United States

Department of

Agriculture

Forest Service

International Institute

of Tropical Forestry

ANNUAL LETTER

1999-2000
November 2001

International Institute of Tropical Forestry

Call Box 25000

Río Piedras, PR 00928-2500
i

File Code: 1630

Date: November 30, 2001

Dear Colleague:

With this Annual Letter we symbolically close our reporting activities for the millennium and poise

ourselves for the 21st century. Notably, this Annual Letter is the first one not to be printed. Instead, it will

be posted on our WEB page. We hope that by using the Internet we will be able to deliver our products

more quickly and effectively. We will do our best to reach those colleagues without access to the Internet

and ask that you help us as well by letting others know of this experiment, which we hope to monitor

carefully. As usual, I ask that you exchange your reprints and research findings. We value these

exchanges and share the information with as many people and institutions as we can.

Sincerely,

ARIAL E. LUGO

Director

United States Forest International Institute P.O. Box 25000

Department of Service of Tropical Forestry Río Piedras, PR 00928

Agriculture Tel: (787) 766-5335 Fax: (787) 766-6263

Caring for the Land and Serving People Printed on Recycled Paper
ii
iii

ANNUAL LETTER

1999–2000

INTERNATIONAL INSTITUTE OF TROPICAL FORESTRY

RÍO PIEDRAS, PUERTO RICO

U.S. DEPARTMENT OF AGRICULTURE

FOREST SERVICE
iv
v

INTERNATIONAL INSTITUTE OF TROPICAL FORESTRY ANNUAL LETTER

1999–2000

Contents

Page

Long-Term Avian Research ...................................................................................................................... 1

Wayne J. Arendt

Big-leaf Mahogany Growth Study: Year 2 Progress Report

October 1, 1999 to September 30, 2000 ................................................................................................... 3

Julio C. Figueroa Colon

I Still Love the Trees, But . . . ................................................................................................................... 5

John K. Francis

Landscape Ecology and Remote Sensing Program ................................................................................. 7

Eileen Helmer

Ecological Research .................................................................................................................................. 8

Ariel E. Lugo

Forest Management and Rehabilitation Research................................................................................ 17

John A. Parrotta

An Evaluation of Direct Seeding for Reforestation of Degraded Lands in

Central São Paulo State, Brazil............................................................................................................... 18

Vera Lex Engel and John A. Parrotta

Ecosystem Research ............................................................................................................................... 19

F.N. Scatena

Continued Studies on Mahogany and Long-term Plots at the

Luquillo Experimental Forest.................................................................................................................. 21

Sheila Ward

Forest Research ...................................................................................................................................... 23

Peter L. Weaver

Wildlife Research .................................................................................................................................... 25

Joseph M. Wunderle, Jr.

State and Private Forestry Accomplishments in Puerto Rico ............................................................... 28

Robin Morgan

State and Private Forestry Accomplishments in the U.S. VI................................................................. 31

Robin Morgan

International Cooperation Highlights ..................................................................................................... 34

Carleen Yocum

Appendix 1: Annual Letter 1999–2000 Publications List ...................................................................... 36
vi
1

PUERTO RICAN PARROT RESTORATION

AND RELATED RESEARCH

With the publication of a parrot conservation and

management article (Arendt 2000), there arose a

renewed interest in developing a safe method for

controlling levels of ectoparasitic botflies (Philornis sp.)

in birds’ nests. Various avenues of cooperative research

are currently being explored involving entomologists and

wildlife researchers at two stateside universities. In lieu

of traditional pesticides with their many well known

disadvantages, researchers are working fervently to

develop all-organic biocides, namely volatile plant

secondary compounds, which they hope eventually can

be safely introduced into nests of the critically

endangered Puerto Rican Parrot and other forest cavitynesting birds to deter botfly ectoparasitism. Also, as a

result of more than 20 years of research into the

reproductive ecology of the Pearly-eyed Thrasher, the

parrot’s prime nest predator, related cooperative

research has begun with Drs. Steven Beissinger and

Mark Cook, professor and postdoctoral candidate,

respectively, of the University of California at Berkeley.

The cooperative research involves a series of egg

experiments to determine the onset of incubation in the

Pearly-eyed Thrasher as well as various ancillary

parameters. Following are some brief excerpts

summarizing our preliminary findings and presented by

the UC-B cooperators:

Egg Hatching Summary

In total, 260 eggs were laid during the 2000 breeding

season in 43 nest boxes, and 114 (43.8 percent)

hatched. Of these 260 eggs, 41 failed to hatch due to

factors unrelated to experimental treatment and were

excluded from analyses. The remaining 219 eggs

comprised 16 controls, 37 unmanipulated and 166

treatment eggs.

Hatching Success

Hatching success of control eggs (14 of 16 hatched or

87.5 percent) and unmanipulated eggs (32 of 37 eggs

or 86.4 percent) was similar suggesting that moving eggs

between sites had little affect on hatchability (G-test of

independence with Yates’s correction: G=0.66, df=1,

P>0.50). Although the transportation period was longer

on average for treatment eggs (151.9 ±113.5 sd) than

for control eggs (101.38 ±64.4 sd), the difference was

not significant (Mann-Whitney test, U16,153=952,

P=0.14). Thus, potential effects on hatchability of the

amount of time an egg was transported were unlikely to

confound comparisons between control and treatment

groups in this study.

Hatching success (40.9 percent, n=166 treatment eggs)

markedly declined with increasing exposure period but

differed little among holding box sites. Hatchability of

experimental eggs after 1 day of exposure (83.3 percent

hatched) was not significantly lower than control eggs

(G-test of independence: G=0.21, n=101, df=1,

P>0.50), but was significantly reduced after exposure

of 3 days (47.6 percent hatched, G=80.05, n=95 df=1,

P<0.001), 5 days (22 percent hatched, G=39.74, n=89

df=1, P<0.001), and 7 days (G=87.54, n=93, df=1,

P<0.001). No 7-day treatments hatched.

Preincubation Development

Initiation of incubation relative to clutch completion in

the thrasher is variable. Full incubation proceeds upon

laying of the penultimate egg in some broods, whereas

in others, full incubation starts upon clutch completion.

Thus, the onset of incubation is variable with respect to

all but last-laid eggs.

Mean incubation periods of all treatment eggs (14.7

days ±0.6 sd, n=64) were slightly, but significantly

(t75 =2.08 P=0.041), longer than those of the pooled

control/unmanipulated eggs (14.2 days ±1.1sd, n=13).

LONG-TERM AVIAN RESEARCH

Wayne J. Arendt

Research Wildlife Biologist
2

Therefore, contrary to prediction, experimental

treatment appeared to increase embryo development

time rather than decrease it. However, caution must be

exercised when interpreting this result since the mean

difference between the two groups (0.5 days) was less

than the sampling period unit (1 day).

Timing of Embryo Mortality

Of the pooled set of 53 control and unmanipulated eggs,

only 7 eggs failed to hatch. Each one was assessed for

stage of embryo mortality. Control-egg embryos mostly

died after no development (57 percent; embryo age 0

to 1d) or little development (43 percent; embryo age 2

to 4d), and none reached an advanced level of

development (embryo age 9 to 13d). Stage of embryo

mortality was also recorded for each of the 98 treatment

eggs that failed to hatch. The pattern of embryo mortality

in treatment eggs was similar to that of control eggs.

The majority of eggs showed no (63 percent) or little

development (16 percent), with only 14 percent and 4

percent reaching moderate (embryo age 5 to 8d) and

advanced levels of development, respectively.

LONG-TERM STUDIES IN THE

GUANICA DRY FOREST

The annual mist netting and banding of Guánica’s forest

birds took place in January 2000, marking the 28th year

of this long-term study of forest-bird populations. We

continue to investigate the various environmental effects

on population dynamics of resident and migratory birds

(Dugger and others, in press) in an effort to develop

appropriate management and conservation strategies to

ensure sustainable avian populations for future

generations (Faaborg and others, in press). By

continuing our long-term research in Guánica, we have

been able to substantiate an earlier preliminary finding;

i.e., that populations of Nearctic-neotropical migrants

that winter in the forest, although fluctuating widely over

the years, have declined slowly but steadily since the

mid-1970's. It is encouraging, however, that the

dominant wintering migrants (black-and-white warbler

Mniotilta varia, American redstart Setophaga

ruticilla, and ovenbird Seiurus aurocapillus, show

stable or only slightly declining populations.

Modeling Avian Survival Constrained by Rainfall

Most recently, we have been modeling the survival of

Guánica’s forest birds by factoring in the constraints of

rainfall. As found previously when we correlated avian

population fluctuations with total and first 6-month

precipitation levels after 9 and then 15 yr (Faaborg

1982, Faaborg and Arendt 1992, Faaborg and others

1984), over a 26-yr period, annual survival of individuals

within and among all foraging guilds, but especially

frugivores, was greatly constrained by total levels of

precipitation, and mainly during the first 6 months of

the year (Dugger and others, in press).

LITERATURE CITED

Arendt, W.J. 2000. Impact of nest predators,

competitors, and ectoparasites on Pearly-eyed

Thrashers, with comments on the potential implications

for Puerto Rican Parrot recovery. Ornitología

Neotropical. 11: 13–63.

Dugger, K.M.; Faaborg, J.; Arendt, W.J. (In press).

Rainfall correlates of bird populations and survival rates

in a Puerto Rican dry forest (Bird Populations). [Place

of publication unknown]: [Publisher unknown].

Faaborg, J. 1982. Avian population fluctuations during

drought conditions in Puerto Rico. Wilson Bull. 94: 20–

30.

Faaborg, J.; Arendt, W.J. 1992. Rainfall correlates of

bird population fluctuations in a Puerto Rican dry forest:

a 15-yr study. Ornitología Caribeña. 3: 10–19.

Faaborg, J.; Arendt, W.J.; Dugger, K.M. (In press).

The Guánica, Puerto Rico, bird monitoring project (Bird

Populations).[Place of publication unknown]: [Publisher

unknown].

Faaborg, J.; Arendt, W.J.; Kaiser, M.S. 1984. Rainfall

correlations of bird population fluctuations in a Puerto

Rican dry forest: a 9-year study. Wilson Bull. 96: 575–

593.
3

Summary

The original objectives of this study remain: (1) to

establish a rangewide network of permanent study plots,

which will allow for the description, comparison, and

monitoring of natural big-leaf mahogany populations

across the entire environmental gradients of climate and

soils in which it occurs; and, (2) to produce much needed

growth data from a variety of sites, which can be used

by conservationists and managers alike.

Accomplishments for this past year and findings to date

are summarized in this report along with a proposed

work schedule for the coming year.

Protocols

Three different sites, each with varied climatic,

topographical, and logistical characteristics, have been

completed, and the study protocols designed for this

study have proved both proficient in the quality of data

collected, as well as efficient in the time and ease of

implementation. Proficiency in the methodology now

allows us, a team of 2 (with 2 to 3 field hands) to locate,

grid, and GPS a 25-ha study block into 25 individual

1-ha plots; locate and GPS an additional 25 individual

0.04-ha subplots; and then measure and GPS all tallied

trees, all in 3 to 4 weeks.

Study Sites

Two new sites were selected, established, and measured

during this past year: Quiringuicharo-Chiapas-Mexico

and Capixaba-Acre-Brazil.

Quiringuicharo—This study site was located in natural

wet tropical forests in the municipality of Benemerito

de Las Americas in southeastern Chiapas, Mexico. In

summary, a total of 37 mahoganies, d.b.h.>= 10 cm,

were encountered and tallied within the 25-ha study

block, for an average of 1.4 trees per ha. Mean d.b.h.

was 63.5 cm, and 54 percent of the stems tallied were

over 60 cm in d.b.h. A total of 14 mahogany individuals,

d.b.h.< 10 cm, were tallied in the regeneration plots (1

sapling and 13 seedlings), for an average of 13.2 per

ha.

Capixaba—This study site was located in natural

tropical moist lowland forests in the municipality of

Capixaba in southeastern Acre, Brazil. In summary, a

total of 8 mahoganies, d.b.h.>= 10.0 cm, were

encountered and tallied within the 25-ha study block

for an average of 0.3 trees per ha. Mean d.b.h. was

86.4 cm, and 63 percent of the stems tallied were over

60 cm in d.b.h. No mahoganies were encountered in

the regeneration sampling, neither saplings nor seedlings.

Workplan for Year 3

The main objectives must continue to be the establishment

and measurement of additional study sites. After three

completed study sites, several combinations within the

environmental gradient matrix of the species’ distribution

have been covered (table 1). Also, results to date

suggest that big-leaf mahogany population structure

appears to be sensitive to disturbance patterns. Figure

1 shows percentages of total mahogany individuals by

diameter-size class within 25-ha blocks. The one site

with a periodic disturbance regime (Bacalar) shows a

distinct preponderance of smaller size classes and few

large individuals when compared to the other two more

environmentally stable sites. The objective for this

coming year is to select three additional differing sites

that will complement those already completed. Particular

emphasis is being put into identifying sites on volcanic

substrates and montane habitats.

