Southern 

Research Station

Research Paper

SRS–47

United States

Department of

Agriculture

Forest Service

Diameter Growth of Subtropical 

Trees in Puerto Rico

Thomas J. Brandeis
November 2009

Southern Research Station 

200 W.T. Weaver Blvd.

Asheville, NC 28804

Author: 

Thomas J. Brandeis, Research Forester, U.S. Forest Service, Southern 

Research Station, Forest Inventory and Analysis, Knoxville, TN 37919.

Cover photo: Plantation of mahoe (Hibiscus elatus Sw.) in the Cambalache Commonwealth Forest, 

Arecibo, Puerto Rico.

DISCLAIMER

The use of trade or firm names in this publication is for reader information and does not imply 

endorsement by the U.S. Department of Agriculture of any product or service.
Abstract 

Puerto Rico’s forests consist of young, secondary stands still recovering 

from a long history of island-wide deforestation that largely abated in the 

mid-20th century. Limited knowledge about growth rates of subtropical tree 

species in these forests makes it difficult to accurately predict forest yield, 

biomass accumulation, and carbon sequestration. This study presents mean 

annual increases (periodic annual increment) in tree diameter at breast 

height among trees measured by the forest inventories of Puerto Rico; this 

information is given for each forested life zone, by species, then by species 

and crown class, and by crown position class. Additionally, the study 

presents mean periodic annual increment values calculated for commercial 

species by tree class (growing stock and cull). From 1980 to 2008, mean 

diameter at breast height periodic annual increment was 0.35 cm/year 

for 4,026 trees remeasured by the forest inventory; growth rate averaged 

0.20 cm/year in subtropical dry forests, 0.37 cm/year in subtropical moist 

forests, 0.36 cm/year in subtropical wet/rain forests, and 0.20 cm/year in 

lower montane forests.

Keywords: Caribbean, crown position, FIA, periodic annual increment, 

secondary forest. 

Introduction 

Puerto Rico’s forests consist of young, secondary stands 

still recovering (Birdsey and Weaver 1982, Brandeis and 

others 2007, Franco and others 1997) from a long history 

of island-wide deforestation that largely abated in the mid20th century (Grau and others 2003, Rudel and others 2000, 

Wadsworth 1950). Such secondary forest ecosystems cover 

an increasing percentage of the tropical and subtropical 

landscape not only in the Caribbean but also globally 

(Brown and Lugo 1990, Finegan 1996, Myster 2004). We 

need to better understand secondary forest development and 

function if resource management and land use planning is to 

be informed and effective. 

Knowledge of tree growth rates is fundamental to 

understanding forest function, and can support estimates 

of biomass accumulation, carbon sequestration, and 

commodity production potential. Previous studies on 

Diameter Growth of Subtropical 

Trees in Puerto Rico

Thomas J. Brandeis

long-term research plots have focused on growth rates of 

many tree species in the subtropical wet, subtropical rain, 

lower montane wet, and lower montane rain forest life 

zones of the Luquillo Mountains of Puerto Rico (Crow and 

Weaver 1977, Schmidt and Weaver 1981, Weaver 1979, 

Weaver and Birdsey 1990). [See Ewel and Whitmore (1973) 

for descriptions of these Holdridge life zones.] Studies also 

have focused—albeit less intensively—on growth rates 

of species in the lower ecological zones and in forests at 

earlier successional stages, including estimated growth 

rates in the subtropical dry forest (Briscoe 1962) and the 

subtropical moist forest (Weaver 1979, Weaver and Nieves 

1978). Weaver and Birdsey (1990) were the first to use 

forest inventory remeasurements (from 1980 to 1985) to 

estimate growth rates for trees across the entire island, but 

that inventory excluded some forest types, particularly in the 

subtropical dry forest life zone, that were not considered to 

have the potential for commercial wood products production 

(Birdsey and Weaver 1982) and therefore might not be 

representative of the full range of growth rates.

The latest forest inventory results greatly increased the 

number of remeasured trees and can provide estimates of 

tree growth over a wider range of environmental conditions 

for more species. The increased sampling allows growth rate 

estimation not only for more species but also for species 

under different levels of competition as reflected by the 

tree’s relative position in the canopy. For Bucida buceras L. 

trees in subtropical dry forests, relative canopy position has 

been found a useful benchmark of growth potential.

The objectives of this study were to calculate growth among 

trees measured by the forest inventories of Puerto Rico, with 

growth represented by annual increases (periodic annual 

increment, or PAI) in tree diameter at breast height (d.b.h.). 

Mean PAI values were calculated for each forested life 

zone, by species, then by species and crown position, and by 

crown class. Additionally, mean PAI values were calculated 

for commercial species by tree class (growing stock 

and cull).
2

Methods

Study Area

The data for this study were collected on the main island 

of the Commonwealth of Puerto Rico, centered on 

18°15’ N. by 66°30’ W. Birdsey and Weaver (1982) and 

Ewel and Whitmore (1973) give excellent descriptions 

of the Holdridge life zones commonly used to describe 

these subtropical forests and the species found in them. 

Tree species nomenclature used here comes from the U.S. 

Department of Agriculture, Natural Resources Conservation 

Service, Plants Database (U.S. Department of Agriculture, 

Natural Resources Conservation Service 2006).

Forest Inventories and Tree Measurements

The tree measurements came from four forest inventories 

of Puerto Rico conducted by the Forest Service, Forest 

Inventory and Analysis (FIA) Program. These inventories 

took place in 1980, 1990, 2001–04, and 2006–09. For details 

on these inventories, see Birdsey and Weaver (1982), Franco 

and others (1997), and Brandeis and others (2007). The first 

two forest inventories (1980 and 1990) were each completed 

in 1 year. More recent inventories spread measurements over 

4 years, with each forest inventory plot remeasured every 5 

years. Forest inventory plots are permanent, and each tree 

in the plot is mapped and remeasured with each revisit to 

the plot. Only a small percentage of the trees have been 

measured since 1980, however, due to changes in the forest 

inventory design between 1990 and 2001, as well as damage 

and mortality on some plots associated with the passages 

of Hurricanes Hugo (1989) and Georges (1998). Therefore, 

while a small number of trees has been measured four times 

for more than 20 years, most of the data used to calculate 

PAI comes from two remeasurements separated by 5 or 

10 years.

Tree diameters were measured at a height of 1.4 m (sensu

U.S. Department of Agriculture Forest Service 2002) for 

stems with d.b.h. ≥ 2.5 cm. Trees with abnormal forms and 

those with estimated diameters were removed from the 

dataset. While bole shrinkage can naturally occur, 250 trees 

with negative growth (decreases in d.b.h. in subsequent 

remeasurements) were excluded from this study’s dataset. 

The exclusion of these data might skew results toward 

greater average growth rates.

Tree Crown Rating

Field crews classified each tree crown class in relation 

to the sunlight received and proximity to neighboring 

trees, using these categories: open grown, dominant, 

codominant, intermediate, and suppressed (or overtopped). 

The crown classifications were based on definitions in the 

forest inventory field manual of the U.S. Department of 

Agriculture Forest Service (2002). Open-grown trees have 

crowns that receive full sunlight from above and all sides 

throughout most of their life. Dominant trees have crowns 

that rise above the general level of the crown cover and get 

full sunlight from above and partly from the sides; these 

trees are taller than the average trees in the stand and their 

crowns are well developed, but they could be somewhat 

crowded by other trees on the sides. Codominant trees 

have medium-sized crowns growing at the general level 

of the crown canopy and receiving full overhead light, but 

surrounding trees restrict some sunlight from the sides. In 

stagnated stands, codominant trees have small crowns and 

are crowded on the sides. Intermediate trees are shorter than 

dominants and codominants, but their crowns extend into 

the canopy of codominant and dominant trees and get little 

direct sunlight from above and none from the sides. As a 

result, intermediates usually have small crowns and are very 

crowded from the sides. Suppressed, or overtopped, trees 

have crowns entirely below the general level of the crown 

canopy and receive no sunlight from above or the sides.

To simplify analysis and increase the number of sample 

trees in each category, two broader categories of relative 

crown position also were created: overstory and understory. 

The overstory crown position consists of open-grown, 

dominant, and codominant trees. The understory crown 

position consists of intermediate and suppressed trees.

Tree Class Rating

To measure trees of commercial species, growing-stock 

classifications also were used. The definition of growing 

stock is a live tree of a commercial species that possesses 

(or has the potential to produce, in the case of poletimbersized trees) at least one-third of the gross board-foot 

volume in sound wood. The logs must meet merchantable 

grade, soundness, and size requirements. Trees that do not 

meet growing-stock specifications are called “cull.” No 

distinctions were made in this study between rough and 

rotten cull.
3

Results

A total of 4,026 trees were measured at least twice for 

growth estimates, and their overall mean d.b.h. PAI was 

0.35 cm/year (table 1). Number of trees measured, mean 

d.b.h. PAI, standard error of the mean, standard deviation 

of the mean, and maximum observed PAI for each life zone 

are presented in table 1. These same statistics are presented 

by species and crown class in appendix table A.1, by 

species and crown position in appendix table A.2, and by 

commercial species and tree class in appendix table A.3.

Individual trees of several species exhibited growth rates 

over 2.5 cm/year in this study. They were Eucalyptus 

robusta Sm. (5.84 cm/year), Pithecellobium unguis-cati

(L.) Benth. (5.74 cm/year), Cordia sulcata DC. (5.08 

cm/year), Cecropia schreberiana Miq. (4.30 cm/year), 

Roystonea borinquena O.F. Cook (4.00 cm/year), Spathodea 

campanulata P. Beauv. (3.76 cm/year), Inga laurina (Sw.) 

Willd. (3.32 cm/year), Ficus citrifolia Mill. (3.05 cm/year), 

Dendropanax arboreus (L.) Decne. & Planch. ex Britton 

(3.03 cm/year), Pouteria multiflora (A. DC.) Eyma (3.00 

cm/year), Guarea guidonia (L.) Sleumer (2.90 cm/year), 

Inga vera Willd. (2.88 cm/year), Pithecellobium dulce

(Roxb.) Benth. (2.74 cm/year), Zanthoxylum martinicense

(Lam.) DC. (2.62 cm/year), Erythrina poeppigiana

(Walp.) O.F. Cook (2.61 cm/year) and Inga vera Willd. 

(2.54 cm/year). 

Conclusions

The mean d.b.h. PAI of 0.35 cm/year for trees remeasured 

by the forest inventory from 1980 to 2008 averages growth 

rates over a broad range of environmental conditions and 

species. While it was higher than the mean of 0.21 cm/

year previously reported for the island by Weaver and 

Birdsey (1990) in their analysis of a partial remeasurement 

of the forest inventory plots in 1985, subsequent growth 

measurements in long-term research plots at several sites 

across the island have shown values that range from 0.13 

to 0.47 cm/year (table 2). For growth rates in specific life 

zones, 0.20 cm/year in subtropical dry forests generally 

agrees with values observed by Briscoe (1962) for naturally 

and artificially regenerated stands on limestone substrate. 

Subtropical moist forests consistently show the highest 

potential mean growth rates on the island, followed by 

subtropical wet forests (Weaver 1979). Growth rates 

decrease moving from subtropical wet forest into the lower 

montane forest, as shown in previous studies. Weaver (1983) 

showed lower growth rates in the lower montane forests 

(0.10 to 0.03 cm/year, depending on the forest type) when 

compared to the adjacent subtropical wet forests (0.15 to 

0.23 cm/year, depending on forest type), and theorized that 

increased cloud cover and poorly drained soils found at 

higher elevations in Puerto Rico limit productivity.

The tree growth estimates of Weaver and Birdsey 

(1990) might be lower than current estimates. The later, 

expanded inventory surveyed more understocked and early 

successional stands where trees were colonizing abandoned 

agricultural land and had less competition for site resources. 

A greater percentage of trees with higher growth rates can 

be expected in the later sampling. Silvicultural research 

in Puerto Rico has shown that some species’ growth rates 

respond positively to competition reduction from stand 

thinning. For example, overstory (codominant) tabonuco 

(Dacryodes excelsa Vahl) trees in thinned stands had growth 

rates of 0.70 cm/year, while growth in undisturbed plots 

ranged from 0.15 to 0.23 cm/year (Weaver 1983). 

Disturbance by hurricanes can have an effect on survivor 

growth rates similar to thinning. Two major hurricanes, 

Hurricane Hugo in 1989 and Hurricane Georges in 1998, 

hit Puerto Rican forest stands during the remeasurement 

period. Authors of growth studies done before the passage of 

Hurricane Hugo in 1989 speculated that a large percentage 

of the trees were suppressed and in “steady state” since 

last released by disturbance caused by Hurricane Ciprián 

in 1932 (Crow and Weaver 1977; Weaver 1979, 1983). 

Table 1—Diameter at breast height (1.4 m) periodic 

annual increments (PAI) by Holdridge life zone with 

number of trees measured, standard error of the mean, 

standard deviation of the mean, and maximum observed 

PAI increase from Puerto Rico forest inventory data

Life zone N

Periodic annual increments

Mean SE SD Max

Subtropical dry 307 0.20 0.03 0.45 5.74

Subtropical moist 2,315 0.37 0.01 0.48 4.30

Subtropical wet/rain 1,292 0.36 0.01 0.51 5.84

Lower montane 112 0.20 0.02 0.24 1.28

All life zones 4,026 0.35 0.01 0.49 5.84

N = number of trees measured; SE = standard error of the mean; 

SD = standard deviation of the mean; Max = maximum observed.
4

Hurricane-force winds more heavily damage crowns of 

larger trees, create gaps for forest regeneration, and reduce 

stand basal area (Pascarella and others 2004, Weaver 1989, 

Zimmerman and others 1994). Weaver (1983) states that 

“the critical element determining whether increment was 

slow or rapid would be the amount of time between 

major storms.”

