55 differs from typical g. patula only in having longer cone length/width ratios. 

Populations 58 and 70 tend to resemble 1. tecunmii. 

These populations may contain P. var. achoternae trees, hybrids between 1. tecunwii and P. ooca a, or both. Such hybrids are y more3= E-2 than in E-1. Populations in E-2 tend to 

&e 1. iecun-ii, uhile those in E-1 tend to resemble z. oocarpa. 

' Hexico, Guatemala, Bonduras, and Nicaragua. 

' No specific data available, but krrett and CDlfari (1962) indicate that the ratios were greater than in 

other areas. 
Appendix 1 

Descriptive data for the 75 yzed for rte 

Population 

Lati- hngi- Trees 

buntry' tude tude Elevation smpled ~uthors~ and species &ivenC 

Pueblo Viejo G 

Dos Aguas M 

Abosola M 

La Cmbre G 

San Juan W 

Cusmapa N 

Siguatepeque H 

Zamorano H 

Dipilto N 

El Corozo B 

Gwimaca B 

Villa Santa-l $3 

Las Camelias N 

G 

Pimien t i lla B 

San Pastor Pine Ridge B 

Ocot ililo H 

Valle de Lepaguare B 

M t . Pine Ridge 13 

anaeas te G 

Sari. eristobald k4 

Rancho Nuevo H 

G 

Wuehue tenango G 

Zmbrano H 

Siguatepeque B 

Gus B 

La illa G 

San Rafael N 

Bucaral G 

Joc6n B 

San Francisco W 

'fucul N 

Las Mangas N 

Villa Santa-2 B 

San Esteban El 

Culni f;-2 

Smta Clara N 

Los Limones B 

Miravelles B 

Culmi W 

Ht . Pine Ridge B 

Guana j a Is land H 

Las Lomi tas B 

Meters 

1800 

1700 

1300 

1300 

1300 

1250 

1200 

1200 

1150 

950 

900 

900 

900 

800 

700 

700 

650 

600 

575 

550 

2450 

(" 1 

2000 

1760 

1550 

1475 

1325 

1300 

1150 

1 loo 

1OOO 

I360 

850 

950 

875 

875 

600 

700 

675 

650 

500 

400 

75 

30 

6 (006) 

1, 3, 6 (m) 

2 (m) 

6 (W) 

6 (W) 

6 (W) 

296 (006) 

6 (W) 

6 (W) 

1 (W) 

2 (W) 

6 (W) 

6 (m) 

7 (W) 

6 (m) 

2 (PAT-t) 

1 (WC) 

1, 6 (006) 

1, 6 (OK); 2 (PAT-t) 

2 (N) 

5 (ooc-0) 

8 dWC) 

7 (WC) 

296 (C)o@) 

2 (PAT-t) 

2 (PAT-t) 

3. (W-0); 2 (PAT-t) 

6 (W) 

2 (PAT-t); 6 (W> 

6 (m) 

2 (PAT-t) 

3. (TEC); 2 (PAT-t) 

2 (PAT-t); 6 (W) 

6 (006) 

r (W,TE@); 2 (PliT-t) 

a (TEC); 2 PAT-^) 

a (PAT-t) 

6 (CAR) 

"1 6 

6, 11 (CAR) 

6 (MI 1, 6, 10, 12 (CAR) 

6, 10 (CAR) 

6 (Qw 

See footnotes at end of table, 
Descriptive data for the 75 tions analyzed for t i t ion4n t in& 

Population 

Lati- Longi- Trees 

Comtrya tude tude Elevation sampled ~uthors~ and species givenC 

He ters 

Alami camba 

Melinda 

Karawala 

Laguna del Pinar 

Zacualtiph 

El Chico 

Xoxoca t la 

Huauchinango 

Llano de Flores 

Ix t lan 

Santa Maiia Papalo 

San Jose 

Las Piedreci tas 

Tlacuache 

Rancho Nuevo 

Napite & Teopisca 

Camino-Chanal 

Montana Sumpul 

La Paz 

Las Trancas 

Pachoc 

La Soledad 

El Manzanal 

San Vicente 

Las Trancas 

Juqui la 

San Jose Pinula 

Guaj iquiro 

San Lorenzo 

San Jeronimo 

Celaque 

6 (a) 

6 (CAR) 

9, 12 (CAR) 

6 (CAR) 

1 (PAT) 

1 (PAT) 

2 (PAT) 

1, 3 (PAT) 

2 (PAT-1) 

1, 4 (PAT-1) 

1 (PAT-1) 

1 (OW-O, PAT-1) 

1 (OOC-09 PAT-1); 2 (PAT-t) 

1 (PAT-1) 

1 (OOC-0, PAT-I) 

1 (OOC-0, PAT-1) 

1 (OK-O, PAT-1) 

2 (PAT-t) 

2 (PAT-t) 

1 (PAT-1) 

2 (PAT-t) 

2 (PAT-t); 4, 5 (OOC-0) 

1 (PAT-1) 

1 (TEC) 

1 (mC> 

1 (TEC, OOC-09 PAT-1); 

2 (PAT-t) 

1 (TEC, OOC-0) 

2 (PAT-t) 

4 

1, 4 (TEC); 2 (PAT-t) 

1 (TEC); 2 (PAT-t) 

' M = Mexico, G = Guatemala, H = Honduras, M = Nicaragua, B = Belize. 

1 = present authors, 2 = Lockhart (19851, 3 = Mirov (1961), 4 = Eguiluz-Piedra (1986), 5 = EguiluzPiedra and Perry (1983), 6 = Burlep and Green (19771, 7 = Coppen and others (19881, 8 = Iloff and Mirov 

(1953), 9 = floff ad Mirov (1954), 10 = Nikles (19661, 11 = Coyne and Critchfield (19741, 12 = Burley 

and Green (1979). 

OOC = _P. oocarpa, OOC-o = P. oocar a var. ochoterenae, PAT = P. patula, PAT-t = P. 

tecunumanii, PAT-1 = P. patuTa &ngipedunculata, TEC = E. tecunumanii, CAR = B. - 

hondurensis. 

Nmbers 21 and 56 may be in the s area, but are kept separate because of major differences in 

terpene composition. 

* Elevation not given. 
Appendix 2