BIG-LEAF MAHOGANY GROWTH STUDY:

YEAR 2 PROGRESS REPORT

October 1, 1999 to September 30, 2000

Julio C. Figueroa Colon

Ecologist
4

Table 1—Environmental gradient matrix for big-leaf mahogany study sites

 Climate

Soils Dry (<1500) Moist (1500–2500) Wet (>2500)

Limestone Bacalaraa Belizeb Belizeb

Belize

Alluvial — Capixabaa Quiringuicharoa

Peru Feijo, Brazilc

Para, Brazilc

Volcanic Oaxaca, Mexicob Oaxaca, Mexicob —

Montana — Oaxaca, Mexicob Perub

a Completed sites.

b Potential unconfirmed sites.

c Potential confirmed sites.











































































































































 0

10

20

30

40

50

60

10-19.9 20-39.9 40-59.9 60-79.9 >80

Stem diameter class (cm)

Stems per ha (%)



Bacalar

 Quiringuicharo

 Capixaba

Figure 1—Comparative summary of diameter-size class distributions for big-leaf mahogany individuals

tallied in study sites in Mexico (Bacalar, Quiringuicharo) and Brazil (Capixaba). Some environmental

characteristics of the study sites may be gleaned from table 1.
5

It is almost the end of an era. Seventeen years ago

scientists of the International Institute of Tropical Forestry

(IITF) and other local institutions began a silvics manual

of tropical trees in Puerto Rico. One hundred and two

species of trees significant to the Caribbean were

described in detail. The English version, printed as

separate short monographs in 3-hole format, has been

available for some time. A Spanish translation has been

completed, editing and formatting have been preformed,

and it is being printed as a bound volume (Francis and

Lowe 1999). Copies should be available at the IITF

library during the first part of January 2001. It may be

viewed on the IITF website (http://www.fs.fed.us/global/

iitf/research/research.html) under “Article: Silvics of

Native and Exotic Trees.” A compendium of information

drawn from the silvics manual and other planting research

preformed over the years in Puerto Rico is also available

in English and Spanish (Francis 1998, 1999). Another

publication concerned with trees’ resistance to

hurricanes in Puerto Rico should also be noted. Percent

defoliation, crown loss, and snap plus throw were

estimated for a large number of individual trees of 24

species in the UPR Botanic Garden area, and

comparisons were made of the relative resistance of

these species. Small leaf mahogany (Swietenia

mahagoni) proved to be the most resistant overall and

African tulip tree (Spathodea campanulata), the least

resistant (table 1). Statistical comparisons and models

are presented in the publication (Francis 2000).

I STILL LOVE THE TREES, BUT . . .

 John K. Francis

Research Forester

Table 1—Species and means of trees sampled and the percent defoliation, percent crown

loss, and percent snap plus throw

Species D.B.H. Ht %Def. %Cl %S-T No.

Swietenia mahagoni 28.5 10.7 70.7 22.7 0 50

Hymenaea courbaril 38.0 17.3 89.6 40.6 2 50

Calophyllum calaba 34.8 15.2 77.3 41.0 6 50

Pinus caribaea 37.2 17.7 51.8 41.8 6 50

Melaleuca quinquinervia 26.6 8.8 52.4 43.2 10 39

Dalonix regia 36.3 10.9 70.2 43.8 2 49

Lagerstromia speciosa 48.5 11.5 77.0 44.0 2 50

Terminalia catappa 46.6 14.9 86.6 46.5 2 41

Tabebuia heterophylla 23.7 10.3 73.2 47.4 4 50

Cassia javonica 43.9 10.5 87.7 47.4 10 31

Casuarina equisetifolia 54.0 20.6 62.0 49.8 4 50

Pterocarpus macrocarpus 76.4 20.7 94.5 50.7 6 50

Bucida buceras 33.6 13.4 80.7 50.7 2 50

Enterolobium cyclocarpum 113.6 17.5 87.6 50.8 9 33

Mangifera indica 64.5 12.6 66.8 51.7 2 50

Clitoria fairchieldiana 26.0 4.7 62.0 51.9 38 32

Ficus benjamina 68.6 9.1 56.5 52.5 30 50

Schefflera morototoni 16.4 10.1 91.3 56.4 30 50

Albizia procera 34.2 18.8 97.3 59.7 22 50

Cecropia sherbertiana 24.5 12.7 94.5 60.6 24 34

Sterculia apetala 33.1 12.9 94.9 63.7 21 34

Peltophorum pterocarpum 38.6 15.4 89.5 65.6 12 33

Senna siamea 30.3 16.2 94.9 66.5 10 50

Spathodea campanulata 41.6 16.9 98.1 81.1 34 50
6

I still love trees, but we woodsmen always have another

ridge to cross and another horizon to view. There are

shrubs growing on the other side of that mountain—

thousands of them and each practically unknown. A

survey of the literature reveals that there are over 500

species of shrubs in Puerto Rico and approximately

2,500 species in the United States and its territories. It

is impossible to fix an exact number of shrub species

for a region because the definition of just what a shrub

is (what characterizes a shrub) cannot be very precise.

I personally believe that as long as a plant is not clearly

a tree or an herb, and as long as we recognize the

ambiguity of its form, it is of no consequence to describe

and understand it.

In cooperation with the Shrub Sciences Laboratory in

Provo, Utah, I have begun a national wildland shrubs

handbook. It is being produced along the lines of the

U.S. Silvics Manual. Each species’ monograph is limited

to two printed pages because of the large number of

species to be treated and the paucity of information

available for most of them. I intend for the manual to be

useful and as easy to read as possible. An introduction,

format for submissions, and eight monographs are

complete. The volume to date and each new monograph

will be posted on the IITF website (early 2001) and

eventually published as a book. Persons interested in

submitting a monograph should view the web page (as

soon as it is available) or contact the coordinator, John

K. Francis, at IITF.

LITERATURE CITED

Francis, John K. 1998. Tree species for planting in forest,

rural, and urban areas of Puerto Rico. Gen. Tech. Rep.

IITF–3. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, International Institute of

Tropical Forestry. 82 p.

Francis, John K. 1999. Especies forestales para plantar

en areas forestales, rurales, y urbanas de Puerto Rico.

Gen. Tech. Rep. IITF–13. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service, International

Institute of Tropical Forestry. 88 p.

Francis, John K. 2000. Comparison of hurricane damage

to several species of urban trees in San Juan, Puerto

Rico. J. Arboriculture. 26(4): 189–197.

Francis, John K.; Lowe, Carol A., eds. 1999.

Bioecologia de especies nativas y exóticas de Puerto

Rico y las Indias Occidentales. Gen. Tech. Rep. IITF–

15. Río Piedras, PR: U. S. Department of Agriculture,

Forest Service, International Institute of Tropical

Forestry. (In press).
7

During 1999-2000, the Landscape Ecology and Remote

Sensing Program at the IITF used Landsat TM satellite

imagery to map land use and natural vegetation for the

entire island of Puerto Rico. The map contains 17 types

of woody vegetation, including 5 successional forest

formations. This is the first such map done for the island

and the first map of land cover completed since 1978.

The IITF has also begun a Caribbean-wide mapping

effort, the Caribbean Mapping Initiative, in partnership

with the U.S. Geological Survey EROS Data Center,

The Nature Conservancy’s International Headquarters,

and the U.S. National Aeronautics and Space

Administration (NASA). Through the Global

Observation of Forest Cover program, NASA has

awarded funds for the purchase and preprocessing of

Landsat 7 satellite imagery for all of the Caribbean

islands and portions of the Guyana Shield. This mapping

effort will be the first of its kind for the Caribbean region

and will provide baseline data for ecosystem management

analyses related to biodiversity conservation, forest

carbon, and hazard mitigation.

LANDSCAPE ECOLOGY AND REMOTE SENSING PROGRAM

Eileen Helmer

Ecologist
8

This year my publications with various collaborators

focused on seven topics. The topics were: (1) ecosystem

management (Lugo 1999d, Lugo and others 1999a,

Lugo and Brown 1997); (2) roads (Lugo and Gucinski

2000); (3) mangroves (Lugo 1999a, 1999b, 1999c;

Lugo and others 1999c); (4) hurricane disturbances

(Lugo and others 2000b); (5) biodiversity (Ewel and

others 1999, Lugo and Brown 2000); (6) mapping of

life zones (Lugo and others 1999b); and (7) the

vegetation of the Caribbean (Lugo and others 2000a).

In addition, I edited a proceedings volume with Carleen

Yocum (Yocum and Lugo 2000) and reviewed a book

on climate change (Lugo 2000). The findings of some

of these publications are highlighted below.

ECOSYSTEM MANAGEMENT

I analyzed arguments against active tropical forest

management in light of available data and new research

that show tropical forests to be more resilient after

disturbances than previously thought. Tropical forest

management involves a diverse array of human activity

embedded in a complex social and natural environment.

Within this milieu, forest structure and composition adjust

to change and reflect the human and natural economy

of regions. Critics of active forest management

overestimate problems and underestimate the human

capacity to solve them. They isolate parts of a complex

issue, i.e., the biodiversity component of tropical forest

management, to generalize about the negatives of logging.

This view of the Tropics is consistent with past treatment

of tropical issues by those that evaluate the situation

from a nontropical perspective. The literature reveals

that conservation of biodiversity can be compatible with

measured use of tropical forests. However, the

conservation of biodiversity could be hurt should society

not approach the tropical forestry issue holistically and

act on misinformation. Action forest management is the

means toward the goal of conservation and the best

available way to simultaneously address human needs

and conservation of biodiversity. However, active forest

management requires the application of emerging

paradigms of ecosystem management (Boxes 1 and 2).

FOREST ROADS AS ECOSYSTEMS

Gucinski and I proposed a unified approach to the

management and analysis of the function and effects of

roads on forested rural landscapes. The approach is

based on considering roads as ecosystems (technoecosystems; fig.1) and conducting analyses of road

ecology prior to making policy or management decisions.

An ecosystem approach to road issues has four

advantages: (1) allows for the analysis of all types of

roads irrespective of geographic location; (2) provides

a holistic framework for analyzing all aspects of roads

from their alignment to their operation and

decommissioning as well as all road functions regardless

of value judgment; (3) provides a holistic focus to their

management; and (4) supplements landscape

management approaches based on spatial concepts.

We present five precautions to be considered when

evaluating road ecosystems: (1) identify the type of road

under consideration; (2) differentiate the effects and

conditions of individual road segments from those of

road networks; (3) be explicit about the phase of road

development to which the argument applies because

different phases of development have different effects

on the landscape; (4) ascertain the age of the road and

evaluate the degree of landscape adjustment to the road

and vice versa; and (5) not to prejudge human-induced

changes in landscapes as automatically good or bad for

the ecology or economy of a region. This ecosystem

focus and guiding principles are applied to several issues

that road policy and management activities must

address.

ECOLOGICAL RESEARCH

Ariel E. Lugo

Ecologist
9

MANGROVES

Mangroves are the subject of a struggle between those

who want to maximize economic benefit through

intensive uses of the system (sometimes at the cost of

eliminating the biota in favor of urban or other

developments) and those who advocate complete

preservation of the ecosystem even at the exclusion of

people. As tropical countries continue to develop

economically, their governments must balance the various

uses of mangroves and other coastal areas to assure

sustainability of development. However, this will not be

possible without a constant flux of scientific information

on these ecosystems. Scientists must provide accurate

information upon which resource managers and policy

makers can base their decisions.

Box 1 

Paradigms of Ecosystem Management 

Advancement in ecosystem science has led to new paradigms in resource management. 

These are listed below with the resource management paradigms they replace. 

• A non-steady-state concept of the ecosystem (as opposed to a focus on climax or 

balance ecosystems). 

• Managing from the perspective of resilience (as opposed to stability). 

• Considering disturbance an integral part of ecosystems, required to maintain them (as 

opposed to suppressing or ignoring catastrophic factors). 

• Considering past land use legacies, all species, and the dead mass (necromass) of 

ecosystems, (as opposed to focusing only on the present state of the site and the live 

component of a few species and populations). 

• Focusing greater attention on the connections within and between ecosystems, 

particularly the interface of land, water, and atmosphere (as opposed to focusing only 

on the site under management). 

• Considering all time and spatial scales (as opposed to focusing only on short-term and 

small geographic scales). 

• Maintaining a global and long-term perspective even when managing at small scales 

(as opposed to a short-term, local perspective). 

• Restoring whole ecosystems (as opposed to only rehabilitating land and water 

productivity). 

Paradigms of Ecosystem Management
10

Box 2 

Some Principles of Ecosystem Management 

• Follow nature’s lead. Mimic natural disturbance patterns and recovery trends in your 

area. 

• Think big. Manage for landscape diversity as well as within-stand or patch diversity, 

and maintain the largest possible contiguous patches of ecosystems of concern. 

• Don’t throw out any of the pieces. Maintain a diverse mix of genes, species, 

biological communities, and regional ecosystems. 

• Side with the underdogs. Set priorities in favor of the species, communities, or 

processes that are threatened or otherwise warrant special attention. 

• Try a different tool. Diversify management approaches and reduce the emphasis on 

complete conversion of ecosystems. 

• Keep your options open. Use existing infrastructure of resources wherever possible. 

• No ecosystem should be an island. Minimize fragmentation of continuous ecosystems 

by exploiting areas near existing clearings and by nibbling away at the edge instead of

creating a new hole. 