Maximum observed growth rates were much higher than 

those reported in previous studies. Growth rates exceeding 

5 cm/year were found, while previous studies only found 

maximum rates of 1 to 2 cm/year, although growth rates 

exceeding 2.5 cm/year have been observed in Puerto Rico’s 

subtropical wet forests [Wadsworth (1958) as cited in 

Wadsworth (1997)]. However, these maximum growth rates 

were observed in individual trees over short periods of time 

(during a 5- to 10-year period between remeasurements) 

and most likely represent trees growing under ideal 

environmental conditions with little to no competition. 

These extreme growth rates are not fully representative of 

the species’ average growth; rather they express potential 

for growth under excellent growing conditions. This study’s 

larger sample size and wider variety of surveyed sites—

particularly open, recently reverted forests where trees 

are still free from neighboring competition—increase the 

probability of sampling trees that grow under such favorable 

environments. 

The results of this study produce a more comprehensive 

assessment of the average and potential growth, biomass 

accumulation, and carbon sequestration of Caribbean 

subtropical forest trees. This study helps address the need 

for information on tree growth among a wide variety of 

species and growing conditions, and such information can 

validate existing growth and yield models as well as create 

new ones. With such models, we gain the capacity to project 

Table 2—Observed subtropical tree growth rates in Puerto Rico by location, Holdridge life zone and forest-type association, time 

period of the measurements, diameter periodic annual increments, relevant notes on the study, and the bibliographic source

Location

Holdridge life zone and 

forest-type association

Time 

IAPdoirep setoN ecruoS

year cm/year

acin‡uG yrD itra dna larutaN63.0 Ð 31.071 fi cial 

regeneration on karst substrate

Briscoe 1962

Luquillo Mtns. Lower montane (elfi n, 

palm, palo colorado, 

tabonuco)

18 0.30 Ð 0.60 Unthinned stands, with some 

prior tree removals

Crow and Weaver 1977

Rio Piedras Moist 32 0.22 Thinned stands Weaver and Nieves 1978

Luquillo Mtns. Wet, rain, and lower 

montane

5 0.03 Ð 0.36 Thinned and unmanaged stands Weaver 1983

tsioMtsuJ .tS ,nauJ naS 9791 revaeWsdnats yradnoces ylrae ,dennihT74.02

Pi–ones Moist (mangrove) 37 0.46 Natural regeneration after 

clearing

Weaver 1979

Toro Negro Lower montane wet 

(palo colorado)

24 0.15 Thinned stands Weaver 1979

Maricao Lower montane wet 24 0.12 Thinnned stands on serpentine 

substrate

Weaver 1979

Luquillo Mtns. Moist (palo colorado) 27 0.25 Thinned stands Schmidt and Weaver 1981

Islandwide All life zones 5 0.21 Partial survey in 1985, with 

some forest types excluded

Weaver and Birdsey 1990

PAI = periodic annual increments.
5

secondary stand development into the future and choose 

management options that move these forests toward the 

structural and compositional conditions that deliver much 

needed forest ecosystem services to island inhabitants.

Acknowledgments

I would like to thank Ariel Lugo and Eileen Helmer of the 

Forest Service’s International Institute of Tropical Forestry; 

Esther Rojas of the Puerto Rican Conservation Foundation; 

and Jonathan Buford, Johanna D’Arcy, Orlando Díaz, 

Christopher Furr, Jeremy Grayson, Humfredo Marcano, 

Omar Monsegur, Luis Ortíz, Humberto Rodriguez, Jim 

Schiller, and Iván Vicéns for field data collection. I would 

also like to thank Dr. Peter Weaver and Dr. Christopher 

Oswalt for their comments and suggestions on the 

draft manuscript.

Literature Cited 

Birdsey, R.A.; Weaver, P.L. 1982. The forest resources of Puerto Rico. 

Resour. Bull. SO–85. New Orleans: U.S. Department of Agriculture 

Forest Service, Southern Forest Experiment Station. 56 p.

Brandeis, T.J.; Helmer, E.H.; Oswalt, S.N. 2007. The status of Puerto 

Rico’s forests, 2003. Resour. Bull. SRS–119. Asheville, NC: U.S. 

Department of Agriculture Forest Service, Southern Research Station. 

75 p.

Briscoe, C.B. 1962. Tree diameter growth in the dry limestone hills. Trop. 

For. Notes 12. Río Piedras, PR: U.S. Department of Agriculture Forest 

Service, Tropical Forest Research Center. 2 p.

Brown, S.; Lugo, A.E. 1990. Tropical secondary forests. Journal of Tropical 

Ecology. 6: 1–32.

Crow, T.R.; Weaver, P.L. 1977. Tree growth in moist tropical forest of 

Puerto Rico. Res. Pap. ITF–22. Río Piedras, PR: U.S. Department of 

Agriculture Forest Service, Institute of Tropical Forestry. 17 p.

Ewel, J.J.; Whitmore, J.L. 1973. The ecological life zones of Puerto Rico 

and the US Virgin Islands. Res. Pap. ITF–18. Río Piedras, PR: U.S. 

Department of Agriculture Forest Service, Institute of Tropical Forestry. 

72 p.

Finegan, B. 1996. Pattern and process in neotropical secondary rain forests: 

the first 100 years of succession. Tree. 11: 119–124.

Franco, P.A.; Weaver, P.L.; Eggen-McIntosh, S. 1997. Forest resources 

of Puerto Rico, 1990. Resour. Bull. SRS–22. Asheville, NC: U.S. 

Department of Agriculture Forest Service, Southern Research Station. 

45 p.

Grau, R.H.; Aide, T.M.; Zimmerman, J.K. [and others]. 2003. The 

ecological consequences of socioeconomic and land-use changes in 

postagriculture Puerto Rico. BioScience. 53: 1159–1168.

Myster, R.W. 2004. Post-agricultural invasion, establishment, and growth of 

neotropical trees. The Botanical Review. 70: 381–402.

Pascarella, J.B.; Aide, T.M.; Zimmerman, J.K. 2004. Short-term response of 

secondary forests to hurricane disturbance in Puerto Rico, USA. Forest 

Ecology and Management. 199: 379–393.

Rudel, T.K.; Pérez-Lugo, M.; Zichal, H. 2000. When fields revert to forest: 

development and spontaneous reforestation in post-war Puerto Rico. The 

Professional Geographer. 52: 386–397.

Schmidt, R.; Weaver, P.L. 1981. Tree diameter increment in the subtropical 

moist life zone of Puerto Rico. Turrialba. 31: 261–263.

U.S. Department of Agriculture Forest Service. 2002. Field procedures for 

Puerto Rico and the Virgin Islands. Suppl. C to SRS Reg. Man. 1.56. 

http://srsfia2.fs.fed.us/data_acquisition/manual.shtml. [Date accessed: 

June, 2006].

Wadsworth, F.H. 1950. Notes on the climax forests of Puerto Rico and 

their destruction and conservation prior to 1900. Caribbean Forester. 11: 

38–47.

Wadsworth, F.H. 1958. Tropical rain forest. In: Proceedings of the 4th 

World Forestry Congress. Dehra Dun, India: Government of India, 

Manager of Publications: 119–129.

Wadsworth, F.H. 1997. Forest production for tropical America. Agric. 

Handb. 710. Washington, DC: U.S. Department of Agriculture. 563 p.

Weaver, P.L. 1979. Tree growth in several tropical forests of Puerto Rico. 

Res. Pap. SO–152. New Orleans: U.S. Department of Agriculture Forest 

Service, Southern Forest Experiment Station. 15 p.

Weaver, P.L. 1983. Tree growth and stand changes in the subtropical life 

zones of the Luquillo Mountains of Puerto Rico. Res. Pap. SO–190. New 

Orleans: U.S. Department of Agriculture Forest Service, Southern Forest 

Experiment Station. 24 p.

Weaver, P.L. 1989. Forest changes after hurricanes in Puerto Rico’s 

Luquillo Mountains. Interciencia. 14: 181–192.

Weaver, P.L.; Birdsey, R.A. 1990. Growth of secondary forest in Puerto 

Rico between 1980 and 1985. Turrialba. 40: 12–22.

Weaver, P.L.; Nieves, L.O. 1978. Periodic annual dbh increment in a 

subtropical moist forest dominated by Syzygium jambos (L) Alston. 

Turrialba. 28: 253–256.

Zimmerman, J.K.; Everham, E.M., III; Waide, R.B. [and others]. 1994. 

Responses of tree species to hurricane winds in subtropical wet forest 

in Puerto Rico: implications for tropical tree life histories. Journal of 

Ecology. 82: 911–922.

7

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Acacia farnesiana .dlliW ).L( 68.043.051.005.05tnanimodoC

Acrocomia media kooC .F.O 40.0ÑÑ40.01tnanimodoC

Adelia ricinella .L 20.0ÑÑ20.01etaidemretnI

Adenanthera pavonina .L 68.124.061.051.17tnanimodoC

Intermediate 1 1.48 Ñ Ñ 1.48

Overtopped 15 0.25 0.09 0.36 1.28

Albizia procera .htneB ).bxoR( 61.246.081.076.031tnanimoD

Codominant 14 0.74 0.17 0.64 2.22

Intermediate 1 0.04 Ñ Ñ 0.04

Overtopped 2 0.14 0.02 0.03 0.16

Alchornea latifolia .wS 82.0ÑÑ82.01nworg nepO

Dominant 3 0.19 0.13 0.22 0.44

Codominant 10 0.32 0.09 0.29 1.04

Intermediate 7 0.17 0.04 0.10 0.32

Overtopped 2 0.33 0.13 0.18 0.46

Overtopped 1 0.00 Ñ Ñ 0.00

Amyris elemifera .L 24.031.040.041.08tnanimodoC

Intermediate 4 0.05 0.02 0.05 0.10

Overtopped 8 0.04 0.01 0.04 0.08

Andira inermis W .W( .CD xe htnuK )thgir 50.1ÑÑ50.11nworg nepO

Dominant 16 0.33 0.08 0.34 1.26

Codominant 72 0.22 0.03 0.23 0.96

Intermediate 32 0.16 0.04 0.22 0.80

Overtopped 12 0.09 0.03 0.10 0.22

Annona muricata .L 84.033.032.052.02tnanimodoC

Intermediate 2 0.36 0.24 0.34 0.60

Codominant 2 0.09 0.09 0.13 0.18

A. squamosa .L 00.0ÑÑ00.01tnanimoD

Codominant 1 0.92 Ñ Ñ 0.92

Antirhea obtusifolia .brU 61.0ÑÑ61.01etaidemretnI

Codominant 11 0.09 0.03 0.09 0.28

Intermediate 3 0.08 0.03 0.05 0.14

Overtopped 8 0.06 0.03 0.08 0.24

Artocarpus altilis grebsoF )nosnikraP( 78.073.051.083.06tnanimoD

Codominant 13 0.22 0.04 0.15 0.58

Intermediate 5 0.17 0.08 0.18 0.44

Overtopped 1 0.01 Ñ Ñ 0.01

Avicennia germinans .L ).L( 41.070.040.040.04tnanimoD

Codominant 10 0.30 0.14 0.46 1.20

continued

Appendix
8

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Avicennia germinans )deunitnoc( .L ).L( 01.0ÑÑ01.01deppotrevO

Banara portoricensis .brU & gurK 23.0ÑÑ23.01etaidemretnI

Dominant 1 0.34 Ñ Ñ 0.34

Bourreria succulenta .qcaJ 80.0ÑÑ80.01tnanimoD

Codominant 27 0.15 0.05 0.24 0.98

Intermediate 6 0.10 0.06 0.14 0.32

Intermediate 2 0.10 0.02 0.03 0.12

Buchenavia tetraphylla drawoH ).lbuA( 54.144.002.097.05tnanimoD

Codominant 1 0.30 Ñ Ñ 0.30

Overtopped 1 0.04 Ñ Ñ 0.04

Bucida buceras .L 45.080.060.084.02nworg nepO

Dominant 4 0.31 0.05 0.10 0.42

Codominant 6 0.28 0.09 0.22 0.60

Intermediate 1 0.00 Ñ Ñ 0.00

Open grown 2 0.56 0.32 0.45 0.88

Dominant 14 0.49 0.18 0.68 2.48

Codominant 46 0.31 0.04 0.27 1.06

Intermediate 1 0.05 Ñ Ñ 0.05

Overtopped 3 0.07 0.04 0.06 0.12

Byrsonima lucida .CD ).lliM( 60.0ÑÑ60.01tnanimoD

Codominant 5 0.11 0.03 0.07 0.22

B. spicata htnuK ).vaC( 25.060.030.054.04tnanimoD

Codominant 7 0.51 0.20 0.53 1.56

Intermediate 2 0.36 0.16 0.23 0.52

B. wadsworthii elttiL 20.0ÑÑ20.01etaidemretnI

Dominant 10 0.49 0.05 0.16 0.80

Codominant 12 0.47 0.13 0.44 1.38

Intermediate 2 0.40 0.37 0.52 0.76

Overtopped 8 0.28 0.09 0.25 0.80

Canella winterana .ntreaG ).L( 61.030.020.031.03tnanimodoC

Overtopped 1 0.16 Ñ Ñ 0.16

Capparis baducca .L 40.0ÑÑ40.01etaidemretnI

Overtopped 1 0.00 Ñ Ñ 0.00

Codominant 1 0.08 Ñ Ñ 0.08

C. fl exuosa .L ).L( 40.030.020.020.02etaidemretnI

Codominant 4 0.05 0.04 0.08 0.16

Intermediate 2 0.03 0.01 0.01 0.04

Codominant 1 0.10 Ñ Ñ 0.10

Casearia arborea .brU ).hciR( 84.002.090.052.05tnanimodoC

Intermediate 11 0.20 0.04 0.13 0.42

Overtopped 6 0.11 0.08 0.18 0.48

continued
9

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Casearia decandra .qcaJ 01.040.020.030.06etaidemretnI