• Encourage free travel. Create a web of connected habitats. Leave broad travel 

connectors for plants and animals, especially along streams and ridge tops. 

• Leave biological legacies. Select what to leave behind as carefully as what to take 

out; specifically, leave standing live and dead trees and fallen trees in managing 

forests. 

• Leave it as nature would. Leave a mixture of tree sizes and species on the site. 

Restore naturally diverse forests after harvest. 

• Be an information hound. Use the latest studies and state-of-the-art technologies to 

design, monitor, and evaluate new approaches. 

• Be a critical thinker. Use only the scientific findings that make sense for your region 

and social setting. 

• Monitor, monitor, monitor. Monitoring is the only sure way to tell if you are really 

conserving biological diversity. 

Some Principles of Ecosystem Management
11

Mangrove forests are tough ecosystems to invade

because few species can tolerate the hydrological and

edaphic conditions that prevail in mangrove habitats.

The small pantropical mangrove species pool is also

the basis for asserting that mangrove forests are easy to

rehabilitate, at least in terms of tree species composition.

The following questions are useful as a guide for

evaluating the invasion of plant species into mangrove

habitats: (1) Is the invading species a halophyte? (2)

What conditions of the environment are the invading

species occupying, and how long will those conditions

last? (3) What is the geographic location of the invasion,

does it penetrate the forest, or is it only at the edge? (4)

Is the invasion a short-term response to changes in

microsite conditions? (5) Is the invasion the result of a

long-term shift in the mangrove habitat?

It appears that the use of models for planning research

is a valuable tool for programs involving one or many

researchers attempting to understand the regional

relationships among ecosystem types. A preliminary

model of a mangrove forest in south Florida has yielded

the following information that was not apparent from

examination of data prior to the present study:

1. Mangrove forests appear to reach a steady state

with respect to their biomass in phase with the frequency

of tropical hurricanes in regions where they occur.

2. The storage of organic detritus in the forest and its

export to the bays are a function of tidal amplitude, but

tides do not seem to affect gross photosynthetic rates

as much as they affect detritus accumulation versus

export.

3. Gross photosynthesis appears to be sensitive to

terrestrial input of nutrients, and the development of

mangrove biomass is dependent on the quantity of

nutrients and the efficiency of nutrient uptake.

4. Mangrove zonation and vigor may thus be a function

of nutrient availability rather than solely salinity as

previously thought.

5. During succession, mangroves exert significant

control over the amount of nutrients in adjacent waters,

but, if terrestrial runoff is reduced, they do not have the

capacity to maintain themselves at the same level of

production. This is due to loss of nutrients in detritus

export to the bays and suggests that there must be

selective pressure for mechanisms of recycling within

the mangrove forest.

HURRICANES, CORAL REEFS,

AND RAINFORESTS

The coexistence of hurricanes, coral reefs, and rainforests

in the Caribbean demonstrates that highly structured

ecosystems with great diversity can flourish in spite of

recurring exposure to intense destructive energy (tables

1, 2). Coral reefs develop in response to wave energy

and resist hurricanes largely by virtue of their structural

strength. Limited fetch also protects some reefs from

fully developed hurricane waves. Whereas storms may

produce dramatic local reef damage, they appear to

have little impact on the ability of coral reefs to provide

food or habitat for fish and other animals. Rainforests

experience an enormous increase in wind energy during

hurricanes with dramatic structural changes in the

vegetation. The resulting changes in forest microclimate

are larger than those on reefs, and the loss of fruit, leaves,

cover, and microclimate has a great impact on animal

populations. Recovery of many aspects of rain forest

structure and function is rapid though there may be longterm changes in species composition. Whereas

resistance and repair have maintained reefs and

rainforests in the past, human impact may threaten their

ability to survive.

INTRODUCTIONS OF SPECIES

Ewel and others (1999) sought scientific consensus on

the controversy of species introductions. Box 3 contains

research questions that emerged from the analysis.

THE LIFE ZONES OF THE UNITED STATES

Aim—Our main goals were to develop a map of the

life zones for the conterminous United States, based on

the Holdridge Life Zone system, as a tool for ecosystem

mapping, and to compare the map of Holdridge life

zones with other global vegetation classification and

mapping efforts.
12

Table 1—Approximate kinetic energy dissipation rates from wind and

waves. For comparison, the 24-hr mean solar energy input to the

rainforest or coral reef is about 185 joules m-2s-1. Explanations are

reported in Lugo and others 2000b

Winds and waves Kinetic Energy

(joules m-2s-1)

Winds

 Global yearly average wind 0.014

 Wind over Lake Mendota, Wisconsin, U.S.A. 0.003 – 0.015

 Winter storm wind, Long Island Sound, U.S.A. 2

 Winds from Hurricane Daisy

 0 to 37 km radius core 1.6

 37 to 74 km radius ring 72

 74 to 111 km radius ring 38

 111 to 148 km radius ring 27

 Area-weighted mean 47

Waves

 Yearly average waves on Grand Cayman Island

 Coral Reef, Caribbean Sea 20 – 25

 Zones of maximum wave energy form trade winds,

 Bikini Atoll Reef, Pacific Ocean 200 – 300

 Waves on exposed rocky coast, Northeast Pacific 3,000

 Maximum waves from Hurricane Allen on Jamaican

 coral reefs 550,000 –750,000

Location—The area of interest is the 48 contiguous

States of the United States.

Methods—We wrote a PERL program for determining

life zones from climatic data and linked it to the image

processing workbench (IPW). The inputs were annual

precipitation (Pann), biotemperature (Tbio), sea-level

biotemperature (T0bio), and the frost line. The spatial

resolution chosen for this study (2.5 arc-minute for

classification, 4-km for mapping) was driving by the

availability of current state-of-the art, accurate, and

reliable precipitation data. We used the Precipitationelevation Regressions on Independent Slopes Model,

or PRISM, output for the contiguous United States

download from the Internet. The accepted standard data

for air temperature surfaces were obtained from the

Vegetation/Ecosystem Modeling and Analysis Project

(VEMAP). This data set, along with station data

obtained from the National Climatic Data Center for

the U.S., was used to develop all temperature surfaces

at the same resolution as the Pann.

Results—The U.S. contains 38 life zones (34 percent

of the word’s life zones and 85 percent of the temperate

zones) including 1 boreal, 12 cool temperate, 20 warm

temperate, 4 subtropical, and 1 tropical. Seventy-four

percent of the U.S. falls in the ‘basal belt’, 18 percent is

montane, 8 percent is subalpine, 1 percent is alpine,

and <0.1 percent is nival. The U.S. ranges from

superarid to superhumid, and the humid province is the

largest (45 percent of the U.S.). The most extensive life

zone is the warm temperate moist forest, which covers

23 percent of the country. We compared the Holdridge

life zone map with output from the BIOME model,
13

Table 2—Some similarities and contrasts between coral reefs and rainforests

Similarities

High diversity of reef and rainforest types Fast development of photosynthetic tissue

Large number of species High respiration

Complex food webs Resistance and resilience strategy for storms

High species dominance Life history traits of dominant species

 reflect responses to disturbances

Day/night shifts in activity Asexual reproduction of corals and

 resprouting of trees after storms

High photosynthesis Delayed mortality after storm damage

Contrasts

Coral reefs Rainforest

Topography is mostly biogenic Topography is tectonic

Carbonate substrate does not provide nutrients Soil provides most of the nutrients

Low biomass of organisms High biomass of organisms

Plants inconspicuous Plants conspicuous

Animals easily visible Animals less visible

Algae controlled by herbivores Producers not controlled by herbivores

High degree of symbiosis Moderate degree of symbiosis

Low coral species turnover at 0.1-ha level High tree species turnover at 0.2-ha level

30 animal phyla represented 15 animal phyla represented

Alien species not important Alien species are a concern

Disease conspicuous Disease not conspicuous

Larvae are pelagic, dispersed by currents Animals disperse seeds

Larvae disperse thousands of km Seeds disperse hundreds of km

Water, nutrients, larvae and fish transported Water, gases, and some nutrients

 by currents transported by wind

Close physical proximity between Photosynthesis and respiration

 photosynthesis and respiration physically distanced

Low nutrient accumulation on sediments High nutrient accumulation belowground

Slow turnover variablesa are wave regime Slow turnover variables are topography

 and calcium carbonate structure and soil

Intermediate turnover variables include large Intermediate turnover variables are large

 fish and organic sediments trees

Fast turnover variables include water, plankton, Past turnover variables include leaves,

 algae, and microbes animals, and microbes

a The rate of turnover is classified relative to other rates within each system. There are also potentially

interesting and important comparisons of rates between systems.

Canal needs Rain forest
14

Box 3

Research Questions About Introductions

Several research questions need to be answered to help ensure that proposed introductions are done wisely and

safely.

Guarding against risks without sacrificing benefits:5

• How can the potential benefits and costs of introductions best be evaluated in economic, environmental, and

social terms?

• Should all introductions be regulated?

• How different must organisms or recipient ecosystems be from those assessed previously to warrant

independent assessment?

• When is it appropriate to assess and regulate taxa other than species?

• What are appropriate ecological and political boundaries for regulation?

Alternative to introductions:

• How and when can indigenous organisms be domesticated so that they can substitute for proposed uses of

nonindigenous organisms?

• How can the retention of indigenous species and natural food webs be integrated into agroecosystems so

that the risk of pest problems is minimized?

Purposeful introductions:

• What common guidelines can be developed for deliberate introductions of all kinds of organisms?

• Have screening procedures differed for introductions that proved successful or harmful?

• How can the potential for nonindigenous organisms to disrupt ecosystem processes be assessed and

reduced?

• Can the demand for introductions be reduced by improving the effectiveness of introductions that are

attempted?

Reducing negative impacts:

• When can reduction of human-caused disturbance within natural areas be used to control nonindigenous

species impacts?

• Can subtle, indirect effects of potential introductions be predicted?

• Can enough be learned from the population growth lags, booms, and crashes of previously introduced

organisms to make useful generalizations?

• Should special guidelines accompany release of sterile forms, which may pose less risk than fertile

organisms?

• Can protocols be developed to predict when an introduced species will hybridize with natives and what the

ecological and economic consequences of such hybridization might be?

• Should special guidelines related to invasion and hybridization potential be added to those that already

regulate release of genetically engineered organisms?
15

Bailey’s ecoregions, Küchler potential vegetation, and

land cover—all aggregated to four cover classes. Despite

differences in the goals and methods for all these

classification systems, there was a very good to excellent

agreement among them for forests but poor for

grasslands, shrublands, and nonvegetated lands.

Main conclusions—We consider the life zone

approach to have many strengths for ecosystem mapping

because it is based on climatic driving factors of

ecosystem processes and recognizes ecophysiological

responses of plants. It is hierarchical and allows for the

use of other mapping criteria at the association and

successional levels of analysis. It can be expanded or

contracted without losing functional continuity among

levels of ecological complexity. It is a relatively simple

system based on sparse empirical data. It uses objective

mapping criteria.

LITERATURE CITED

Ewel, J.J.; O’Dowd, D.J.; Bergelson, J. [and others].

1999. Deliberate introductions of species: research

needs. BioScience. 49(8): 619–630.

Lugo, A.E. 1999a. El manglar: un ecosistema al servicio

del ser humano. In: Matteucci, S.D.; Solbrig, O.;

Morello, J.; Halffter, G., eds. Biodiversidad y uso de la

tierra. Conceptos y ejemplos de Latinoamerica. Buenos

Aires, Argentina: Universidad de Buenos Aires,

Ceudeba, Centro de Estudios Avanzados: 361–382.

Lugo, A.E. 1999b. Mangrove ecosystem research with

emphasis on nutrient cycling. In: Yañez-Arrancibia, A.;

Lara-Domínguez, A.L., eds. Ecosistemas de manglar

en América tropical. Instituto de Ecología. A.C. Xalapa,

Veracruz; IUCN Centro América; Silver Springs, MD:

NOAA–NMFS: 17–38.

Lugo, A.E. 1999c. Mangrove forests: a tough system

to invade but an easy one to restore. Marine Pollution

Bulletin. 37(8–12): 427–430.

Lugo, AE. 1999d. Will concern for biodiversity spell

doom to tropical forest management? The Science of

the Total Environment. 240(1–3): 123–131.

Lugo, A.E. 2000. Climate change and tropical forests—

uncertainty and fear. Journal of Environmental Quality.

29(1): (book review).

Lugo, A.E.; Baron, J.S.; Frost, T.P. [and others]. 1999a.

Ecosystem processes and functioning. In: Szaro, R.C.;

Johnson, N.C.; Sexton, W.T.; Malk, A.J., eds.

Ecological stewardship: a common reference for

ecosystem management. Oxford, UK: Elsevier Science:

219–254. Vol. 2.

Lugo, A.E.; Brown, S. 1997. Tropical forests:

supporting Latin America and the world into the 21st

century. Acta Científica. 11(1–3): 97–102.

Lugo, A.E.; Brown, S. 2000. Biodiversity in the

Caribbean: concepts, assessments, management and

monitoring at the landscape scale In: Yocum, C.; Lugo,

A.E., eds. Biodiversity in the Caribbean: management

and benefits. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, International Institute of

Tropical Forestry: 92–98.

Lugo, A.E.; Brown, S.L.; Dodson, R. [and others].

1999b. The Holdridge life zone system of the

conterminous United States in relation to ecosystem

mapping. Journal of Biogeography. 26: 1025–1038.

Lugo, A.E.; Figueroa Colón, J.; Scatena, F.N. 2000a.

The Caribbean. In: Barbour, M.G.; Billings, W.D.,eds.

North American terrestrial vegetation. England:

Cambridge Press: 593–622.

Lugo, A.E.; Gucinski, H. 2000. Function, effects, and

management of forest roads. Forest Ecology and

Management. 133: 249–262.

Lugo, A.E.; Rogers, C.S.; Nixon, S.W. 2000b.

Hurricanes, coral reefs and rainforests: resistance, ruin

and recovery in the Caribbean. Ambio. 29: 106–114.
16

Lugo, A.E.; Sell, M.; Snedaker, S.C. 1999c. Mangrove

ecosystem analysis. In: Yañez-Arrancibia, A.; LaraDomínguez, A.L., eds. Ecosistemas de manglar en

América tropical. Instituto de Ecología. A.C. Xalapa,

Veracruz; IUCN Centro América; Silver Springs, MD:

NOAA–NMFS: 343–364.

Yocum, C.; Lugo, A.E., eds. 2000. Biodiversity in the

Caribbean: management and benefits. Río Piedras, PR:

U.S. Department of Agriculture, Forest Service,

International Institute of Tropical Forestry. 130 p.
17

During FY2000, work was completed on a project

initiated in 1994 to describe and document the traditional

medicinal uses of forest plants in peninsular India. The

primary purpose of this project has been to synthesize

existing knowledge on the botany, ecology, and

traditional uses of Indian medicinal plants in a form that

is accessible to a broad spectrum of potential users and

the general public. This resulting book, to be published

by CAB International in early 2001, focuses on the flora

characteristic of the deciduous forest zone that includes

nearly all of central India south of the Indo-Gangetic

Plain and interior regions of Southern India. It includes

detailed information on the botany, geographical ranges,

habitat preferences, and traditional medicinal uses (in

FOREST MANAGEMENT AND REHABILITATION RESEARCH

Healing Plants of Peninsular India

 John A. Parrotta

 Reseach Forester

Ayurveda, Unani, Siddha, folk, and tribal medicine) of

545 plant species, each illustrated with color photographs

taken during field excursions in several Indian States in

1994 and 1997. The book’s introduction examines the

cultural and historical development of traditional Indian

medicine and research on plant drugs in an international

context, as well as the current distribution and status of

India’s forests and medicinal plant resources. Species

descriptions include their common names in the 14 major

languages of the sub-Himalayan region (including English

and Sanskrit). Specialized glossaries and indices of

medicinal uses as well as common names in English,

Sanskrit, and regional Indian languages are also

provided.
18

As part of a larger study evaluating several silvicultural

techniques for restoring tropical moist forests on

abandoned agricultural lands in southeastern Brazil,

direct seeding with five early successional Atlantic forest

species was tested at three degraded sites, characterized

by different soil types and land use histories, within the

Environmental Protection Area at Botucatu, SP. The

species used in this study were Chorisia speciosa,

Croton floribundus, Enterolobium contorstisiliquum,

Mimosa scabrella and Schizolobium parahyba.

Scarified seeds of each of these species were sown in

prepared seed spots in replicated, 0.25-ha mixedspecies plots at an initial spacing of 1 by 1 m at each

site. Of the five species planted, only two, Enterolobium

and Schizolobium, showed good seed germination,

seedling survival, and early growth rates, averaging 4.1

to 4.6 cm stem diameter and 0.8 to 0.9 m height growth

during the first 2 years after sowing. These two species

constituted 88 to 100 percent of the total stand density,

which ranged from 1050 to 1790 stems ha-1 at 2 years.

Despite the poor performance of the other species

tested, we observed that the natural regeneration of

native forest species originating from remnant forests in

the general vicinity of our study sites was significantly

greater within the direct-seeded plots than in unplanted

control plots that were protected from fire and other

disturbances.

FY2000 PUBLICATIONS

Engel, V.L.; Parrotta, J.A. 2000. Restauração de

ecossistemas florestais. Agroecologia Hoje.1(4):

22–23.

Keenan, R.J.; Lamb, D.; Parrotta, J.; Kikkawa, J. 1999.

Ecosystem management in tropical timber plantations:

AN EVALUATION OF DIRECT SEEDING FOR REFORESTATION OF

DEGRADED LANDS IN CENTRAL SÃO PAULO STATE, BRAZIL

Vera Lex Engel1

 and John A. Parrotta

1

 Natural Resources Department, Universidade Estadual PaulistaUNESP/FCA, P.O. Box 237, 18603–970 Botucatu, SP, Brazil.

E-mail: veralex@fca.unesp.br

satisfying economic, conservation and social objectives.

Journal of Sustainable Forestry. 9(1/2): 117–134.

Parrotta, J.A. 1999. Productivity, nutrient cycling and

succession in single- and mixed-species plantations of

Casuarina equisetifolia, Eucalyptus robusta and

Leucaena leucocephala in Puerto Rico. Forest Ecology

& Management. 124(1): 45–77.

IN PRESS

Engel, V.L.; Parrotta, J.A. [In press]. An evaluation of

direct seeding for reforestation of degraded lands in

central São Paulo State, Brazil. Forest Ecology &

Management. [2001].

Parrotta, J.A. [In press]. Healing Plants of Peninsular

India. CAB International, UK. [2001].

Parrotta, J.A.; Knowles, O.H. [In press]. Restoring

tropical forests on lands mined for bauxite: examples

from the Brazilian Amazon. Ecological Engineering.

[2001].

CONFERENCE PRESENTATIONS

Catalyzing natural forest restoration on degraded tropical

landscapes. Invited keynote paper presented at Forest

Restoration for Wildlife Conservation, Chiang Mai,

Thailand; 2000 January 30–February 4.

Restoration forestry for multiple objectives. Invited lead

paper presented at Tropical Forestry Research:

Challenges in the New Millennium international

syposium, Kerala Forest Research Institute, Peechi,

Kerala, India; 2000 August 2–4.

Native tropical forest rehabilitation: a case study from

Brazil. Invited paper presented at XXI IUFRO World

Congress, Kuala Lumpur, Malaysia; 2000 August

7–12.
19

The past year was as busy as ever but will probably

be remembered for four important events:

• No hurricanes passed the island, and we had only

one hurricane warning. After several years of

hurricane worries, this was a great relief for all at

IITF.

• Dozens of healthy tabonuco (Dacryodes excelsa

Vahl.) seedlings were found in and around the

Sabana field station. During the past year seed,

rain, and regeneration of this species have been

abundant within the forest. However, this is the

first time we have seen this important species

regenerating under advanced secondary vegetation

so far away from closed tabonuco-type forest. We

are keeping a close watch on these true pioneers.

• Massive upstream migrations of fish and snails

were observed in the rivers of the Luquillo

Experimental Forest. In June and July, we saw

several large schools of the mountain mullet

(Agonostomus monticola) “jumping like salmon”

as they migrated upstream. Large schools were

seen in the Mameyes at Puente Roto in mid-June

and in the Espirtu Santo River near the Girl Scout

camp in mid-July. When questioned, several longterm residents of the area who are also avid

fisherman said they have never seen anything like

this before. Starting in September and continuing

until December 2000, we also witnessed 5 massive

upstream migrations of the freshwater snail

Neritina in the Río Mameyes near Route 3.

Although these migrations have been described

before, this is the first time we have been able to

follow them and carefully document the migrations.

Based on our initial information of their rate of

migration, we estimate it may take more than 5

years for these snails to make it all the way to

Puente Roto.

• The construction of the new IITF laboratory and

sample storage facilities in Río Piedras began. This

has been in the planning for several years and will

greatly improve our ability to conduct research and

provide services to our collaborators.

In addition to these historic events, we continued to

have an active collaborative research program.

Moreover, the Sabana Field Station has been occupied

throughout the year with students and collaborators from

the University of New Hampshire, University of

California at Berkeley, University of Puerto Rico,

SUNY-Stony Brook, SUNY-Environmental Science

and Forestry, and the Vrije Universiteit of Amsterdam.

In addition to investigators from the United States and

Puerto Rico, students and investigators from Singapore,

Holland, Germany, Panama, Colombia, and Guatemala

worked with us in the Luquillo Experimental Forest.

During the fiscal year, two new projects were also

established. The first was under the direction of Dr.

William McDowell of the University of New Hampshire.

This project, which is funded by a U.S. Department of

Agriculture competitive grant, is focused on

understanding nitrogen dynamics in the forests of Bisley

and the Río Icacos. The second large project was a

year-long micrometeorological study of the Luquillo

climatic gradient under the direction of Prof. Hans Vugts

and students from the Vrije Universiteit of Amsterdam.

In our continuing effort to promote the transfer of

scientific knowledge for the management of aquatic

resources, we co-sponsored a meeting on the

management of Puerto Rican streams at the Society of

Puerto Rican Engineers. The meeting was organized with

the U.S. Fish and Wildlife Service and the local

engineering firm Gregg Morris and Associates. The

meeting was attended by over 85 engineers and natural

resource managers from throughout the island.

ECOSYSTEM RESEARCH

F.N. Scatena

Hydrologist
20

Participants from the U.S. Geological Survey, the Army

Corps of Engineers, the Natural Resource and

Conservation Service, the Puerto Rican Departments

of Highways and Natural Resources, and environmental

officers from several municipalities attended the

conference.

This year also marks the first year of the Caribbean

AMIGO project on Regional Hydrology. This is a

multinational volunteer effort aimed at developing

integrated hydrological analysis. During the year, the

AMIGO web page was established (http://

biblioteca.imta.mx/amigo/), and several projects on

ecohydrology, extreme events, droughts, and data

management were started. At an international meeting

sponsored by the International Hydrological Program,

the AMIGO group presented a paper that will be

published shortly on natural disturbances and the

hydrology of humid tropical forests.
21

The emphasis of this year’s work was again on

mahogany. The mahogany provenance study in Puerto

Rico was planted in the 1960’s at 14 sites in Puerto

Rico and St. Croix, U.S.VI, from dry to wet forest sites,

using 20 provenances collected from Mexico, Central

America, and the Caribbean. This year, material from

this study is being included in a phylogeographic analysis

of population differentiation of Swietenia macrophylla

using restriction fragment length polymorphisms in

chloroplast DNA. The Institute of Terrestrial Ecology

(ITE, Scotland) and CATIE (Costa Rica) are heading

up this project. Puerto Rican Cedrela odorata, and

representatives of the genus Toona that are in the IITF

Arboretum at the Luquillo Experimental Forest are also

being included in a genetic marker taxonomic survey of

the Swietenoideae of the family Meliaceae (project

headed by ITE, Scotland). The survey of wood density

in the surviving trees of the Puerto Rican provenances

has been completed for comparing the effects of

environment and genetic background on this component

of wood quality. A perspective on the causes and effects

of shootborer attack in the early years of this provenance

trial were presented at the North American Forestry

Commission in Merida, Mexico, and has been submitted

for publication.

The collaborative work funded by FAS with CATIE on

genetic trials of S. macrophylla and C. odorata in

Mexico and Costa Rica continue, with analysis of early

measurements. Preliminary results of the S. macrophylla

trials in Mexico were presented at a forest genetics

conference this year in French Guiana. A study

comparing quantitative traits in seedlings of C. odorata

from ecologically distinct zones of Costa Rica has been

submitted for publication. I assisted with the planning

for the collection of S. macrophylla germ plasm

completed in eastern Bolivia this summer by C. Navarro

of CATIE in a joint project funded by FAS. This

collection was part of a larger objective to complete

the collection of germplasm of Swietenia from the

ranges of the three species for purposes of genetic

characterization (using genetic markers and quantitative

traits), performance assessment, and conservation.

The history of U.S. Forest Service research at Estate

Thomas, St. Croix, U.S.VI, was reviewed and presented

at the Urban Forestry Conference in St. Croix. The

proceedings are to be published (Ward and others

2000). I also provided the research component for the

future planning for Estate Thomas.

Arranging the remeasurement data sets of the four

hundred 0.089-ha plots that Frank Wadsworth

established in the 1950’s at the Luquillo Experimental

Forest continues under UPR student Vanessa Rivera

with funding from the NASA INRA program. GPS of

representative plots has been obtained, and mapping of

these plots is underway. The remeasurement data sets

will provide information on forest dynamics that can be

interpreted in relation to land use history and other spatial

environmental variables for the mapped plots.

LITERATURE AND

PRESENTATIONS CITED

Navarro, Carlos; Ward, Sheila; Hernández, Marvin.

Cedrela odorata (Meliaceae): a subdivided species in

Costa Rica. To Revista de Biología Tropical. [number

of pages unknown].

Ward, S.E; Rodriguez, A.; Morgan, R. [In press]. Estate

Thomas Experimental Forest: past, present, and future.

Proceedings of the Fifth Caribbean Urban Forestry

conference;[date of conference unknown]; St. Croix,

U.S.VI. [Place of publication unknown]: [Publisher

unknown]. [2000].