Overtopped 7 0.02 0.01 0.02 0.05

C. guianensis .brU ).lbuA( 60.0——60.01tnanimoD

Codominant 7 0.18 0.06 0.15 0.42

Intermediate 14 0.17 0.04 0.13 0.36

Overtopped 50 0.08 0.01 0.10 0.50

Codominant 5 0.19 0.09 0.19 0.52

Intermediate 19 0.11 0.03 0.15 0.48

Overtopped 23 0.09 0.02 0.10 0.36

Cassine xylocarpa .tneV 01.040.020.030.05tnanimodoC

Castilla elastica ésseS 42.0——42.01tnanimodoC

Open grown 1 0.68 — — 0.68

Dominant 33 0.82 0.13 0.75 2.74

Codominant 57 0.86 0.11 0.81 4.30

Intermediate 6 0.72 0.47 1.16 3.04

Overtopped 2 1.79 1.70 2.40 3.48

Dominant 2 0.41 0.14 0.19 0.54

Codominant 3 0.06 0.03 0.06 0.12

Intermediate 1 0.00 — — 0.00

Overtopped 1 0.03 — — 0.03

Cestrum laurifolium .réH'L 41.0——41.01tnanimodoC

Dominant 1 0.02 — — 0.02

Codominant 1 0.16 — — 0.16

Cinnamomum elongatum (Vahl ex Nees) Kosterm. Dominant 5 0.58 0.24 0.53 1.48

Codominant 10 0.85 0.17 0.55 1.74

Intermediate 1 0.44 — — 0.44

Overtopped 3 0.31 0.17 0.30 0.62

C. montanum lserP .J & .thcreB ).wS( 24.1——24.11tnanimodoC

Codominant 1 0.04 — — 0.04

Citharexylum spinosum .L 23.0——23.01nworg nepO

Dominant 1 0.50 — — 0.50

Codominant 12 0.17 0.03 0.11 0.34

Intermediate 5 0.36 0.16 0.37 0.98

Overtopped 4 0.03 0.01 0.03 0.06

Citrus paradisi Macfad. (pro sp.) [maxima sinensis] Codominant 3 0.09 0.03 0.05 0.14

Dominant 1 0.00 — — 0.00

Codominant 5 0.10 0.05 0.11 0.22

Intermediate 11 0.19 0.04 0.14 0.46

Overtopped 10 0.05 0.01 0.03 0.11

Clibadium erosum .CD ).wS( 70.0——70.01tnanimodoC

Dominant 1 0.04 — — 0.04

continued

× ×
10

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Clibadium erosum )deunitnoc( .CD ).wS( 61.0ÑÑ61.01tnanimodoC

Intermediate 1 0.00 Ñ Ñ 0.00

Open grown 1 0.64 Ñ Ñ 0.64

Dominant 3 0.29 0.21 0.37 0.71

Codominant 7 0.12 0.05 0.13 0.36

Intermediate 3 0.07 0.05 0.08 0.16

Overtopped 1 0.12 Ñ Ñ 0.12

Dominant 1 1.51 Ñ Ñ 1.51

Coccoloba costata ellavuaS xe thgirW .C 00.0ÑÑ00.01etaidemretnI

Overtopped 3 0.03 0.02 0.04 0.07

Dominant 4 0.18 0.12 0.25 0.54

Codominant 9 0.16 0.05 0.16 0.44

Intermediate 3 0.02 0.01 0.02 0.04

Overtopped 2 0.00 0.00 0.00 0.00

Codominant 8 0.02 0.01 0.03 0.10

C. microstachya .dlliW 61.070.050.011.02tnanimoD

Codominant 15 0.09 0.02 0.08 0.30

Intermediate 1 0.00 Ñ Ñ 0.00

C. swartzii .nsieM 13.0ÑÑ13.01tnanimodoC

Intermediate 1 0.17 Ñ Ñ 0.17

C. venosa .L 43.0ÑÑ43.01etaidemretnI

Codominant 1 1.66 Ñ Ñ 1.66

Cocos nucifera .L 00.0ÑÑ00.01nworg nepO

Dominant 4 0.07 0.03 0.05 0.12

Codominant 2 0.14 0.14 0.20 0.28

Coffea arabica .L nanimodoC 21.0ÑÑ21.01t

Intermediate 13 0.06 0.03 0.10 0.34

Overtopped 16 0.02 0.01 0.04 0.10

Codominant 5 0.25 0.10 0.23 0.53

Overtopped 2 0.20 0.12 0.17 0.32

Overtopped 1 0.14 Ñ Ñ 0.14

Colubrina arborescens .graS ).lliM( 60.0ÑÑ60.01etaidemretnI

Codominant 1 0.02 Ñ Ñ 0.02

Conocarpus erectus .L 49.066.074.074.02tnanimodoC

Intermediate 1 0.00 Ñ Ñ 0.00

Overtopped 1 0.06 Ñ Ñ 0.06

Cordia alliodora nekO ).vaP & ziuR( 27.003.031.072.05tnanimoD

Codominant 9 0.45 0.12 0.36 1.16

Intermediate 2 0.26 0.10 0.14 0.36

Overtopped 1 0.00 Ñ Ñ 0.00

Codominant 1 0.62 Ñ Ñ 0.62

continued
11

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Cordia alliodora (Ruiz & Pav.) Oken (continued) Intermediate 2 0.05 0.03 0.04 0.08

Overtopped 1 0.22 Ñ Ñ 0.22

C. sulcata .CD 67.043.071.034.04tnanimoD

Codominant 10 0.38 0.13 0.41 1.28

Intermediate 4 1.60 1.16 2.32 5.08

Overtopped 3 0.60 0.56 0.97 1.72

Codominant 7 0.07 0.02 0.06 0.18

Croton astroites .dnayrD 23.0ÑÑ23.01etaidemretnI

C. poecilanthus .brU 20.0ÑÑ20.01tnanimoD

Codominant 2 0.22 0.02 0.03 0.24

Intermediate 3 0.05 0.03 0.05 0.10

Cupania americana .L 86.0ÑÑ86.01tnanimoD

Codominant 5 0.67 0.14 0.32 1.00

Intermediate 4 0.32 0.12 0.24 0.62

Overtopped 2 0.78 0.10 0.14 0.88

Dominant 1 0.04 Ñ Ñ 0.04

Codominant 1 0.36 Ñ Ñ 0.36

Codominant 2 0.52 0.52 0.74 1.04

Intermediate 3 0.15 0.12 0.22 0.40

Dominant 1 0.08 Ñ Ñ 0.08

Dacryodes excelsa lhaV 87.003.041.074.05tnanimoD

Codominant 6 0.56 0.11 0.26 0.90

Intermediate 1 0.18 Ñ Ñ 0.18

Overtopped 1 0.36 Ñ Ñ 0.36

Daphnopsis americana .tsnhoJ .R.J ).lliM( 86.0ÑÑ86.01tnanimoD

Overtopped 1 0.28 Ñ Ñ 0.28

Delonix regia .faR ).kooH xe rejoB( nimodoC 24.124.042.030.13tna

Dominant 4 0.86 0.72 1.45 3.03

Codominant 11 0.43 0.19 0.65 2.28

Intermediate 8 0.16 0.03 0.09 0.28

Overtopped 6 0.16 0.08 0.20 0.56

Ditta myricoides .besirG 21.010.010.011.02deppotrevO

Intermediate 1 0.13 Ñ Ñ 0.13

Drypetes glauca lhaV 61.0ÑÑ61.01tnanimodoC

Overtopped 2 0.14 0.02 0.03 0.16

Codominant 2 0.06 0.02 0.03 0.08

Intermediate 1 0.07 Ñ Ñ 0.07

Erythrina berteriana .brU 01.0ÑÑ01.01tnanimoD

Codominant 1 0.64 Ñ Ñ 0.64

E. poeppigiana kooC .F.O ).plaW( 16.266.071.059.051tnanimoD

Codominant 6 0.39 0.16 0.40 1.14

continued
12

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Erythrina poeppigiana (Walp.) O.F. Cook (continued) Intermediate 6 0.56 0.32 0.78 2.10

Overtopped 1 0.04 Ñ Ñ 0.04

Intermediate 3 0.07 0.03 0.05 0.12

Overtopped 3 0.10 0.03 0.06 0.16

Dominant 12 0.61 0.48 1.67 5.84

Codominant 17 0.50 0.13 0.55 1.82

Codominant 4 0.12 0.03 0.06 0.20

Intermediate 6 0.06 0.01 0.04 0.10

Overtopped 5 0.10 0.04 0.09 0.20

Codominant 1 0.52 Ñ Ñ 0.52

Eugenia confusa .CD 40.0ÑÑ40.01etaidemretnI

Overtopped 1 0.14 Ñ Ñ 0.14

Dominant 1 0.38 Ñ Ñ 0.38

E. ligustrina .dlliW ).wS( 80.040.030.050.02tnanimodoC

Intermediate 1 0.00 Ñ Ñ 0.00

E. monticola .CD ).wS( 42.001.050.031.04tnanimodoC

Intermediate 12 0.12 0.03 0.11 0.38

Overtopped 5 0.03 0.02 0.05 0.12

E. pseudopsidium .qcaJ 66.031.090.075.02tnanimoD

Codominant 1 0.00 Ñ Ñ 0.00

Intermediate 5 0.05 0.01 0.02 0.08

Overtopped 1 0.10 Ñ Ñ 0.10

E. xerophytica nottirB 21.070.050.070.02tnanimodoC

Euphorbia cotinifolia .L 00.0ÑÑ00.01tnanimodoC

Dominant 2 0.11 0.01 0.01 0.12

Codominant 7 0.09 0.02 0.06 0.18

Intermediate 5 0.13 0.04 0.09 0.22

Overtopped 2 0.04 0.04 0.06 0.08

Overtopped 2 0.04 0.00 0.00 0.04

Ficus americana .lbuA 56.0ÑÑ56.01tnanimodoC

Dominant 2 1.71 1.35 1.90 3.05

Codominant 5 0.25 0.13 0.29 0.61

Intermediate 2 0.04 0.04 0.05 0.07

Overtopped 1 0.13 Ñ Ñ 0.13

F. trigonata .L 81.0ÑÑ81.01tnanimoD

Codominant 1 0.10 Ñ Ñ 0.10

Codominant 3 0.20 0.16 0.28 0.52

Guajacum offi cinale .L 83.002.041.042.02tnanimodoC

Open grown 2 1.50 0.00 0.00 1.50

Dominant 3 0.18 0.08 0.14 0.30

Codominant 23 0.20 0.05 0.22 0.82

continued
13

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Guajacum offi cinale )deunitnoc( L 65.091.070.062.07etaidemretnI

Overtopped 4 0.32 0.18 0.36 0.68

Codominant 2 0.06 0.04 0.06 0.10

Guarea glabra lhaV 20.0ÑÑ20.01etaidemretnI

Overtopped 3 0.02 0.02 0.03 0.06

G. guidonia remuelS ).L( 21.265.090.029.004tnanimoD

Codominant 67 0.54 0.07 0.55 2.90

Intermediate 52 0.53 0.07 0.49 2.04

Overtopped 71 0.25 0.04 0.30 1.34

Guazuma ulmifolia .maL 24.0ÑÑ24.01nworg nepO

Dominant 5 0.82 0.26 0.58 1.80

Codominant 11 0.44 0.11 0.36 1.18

Intermediate 1 0.04 Ñ Ñ 0.04

Overtopped 1 0.28 Ñ Ñ 0.28

Guettarda scabra .tneV ).L( 60.020.010.040.03tnanimoD

Codominant 15 0.06 0.02 0.07 0.28

Intermediate 14 0.01 0.01 0.02 0.06

Overtopped 9 0.01 0.01 0.02 0.04

Gyminda latifolia .brU ).wS( 80.0ÑÑ80.01tnanimoD

Intermediate 2 0.11 0.05 0.07 0.16

Overtopped 3 0.07 0.03 0.06 0.13

Codominant 13 0.04 0.01 0.04 0.10

Intermediate 8 0.05 0.01 0.03 0.08

Overtopped 2 0.03 0.03 0.04 0.06

Codominant 2 0.53 0.01 0.01 0.54

Henriettea squamulosum dduJ .S.W ).ngoC( 82.0ÑÑ82.01tnanimoD

Codominant 1 0.32 Ñ Ñ 0.32

Intermediate 2 0.06 0.06 0.08 0.12

Intermediate 1 0.02 Ñ Ñ 0.02

Hibiscus elatus .wS 60.0ÑÑ60.01deppotrevO

Hirtella rugosa .sreP xe .lliuhT 80.060.040.040.02etaidemretnI

Codominant 7 0.21 0.08 0.21 0.68

Intermediate 1 0.02 Ñ Ñ 0.02

Overtopped 1 0.14 Ñ Ñ 0.14

Dominant 6 0.36 0.06 0.14 0.62

Codominant 7 0.31 0.06 0.16 0.62

Intermediate 6 0.25 0.10 0.24 0.60

Overtopped 3 0.10 0.04 0.07 0.18

Ilex nitida .mixaM )lhaV( 00.0ÑÑ00.01etaidemretnI

Inga laurina .dlliW ).wS( 23.380.133.079.011tnanimoD

Codominant 24 0.50 0.11 0.51 2.22

continued
14

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Inga laurina )deunitnoc( .dlliW ).wS( 44.144.051.063.09etaidemretnI

Overtopped 2 0.11 0.11 0.15 0.21

Dominant 1 0.13 Ñ Ñ 0.13

Intermediate 1 0.54 Ñ Ñ 0.54

Dominant 24 0.62 0.10 0.48 2.18

Codominant 46 0.42 0.08 0.54 2.54

Intermediate 7 0.30 0.07 0.19 0.56

Overtopped 10 0.37 0.12 0.37 1.26

Ixora ferrea .htneB ).qcaJ( 41.1ÑÑ41.11tnanimoD

Codominant 1 0.64 Ñ Ñ 0.64

Krugiodendron ferreum .brU )lhaV( 82.021.060.011.04tnanimodoC

Intermediate 6 0.03 0.01 0.02 0.06

Laguncularia racemosa .ntreaG .F.C ).L( 00.0ÑÑ00.01tnanimoD

Intermediate 4 0.15 0.04 0.07 0.24

Overtopped 2 0.15 0.03 0.04 0.18

Leucaena leucocephala tiW ed ).maL( 06.081.060.063.08tnanimoD

Codominant 43 0.22 0.03 0.19 0.84

Intermediate 12 0.10 0.02 0.07 0.20

Overtopped 1 0.02 Ñ Ñ 0.02

Intermediate 1 0.18 Ñ Ñ 0.18

Licaria parvifolia .mretsoK ).maL( 84.0ÑÑ84.01tnanimodoC

Intermediate 1 0.00 Ñ Ñ 0.00

Overtopped 1 0.02 Ñ Ñ 0.02

Lonchocarpus glaucifolius .brU 60.0ÑÑ60.01tnanimodoC

Intermediate 1 0.10 Ñ Ñ 0.10

L. heptaphyllus .CD ).rioP( 22.060.050.081.02tnanimoD

Codominant 1 0.36 Ñ Ñ 0.36

Dominant 1 0.90 Ñ Ñ 0.90

Codominant 1 0.15 Ñ Ñ 0.15

Mammea americana .L 11.1ÑÑ11.11tnanimoD

Codominant 1 0.31 Ñ Ñ 0.31

Overtopped 1 0.02 Ñ Ñ 0.02

Open grown 6 0.41 0.12 0.29 0.75

Dominant 14 0.53 0.10 0.38 1.38

Codominant 30 0.35 0.08 0.45 2.28

Intermediate 6 0.29 0.15 0.37 0.94

Overtopped 2 0.16 0.10 0.14 0.26

Manilkara bidentata vehC .A ).CD .A( 50.1ÑÑ50.11tnanimoD

Codominant 1 0.61 Ñ Ñ 0.61

Margaritaria nobilis .f .L 20.0ÑÑ20.01tnanimodoC

Intermediate 1 0.00 Ñ Ñ 0.00

continued
15

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Margaritaria nobilis )deunitnoc( .f .L 80.0ÑÑ80.01deppotrevO

Matayba domingensis .kldaR ).CD( 21.0ÑÑ21.01tnanimoD

Codominant 2 0.12 0.04 0.06 0.16

Maytenus ponceana nottirB 61.0ÑÑ61.01etaidemretnI

Overtopped 1 0.00 Ñ Ñ 0.00

Codominant 8 0.63 0.13 0.37 1.16

Intermediate 3 0.36 0.16 0.27 0.58

Overtopped 2 0.11 0.01 0.01 0.12

Dominant 1 0.22 Ñ Ñ 0.22

Codominant 5 0.11 0.03 0.07 0.22

Intermediate 1 0.14 Ñ Ñ 0.14

Miconia impetiolaris .CD xe noD .D ).wS( 94.0ÑÑ94.01tnanimoD

Intermediate 1 0.08 Ñ Ñ 0.08

M. laevigata noD .D ).L( 86.081.080.093.05tnanimodoC

Intermediate 2 0.34 0.14 0.20 0.48

Dominant 1 0.02 Ñ Ñ 0.02

Codominant 2 0.63 0.12 0.16 0.74

Intermediate 3 0.29 0.13 0.22 0.52

Overtopped 5 0.08 0.06 0.13 0.30

M. pycnoneura .brU 21.0ÑÑ21.01deppotrevO

Open grown 1 0.33 Ñ Ñ 0.33

M. tetrandra noD .D ).wS( 01.0ÑÑ01.01tnanimoD

Codominant 1 0.12 Ñ Ñ 0.12

Intermediate 2 0.51 0.17 0.24 0.68

Overtopped 1 0.50 Ñ Ñ 0.50

Dominant 5 0.05 0.01 0.03 0.10

Codominant 8 0.16 0.09 0.25 0.72

Dominant 1 0.05 Ñ Ñ 0.05

Codominant 21 0.18 0.03 0.13 0.46

Intermediate 3 0.13 0.08 0.14 0.28

Overtopped 1 0.15 Ñ Ñ 0.15

Myrcia citrifolia .brU ).lbuA( 01.0ÑÑ01.01tnanimodoC

M. defl exa .CD ).rioP( 13.0ÑÑ13.01tnanimodoC

Intermediate 2 0.20 0.04 0.06 0.24

Overtopped 2 0.06 0.00 0.01 0.06

M. fallax .CD ).hciR( 40.010.010.030.02etaidemretnI

Codominant 3 0.28 0.14 0.24 0.54

Intermediate 7 0.33 0.10 0.26 0.62

Overtopped 6 0.07 0.04 0.10 0.24

Codominant 4 0.47 0.04 0.09 0.54

Intermediate 1 0.32 Ñ Ñ 0.32

continued
16

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Myrcia fallax )deunitnoc( .CD ).hciR( 61.0ÑÑ61.01deppotrevO

Dominant 1 0.34 Ñ Ñ 0.34

Codominant 3 0.43 0.07 0.12 0.52

Intermediate 1 0.02 Ñ Ñ 0.02

Nectandra coriacea .besirG ).wS( 22.050.030.051.04tnanimodoC

Intermediate 2 0.36 0.04 0.06 0.40

Overtopped 2 0.08 0.08 0.11 0.16

N. hihua rewhoR ).vaP & ziuR( 05.1ÑÑ05.11tnanimoD

Dominant 2 0.78 0.04 0.06 0.82

Codominant 1 0.73 Ñ Ñ 0.73

Overtopped 2 0.08 0.02 0.03 0.10

Neea buxifolia lremieH ).f .kooH( 20.0ÑÑ20.01etaidemretnI

Neolaugeria resinosa noslociN )lhaV( 07.0ÑÑ07.01tnanimoD

Codominant 12 0.27 0.07 0.24 0.70

Intermediate 8 0.24 0.08 0.22 0.74

Overtopped 2 0.11 0.09 0.13 0.20

Ocotea fl oribunda zeM ).wS( 43.0ÑÑ43.01tnanimoD

Codominant 1 0.28 Ñ Ñ 0.28

Dominant 4 0.46 0.07 0.15 0.61

Codominant 15 0.25 0.05 0.21 0.66

Intermediate 11 0.17 0.08 0.26 0.76

Overtopped 11 0.26 0.09 0.29 0.78

O. moschata zeM ).nsieM xe .vaP( 25.0ÑÑ25.01tnanimoD

Codominant 1 0.18 Ñ Ñ 0.18

O. wrightii zeM ).nsieM( 85.0ÑÑ85.01tnanimoD

Ormosia krugii .brU 23.060.040.082.02tnanimoD

Codominant 9 0.22 0.07 0.22 0.66

Intermediate 3 0.07 0.03 0.05 0.12

Overtopped 1 0.00 Ñ Ñ 0.00

Ouratea littoralis .brU 80.0ÑÑ80.01etaidemretnI

Open grown 1 5.74 Ñ Ñ 5.74

Codominant 4 0.04 0.03 0.06 0.12

Palicourea croceoides .maH 11.040.020.070.03tnanimodoC

Overtopped 2 0.11 0.03 0.04 0.13

Open grown 1 0.58 Ñ Ñ 0.58

Peltophorum pterocarpum (DC.) Backer ex K. Heyne Codominant 5 0.64 0.04 0.09 0.72

Dominant 1 0.60 Ñ Ñ 0.60

Codominant 5 0.64 0.31 0.70 1.80

Intermediate 3 1.26 0.52 0.90 2.28

Dominant 1 0.20 Ñ Ñ 0.20

continued
17

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Peltophorum pterocarpum (DC.) Backer ex K. Heyne (continued) Codominant 3 0.21 0.04 0.08 0.28

Intermediate 1 0.30 Ñ Ñ 0.30

Overtopped 1 0.06 Ñ Ñ 0.06

Picramnia pentandra .wS 80.0ÑÑ80.01etaidemretnI

Dominant 1 0.14 Ñ Ñ 0.14

Codominant 12 0.05 0.02 0.06 0.16

Intermediate 1 0.00 Ñ Ñ 0.00

Overtopped 2 0.13 0.03 0.04 0.16

Pilosocereus royenii yelwoR & selyB ).L( 26.024.003.023.02tnanimodoC

Codominant 7 0.28 0.08 0.22 0.66

Piper aduncum .L 51.0ÑÑ51.01etaidemretnI

Codominant 2 0.05 0.01 0.01 0.06

Intermediate 3 0.25 0.16 0.27 0.56

Intermediate 1 0.14 Ñ Ñ 0.14

Pithecellobium dulce .htneB ).bxoR( 60.2ÑÑ60.21tnanimoD

Codominant 4 1.53 0.47 0.94 2.74

Plumeria obtusa .L 42.040.020.012.03tnanimodoC

Podocarpus coriaceus .hciR 50.0ÑÑ50.01tnanimodoC

Poitea fl orida nivaL )lhaV( 40.0ÑÑ40.01tnanimoD

Pouteria multifl ora amyE ).CD .A( 00.313.195.066.05tnanimoD

Codominant 2 0.22 0.20 0.29 0.42

Overtopped 2 0.54 0.00 0.00 0.54

P. sapota nraetS & erooM .E.H ).qcaJ( 42.0ÑÑ42.01tnanimoD

Codominant 1 0.46 Ñ Ñ 0.46

Prestoea acuminata (Willd.) H.E. Moore var. 

montana onaelaG .G & .dneH .A )maharG( 80.030.010.020.07tnanimoD

Codominant 34 0.04 0.01 0.06 0.20

Intermediate 27 0.16 0.03 0.18 0.74

Overtopped 20 0.19 0.05 0.25 1.00

Prosopis pallida (Humb. & Bonpl. ex Willd.) Kunth Codominant 3 0.11 0.11 0.18 0.32

Prunus myrtifolia .brU ).L( 90.0ÑÑ90.01deppotrevO

Pseudolmedia spuria .besirG ).wS( 81.0ÑÑ81.01tnanimodoC

Psidium amplexicaule .sreP 50.0ÑÑ50.01etaidemretnI

Overtopped 4 0.07 0.03 0.07 0.16

P. guajava .L 13.021.050.070.06tnanimodoC

Intermediate 3 0.04 0.02 0.03 0.06

Overtopped 2 0.00 0.00 0.00 0.00

Psychotria berteriana .CD 40.0ÑÑ40.01deppotrevO

continued
18

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Psychotria brachiata .wS 20.010.010.010.02deppotrevO

Quararibea turbinata .rioP ).wS( 04.051.070.012.04etaidemretnI

Overtopped 3 0.08 0.03 0.05 0.12

Randia aculeata .L 62.080.040.071.03tnanimodoC

Intermediate 1 0.24 Ñ Ñ 0.24

Overtopped 1 0.08 Ñ Ñ 0.08

Rhizophora mangle .L 00.0ÑÑ00.01tnanimoD

Codominant 1 0.00 Ñ Ñ 0.00

Intermediate 1 0.00 Ñ Ñ 0.00

Roystonea borinquena kooC .F.O 00.0ÑÑ00.01nworg nepO

Dominant 10 0.25 0.13 0.42 1.25

Codominant 6 0.46 0.21 0.52 1.28

Intermediate 2 3.40 0.60 0.85 4.00

Overtopped 1 0.44 Ñ Ñ 0.44

Sagraea umbrosa .CD ).wS( 72.070.040.002.03tnanimoD

Codominant 2 0.20 0.09 0.12 0.28

Intermediate 1 0.45 Ñ Ñ 0.45

Samanea saman .rreM ).qcaJ( 03.2ÑÑ03.21tnanimodoC

Sapindus saponaria .L 52.0ÑÑ52.01tnanimodoC

Sapium laurocerasus .fseD 43.0ÑÑ43.01tnanimodoC

Intermediate 1 0.88 Ñ Ñ 0.88

Savia sessilifl ora .dlliW ).wS( 01.060.040.060.02deppotrevO

Scheffl era morototonii (Aubl.) Maguire, Steyerm. & Frodin Open grown 1 0.32 Ñ Ñ 0.32