CONTINUED STUDIES ON MAHOGANY AND LONG-TERM PLOTS AT

THE LUQUILLO EXPERIMENTAL FOREST

Sheila Ward

Ecologist
22

Ward, S.E.; Waterman, D. 2000. Causes and

consequences of shoot borer (Hypsiphyla

grandella)attack in a mahogany provenance study in

Puerto Rico. NAFC Forest Biology Workshop; [date

of workshop unknown]; Merida, México.[Place of

publication unknown]: [Publisher unknown]. [number

of pages unknown].

Ward, Sheila; Wightman, Kevyn; Haggar, Jeremy;

Santiago, Bartolo. 2000. Preliminary results from

Swietenia macrophylla genetic trials in the Yucatan,

Mexico. International symposium on modeling and

experimental research on genetic processes in tropical

and temperate forest; [date of symposium unknown];

Kourou, French Guiana. [Place of publication

unknown]: [Publisher unknown]. [number of pages

unknown].
23

INTERNATIONAL COOPERATION

Mahogany in Nicaragua

Nicaragua’s Indians, originally from Mexico and

northern South America, were decimated by the Spanish

soon after Columbus landed in 1502 (Weaver and Bauer

1999). By the 1530's, the new Spanish settlers were

cutting mahogany along with other timbers for

shipbuilding at Realejo [now Corinto] on the Pacific

coast. By the 1600’s, the Spanish occupied much of

the Pacific coast, and the British exploited forest

resources along the Caribbean coast. The major

Meliaceae, big-leaf mahogany, Pacific coast mahogany,

and Spanish cedar were selectively logged throughout

the colonial period, a practice that continued during the

20th century when a few more species were added to

the commercial harvest. Land clearing for agriculture

also eliminated forest resources and reduced the major

Meliaceae. Important agricultural activities included

coffee plantations from the late 19th century through

1950 and cotton farming from 1950 through 1980, both

in the Pacific region, and expansion of the agricultural

frontier eastward toward the Caribbean coast from the

1970’s to the present. Forest cover, occupying slightly

more than 50 percent of the country in 1950, declined

to 33 percent in 1990; it is estimated to be 25 percent

for the year 2000. Big-leaf mahogany and Spanish cedar

plantations occupy only about 20 ha, and forest

management is limited to experimental work on only 15

ha. A total of 73 conservation areas cover nearly 17

percent of the entire country, and forest resources in

these areas should remain protected. Currently, Pacific

coast mahogany is listed in Appendix II and big-leaf

mahogany in Appendix III of Nicaragua’s Redbook of

Endangered and Threatened Species.

Species Trials in the Dominican Republic

The growth and acclimatization of nine species (Acacia

mangium, Azadirachta indica, Catalpa longissima,

Cedrela odorata, Eucalyptus grandis, Pinus

caribaea, Simarouba glauca, Swietenia

macrophylla, and S. mahagoni) established either as

line plantings or block plantations, were evaluated 18

to 44 months after planting in Tocoa, Dominican

Republic (Sanchez 1999). Different methods of planting

did not result in marked differences in growth except

for A. mangium in dense, block plantations, (e.g., 2,667

trees per ha) where the initial rapid growth rate and

density of the stand led to reduced diameter growth at

an early age. The mean annual increment in height and

diameter on the best sites were A. mangium (4.25 m

and 3.95 cm), E. grandis (3.38 m and 2.54 cm), S.

macrophylla (2.20 m and 2.31 cm), S. mahagoni

(1.34 m and 1.26 cm), A. indica (2.06 m and 1.90

cm), and S. glauca (1.46 m and 1.57 cm). The mean

annual increments in height of the poorly developed

species were P. caribaea (0.23 m), C. longissima

(0.81 m) and C. odorata (0.53 m). It is concluded that

the growth of the nine species was influenced by soil

conditions, maintenance practices (weed control,

especially grasses), and the capacity of each species to

adjust to extremely degraded soils. Acacia mangium,

E. grandis, S. glauca, S. macrophylla, and S.

mahagoni were the most promising species for

reforestation under the conditions tested.

San Lorenzo Protected Area (SLPA), Panama

Fort Sherman, at the northwest entrance to the Panama

Canal, reverted to the Panamanian Goverment at the

end of June 2000. Currently, the International

Cooperation program of the IITF is cooperating with

USAID and CEASPA (a local NGO) in the preparation

of a slide program and technical report on the SLPA.

Both highlight the SLPA’s setting as a major crossroad

and briefly describe pre-Columbian activities, the

Spanish conquest, the legacy of fortune seekers and

the Chagres River, the building of the Panama Canal,

and early agricultural activities. The military history of

FOREST RESEARCH

Peter L. Weaver

Research Forester
24

both Fort San Lorenzo and Fort Sherman will be

discussed along with current knowledge on the geology,

soils, flora, fauna, marine resources, ecological research,

and proposed conservation of the SLPA. A chronology

of major historical events related to the SLPA will also

be included.

RESEARCH

Elfin Woodland in Puerto Rico

Grasses and ferns characterized the recovery of elfin

woodland for the first 18 years after a December 1968

airplane crash (Weaver 2000). From 1986 to 1998,

ferns and woody dicots were prominent, and the total

aboveground dry weight biomass increased from 775

to 2210 g/m2

. Woody dicots increased 3.5 times, palms

1.3 times, and ferns 4.4 times above their 1986 levels.

Grasses and herbs decreased by nearly 10 percent.

Recovery was patchy, with some areas dominated by

trees 34 m in height and others by grasses and scattered

ferns. Puerto Rican endemic trees are playing a critical

role in the recovery of the elfin woodland and are

prominent in mature elfin woodland as well. Endemics

account for 88 to 94 percent of the woody dicot stems

18 to 30 years after the wreck, and from 55 to 72

percent of the stems in mature elfin woodland. The

largest stems on the wreck site average about half of

the typical heights and d.b.h.’s of mature elfin woodland

trees and biomass are about a fourth of that found in

mature elfin woodland. Species composition is similar.

It is estimated that complete recovery will take almost

two centuries.

Montane Rain Forest in Puerto Rico

Forest structure and species richness vary along

environmental gradients in the colorado forest [montane

rain forest] of Puerto Rico’s Luquillo Mountains (Weaver

2000). Rainfall and temperature vary with aspect and

elevation resulting in wetter conditions to the windward

than to the leeward. A covariance model showed that:

stem density increases significantly with elevation to the

windward but not to the leeward; tree height decreases

significantly with elevation on both windward and

leeward exposures; total aboveground woody biomass

decreases significantly with elevation on ridges to the

windward; and organic matter increases significantly with

elevation to the leeward. ANOVAs disclosed

interactions between aspect and topography for species

diversity and biomass. Species-area curves show the

greatest number of species on ridges and the fewest in

ravines.

LITERATURE CITED

Sanchez, A.; with Rodriguez, A.; Valenzuela, O.;

Weaver, P.L. 1999. Growth and acclimatization of nine

forest species in the Dominican Republic. In: Biodiversity

in the Caribbean: management and benefits.

Proceedings of the ninth meeting of Caribbean Foresters;

1998 June 1–5; Dominican Republic. [Location of

publisher unknown]: [Publisher unknown]: 99–106.

Weaver, P.L.; Bauer, G.P. 1999. Major Meliaceae in

Nicaragua. Gen. Tech. Rep.–10. Río Piedras, PR:

International Institute of Tropical Forestry. 38 p.

Weaver, P.L. 2000. Elfin woodland recovery 30 years

after a plane wreck in Puerto Rico’s Luquillo Mountains.

Caribbean Journal of Science. 36(1–2): 1–9.

Weaver, P.L. 2000. Environmental gradients affect

forest structure in Puerto Rico’s Luquillo Mountains.

Interciencia. 25(5): 254–259.
25

During FY-2000, I continued four studies, supported

cooperative studies, and published the results of previous

wildlife investigations. These activities are summarized

below:

CURRENT FIELD PROJECTS

AND REVIEWS

Effects of Low-Impact Logging on Birds and Bats

of the Forest Understory of the Tapajós National

Forest

Continental tropical forests are characterized by a high

diversity of species, most of which are rare. Nowhere

is this more evident than in the Amazon forests of Brazil,

where our current collaborative studies with researchers

from the Goeldi Museum and Dr. Michael Willig of Texas

Tech University are helping to characterize the

understory bird and bat assemblages in the Tapajós

National Forest near Santarem. Our collaborative studies

based on mist netting enable us to describe and

characterize the avian and bat diversity in the forest

understory, and more importantly, serve as a control

for examining the effects of low-impact logging on these

components of the fauna in an Amazon forest. Such

studies are beneficial for devising tropical logging and

silvicultural methods, which minimize biodiversity loss

and ensure that animal seed dispersers remain on

managed sites to provide adequate seed dispersal. Our

current field study will end in June 2001.

Movements, Home Range, and Habitat Use by

the Puerto Rican Boa in the Luquillo Experimental

Forest

We are now in the fourth year of a study of the

endangered Puerto Rican Boa (Epicrates inornatus)

based on surgically-implanted radio transmitters, which

enable us to use radio telemetry to locate and follow

snakes in the LEF. The locations where the radio-tagged

snakes are found by telemetry are mapped by the use

of GPS, which enables us to quantify movements and

determines the size of a boa’s home range. The passage

of Hurricane Georges in September 1998 appeared to

have little or no effect on the five radio-tagged boas we

were following at the time. Field work will continue until

June 2001 to further document the natural history and

movements of these snakes.

Phenology of Some Common Fruits Consumed by

the Puerto Rican Parrot

We are now in the fourth year of a phenology study

designed to monitor fruit production of the more common

tree and vine species in the Palo Colorado forest of the

LEF. The study involves monthly sampling of marked

plants along trails at Camitillo (10 species; 122

individuals originally) and Palo Hueco (11 species, 144

individuals originally). The study is designed to

characterize variation in fruit production to help

understand the behavior of the Puerto Rican parrot to

identify the most important time periods to release

aviary-produced parrots.

Hurricane Georges struck the LEF in September 1998,

just as we completed 2 years of phenology sampling,

providing baseline for posthurricane comparisons. As

we found previously in the aftermath of Hurricane Hugo,

little fruit remained on the marked plants after the storm,

and phenological patterns shifted out of phase from

normal prehurricane patterns. We will continue to

monthly monitor the recovery of fruit production at these

sites.

A Review of the Puerto Rican Parrot and Its

Recovery: 1973–2000

I am currently working with participants from the Puerto

Rican parrot (Amazona vittata.) workshop and

members of the recovery effort to summarize and

evaluate the parrot’s population growth, nesting success,

survival, and the overall success of the recovery effort.

This involves analyses of wild and captive data, much

of which has never been summarized previously. The

WILDLIFE RESEARCH

Joseph M. Wunderle, Jr.

Wildlife Team Leader & Research Wildlife Biologist
26

goal of the project is to identify critical research needs

and areas in which increased management intervention

could increase population growth.

PUBLICATIONS

As part of our contribution to the recovery of the

endangered Puerto Rican parrot, I published the results

of a 5-year fruiting phenology study (Wunderle 1999b).

In this work, I quantified fruit availability on 25 plant

species consumed or potentially consumed by the Puerto

Rican Parrot to document the seasonal and annual

variation in fruit production in the Luquillo Mountains.

In the 33 months before Hurricane Hugo, an annual

cycle in the number of species with ripe fruit was evident,

with a peak in October–February and a trough during

June–July. About half the plant species showed this

annual fruiting cycle. Irregular noncyclic fruiting was

found in the other half and varied among species in annual

duration. Fruit production reached its lowest point in

October 1989, just after Hurricane Hugo, when 72

percent of the broadleaf foliage was lost and only one

species had ripe fruit. The number of fruiting species

subsequently increased, but the cyclic fruiting pattern,

evident in the number of fruiting species before the storm,

disappeared and was not observed during 27 months

after the storm. This noncyclic pattern was attributed

mostly to species with annual fruiting cycles in which

annual fruiting shifted out of phase, was suppressed after

the hurricane, or both. Parrot breeding was associated

with fruiting, as breeding occurred during fruiting peaks

before the storm, and was delayed in the first season

after the storm, but returned to normal by the second

season. Thus, parrots faced considerable annual and

year-to-year variation in fruit availability prior to the

hurricane, and substantial fruit loss afterwards followed

by a recovery involving changes in fruiting phenology of

individual species and the overall community.

As part of our studies on the role of traditional shade

coffee plantations as potential refugia for forest-dwelling

birds in deforested or degraded landscapes in the

Tropics, we published two papers. I summarized the

role of shade coffee plantations in conserving bird

diversity in a presentation and publication for the Society

of Caribbean Foresters (Wunderle 1999a). In addition,

we published a paper on winter-site fidelity of Nearctic

migrants in shade coffee plantations of different sizes in

the Dominican Republic (Wunderle and Latta 2000).

Here we studied three Nearctic migrant species

(American redstart [Setophaga ruticilla ], AMRE;

black-and-white warbler [Mniotilta varia ], BAWW;

black-throated blue warbler [Dendroica caerulescens

], BTBW) wintering in 14 isolated shade coffee

plantations (0.1 to 8.7 ha) to determine if site fidelity in

shade plantations was comparable to that in tropical

forests and if it decreased with plantation size. Site

fidelity was measured as the percentage of wandering

birds captured in mist nets, as overwinter site persistence

of uniquely marked birds observed on the same sites

(November to March), and as the annual return of

marked individuals to previously occupied sites (January

to January). The percentages of wanderers in net

captures were mostly lower than values reported for

natural forests (AMRE 21 percent, BAWW 12 percent,

BTBW 41 percent) and did not vary with plantation

size. Overwinter site persistence (AMRE 65 percent,

BAWW 65 percent, BTBW 76 percent) and annual

return (AMRE, 34 percent; BAWW, 40 percent;

BTBW, 31 percent) in the plantations fell within the range

of values reported for natural forests. Overwinter site

persistence decreased with plantation size only in

AMRE, although BAWW showed lower persistence in

small plantations from early to midwinter. Annual return

decreased with plantation size only in AMRE. Despite

diminished site fidelity in small plantations, these birds

still showed some fidelity to small plantations, many

smaller than the mean winter home range sizes.

Several minor publications also appeared in this period.

For example, I provided brief summaries of hurricanes

and wildlife, island exotics, and bird migration in the

Caribbean (Wunderle 1999c) for a wildlife viewing

guide. In addition, we published a brief review of the

Puerto Rican parrot (Wiley and others 2000) in a parrot

conservation review. Finally, as a member of the

Conservation Committee of the American Ornithologists

Union, we evaluated the Partners In Flight species

prioritization scheme for North American birds

(Beissinger and others 2000).
27

LITERATURE CITED

Beissinger, S.R.; Reed, J.M.; Wunderle, J.M., Jr. [and

others]. 2000. Report of the AOU Conservation

Committee on the Partners in Flight species prioritization

plan. Auk. 117: 549–561.

Wiley, J.; Wunderle, J.; Smith, A.; [and others]. 2000.

Puerto Rican Parrot (Amazona vittate). In: Snyder,

N.; McGowan, P.; Gilardi, J.; Grajal, A., eds. Parrots:

status survey and conservation action plan 2000–2004.

IUCN. Oxford, UK: Information Press: 117–118.

Wunderle, J.M. 1999a. Conservation of Caribbean

wildlife diversity: the role of agriforestry plantations. In:

Yocum, C.; Lugo, A.E., eds. Proceedings of the ninth

meeting of Caribbean Foresters;1998 June 1–5;

Dominican Republic. Río Piedras, PR. International

Institute of Tropcial Forestry, U.S. Department of

Agriculture, Forest Service: 115–120.

Wunderle, J.M. 1999b. Pre- and posthurricane fruit

availability: implications for Puerto Rican Parrots in the

Luquillo Mountains. Caribbean Journal of Science.

35(3–4): 1–14.

Wunderle, J.M. 1999c. Short introductions to:

hurricanes and wildlife; migrant birds in the Caribbean;

exotic birds. In: Nellis, D.W., ed. Wildlife Viewing Guide

for Puerto Rico & U.S. Virgin Islands. Helena, MT:

Falcon Press: 22, 27, 41.

Wunderle, J.M.; Latta, S.C. 2000. Winter site fidelity

of Nearctic migrant birds in shade coffee plantations of

different sizes in the Dominican Republic. Auk. 117:

596–614.
28

The cooperative programs are administered and

implemented through a partnership between the

Government of Puerto Rico, the U.S. Department of

Agriculture, Forest Service, and many other private and

government entities. Emphasis focuses on agroforestry,

timber and other forest products, wildlife, water

resources, rural economies, and conservation practices.

The goal is to maintain and improve the health of Puerto

Rico’s urban and rural forests and related economies.

These programs:

• reduce costs through the use of partnerships;

• increase values through sustained productivity of

forests; and

• are voluntary and nonregulatory in their delivery.

STATE AND PRIVATE FORESTRY ACCOMPLISHMENTS IN PUERTO RICO

Robin Morgan

Silviculturist

Investment in State’s Cooperative Programs

Puerto Rico has almost 700,000 acres of privately

owned forest land and 28,000 acres of national forests.

Of the 368,668 individuals owning forest lands, most

have parcels less than 20 acres in size. Almost all of

these landowners report that conservation, esthetic

enjoyment, and recreation are primary reasons for

owning forest land. The State‘s 3.9 million people include

78 municipalities (approximately 274 communities) with

populations over 100; all of these have initiated some

level of community forestry program.

Key Issues

Key issues, which the State will address with the 2001

budget include:

• protection and restoration of natural areas;

Programs FY2000 FY2000 FY2001 FY2001 FY2001 FY2001

Title II, VTitle II, V

& VIII & VIII Title IV Title IV

Federal State Federal State

Federal State estimate estimate estimate estimate

(U.S.$) (U.S.$) (U.S.$) (U.S.$) (U.S.$) (U.S.$)

Forest health management 68,750 118,348 16,000 32,000 50,000 50,000

Volunteer fire assistance 14,400 14,400 14,400 14,400 50,000 50,000

Forest stewardship program 158,882 233,529 162,888 233,529

Stewardship incentives program 0 0 0 0

Forest legacy program 729,000 729,000 1,330,000 1,330,000

Urban and community forestry 177,661 280,131 177,661 280,131

Economic action program 123,705 166,809 123,705 166,809

Forest resource and analysis

 program 0 0 0 0

Natural resource conservation

 Education 18,000 120,480 18,000 120,480

 Total 1,378,388 2,260,697 1,966,639 2,495,218
29

Forest Facts and Accomplishments

Selected facts Selected results

Population 3.9 million Stewaardship plans prepared 25

 (current year)

Population density, people per square mile 1,000 people Area under stewardship plans 449 acres

 (current year)

People per square mile, San Juan 20,000 people Area under stewardship plans 3,438 acres

metro area (all years)

Tree species on the island, number 570 Rural acres planted 3,895 acres

Exotic or naturalized tree species, number 225–250 Technical assists to private 1,344

 landowners

Extent of protected lands < 5% Rural fire districts assisted N/A

Miles of roads per square mile 4 miles Rural fire department N/A

 volunteers trained

Acres of forest land 709,177 acres Acres surveyed for forest 55,723

 health acres

Acres of nonindustrial private forest land 689,409 acres Forest health assistance visits 140

Number of NIPF landowners 368,668 Communities assisted 274

Acres of land use under state fire protection 2,226,400 acres Economic action grants to 7

 rural areas

Number of municipalities 78 Technology transfer and 4,289 days

 workshops

State forestry budget $13,686,000 Acres protected through 529 acres

 acres program
30

• soil protection and watershed management;

• sustainable urban forestry programs at the local level;

• creation of jobs and income utilizing natural resource

based opportunities.

PROGRAM HIGHLIGHTS

Forest Health Protection

Prominent forest health problems identified in a survey

of State forest lands include: hurricane damage,

construction pressure, and general insects and disease.

Planning for periodic (vs. annualized) FIA was

conducted with the PR DNR, and implementation of

the Urban Forest Effects Model as part of FIA was

planned and budgeted in the 2000 annual budget.

Cooperative Fire Protection

Development of a Wildfire Assessment was begun,

overlaying the mapped history of wildlife on the island

(risk) with fuel types (hazard) and structures (value).

The assessment will not be utilized in future resource

placement and training.

Forest Stewardship

Hispanic landowners numbering 1,360 have become

active in reforestation and management activities on

privately owned forest land. A watershed approach was

taken this past year in that an NRCS-rated priority

watershed received focused technical assistance for the

development of landowner plans for adjacent properties

along the river’s edge. The management plans together

form the basis for a subwatershed management

approach, and the landowners form a coalition for

management of the land as a landscape verses individual

parcels.

Forest Legacy

The priorities for protection are: to conserve and restore

forested watersheds, which are the island’s primary

water supply; buffer zones of existing forest reserves,

natural reserves, and wildlife refuges; conserve forested

areas in primary or close to primary conditions; and

conserve bio-diversity and unique features. Five hundred

twenty-nine acres were acquired fee simple in the

Guanica Legacy Area. In an effort to promote the Forest

Legacy Program, promotional materials were

developed, a landowner survey was conducted, and

contact was made with landowners in each of the forest

legacy areas.

Urban and Community Forestry

A major reforestation program was undertaken by the

central government in Puerto Rico in collaboration with

many partners. Community action in the planting and

care of trees in urban areas was effectively conducted

in every municipality on the island and involved over

19,000 volunteers.

Economic Action

A very strong ecotourism network and local community

planning have been facilitated through the RCA

programs. Support for sustainable management of rural

natural resources has been increased through Rural

Community Assistance projects focused on ecotourism,

planting, and development.

Natural Resource Conservation Education

An ambitious educational component has been

developed that offers to the participants printed materials

written in Spanish (fact sheets and a manual for school

children), planting materials, and the technical and

organizational assistance for school planting projects.

In addition, there are six high schools implementing an

education reform.
31

The cooperative programs are administered and

implemented through a partnership between the U.S.

Virgin Islands, the U.S. Department of Agriculture,

Forest Service, and many other private and government

entities. These programs promote the health and

productivity of the Virgin Island’s forest lands and rural

economies. Emphasis focuses on agroforestry and other

forest products, wildlife, water resources, rural

economies, and conservation practices. The goal is to

maintain and improve the health of the Virgin Islands’

urban and rural forests and related economies. These

programs:

• reduce costs through the use of partnerships;

• increase values through sustained productivity of

forests; and

• are voluntary and nonregulatory in their delivery.

U.S.VI has 21,702.73 acres of privately owned forest

land and 140 acres of experimental forest. Of the 1,000

individuals owning forest lands, most have parcels less

than 20 acres in size. Almost all of these landowners

report that agriculture and aesthetics are primary reasons

for owning forest land. The State’s 112,863 people

include 46 communities with populations over 100; 41

percent of these have initiated some level of community

forestry program.

STATE AND PRIVATE FORESTRY ACCOMPLISHMENTS IN THE U.S.VI

Robin Morgan

Silviculturist

Investment in State’s Cooperative Programs

Programs FY2000 FY2000 FY2001 FY2001 FY2001 FY2001

Title II, VTitle II, V

& VIII & VIII Title IV Title IV

Federal State Federal State

Federal State estimate estimate estimate estimate

(U.S.$) (U.S.$) (U.S.$) (U.S.$) (U.S.$) (U.S.$)

Forest health management 0 0 15,000 15,000

Forest health monitoring 0 0 20,000 0

State fire assistance 70,000 70,000 70,000 70,000 27,000 27,000

Volunteer fire assistance 9,000 9,000 9,000 9,000 21,000 21,000

Forest stewardship program 60,000 60,000 60,000 60,000

Stewardship incentives program 0 0 0 0

Forestry legacy program 0 0 50,000 50,000

Urban and community forestry 138,000 138,000 138,000 138,000

Economic action program 0 0 10,000 10,000

Forest resource information and 0 0 0 0

 analysis

Natural resource conservation 8,000 8,000 8,000 8,000

 education

 Total 285,000 285,000 380,000 360,000
32

Key Issues

Key issues, which the State will address with the 2001

budget, include:

• ecological restoration of natural and built-up areas;

• soil protection and watershed management;

• sustainable urban forestry programs; and

• creation of jobs and income utilizing natural resourcebased opportunities.

PROGRAM HIGHLIGHTS

Forest Health Protection

The pink hibiscus mealybug has infested all lands in the

U.S. Virgin Islands (St. Croix, St. Thomas, and St.

John). Damage was also reported from bark beetles,

aphids, white flies, caterpillars, wilt, shoot borers,

hurricanes, and construction stress. The Department of

Agriculture is working closely with the University of the

Virgin Islands to assess the forest health of all forest

communities in the islands.

Cooperative Fire Protection

The entire territory can be classified as an urban/wildland

interface. There are very few rivers, no lakes, and limited

number of fire hydrants in these islands’ environment.

Variable trade winds, fuel build up associated with dry

tropical forests, narrow and steep roads, and difficult

building entrances all combine to create a wildfire

challenge. The VI Fire Service is working with the VI

Department of Agriculture and the Cooperative

Forest Facts and Accomplishments

Selected facts Selected results

Population (1997) 112,863 Stewardship plans prepared 12

 (current year)

Acres of forest land 21,702 Area under stewardship plans 428

 (current year)

Acres of nonindustrial private forest land 13,824 Area under stewardship plans 428

 (all years)

Number of NIPF landowners, estimated 1,000 Rural acres planted 3

Acres of Federal land under State fire 7,500 Technical assists to private 16

 protection landowners

Acres of private land under State fire 13,824 Acres surveyed for forest 13,824

 protection health

Number of communities 46 Forest health assistance visits 20

State forestry budget $217,000 Communities assisted 46

Technology transfer and 150

 workshops (days)
33

Extension Service to prepare a wildlife protection

assessment.

Forest Stewardship

Matching natural resource professionals with farmers

and other landowners is expanding the vision of forest

stewardship opportunities in the U.S.VI. Recognizing

the importance of protecting habitat for wildlife, soil for

water quality, and aesthetics for tourism and economic

development changes the way people view their

responsibility to utilize the opportunities for maintaining

a healthier island.