Dominant 5 0.41 0.11 0.25 0.72

Codominant 18 0.33 0.06 0.27 0.88

Intermediate 4 0.82 0.35 0.70 1.70

Overtopped 3 0.21 0.04 0.06 0.28

Schoepfi a obovata thgirW .C 00.0ÑÑ00.01tnanimodoC

Senna siamea ybenraB & niwrI ).maL( 20.207.072.066.07tnanimodoC

Intermediate 2 0.98 0.50 0.71 1.48

Sideroxylon cubense .nneP .D.T ).besirG( 20.0ÑÑ20.01tnanimodoC

Intermediate 1 0.11 Ñ Ñ 0.11

S. salicifolium .maL ).L( 14.051.050.022.07tnanimodoC

Intermediate 2 0.08 0.04 0.06 0.12

Sloanea berteriana .CD xe ysiohC 22.0ÑÑ22.01tnanimoD

Codominant 4 0.13 0.02 0.05 0.20

Intermediate 5 0.05 0.02 0.04 0.10

Solanum rugosum lanuD 81.031.090.090.02deppotrevO

continued
19

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Spathodea campanulata .vuaeB .P 42.366.001.059.064tnanimoD

Codominant 137 0.77 0.06 0.67 3.76

Intermediate 79 0.44 0.05 0.44 2.14

Overtopped 69 0.22 0.03 0.28 1.46

Spondias dulcis nosnikraP 74.0ÑÑ74.01tnanimoD

Codominant 1 0.00 Ñ Ñ 0.00

S. mombin .L 20.103.021.046.06tnanimoD

Codominant 7 0.59 0.16 0.43 1.18

Intermediate 3 0.35 0.07 0.12 0.43

Swietenia macrophylla gniK 27.043.002.033.03tnanimodoC

S. mahagoni .qcaJ ).L( 84.032.031.072.03tnanimoD

Codominant 7 0.16 0.05 0.13 0.30

Intermediate 1 0.16 Ñ Ñ 0.16

Symplocos martinicensis .qcaJ 46.022.011.093.04tnanimodoC

Syzygium jambos notslA ).L( 00.000.000.000.02tnanimoD

Codominant 27 0.12 0.03 0.14 0.54

Intermediate 30 0.15 0.04 0.20 0.90

Overtopped 26 0.10 0.04 0.18 0.70

Tabebuia haemantha (Bertol. ex Spreng.) DC. Dominant 1 0.12 Ñ Ñ 0.12

Codominant 5 0.03 0.02 0.05 0.12

Intermediate 1 0.04 Ñ Ñ 0.04

Overtopped 1 0.02 Ñ Ñ 0.02

T. heterophylla nottirB ).CD( 052.050.072.072tnanimoD 48.

Codominant 50 0.38 0.06 0.39 1.46

Intermediate 22 0.20 0.05 0.23 0.78

Overtopped 12 0.08 0.02 0.08 0.30

T. rigida .brU 00.0ÑÑ00.01tnanimodoC

Tamarindus indica .L 25.1ÑÑ25.11nworg nepO

Codominant 1 1.08 Ñ Ñ 1.08

Terminalia catappa .L 38.091.080.016.05tnanimoD

Codominant 1 0.68 Ñ Ñ 0.68

Intermediate 1 0.42 Ñ Ñ 0.42

Tetragastris balsamifera nekO ).wS( 80.0ÑÑ80.01tnanimoD

Codominant 3 0.38 0.16 0.28 0.60

Intermediate 2 0.29 0.01 0.01 0.30

Overtopped 1 0.80 Ñ Ñ 0.80

Tetrazygia elaeagnoides .CD ).wS( 69.072.080.012.011tnanimodoC

Intermediate 4 0.19 0.06 0.11 0.34

Overtopped 3 0.07 0.05 0.08 0.16

continued
20

Table A.1—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown class with number 

of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI increase from 

Puerto Rico forest inventory data (continued)

Speciesa Crown class N

Periodic annual increments

Mean SE SD Max

Thespesia grandifl ora .CD 63.041.080.002.03tnanimoD

Codominant 6 0.38 0.16 0.39 1.14

Intermediate 3 0.11 0.05 0.08 0.20

Overtopped 1 0.22 Ñ Ñ 0.22

Thouinia striata .kldaR 48.0ÑÑ48.01tnanimoD

Codominant 9 0.10 0.04 0.13 0.38

Intermediate 5 0.19 0.09 0.19 0.50

T. striata Radlk. var. portoricensis (Radlk.) Votava & Alain Dominant 2 0.06 0.00 0.00 0.06

Codominant 5 0.02 0.02 0.04 0.10

Intermediate 5 0.02 0.02 0.03 0.08

Trema micrantha emulB ).L( 45.0ÑÑ45.01tnanimodoC

Overtopped 1 0.06 Ñ Ñ 0.06

Trichilia hirta .L 47.043.042.005.02tnanimodoC

Overtopped 1 0.00 Ñ Ñ 0.00

T. pallida .wS 44.081.001.052.03etaidemretnI

Overtopped 9 0.05 0.02 0.05 0.16

Turpinia occidentalis noD .G ).wS( 87.072.090.062.001tnanimodoC

Overtopped 1 0.01 Ñ Ñ 0.01

Urera baccifera .hciduaG ).L( 40.0ÑÑ40.01deppotrevO

Vitex divaricata .wS 67.043.071.013.04tnanimodoC

Intermediate 1 0.12 Ñ Ñ 0.12

Xylosma buxifolia yarG .A 50.0ÑÑ50.01etaidemretnI

X. pachyphylla .brU ).brU & gurK( 41.0ÑÑ41.01tnanimodoC

Zanthoxylum martinicense .CD ).maL( 26.228.092.048.08tnanimoD

Codominant 15 0.43 0.09 0.35 1.38

Intermediate 3 0.31 0.11 0.19 0.48

Overtopped 5 0.21 0.09 0.19 0.42

Z. monophyllum nosliW .P ).maL( 84.071.021.063.02tnanimodoC

Overtopped 2 0.03 0.01 0.01 0.04

— = insuffi cient sample; N = number of trees measured; SE = standard error of the mean; SD = standard deviation of the mean; Max = maximum 

observed.

a

 USDA Natural Resources Conservation Service (2006).
21

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Acacia farnesiana .dlliW ).L( 68.043.051.005.05yrotsrevO

All 5 0.50 0.15 0.34 0.86

Acrocomia media kooC .F.O 40.0ÑÑ40.01yrotsrevO

All 1 0.04 Ñ Ñ 0.04

Adelia ricinella .L 20.0ÑÑ20.01yrotsrednU

All 1 0.02 Ñ Ñ 0.02

Adenanthera pavonina .L 68.124.061.051.17yrotsrevO

Understory 16 0.33 0.12 0.47 1.48

All 23 0.58 0.12 0.59 1.86

Albizia procera .htneB ).bxoR( 22.236.021.017.072yrotsrevO

Understory 3 0.11 0.04 0.06 0.16

All 30 0.65 0.11 0.62 2.22

Alchornea latifolia .wS 40.172.070.072.031yrotsrevO

Understory 9 0.21 0.04 0.13 0.46

All 38 0.27 0.04 0.23 1.04

Alsophila portoricensis (Spreng. ex Kuhn) Conant Understory 1 0.00 Ñ Ñ 0.00

All 1 0.00 Ñ Ñ 0.00

Amyris elemifera .L 24.031.040.041.08yrotsrevO

Understory 12 0.04 0.01 0.04 0.10

All 21 0.08 0.02 0.10 0.42

Andira inermis .CD xe htnuK )thgirW .W( tsrevO 62.152.030.042.088yro

Understory 44 0.14 0.03 0.20 0.80

All 169 0.23 0.02 0.25 1.26

Annona muricata .L 84.033.032.052.02yrotsrevO

Understory 2 0.36 0.24 0.34 0.60

All 4 0.31 0.14 0.28 0.60

A. reticulata .L 81.031.090.090.02yrotsrevO

All 2 0.09 0.09 0.13 0.18

A. squamosa .L 29.056.064.064.02yrotsrevO

All 2 0.46 0.46 0.65 0.92

Antirhea obtusifolia .brU 61.0ÑÑ61.01yrotsrednU

All 1 0.16 Ñ Ñ 0.16

Ardisia obovata .maH xe .vseD 82.090.030.090.011yrotsrevO

Understory 11 0.07 0.02 0.07 0.24

All 27 0.07 0.01 0.07 0.28

Artocarpus altilis grebsoF )nosnikraP( 86.091.050.052.041yrotsrevO

Understory 5 0.17 0.08 0.18 0.44

All 29 0.23 0.04 0.23 0.81

continued
22

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Avicennia germinans .L ).L( 02.104.011.032.041yrotsrevO

Understory 1 0.10 Ñ Ñ 0.10

All 15 0.22 0.10 0.39 1.20

Banara portoricensis .brU & gurK 23.0ÑÑ23.01yrotsrednU

All 1 0.28 Ñ Ñ 0.28

Beilschmiedia pendula .lsmeH ).wS( 43.0ÑÑ43.01yrotsrevO

All 1 0.34 Ñ Ñ 0.34

Bourreria succulenta .qcaJ 89.032.040.051.082yrotsrevO

Understory 6 0.10 0.06 0.14 0.32

All 34 0.14 0.04 0.22 0.98

B. virgata noD .G ).wS( 21.030.020.001.02yrotsrednU

All 3 0.08 0.02 0.04 0.12

Buchenavia tetraphylla drawoH ).lbuA( 54.144.081.017.06yrotsrevO

Understory 1 0.04 Ñ Ñ 0.04

All 7 0.62 0.18 0.48 1.45

Bucida buceras .L 06.081.050.023.021yrotsrevO

Understory 1 0.00 Ñ Ñ 0.00

All 15 0.34 0.07 0.27 1.06

Bursera simaruba graS ).L( . 84.204.050.063.026yrotsrevO

Understory 4 0.05 0.03 0.06 0.12

All 67 0.34 0.05 0.40 2.48

Byrsonima lucida .CD ).lliM( 22.070.030.001.06yrotsrevO

All 6 0.10 0.03 0.07 0.22

B. spicata htnuK ).vaC( 65.124.031.094.011yrotsrevO

Understory 2 0.36 0.16 0.23 0.52

All 19 0.44 0.09 0.39 1.56

B. wadsworthii elttiL 20.0ÑÑ20.01yrotsrednU

All 1 0.02 Ñ Ñ 0.02

Calophyllum antillanum nottirB 83.163.080.084.091yrotsrevO

Understory 8 0.28 0.09 0.25 0.80

All 41 0.41 0.05 0.32 1.38

Canella winterana .ntreaG ).L( 61.030.020.031.03yrotsrevO

Understory 1 0.16 Ñ Ñ 0.16

All 4 0.14 0.01 0.03 0.16

Capparis baducca .L 40.030.020.020.02yrotsrednU

All 2 0.02 0.02 0.03 0.04

C. cynophallophora .L 80.0ÑÑ80.01yrotsrevO

All 1 0.08 Ñ Ñ 0.08

continued
23

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Capparis fl exuosa .L ).L( 40.030.020.020.02yrotsrednU

All 2 0.02 0.02 0.03 0.04

C. hastata .qcaJ 61.080.040.050.04yrotsrevO

Understory 2 0.03 0.01 0.01 0.04

All 6 0.04 0.02 0.06 0.16

Carapa guianensis .lbuA 01.0ÑÑ01.01yrotsrevO

All 1 0.10 Ñ Ñ 0.10

Casearia arborea .brU ).hciR( 84.002.090.052.05yrotsrevO

Understory 17 0.17 0.04 0.15 0.48

All 32 0.20 0.03 0.16 0.49

C. decandra .qcaJ 01.030.010.030.031yrotsrednU

All 17 0.03 0.01 0.03 0.10

C. guianensis .brU ).lbuA( 24.041.050.071.08yrotsrevO

Understory 64 0.10 0.01 0.12 0.50

All 99 0.11 0.01 0.11 0.50

C. sylvestris .wS 25.091.090.091.05yrotsrevO

Understory 42 0.10 0.02 0.12 0.48

All 67 0.11 0.02 0.13 0.56

Cassine xylocarpa .tneV evO 01.040.020.030.05yrotsr

All 5 0.03 0.02 0.04 0.10

Castilla elastica ŽsseS 42.0ÑÑ42.01yrotsrevO

All 3 0.82 0.68 1.17 2.17

Cecropia schreberiana .qiM 03.418.090.048.067yrotsrevO

Understory 8 0.91 0.52 1.46 3.48

All 137 0.80 0.07 0.77 4.30

Cedrela odorata .L 82.011.050.031.05yrotsrevO

Understory 2 0.02 0.02 0.02 0.03

All 11 0.18 0.05 0.18 0.54

Cestrum laurifolium .rŽH'L 41.0ÑÑ41.01yrotsrevO

All 1 0.14 Ñ Ñ 0.14

Chrysophyllum cainito .L 61.001.070.090.02yrotsrevO

All 2 0.09 0.07 0.10 0.16

Cinnamomum elongatum (Vahl ex Nees) Kosterm. Overstory 15 0.76 0.14 0.54 1.74

Understory 4 0.34 0.13 0.25 0.62

All 26 0.60 0.09 0.48 1.74

C. montanum lserP .J & .thcreB ).wS( 24.1ÑÑ24.11yrotsrevO

All 1 1.42 Ñ Ñ 1.42

Citharexylum caudatum .L 40.0ÑÑ40.01yrotsrevO

All 1 0.04 Ñ Ñ 0.04

continued
24

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Citharexylum spinosum .L 05.041.040.002.041yrotsrevO