Urban and Community Forestry

The urban and community forestry assistance is well

positioned to confer benefits to the people of the Virgin

Islands. The program inspires hope in the community

through the activities of grant subrecipients and

motivated citizens. The traditional tendency to top trees

at the approach of a hurricane is deep rooted. This

program seeks to address these concerns at the local

level by offering training opportunities and technical

assistance in proper tree management.

Economic Action

The VI Resource Conservation and Development

Council is an active partner in delivering this program.

Supporting the existing resources for helping present

and newly developed business, the VI RC&D Council

strives to enhance job and income opportunities, which

utilize the islands’ natural resources.

Natural Resource Conservation Education

The VI ReLeaf program addresses the territory’s need

for new tree planting, tree care information, and

education. Lack of an organized governmental tree-care

policy and land use planning leads to tree removal

without replacement and improper and neglected tree

care on public lands. VI ReLeaf provides trees for

schools and reaches at least 2,000 students annually.

The “Trees for Schools” program teaches students the

benefits of trees and proper tree care, thus encouraging

greater pride in their schools and in their islands.
34

During the course of fiscal year 2000, the International

Cooperation (IC) unit of the IITF was involved in many

projects of benefit to a variety of individuals and

organizations in many countries. Some of the major

highlights of this year follow:

• Developed an agreement with FAO (Food and

Agriculture Organization of the United Nations) for

participation in delivering a training course in

Management of Secondary Forests, held in April in

Dominica.

• Continued to work in Hurricane Georges

Reconstruction Program in the Dominican Republic.

A U.S. $410,000 Inter-Agency Agreement (IAA)

with USAID/WO (United States Agency for

International Development, Washington Office) was

formalized, and the first portion of money transferred

to the IITF. This IAA enabled, among other things,

the sponsorship of 14 Dominicans to an urban

forestry workshop at the University of Puerto Rico

at Mayaguez. Also of note during this term was the

assistance of four Forest Service short-term detailers

that provided technical assistance to Dominican

counterparts in wildfire reconnaissance, firefighting

organization and actual suppression, and timber

salvaging. A report was prepared in English and

translated into Spanish regarding the latter subject

to serve as a case study for future timber salvaging

operations in developing countries in the face of

natural disasters such as hurricanes.

• Developed agreements with FAO and Puerto Rico

Conservation Foundation for preparations of the

10th Caribbean Foresters Meeting. This meeting was

held June 13–16, 2000, in Guyana. The theme for

this year’s meeting was Possibilities and Approaches

to Community Forestry in the Caribbean. There

was representation from 14 countries and a total of

54 participants. IC received requests, and the library

distributed approximately 167 publications to 25

participants of the meeting.

• IC published, and the IITF library distributed, the

proceedings of the ninth Caribbean Foresters

meeting, which was held in the Dominican Republic

in 1998.

• A U.S. Department of Agriculture, Forest Service,

IITF employee, assigned to the IC unit and stationed

in Panama, facilitated the transfer of the canal, and

continues to work on watershed issues of this zone

as well as protected areas management. Also of note

was the development of a signage and interpretation

plan for one of the national parks; partnership

development; production of a research paper for the

San Lorenzo protected area; development of a

mahogany line planting study plan; management of

the AID/Panama environmental portfolio; technical

assistance to the Nicaragua Mission; one-day

workshop for 70 counterparts on the development

of signage plans for national parks; assistance to a

local NGO in preparing a proposal for conservation

training for journalists; assistance to a local NGO in

developing a proposal for an environmental

education program; providing technical assistance to

develop a national campaign for conservation for the

Panama Canal watershed; and conducting an

agroforestry assessment of the AID/Panama

program.

• The IC unit received funds from USAID to fund

followup activities to the low impact logging training

conducted by the Tropical Forest Foundation in

Brazil for Guyanese participants held in November

1999.

• Awarded and managed a U.S. $72,000 contract for

the study for increasing marketing opportunities of

lesser-known wood species and secondary wood

INTERNATIONAL COOPERATION HIGHLIGHTS

Carleen Yocum

Caribbean Environmental Advisor
35

products in Central America and Mexico. During

the fiscal year, this contract produced a report in

Spanish regarding the production potential of smalldiameter trees in Quintana Roo, Mexico.

• IC performed a field review and generated a full

report for an evaluation of the forest concessions

portion of the AID-funded Maya Biosphere Reserve

Project, managed in Guatemala. The report is titled,

1999 Review of Forest Management in

Cooperatives and Community Concessions in the

Maya Biosphere Reserve.

• Completed analysis and input to FAO’s Global

Resource Assessment 2000, and participated in

related meeting in Trinidad.

• Participated in a workshop in Honduras for

environmental considerations in designing rural roads.

• Performed an evaluation for USAID/Nicaragua of

the national forestry agency and developed possible

alternatives for collaboration with the U.S.

Department of Agriculture, Forest Service, and other

opportunities for AID assistance.

• Developed possible options for resolving the Vieques

Island land issue.
36

A. Recent publications of the International

Institute of Tropical Forestry (Numbers

indicate reprints available for distribution).

Publicaciones recientes del Instituto

Internacional de Dasonomía Tropical (Los

números indican disponibilidad de separatas

para distribución).

Alvarez Ruiz, M.; Acevedo Rodriguez, P.;

Vazquez, M. 1997. Quantitative description of

the structure and diversity of the vegetation in

the limestone forest of Río Abajo, AreciboUtuado, Puerto Rico. Acta Científica. 11(1–

3): 21–66.

Arendt, W.J. 1999. Advantages of long-term

avian studies: examples from Puerto Rico. El

Pitirre. 12(2): 62–63.

Arendt, W.J. 2000. Impact of nest predators,

competitors, and ectoparasites on Pearly-eyed

Thrashers, with comments on the potential

implications for Puerto Rican Parrot recovery.

Ornitología Neotropical. 11: 13–63.

Arendt, W.J.; Gibbons, D.W.; Gray, G. 2000.

Status of the volcanically-threatened Montserrat

Oriole, Icterus oberi. Bird Conservation

International. 9: 351–372.

Beissinger, S.R.; Reed, J.M.; Wunderle, J.M. [and

others]. 2000. Report of the AOU conservation

committee on the partners in flight species

prioritization plan. The Auk. 117(2): 549–561.

Chestnut, T.J.; McDowell, W.H. 2000. C and N

dynamics in the riparian and hyporheic zones

of a tropical stream, Luquillo Mountains, Puerto

Rico. Journal of the North American

Benthological Society. 19(2): 199–214.

Chestnut, T.J.; Zarin, D.J.; McDowell, W.H.;

Keller, M. 1999. A nitrogen budget for latesuccessional hillslope tabonuco forest, Puerto

Rico. Biogeochemistry. 46(1–3): 85–108.

001 Chinea, J.D. 1999. Changes in the herbaceous and

vine communities at the Bisley Experimental

Watersheds, Puerto Rico, following Hurricane

Hugo. Canadian Journal of Forestry Research.

29(9): 1433–1437.

DeJaco, C.E. 2000. Characterization and analysis

of algal communities throughout a watershed in

Puerto Rico. Louisville, KY: University of

Louisville. [Number of pages unknown]. M.S.

thesis.

Despiau Batista, A. 1997. Distribución de las

especies arbóreas de acuerdo al gradiente en

topografía en el bosque de Río Piedras, Puerto

Rico, luego de 60 años de abandono agrícola.

Acta Científica. 11(1–3): 3–20.

Domínguez Cristóbal, C.M. 1997. El inventario

de los montes públicos de Puerto Rico (1870).

Acta Científica. 11(1–3): 93–95.

Domínguez Cristóbal, C.M. 1998. El traspaso del

manglar: de la Comandancia de la Marina a la

Inspección de Montes. Acta Científica. 12(1–

3): 67–72.

Domínguez Cristóbal, C.M. 2000. Panorama

histórico forestal de Puerto Rico. San Juan,

PR: Editorial de la Universidad de Puerto Rico.

680 p.

Engel, V.L.; Parrotta, J.A. 2000. Restauraçao de

ecossistemas florestais. Agroecologia (Brazil).

1(4): 22–23.

Eusse, A.M.; Aide, T.M. 1999. Patterns of litter

production across a salinity gradient in a

Pterocarpus officinalis tropical wetland. Plant

Ecology. 145(2): 307–315.

APPENDIX 1: ANNUAL LETTER 1999-2000 PUBLICATIONS LIST
37

Ewel, J.J.; O’Dowd, D.J.; Bergelson, J. [and

others]. 1999. Deliberate introductions of

species: research needs. BioScience. 49(8):

619–630.

Fernández, D.S.; Myster, R.W. 1995. Temporal

variation and frequency distribution of

photosynthetic photon flux densities on

landslides in Puerto Rico. Tropical Ecology.

36(1): 73–87.

Fetcher, N.; Cordero, R.A.; Votzow, J. 2000.

Lack of ecotypic differentiation: plant response

to elevation, population origin, and wind in the

Luquillo Mountains, Puerto Rico. Biotropica.

32(2): 225–234.

Francis, J.K. 2000. Comparison of hurricane

damage to several species of urban trees in San

Juan, Puerto Rico.Journal of Arboriculture.

26(4): 189–197.

Francis, J.K. 2000. Hibiscus elatus Sw., 1788,

Hoher Linden-Eibisch, Familie: Malvaceae, III–

4. In: Enzyklopadie der Holzgewachse.

Landsberg, Germany: Ecomed

Verlagsgesellschaft. 6 p. Vol. 19.

Genet, J.A., III. 1999. The diversity and

abundance of termites (Isoptera) in a

fragmented subtropical dry forest landscape.

Lansing, MI: Michigan State University,

Department of Zoology. [Number of pages

unknown]. M.S. thesis.

Genet, K.S. 1999. The resiliency of lizard

communities to habitat fragmentation in dry

forests of southwestern Puerto Rico. Lansing,

MI: Michigan State University, Department of

Zoology. [Number of pages unknown]. M.S.

thesis.

002 González-Caban, A.; Loomis, J. 2000. Measuring

the economic benefit of maintaining the

ecological integrity of the Río Mameyes in

Puerto Rico. Res. Pap. PSW–RP–240.

Berkeley, CA: U.S. Department of Agriculture,

Forest Service, Pacific Southwest Research

Station. 61 p.

003 González-Caban, A.; Loomis, J. 2000. Medida

de los beneficios económicos de la integridad

ecológica del Río Mameyes en Puerto Rico.

Res. Pap. PSW–RP–240. Berkeley, CA: U.S.

Department of Agriculture, Forest Service,

Pacific Southwest Research Station. 62 p.

González, G.; Zou, X. 1999. Plant and litter

influences on earthworm abundance and

community structure in a tropical wet forest.

Biotropica. 31(3): 486–493.

004 Helmer, E.H. 2000. The landscape ecology of

tropical secondary forest in montane Costa Rica.

Ecosystems. 3: 98–114.

Helmer, E.H.; Brown, S. 2000. Gradient analysis

of biomass in Costa Rica and a first estimate of

trace gas emissions from biomass burning. In:

Hall, C.A.S., ed. Quantifying Sustainable

Development. New York: Academic Press:

503–526.

005 Helmer, E.H., Brown, S.; Cohen, W.B. 2000.

Mapping montane tropical forest successional

stage and land use with multi-date Landsat

imagery. International Journal of Remote

Sensing. 21(11): 2163–2183.

Hendrix, P.F.; Lachnicht, S.L.; Callaham, M.A.,

Jr.; Zou, X. 1999. Stable isotopic studies of

earthworm feeding ecology in tropical

ecosystems of Puerto Rico. Rapid

Communications in Mass Spectrometer. 13:

1295–1299.

Johnson, S.L.; Covich, A.P. 2000. The importance

of night-time observations for determining

habitat preferences of stream biota. Regulated

Rivers: Research andManagement. 16: 91–99.

Jones, T.H.; Flournoy, R.C.; Torres, J.A. [and

others]. 1999. 3-Methyl-4-phenylpyrrole from

the ants Anochetus kempfi and Anochetes
38

mayri. Journal of Natural Products. 62(9):

1343–1345.

Jones, T.H.; Wojciechowski, T.J.; Snelling; R.R.

[and others]. 1999. Dialkylpyrrolidines from the

ants Megalomyrmex cyendyra Brandao and

M. latreillei Emery. Caribbean Journal of

Science. 35(3–4): 310–311.

Keenan, R. J.; Lamb, D.J.; Parrotta, J.A.;

Kikkawa, J. 1999. Ecosystem management in

tropical timber plantations: satisfying economic,

conservation, and social objectives. Journal of

Sustainable Forestry. 9(1–2): 117–134.

Keller, M.; Weitz, A.M.; Bryan, B. [and others].

2000. Soil-atmosphere nitrogen oxide fluxes:

effects of root disturbance. Journal of

Geophysical Research. 105(D14): 17,693–

17,698.

Lewis, W.M, Jr.; Melack, J.M.; McDowell, W.H.

[and others]. 1999. Nitrogen yields from

undisturbed watersheds in the Americas.

Biogeochemistry. 46: 149–162.

Lugo, A.E., ed. 1997. Acta Científica. 11(1–3).