Understory 9 0.22 0.10 0.31 0.98

All 23 0.21 0.05 0.22 0.98

Citrus paradisi Macfad. (pro sp.) [maxima ×

×

×

×

sinensis] Overstory 3 0.09 0.03 0.05 0.14

All 3 0.09 0.03 0.05 0.14

C. sinensis (L.) Osbeck (pro sp.) [maxima reticulata] Overstory 6 0.08 0.04 0.10 0.22

Understory 21 0.07 0.01 0.07 0.24

All 38 0.13 0.03 0.21 1.19

Clibadium erosum .CD ).wS( 70.0——70.01yrotsrevO

All 1 0.07 — — 0.07

Clusia clusioides ycrA'D ).besirG( 61.080.060.001.02yrotsrevO

Understory 1 0.00 — — 0.00

All 3 0.07 0.05 0.08 0.16

C. rosea .qcaJ 46.022.080.081.08yrotsrevO

Understory 4 0.08 0.04 0.07 0.16

All 23 0.29 0.07 0.31 0.98

Cnemidaria horrida lserP .C ).L( 15.1——15.11yrotsrevO

All 1 1.51 — — 1.51

Coccoloba costata ellavuaS xe thgirW .C 70.030.020.020.04yrotsrednU

All 4 0.02 0.02 0.03 0.07

C. diversifolia .qcaJ 45.081.050.071.031yrotsrevO

Understory 5 0.01 0.01 0.02 0.04

All 23 0.12 0.03 0.16 0.54

C. krugii uadniL 01.030.010.020.08yrotsrevO

All 8 0.02 0.01 0.03 0.10

C. microstachya .dlliW 03.080.020.090.071yrotsrevO

All 18 0.11 0.02 0.09 0.31

C. pyrifolia .fseD 00.0——00.01yrotsrednU

All 1 0.00 — — 0.00

C. swartzii .nsieM 3.0——13.01yrotsrevO 1

Understory 1 0.17 — — 0.17

All 2 0.24 0.07 0.10 0.31

C. venosa .L 43.0——43.01yrotsrednU

All 1 0.34 — — 0.34

Cochlospermum vitifolium (Willd.) Willd. ex Spreng. Overstory 1 1.66 — — 1.66

All 1 1.66 — — 1.66

Cocos nucifera .L 82.001.040.090.06yrotsrevO

All 7 0.08 0.04 0.10 0.28

continued
25

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Coffea arabica .L 21.0ÑÑ21.01yrotsrevO

Understory 29 0.04 0.01 0.08 0.34

All 36 0.06 0.02 0.10 0.38

C. liberica .nreiH xe lluB .W 35.032.001.052.05yrotsrevO

Understory 2 0.20 0.12 0.17 0.32

All 9 0.28 0.06 0.19 0.53

Cojoba arborea esoR & nottirB ).L( 41.0ÑÑ41.01yrotsrednU

All 1 0.14 Ñ Ñ 0.14

Colubrina arborescens .graS ).lliM( 60.0ÑÑ60.01yrotsrednU

All 1 0.06 Ñ Ñ 0.06

Comocladia dodonaea .brU ).L( 20.0ÑÑ20.01yrotsrevO

All 1 0.02 Ñ Ñ 0.02

Conocarpus erectus .L 49.066.074.074.02yrotsrevO

Understory 2 0.03 0.03 0.04 0.06

All 4 0.25 0.23 0.46 0.94

Cordia alliodora nekO ).vaP & ziuR( 61.133.090.054.021yrotsrevO

Understory 2 0.26 0.10 0.14 0.36

All 26 0.32 0.06 0.30 1.16

C. borinquensis brU . 26.0ÑÑ26.01yrotsrevO

Understory 3 0.11 0.06 0.10 0.22

All 5 0.26 0.10 0.22 0.51

C. sulcata .CD 82.183.001.004.041yrotsrevO

Understory 6 1.36 0.78 1.91 5.08

All 31 0.56 0.17 0.94 5.08

Crossopetalum rhacoma ztnarC 81.060.020.070.07yrotsrevO

All 7 0.07 0.02 0.06 0.18

Croton astroites .dnayrD 23.0ÑÑ23.01yrotsrednU

All 1 0.32 Ñ Ñ 0.32

C. poecilanthus .brU 42.021.070.051.03yrotsrevO

Understory 3 0.05 0.03 0.05 0.10

All 6 0.10 0.04 0.10 0.24

Cupania americana .L 00.182.021.076.06yrotsrevO

Understory 6 0.47 0.13 0.31 0.88

All 21 0.48 0.07 0.31 1.00

C. triquetra .hciR .A 63.032.061.002.02yrotsrevO

All 2 0.20 0.16 0.23 0.36

Cyathea arborea .mS ).L( 40.147.025.025.02yrotsrevO

Understory 3 0.15 0.12 0.22 0.40

All 9 0.18 0.12 0.35 1.04

continued
26

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Cyrilla racemifl ora .L 80.0ÑÑ80.01yrotsrevO

All 1 0.08 Ñ Ñ 0.08

Dacryodes excelsa lhaV 09.082.090.035.001yrotsrevO

Understory 2 0.27 0.09 0.13 0.36

All 19 0.46 0.07 0.28 1.08

Daphnopsis americana .tsnhoJ .R.J ).lliM( 86.0ÑÑ86.01yrotsrevO

Understory 1 0.28 Ñ Ñ 0.28

All 2 0.48 0.20 0.28 0.68

Delonix regia .faR ).kooH xe rejoB( 24.124.042.030.13yrotsrevO

All 3 1.03 0.24 0.42 1.42

Dendropanax arboreus (L.) Decne. & Planch. ex Britton Overstory 13 0.37 0.17 0.61 2.28

Understory 14 0.16 0.04 0.14 0.56

All 51 0.35 0.08 0.58 3.03

Ditta myricoides .besirG 21.010.010.011.02yrotsrednU

All 2 0.11 0.01 0.01 0.12

Drypetes alba .tioP 31.0ÑÑ31.01yrotsrednU

All 1 0.13 Ñ Ñ 0.13

D. glauca lhaV 61.0ÑÑ61.01yrotsrevO

Understory 2 0.14 0.02 0.03 0.16

All 3 0.15 0.01 0.02 0.16

Erithalis fruticosa .L 0.02yrotsrevO 80.030.020.06

Understory 1 0.07 Ñ Ñ 0.07

All 3 0.06 0.01 0.02 0.08

Erythrina berteriana .brU 46.083.072.073.02yrotsrevO

All 2 0.37 0.27 0.38 0.64

E. poeppigiana kooC .F.O ).plaW( 16.256.041.067.012yrotsrevO

Understory 7 0.20 0.07 0.19 0.52

All 34 0.73 0.12 0.71 2.61

Erythroxylum rotundifolium nanuL 61.050.020.090.06yrotsrednU

All 6 0.09 0.02 0.05 0.16

Eucalyptus robusta .mS 48.531.112.045.092yrotsrevO

All 29 0.54 0.21 1.13 5.84

Eugenia bifl ora .CD ).L( 02.060.030.021.04yrotsrevO

Understory 11 0.08 0.02 0.07 0.20

All 16 0.08 0.02 0.07 0.20

E. boqueronensis nottirB 25.0ÑÑ25.01yrotsrevO

All 1 0.52 Ñ Ñ 0.52

E. confusa .CD 41.070.050.090.02yrotsrednU

All 2 0.09 0.05 0.07 0.14

continued
27

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Eugenia domingensis greB 83.0——83.01yrotsrevO

All 1 0.38 — — 0.38

E. ligustrina .dlliW ).wS( 80.040.030.050.02yrotsrevO

Understory 1 0.00 — — 0.00

All 3 0.03 0.02 0.04 0.08

E. monticola .CD ).wS( 42.001.050.031.04yrotsrevO

Understory 17 0.09 0.02 0.10 0.38

All 24 0.12 0.02 0.11 0.38

E. pseudopsidium .qcaJ 66.031.090.075.02yrotsrevO

All 2 0.57 0.09 0.13 0.66

E. rhombea .brU & gurK )greB( 00.0——00.01yrotsrevO

Understory 6 0.06 0.01 0.03 0.10

All 7 0.05 0.01 0.03 0.10

E. xerophytica nottirB 21.070.050.070.02yrotsrevO

All 2 0.07 0.05 0.07 0.12

Euphorbia cotinifolia .L 00.0——00.01yrotsrevO

All 1 0.00 — — 0.00

Exostema caribaeum .tluhcS ).qcaJ( 81.050.020.090.09yrotsrevO

All 9 0.09 0.02 0.05 0.18

Exothea paniculata .kldaR ).ssuJ( 22.090.040.031.05yrotsrednU

All 8 0.12 0.03 0.10 0.25

Faramea occidentalis .hciR .A ).L( 40.000.000.040.02yrotsrednU

All 2 0.04 0.00 0.00 0.04

Ficus americana .lbuA 56.0——56.01yrotsrevO

All 1 0.65 — — 0.65

F. citrifolia .lliM 16.062.001.032.07yrotsrevO

Understory 3 0.49 0.43 0.74 1.35

All 16 0.58 0.21 0.83 3.05

F. trigonata .L 81.060.040.041.02yrotsrevO

All 2 0.14 0.04 0.06 0.18

Gliricidia sepium .plaW xe htnuK ).qcaJ( 25.082.061.002.03yrotsrevO

All 3 0.20 0.16 0.28 0.52

Guajacum offi cinale .L 83.002.041.042.02yrotsrevO

All 2 0.24 0.14 0.20 0.38

Guapira fragrans elttiL ).sruoC .muD( 28.012.040.002.062yrotsrevO

Understory 11 0.28 0.07 0.24 0.68

All 44 0.28 0.05 0.34 1.50

G. obtusata elttiL ).qcaJ( 01.060.040.060.02yrotsrevO

All 2 0.06 0.04 0.06 0.10

continued
28

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Guarea glabra lhaV 60.030.020.030.03yrotsrednU

All 4 0.02 0.01 0.03 0.06

G. guidonia remuelS ).L( 09.245.050.085.0701yrotsrevO

Understory 123 0.37 0.04 0.41 2.04

All 352 0.54 0.03 0.53 2.90

Guazuma ulmifolia .maL 08.154.011.065.061yrotsrevO

Understory 1 0.04 Ñ Ñ 0.04

All 21 0.53 0.09 0.42 1.80

Guettarda scabra .tneV ).L( 82.070.020.050.081yrotsrevO

Understory 15 0.02 0.01 0.02 0.06

All 47 0.03 0.01 0.05 0.28

Gyminda latifolia .brU ).wS( 80.0ÑÑ80.01yrotsrevO

Understory 4 0.09 0.03 0.06 0.16

All 7 0.08 0.02 0.05 0.16

Gymnanthes lucida .wS 01.040.010.040.031yrotsrevO

Understory 10 0.04 0.01 0.03 0.08

All 28 0.05 0.01 0.05 0.20

Henriettea macfadyenii nialA )anairT( 45.010.010.035.02yrotsrevO

All 2 0.53 0.01 0.01 0.54

H. squamulosum dduJ .S.W ).ngoC( 020.003.02yrotsrevO 23.030.

Understory 2 0.06 0.06 0.08 0.12

All 4 0.18 0.07 0.15 0.32

Hernandia sonora .L 20.0ÑÑ20.01yrotsrednU

All 1 0.02 Ñ Ñ 0.02

Hibiscus elatus .wS 60.0ÑÑ60.01yrotsrednU

All 1 0.06 Ñ Ñ 0.06

Hirtella rugosa .sreP xe .lliuhT 80.060.040.040.02yrotsrednU

All 2 0.04 0.04 0.06 0.08

Homalium racemosum .qcaJ 86.012.080.012.07yrotsrevO

Understory 2 0.08 0.06 0.08 0.14

All 10 0.21 0.07 0.21 0.68

Hymenaea courbaril .L 26.051.040.033.031yrotsrevO

Understory 9 0.20 0.07 0.21 0.60

All 26 0.36 0.05 0.24 0.91

Ilex nitida .mixaM )lhaV( 00.0ÑÑ00.01yrotsrednU

All 1 0.00 Ñ Ñ 0.00

Inga laurina .dlliW ).wS( 23.328.051.096.092yrotsrevO

Understory 10 0.32 0.14 0.43 1.44

All 70 0.50 0.07 0.60 3.32

continued
29

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Inga nobilis Willd. ssp. quaternata (Poepp. & Endl.) T.D. Penn. Overstory 1 0.13 Ñ Ñ 0.13

Understory 1 0.54 Ñ Ñ 0.54

All 2 0.34 0.21 0.29 0.54

I. vera .dlliW 45.235.060.024.007yrotsrevO

Understory 16 0.35 0.08 0.31 1.26

All 129 0.46 0.05 0.56 2.88

Ixora ferrea .htneB ).qcaJ( 41.153.052.098.02yrotsrevO

All 2 0.89 0.25 0.35 1.14

Krugiodendron ferreum .brU )lhaV( 82.021.060.011.04yrotsrevO

Understory 6 0.03 0.01 0.02 0.06

All 10 0.06 0.03 0.08 0.28

Laguncularia racemosa .ntreaG .F.C ).L( 00.0ÑÑ00.01yrotsrevO

Understory 6 0.15 0.02 0.06 0.24

All 7 0.13 0.03 0.08 0.24

Leucaena leucocephala tiW ed ).maL( 48.091.030.042.015yrotsrevO

Understory 13 0.10 0.02 0.07 0.20

All 64 0.21 0.02 0.18 0.84

Licaria brittoniana yrogerG & nellA 81.0ÑÑ81.01yrotsrednU

All 1 0.18 Ñ Ñ 0.18

L. parvifolia .mretsoK ).maL( revO 84.0ÑÑ84.01yrots

Understory 2 0.01 0.01 0.01 0.02

All 4 0.18 0.11 0.22 0.48

Lonchocarpus glaucifolius .brU 60.0ÑÑ60.01yrotsrevO

Understory 1 0.10 Ñ Ñ 0.10

All 2 0.08 0.02 0.03 0.10

L. heptaphyllus .CD ).rioP( 63.021.070.042.03yrotsrevO

All 3 0.24 0.07 0.12 0.36

Magnolia portoricensis olleB 09.035.083.035.02yrotsrevO

All 3 0.35 0.28 0.48 0.90

Mammea americana .L 11.175.004.017.02yrotsrevO

Understory 1 0.02 Ñ Ñ 0.02

All 3 0.48 0.33 0.56 1.11

Mangifera indica .L 82.284.090.083.013yrotsrevO

Understory 8 0.26 0.11 0.32 0.94

All 68 0.40 0.05 0.39 2.28

Manilkara bidentata vehC .A ).CD .A( 50.113.022.038.02yrotsrevO

All 2 0.83 0.22 0.31 1.05

Margaritaria nobilis .f .L 20.0ÑÑ20.01yrotsrevO

Understory 2 0.04 0.04 0.06 0.08

All 3 0.03 0.02 0.04 0.08

continued
30

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Matayba domingensis .kldaR ).CD( 61.040.020.021.03yrotsrevO

All 4 0.24 0.08 0.17 0.47

Maytenus ponceana nottirB 61.011.080.080.02yrotsrednU

All 2 0.08 0.08 0.11 0.16

Melicoccus bijugatus .qcaJ 61.173.031.036.08yrotsrevO

Understory 5 0.26 0.10 0.23 0.58

All 13 0.49 0.10 0.37 1.16

Meliosma herbertii efloR 22.080.030.031.06yrotsrevO

Understory 1 0.14 Ñ Ñ 0.14

All 8 0.14 0.02 0.07 0.22

Miconia impetiolaris .CD xe noD .D ).wS( 94.0ÑÑ94.01yrotsrevO

Understory 1 0.08 Ñ Ñ 0.08

All 2 0.29 0.21 0.29 0.49

M. laevigata noD .D ).L( 86.081.080.093.05yrotsrevO

Understory 2 0.34 0.14 0.20 0.48

All 7 0.38 0.06 0.17 0.68

M. prasina .CD ).wS( 47.022.021.025.03yrotsrevO

Understory 6 0.15 0.09 0.21 0.52

All 16 0.22 0.06 0.26 0.74

M. pycnoneura .brU Ñ21.01yrotsrednU 21.0Ñ

All 1 0.12 Ñ Ñ 0.12

M. subcorymbosa nottirB 33.0ÑÑ33.01yrotsrevO

All 1 0.33 Ñ Ñ 0.33

M. tetrandra noD .D ).wS( 21.010.010.011.02yrotsrevO

Understory 3 0.51 0.10 0.17 0.68

All 5 0.35 0.11 0.25 0.68

Micropholis garciniifolia erreiP 27.002.050.021.031yrotsrevO

All 17 0.28 0.08 0.35 0.96

M. guyanensis erreiP ).CD .A( 64.031.030.081.012yrotsrevO

Understory 4 0.13 0.06 0.11 0.28

All 33 0.16 0.02 0.12 0.46

Myrcia citrifolia .brU ).lbuA( 01.0ÑÑ01.01yrotsrevO

All 1 0.10 Ñ Ñ 0.10

M. defl exa .CD ).rioP( 13.0ÑÑ13.01yrotsrevO

Understory 3 0.15 0.05 0.09 0.24

All 7 0.16 0.05 0.12 0.31

M. fallax .CD ).hciR( 40.010.010.030.02yrotsrednU

All 2 0.03 0.01 0.01 0.04

continued
31

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Myrcia splendens .CD ).wS( 45.042.041.082.03yrotsrevO

Understory 13 0.21 0.07 0.24 0.62

All 29 0.20 0.04 0.19 0.62

Myrsine coriacea (Sw.) R. Br. ex Roem. & J.A. Schult. Overstory 4 0.47 0.04 0.09 0.54

Understory 2 0.24 0.08 0.11 0.32

All 7 0.36 0.06 0.16 0.54

M. cubana .CD .A 25.021.070.034.03yrotsrevO

Understory 1 0.02 Ñ Ñ 0.02

All 5 0.33 0.09 0.20 0.52

Nectandra coriacea .besirG ).wS( 22.050.030.051.04yrotsrevO

Understory 4 0.22 0.09 0.18 0.40

All 9 0.20 0.05 0.14 0.40

N. hihua rewhoR ).vaP & ziuR( 05.1ÑÑ05.11yrotsrevO

All 1 1.50 Ñ Ñ 1.50

N. turbacensis seeN )htnuK( 37.000.000.037.03yrotsrevO

Understory 2 0.08 0.02 0.03 0.10

All 6 0.53 0.14 0.35 0.82

Neea buxifolia lremieH ).f .kooH( 20.0ÑÑ20.01yrotsrednU

All 1 0.02 Ñ Ñ 0.02

Neolaugeria resinosa noslociN )lhaV( srevO 07.062.070.003.031yrot

Understory 10 0.21 0.06 0.20 0.74

All 30 0.28 0.04 0.23 0.74

Ocotea fl oribunda zeM ).wS( 43.040.030.013.02yrotsrevO

All 2 0.31 0.03 0.04 0.34

O. leucoxylon .ssenaL eD ).wS( 66.002.050.072.081yrotsrevO

Understory 22 0.21 0.06 0.27 0.78

All 61 0.27 0.04 0.30 1.69

O. moschata zeM ).nsieM xe .vaP( 25.042.071.053.02yrotsrevO

All 2 0.35 0.17 0.24 0.52

O. wrightii zeM ).nsieM( 85.0ÑÑ85.01yrotsrevO

All 1 0.58 Ñ Ñ 0.58

Ormosia krugii .brU 66.002.060.032.011yrotsrevO

Understory 4 0.05 0.03 0.05 0.12

All 18 0.21 0.04 0.16 0.66

Ouratea littoralis .brU 80.0ÑÑ80.01yrotsrednU

All 1 0.08 Ñ Ñ 0.08

Palicourea croceoides .maH 11.040.020.070.03yrotsrevO

Understory 2 0.11 0.03 0.04 0.13

All 5 0.08 0.02 0.04 0.13

continued
32

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Parathesis crenulata .f .kooH ).tneV( 85.0ÑÑ85.01yrotsrevO

All 1 0.58 Ñ Ñ 0.58

Peltophorum pterocarpum (DC.) Backer ex K. Heyne Overstory 5 0.64 0.04 0.09 0.72

All 5 0.64 0.04 0.09 0.72

Persea americana .lliM 08.126.052.036.06yrotsrevO

Understory 3 1.26 0.52 0.90 2.28

All 11 0.70 0.22 0.73 2.28

Petitia domingensis .qcaJ 82.080.040.002.03yrotsrevO

Understory 2 0.04 0.02 0.03 0.06

All 7 0.19 0.05 0.13 0.37

Picramnia pentandra .wS 80.0ÑÑ80.01yrotsrednU

All 1 0.08 Ñ Ñ 0.08

Pictetia aculeata .brU )lhaV( 61.060.020.060.031yrotsrevO

Understory 3 0.09 0.05 0.08 0.16

All 16 0.06 0.02 0.06 0.16

Pilosocereus royenii yelwoR & selyB ).L( 26.024.003.023.02yrotsrevO

All 2 0.32 0.30 0.42 0.62

Pimenta racemosa erooM .W.J ).lliM( 66.022.080.082.07yrotsrevO

All 7 0.28 0.08 0.22 0.66

Piper aduncum .L 51.0ÑÑ51.01yrotsrednU

All 1 0.15 Ñ Ñ 0.15

Pisonia albida .ldnatS xe nottirB )lremieH( 60.010.010.050.02yrotsrevO

Understory 3 0.25 0.16 0.27 0.56

All 5 0.17 0.10 0.22 0.56

P. subcordata .wS 41.0ÑÑ41.01yrotsrednU

All 1 0.14 Ñ Ñ 0.14

Pithecellobium dulce .htneB ).bxoR( 47.248.083.046.15yrotsrevO

All 5 1.64 0.38 0.84 2.74

P. unguis-cati .htneB ).L( 47.555.241.181.15yrotsrevO

All 5 1.18 1.14 2.55 5.74

Plumeria obtusa .L 42.040.020.012.03yrotsrevO

All 3 0.21 0.02 0.04 0.24

Podocarpus coriaceus .hciR 50.0ÑÑ50.01yrotsrevO

All 1 0.05 Ñ Ñ 0.05

Poitea fl orida nivaL )lhaV( 40.0ÑÑ40.01yrotsrevO

All 1 0.04 Ñ Ñ 0.04

Pouteria multifl ora A( amyE ).CD . 00.390.114.055.07yrotsrevO

Understory 2 0.54 0.00 0.00 0.54

All 9 0.55 0.31 0.94 3.00

continued
33

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Pouteria sapota (Jacq.) H.E. Moore & Stearn Overstory 2 0.35 0.11 0.16 0.46

All 2 0.35 0.11 0.16 0.46

Prestoea acuminata (Willd.) H.E. Moore var. 

montana (Graham) A. Hend. & G. Galeano Overstory 41 0.04 0.01 0.06 0.20

Understory 47 0.17 0.03 0.21 1.00

All 110 0.10 0.02 0.16 1.00

Prosopis pallida (Humb. & Bonpl. ex Willd.) Kunth Overstory 3 0.11 0.11 0.18 0.32

All 3 0.11 0.11 0.18 0.32

Prunus myrtifolia .brU ).L( 90.0ÑÑ90.01yrotsrednU

All 1 0.09 Ñ Ñ 0.09

Pseudolmedia spuria .besirG ).wS( 81.0ÑÑ81.01yrotsrevO

All 1 0.18 Ñ Ñ 0.18

Psidium amplexicaule .sreP 61.060.030.070.05yrotsrednU

All 5 0.07 0.03 0.06 0.16

P. guajava .L 13.021.050.070.06yrotsrevO

Understory 5 0.02 0.01 0.03 0.06

All 14 0.05 0.02 0.09 0.31

Psychotria berteriana .CD 40.0ÑÑ40.01yrotsrednU

All 1 0.04 Ñ Ñ 0.04

Psychotria brachiata .wS 20.010.010.010.02yrotsrednU

All 2 0.01 0.01 0.01 0.02

Quararibea turbinata .rioP ).wS( 04.031.050.051.07yrotsrednU

All 8 0.15 0.04 0.12 0.40

Randia aculeata .L 62.080.040.071.03yrotsrevO

Understory 2 0.16 0.08 0.11 0.24

All 5 0.17 0.03 0.08 0.26

Rhizophora mangle .L 00.000.000.000.02yrotsrevO

Understory 1 0.00 Ñ Ñ 0.00

All 3 0.00 0.00 0.00 0.00

Roystonea borinquena kooC .F.O 82.154.011.062.061yrotsrevO

Understory 3 2.41 1.05 1.81 4.00

All 27 0.46 0.18 0.94 4.00

Sagraea umbrosa .CD ).wS( 82.080.030.002.05yrotsrevO

Understory 1 0.45 Ñ Ñ 0.45

All 6 0.24 0.05 0.12 0.45

Samanea saman .rreM ).qcaJ( 03.2ÑÑ03.21yrotsrevO

All 1 2.30 Ñ Ñ 2.30

Sapindus saponaria .L 52.0ÑÑ52.01yrotsrevO

All 1 0.75 Ñ Ñ 0.75

continued
34

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Sapium laurocerasus .fseD 43.0ÑÑ43.01yrotsrevO

Understory 1 0.88 Ñ Ñ 0.88

All 3 0.50 0.19 0.33 0.88

Savia sessilifl ora .dlliW ).wS( 01.060.040.060.02yrotsrednU

All 2 0.06 0.04 0.06 0.10

Scheffl era morototonii (Aubl.) Maguire, Steyerm. & Frodin Overstory 23 0.33 0.05 0.25 0.88

Understory 7 0.55 0.22 0.59 1.70

All 53 0.42 0.04 0.32 1.70

Schoepfi a obovata thgirW .C 00.0ÑÑ00.01yrotsrevO

All 1 0.00 Ñ Ñ 0.00

Senna siamea ybenraB & niwrI ).maL( 20.207.072.066.07yrotsrevO

Understory 2 0.98 0.50 0.71 1.48

All 9 0.64 0.20 0.61 2.02

Sideroxylon cubense .nneP .D.T ).besirG( 20.0ÑÑ20.01yrotsrevO

Understory 1 0.11 Ñ Ñ 0.11

All 3 0.15 0.09 0.16 0.33

S. salicifolium .maL ).L( 14.041.050.002.07yrotsrevO

Understory 2 0.08 0.04 0.06 0.12

All 11 0.22 0.05 0.16 0.57

Sloanea berteriana .CD xe ysiohC otsrevO 02.050.020.031.04yr

Understory 5 0.05 0.02 0.04 0.10

All 13 0.17 0.05 0.16 0.56

Solanum rugosum lanuD 81.031.090.090.02yrotsrednU

All 2 0.09 0.09 0.13 0.18

Spathodea campanulata .vuaeB .P 67.376.050.028.0381yrotsrevO

Understory 148 0.33 0.03 0.39 2.14

All 371 0.64 0.03 0.63 3.76

Spondias dulcis nosnikraP 74.033.042.042.02yrotsrevO

All 2 0.24 0.24 0.33 0.47

S. mombin .L 81.163.001.016.031yrotsrevO

Understory 3 0.35 0.07 0.12 0.43

All 21 0.48 0.08 0.37 1.18

Swietenia macrophylla gniK 27.043.002.033.03yrotsrevO

All 3 0.33 0.20 0.34 0.72

S. mahagoni .qcaJ ).L( 84.061.050.091.001yrotsrevO

Understory 1 0.16 Ñ Ñ 0.16

All 11 0.19 0.05 0.15 0.48

Symplocos martinicensis .qcaJ 46.022.011.093.04yrotsrevO

All 4 0.36 0.12 0.24 0.64

continued
35

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Syzygium jambos notslA ).L( 45.041.030.021.072yrotsrevO

Understory 56 0.13 0.03 0.19 0.90

All 120 0.14 0.02 0.20 0.90

Tabebuia heterophylla nottirB ).CD( 64.153.040.043.077yrotsrevO

Understory 27 0.18 0.04 0.21 0.78

All 142 0.29 0.03 0.30 1.46

T. rigida .brU 00.0ÑÑ00.01yrotsrevO

All 1 0.00 Ñ Ñ 0.00

Tamarindus indica .L 80.1ÑÑ80.11yrotsrevO

All 2 1.30 0.22 0.31 1.52

Terminalia catappa .L 38.003.021.045.06yrotsrevO

Understory 1 0.42 Ñ Ñ 0.42

All 8 0.53 0.09 0.25 0.83

Tetragastris balsamifera nekO ).wS( 06.082.041.013.04yrotsrevO

Understory 3 0.46 0.17 0.29 0.80

All 7 0.37 0.10 0.27 0.80

Tetrazygia elaeagnoides ).wS( .CD 69.072.080.012.011yrotsrevO

Understory 5 0.18 0.04 0.10 0.34

All 22 0.16 0.04 0.20 0.96

Thespesia grandifl ora .CD 41.193.061.083.06yrotsrevO

Understory 3 0.11 0.05 0.08 0.20

All 14 0.24 0.07 0.28 1.14

Thouinia striata .kldaR 48.062.080.071.001yrotsrevO

Understory 5 0.19 0.09 0.19 0.50

All 17 0.20 0.05 0.23 0.84

T. striata Radlk. var. portoricensis

nialA & avatoV ).kldaR( 01.040.010.030.07yrotsrevO

Understory 5 0.02 0.02 0.03 0.08

All 12 0.03 0.01 0.04 0.10

Trema micrantha emulB ).L( 45.0ÑÑ45.01yrotsrevO

Understory 1 0.06 Ñ Ñ 0.06

All 2 0.30 0.24 0.34 0.54

Trichilia hirta .L 47.043.042.005.02yrotsrevO

Understory 1 0.00 Ñ Ñ 0.00

All 3 0.33 0.22 0.38 0.74

T. pallida .wS 44.031.040.001.011yrotsrednU

All 21 0.10 0.03 0.12 0.44

Turpinia occidentalis .G ).wS( noD 87.072.090.062.001yrotsrevO

Understory 1 0.01 Ñ Ñ 0.01

All 11 0.27 0.09 0.29 0.78

continued
36

Table A.2—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) by species and crown position, with 

number of trees measured, standard error of the mean, standard deviation of the mean, and maximum observed PAI 

increase from Puerto Rico forest inventory data (continued)

Speciesa

Crown 

positionb N

Periodic annual increments

Mean SE SD Max

Urera baccifera .hciduaG ).L( 40.0ÑÑ40.01yrotsrednU

All 1 0.04 Ñ Ñ 0.04

Vitex divaricata .wS 67.043.071.013.04yrotsrevO

Understory 1 0.12 Ñ Ñ 0.12

All 6 0.23 0.12 0.29 0.76

Xylosma buxifolia yarG .A 50.0ÑÑ50.01yrotsrednU

All 1 0.05 Ñ Ñ 0.05

X. pachyphylla .brU ).brU & gurK( 41.0ÑÑ41.01yrotsrevO

All 1 0.14 Ñ Ñ 0.14

Zanthoxylum martinicense .CD ).maL( 26.275.021.075.032yrotsrevO

Understory 8 0.25 0.07 0.18 0.48

All 40 0.40 0.08 0.49 2.62

Z. monophyllum nosliW .P ).maL( 84.071.021.063.02yrotsrevO

Understory 2 0.03 0.01 0.01 0.04

All 4 0.20 0.11 0.21 0.48

— = insuffi cient sample; N = number of trees measured; SE = standard error of the mean; SD = standard deviation of the mean; Max = 

maximum observed.

a

 USDA Natural Resources Conservation Service (2006).

b

 Trees in the overstory crown positon have crowns that are exposed to direct sunlight from above and some light from the side, while 

understory trees receive little or no direct sunlight.
37

Table A.3—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) for commercial species 

by tree class with number of trees measured, standard error of the mean, standard deviation of the mean, 

and maximum observed PAI increase from Puerto Rico forest inventory data

Speciesa Tree classb N

Periodic annual increments

Mean SE SD Max

Albizia procera (Roxb.) Benth. Growing stock 10 0.99 0.24 0.75 2.22

Cull 20 0.48 0.11 0.48 2.16

Andira inermis (W. Wright) Kunth ex DC. Growing stock 54 0.32 0.04 0.31 1.26

Cull 115 0.19 0.02 0.20 1.05

Avicennia germinans (L.) L. Growing stock 2 0.60 0.60 0.85 1.20

Cull 13 0.16 0.08 0.30 1.12

Buchenavia tetraphylla (Aubl.) Howard Growing stock 5 0.37 0.10 0.23 0.63

Cull 2 1.23 0.23 0.32 1.45

Bursera simaruba (L.) Sarg. Growing stock 9 0.54 0.26 0.78 2.48

Cull 58 0.30 0.04 0.30 1.44

Byrsonima spicata (Cav.) Kunth Growing stock 7 0.46 0.13 0.34 1.01

Cull 12 0.42 0.13 0.44 1.56

Calophyllum antillanum Britton Growing stock 27 0.42 0.06 0.33 1.38

Cull 14 0.41 0.08 0.32 0.88

Cecropia schreberiana Miq. Growing stock 86 0.73 0.07 0.65 2.96

Cull 51 0.93 0.13 0.94 4.30

Cedrela odorata .L kcots gniworG 45.071.060.022.07

Cull 4 0.12 0.10 0.20 0.41

Conocarpus erectus L. Cull 4 0.25 0.23 0.46 0.94

Cordia alliodora (Ruiz & Pav.) Oken Growing stock 17 0.34 0.08 0.32 1.16

Cull 9 0.28 0.09 0.26 0.72

Cupania americana .L kcots gniworG 38.033.051.034.05

Cull 16 0.50 0.08 0.31 1.00

Dacryodes excelsa Vahl Growing stock 14 0.44 0.08 0.28 1.08

Cull 5 0.50 0.14 0.32 0.90

Eucalyptus robusta Sm. Growing stock 16 0.28 0.08 0.33 0.96

Cull 13 0.87 0.45 1.62 5.84

Ficus citrifolia .lliM kcots gniworG 5.03 55.188.015.04

Cull 13 0.58 0.24 0.85 3.05

Guapira fragrans (Dum. Cours.) Little Growing stock 6 0.20 0.06 0.15 0.44

Cull 38 0.29 0.06 0.36 1.50

Guarea guidonia (L.) Sleumer Growing stock 104 0.69 0.05 0.55 2.90

Cull 248 0.48 0.03 0.51 2.85

Guazuma ulmifolia Lam. Growing stock 2 0.38 0.26 0.36 0.63

Cull 19 0.54 0.10 0.43 1.80

Gymnanthes lucida Sw. Growing stock 3 0.13 0.04 0.07 0.20

Cull 25 0.04 0.01 0.03 0.10

continued
38

Table A.3—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) for commercial species 

by tree class with number of trees measured, standard error of the mean, standard deviation of the mean, 

and maximum observed PAI increase from Puerto Rico forest inventory data (continued)

Speciesa Tree classb N

Periodic annual increments

Mean SE SD Max

Hibiscus elatus .wS lluC 60.0ÑÑ60.01

Homalium racemosum Jacq. Growing stock 1 0.68 Ñ Ñ 0.68

Cull 9 0.16 0.04 0.13 0.49

Hymenaea courbaril L. Growing stock 8 0.41 0.11 0.32 0.91

Cull 18 0.33 0.05 0.20 0.75

Inga laurina (Sw.) Willd. Growing stock 31 0.63 0.13 0.74 3.32

Cull 39 0.40 0.07 0.44 2.22

Inga vera .dlliW 09.134.060.024.045kcots gniworG

Cull 75 0.49 0.07 0.65 2.88

Laguncularia racemosa (L.) C.F. Gaertn. Cull 7 0.13 0.03 0.08 0.24

Mangifera indica .L kcots gniworG 98.022.070.073.09

Cull 59 0.40 0.05 0.41 2.28

Manilkara bidentata (A. DC.) A. Chev Growing stock 2 0.83 0.22 0.31 1.05

Neolaugeria resinosa (Vahl) Nicolson Growing stock 12 0.46 0.07 0.23 0.74

Cull 18 0.16 0.03 0.13 0.60

Ocotea fl oribunda (Sw.) Mez Cull 2 0.31 0.03 0.04 0.34

O. leucoxylon (Sw.) De Laness. Growing stock 20 0.27 0.05 0.23 0.78

Cull 41 0.27 0.05 0.33 1.69

Ormosia krugii .brU 66.071.050.042.011kcots gniworG

Cull 7 0.16 0.05 0.14 0.38

Petitia domingensis Jacq. Growing stock 3 0.20 0.10 0.18 0.37

Cull 4 0.18 0.05 0.11 0.30

Pimenta racemosa (Mill.) J.W. Moore Cull 7 0.28 0.08 0.22 0.66

Pouteria multifl ora (A. DC.) Eyma Growing stock 7 0.69 0.39 1.04 3.00

Cull 2 0.05 0.05 0.07 0.10

Senna siamea (Lam.) Irwin & Barneby Growing stock 3 0.68 0.24 0.41 1.12

Cull 6 0.61 0.30 0.73 2.02

Spondias mombin .L kcots gniworG 74.002.080.091.06

Cull 15 0.59 0.09 0.36 1.18

Swietenia macrophylla King Cull 3 0.33 0.20 0.34 0.72

S.mahagoni .qcaJ ).L( kcots gniworG 03.050.030.072.03

Cull 8 0.16 0.06 0.17 0.48

Terminalia catappa .L kcots gniworG 38.092.021.035.06

Cull 2 0.51 0.08 0.12 0.59

continued
39

Table A.3—Diameter at breast height (1.4 m) periodic annual increments (PAI, cm) for commercial species 

by tree class with number of trees measured, standard error of the mean, standard deviation of the mean, 

and maximum observed PAI increase from Puerto Rico forest inventory data (continued)

Speciesa Tree classb N

Periodic annual increments

Mean SE SD Max

Thespesia grandifl ora DC. Growing stock 3 0.11 0.01 0.03 0.13

Cull 11 0.28 0.09 0.31 1.14

Vitex divaricata .wS kcots gniworG 63.002.021.031.03

Cull 3 0.33 0.22 0.38 0.76

Zanthoxylum martinicense (Lam.) DC. Growing stock 11 0.34 0.11 0.37 1.04

Cull 29 0.41 0.10 0.53 2.62

All commerical species combined Growing stock 570 0.51 0.02 0.52 3.32

Cull 1,045 0.40 0.02 0.52 5.84

— = insuffi cient sample; N = number of trees measured; SE = standard error of the mean; SD = standard deviation of the 

mean; Max = maximum observed.

a USDA Natural Resources Conservation Service (2006).

b To meet requirements for the growing-stock tree class, trees must have one-third or more of the gross board-foot volume in 

the entire saw-log section with commercial logs meeting grade, soundness, and size requirements or the potential to do so for 

poletimber-sized trees. A growing-stock tree must have one 3.5-m or two 2.5-m logs, now or prospectively, for live poletimbersized trees to qualify as growing stock. Trees that do not meet these requirements are considered cull.

Brandeis, Thomas J. 2009. Diameter growth of subtropical trees in Puerto Rico. Res. Pap. 

 SRS–47. Asheville, NC: U.S. Department of Agriculture Forest Service, Southern 

 Research Station. 39 p.

Puerto Rico’s forests consist of young, secondary stands still recovering from a long history 

of island-wide deforestation that largely abated in the mid-20th century. Limited knowledge 

about growth rates of subtropical tree species in these forests makes it difficult to accurately 

predict forest yield, biomass accumulation, and carbon sequestration. This study presents 

mean annual increases (periodic annual increment) in tree diameter at breast height among 

trees measured by the forest inventories of Puerto Rico; this information is given for each 

forested life zone, by species, then by species and crown class, and by crown position class. 

Additionally, the study presents mean periodic annual increment values calculated for 

commercial species by tree class (growing stock and cull). From 1980 to 2008, mean diameter 

at breast height periodic annual increment was 0.35 cm/year for 4,026 trees remeasured by the 

forest inventory; growth rate averaged 0.20 cm/year in subtropical dry forests, 0.37 cm/year 

in subtropical moist forests, 0.36 cm/year in subtropical wet/rain forests, and 0.20 cm/year in 

lower montane forests.

Keywords: Caribbean, crown position, FIA, periodic annual increment, secondary forest. 
The Forest Service, United States Department of 

Agriculture (USDA), is dedicated to the principle of 

multiple use management of the Nation’s forest 

resources for sustained yields of wood, water, forage, 

wildlife, and recreation. Through forestry research, cooperation 

with the States and private forest owners, and management of the 

National Forests and National Grasslands, it strives—as directed 

by Congress—to provide increasingly greater service to a growing 

Nation.

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