San Juan, PR: Asociación de Maestros de

Ciencia de Puerto Rico. 102 p.

Lugo, A.E., ed. 1998. Acta Científica. 12(1–3).

San Juan, PR: Asociación de Maestros de

Ciencia de Puerto Rico. 72 p.

Lugo, A.E. 1998. Mangrove forests: a tough

system to invade but an easy one to rehabilitate.

Marine Pollution Bulletin. 37: 8–12.

Lugo, A.E. 1999. Mangroves. In: Nellis, D.W.,

comp., ed. Puerto Rico & Virgin Islandswildlife

viewing guide. Helena, Montana: Falcon Press.

21 p.

Lugo, A.E. 1999. El manglar: un ecosistema al

servicio del ser humano. In: Colección CEA,

Universidad de Buenos Aires. Buenos Aires,

Argentina: Editorial Universitaria de Buenos

Aires (EUDEBA), Centro de Estudios

Avanzados, Universidad de Buenos Aires:

361–382. Vol. 24.

Lugo, A.E. 1999. Mangrove ecosystem research

with emphasis on nutrient cycling. In: YáñezArancibia, A.; Lara-Domínguez, A.L.,comps.,

eds. Ecosistemas de manglar en América

tropical [Mangrove ecosystems in tropical

America]. Veracruz, México: Instituto de

Ecología, A.C., also UICN/ORMA Costa Rica

and NOAA/NMFS Silver Spring, MD: 17–38.

Lugo, A.E. 1999. Will concern for biodiversity spell

doom to tropical forest management? The

Science of the Total Environment. 240: 123–

131.

Lugo, A.E., ed. 2000. Annual letter 1996–97. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, International Institute of Tropical

Forestry. 185 p.

006 Lugo, A.E., ed. 2000. Annual letter 1997–1998.

Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, International

Institute of Tropical Forestry. 84 p.

007 Lugo, A.E., ed. 2000. Annual Letter 1998–99.

Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, International

Institute of Tropical Forestry. 143 p.

Lugo, A.E. 2000. Book review: Climate change

and tropical forests—uncertainty and fear.

Journal of Environmental Quality. 29(1): 257.

Lugo, A.E.; Brown, S. 1997. Tropical forests:

supporting Latin America and the world into

the 21st century. Acta Científica. 11(1–3): 97–

102.

Lugo, A.E., Baron, J.S.; Frost, T.P. [and others].

1999. Ecosystem processes and functioning. In:
39

Johnson, N.C.; Molk, A.J.; Sexton, W.T.; Zaro,

R.C., eds. Ecological stewardship: a common

reference for ecosystem management. New

York: Elsevier Science: 219–254. Vol. 2.

Lugo, A.E.; Brown. S. 1999. Biodiversity in the

Caribbean: concepts, assessment, management,

and monitoring at the landscape level. In:

Yocum, C.; Lugo, A.E., comps., eds.

Proceedings of the ninth meeting of Caribbean

foresters: Biodiversity in the Caribbean:

management and benefits; 1998 June 1–5;

Santo Domingo, DR. San Juan, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry: 92–

98.

Lugo, A.E.; Brown, S.L.; Dodson, R. [and others].

1999. The Holdridge life zones of the

conterminous United States in relation to

ecosystem mapping. Journal ofBiogeography.

26: 1025–1038.

Lugo, A.E.; Sell, M.; Snedaker, S.C. 1999.

Mangrove ecosystem analysis. In: YáñezArancibia, A.; Lara-Domínguez, A.L.,comps.,

eds. Ecosistemas de manglar en América

tropical [Mangrove ecosystems in tropical

America]. Veracruz, México: Instituto de

Ecología, A.C., also UICN/ORMA Costa Rica

and NOAA/NMFS Silver Spring, MD: 343–

364.

Lugo, A.E.; Figueroa Colon, J.; Scatena, F.N.

2000. The Caribbean. In: Barbour, M.G.;

Billings, W.D., eds. North American Terrestrial

Vegetation. 2nd ed. New York: Cambridge

University Press: 594–622. Chapter 16:

Lugo, A.E.; Gucinski, H. 2000. Function, effects,

and management of forest roads. Forest

Ecology and Management. 133: 249–262.

 Lugo, A.E.; Rogers, C.S.; Nixon, S.W. 2000.

Hurricanes, coral reefs and rainforests:

resistance, ruin and recovery in the Caribbean.

Ambio. 29(2): 106–114.

 Marley, D.P. 1998. Spatial modeling of climate

and photosynthesis in the Luquillo Mountains,

Puerto Rico. Syracuse, New York: State

University of New York, College of

Environmental Science and Forestry. [Number

of pages unknown]. M.S. thesis.

 Matson, P.A.; McDowell, W.H.; Townsend,

A.R.; Vitousek, P.M. 1999. The globalization

of N deposition: ecosystem consequences in

tropical environments. Biogeochemistry. 46(1–

3): 67–83.

 Medina, E. 1998. Plasticity of crassulacean acid

metabolism (CAM) and biomass allocation of

the pantropical Kalanchoe pinnata (Lam.)

Pers. Acta Científica. 12(1–3): 59–66.

 Morgan, N.R. 1998. Aspectos financieros y

económicos del enverdecimiento urbano. In:

Krishnamurthy, L.; Nascimento, J.R., comps.,

eds. Áreas verdes urbanas enLatinoamérica y

el Caribe. Chapingo, México: Universidad

Autónoma de Chapingo, Centro de

Agroforestería para el Desarrollo Sostenible:

139–178.

Murphy, P.G. 1997. Footprints on the forest trail:

a tribute to the career and accomplishments of

Dr. Frank H. Wadsworth. Acta Científica. 11(1–

3): 85–87.

 Parresol, B.R.; Alemañy, S. 1998. Analysis of tree

damage from Hurricane Hugo in the Caribbean

National Forest, Puerto Rico. In: Waldrop, T.A.,

comp., ed. Proceedings of the ninth biennial

southern silvicultural research conference; [Date

of meeting unknown]; [Place of meeting

unknown]. Asheville, NC: U.S. Department of

Agriculture, Forest Service, Southern Research

Station: 599–603.

008 Parrotta, J.A. 1999. Productivity, nutrient cycling,

and succession in single-and mixed-species

plantations of Casuarina equisetifolia,

Eucalyptus robusta, and Leucaena
40

leucocephala in Puerto Rico. Forest Ecology

and Management. 124(1): 45–77.

Pringle, C.M.; Hemphill, N.; McDowell, W.H.

[and others]. 1999. Linking species and

ecosystems: different biotic assemblages cause

interstream differences in organic matter.

Ecology. 80(6): 1860–1872.

 Reagan, D.P.; Waide, R.B., comps., eds. 1996.

The food web of a tropical rain forest. Chicago:

University of Chicago Press.616 p. [The works

herein were done with substantial funding from

an NSF grant for an LTER project where the

IITF is one of the two recipients.]

 Richardson, B.A.; Richardson, M.J.; Scatena,

F.N.; McDowell, W.H. 2000. Effects of nutrient

availability and other elevational changes on

bromeliad populations and their invertebrate

communities in a humid tropical forest in Puerto

Rico. Journal ofTropical Ecology. 16(2): 167–

188.

Richardson, B.A.; Rogers, C.; Richardson, M.J.

2000. Nutrients, diversity, and community

structure of two phytotelm systems in a lower

montane forest, Puerto Rico. Ecological

Entomology. 25: 348–356.

 Sánchez, A.; Rodríguez, A.; Valenzuela, O.;

Weaver, P.L. 1999. Growth and acclimatization

of nine forest species in the Dominican

Republic. In: Yocum, C.; Lugo, A.E., comps.,

eds. Proceedings of the ninth meeting of

Caribbean foresters. Biodiversity in the

Caribbean: management and benefits; 1998

June 1–5; Santo Domingo, DR. San Juan, PR:

U.S. Department of Agriculture, Forest Service,

International Institute of Tropical Forestry: 96–

106.

Santiago, M.; Ramírez, J.L.; Francis, J. 2000.

Proyecto: Inventario de flora en el sector Cueva

del Indio, Barrio Quebrada Arena, Las Piedras,

PR. San Juan, PR: U.S. Department of

Agriculture, Forest Service, International

Institute of Tropical Forestry. [number of pages

unknown].

 Santos-Román, D.M. 1999. Multivariate analysis

of water quality and physical characteristics of

selected watersheds in Puerto Rico. Storrs, CT:

University of Connecticut. [Number of pages

unknown]. M.S. thesis.

Scatena, F.N.; Rivera-Costa, M.M.; Johnson, S.

[and others]. 2000. Bibliography of tropical

stream ecology and instream flow related to the

Caribbean region. San Juan, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry and

Long Term Ecological Research (LTER) and

UNESCO - ihp/phi, FRIEND - AMIGO,

CARIBBEAN – CARIBE. 31 p.

 Schellekens, J.; Bruijnzeel, L.A.; Scatena, F.N.

[and others]. 2000. Evaporation from a tropical

rain forest, Luquillo Experimental Forest,

eastern Puerto Rico. Water Resources

Research. 36(8): 2183–2196.

 Schellekens, J.; Scatena, F.N.; Bruijnzeel, L.A.;

Wickel, A.J. 1999. Modeling rainfall

interception by a lowland tropical rain forest in

northeastern Puerto Rico. Journal ofHydrology.

225(3–4): 168–184.

Schellekens, J. 2000. Hydrological processes in

a humid tropical rain forest: a combined

experimental and modeling approach.

Amsterdam: Vrije Universiteit. [Number of

pages unknown]. Ph.D. thesis.

 Schellekens, J.; Scatena, F.N.; Bruijnzeel, L.A.

[and others]. 2000. Stormflow generation in a

small rain forest catchment in the Luquillo

Experimental Forest, Puerto Rico. In:

Schellekens, J., ed. Hydrological processes in

a humid tropical rain forest: a combined

experimental and modeling approach.

Amsterdam: Vrije Universiteit: 63–92.
41

Silver, W.L.; Neff, J.; McGroddy, M. [and others].

2000. Effects of soil texture on belowground

carbon and nutrient storage in a lowland

Amazonian forest ecosystem. Ecosystems. 3:

193–209.

 Sullivan, N.H.; Bowden, W.B.; McDowell, W.H.

1999. Short-term disappearance of foliar litter

in three species before and after a hurricane.

Biotropica. 31(3): 382–393.

009 Torres, J.A.; Medina Gaud, S. 1998. Los insectos

de Puerto Rico. Acta Cientifica. 12(1–3): 3–

41.

010 Torres, J.A.; Santiago, M.; Salgado, M. 1999. The

effects of the fungus-growing ant,

Trachymyrmex jamaicensis, on soil fertility

and seed germination in a subtropical dry forest.

Tropical Ecology. 40(2): 237–245.

011 Torres, J.A.; Thomas, R.; Leal, M.; Gush, T. 2000.

Ant and termite predation by the tropical

blindsnake Typhlops platycephalus. Insectes

Socieaux. 47: 1–6.

Wadsworth, F.H., ed. 1998. ISTF Noticias 19(4).

Bethesda, MD: International Society of Tropical

Foresters. 7 p.

 Wadsworth, F.H., ed. 1999. ISTF News 20(2).

Bethesda, MD: International Society of Tropical

Foresters. 7 p.

 Wadsworth, F.H., ed. 1999. ISTF News 20(3).

Bethesda, MD: International Society of Tropical

Foresters. 6 p.

 Wadsworth, F.H. 2000. Book review: Managing

multiple uses on national forests, 1905 to 1995:

a 90-year learning experience and it isn’t finished

yet. Environmental History. 5(1): 112–113.

 Wadsworth, F.H. 2000. Book review: Reduced

impact logging in the tropical rain forest of

Guyana. ISTF News 21(1). p.10.

Waide, R.B.; Willig, M.R.; Steiner, C.F. [and

others]. 1999. The relationship between

productivity and species richness. Annual

Review of Ecology and Systematics. 30: 257–

300.

Walker, L.R. 2000. Seedling and sapling dynamics

of treefall pits in Puerto Rico. Biotropica. 32(2):

262–275.

 Walker, R.; Moran, E.; Anselin, L. 2000.

Deforestation and cattle ranching in the Brazilian

Amazon: external capital and household

processes. World Development. 28(4): 683–

699.

 Walsh-McGehee, M.; Lee, D.S.; Claridge, D.

1999. Distribution and population status of

White-tailed Tropicbirds nesting in the

Bahamas. Bahama Journal of Science. 6(2): 44–

48.

 Walsh-McGehee, M.; Lee, D.S.; Haney, J.C.

1998. Kirtland’s warblers in the Bahama

archipelago: a compilation of records and

reports and a bibliography of the species on its

wintering grounds. In: Proceedings of the

seventh symposium on the naturalhistory of the

Bahamas; [Date unknown]; [Location of

proceedings unknown].San Salvador, Bahamas:

Bahamian Field Station: 199–131.

012 Weaver, P.L. 2000. Elfin woodland recovery 30

years after a plane wreck in Puerto Rico’s

Luquillo Mountains. Caribbean Journal of

Science. 36(1–2): 1–9.

013 Weaver, P.L. 2000. Environmental gradients affect

forest structure in Puerto Rico’s Luquillo

Mountains. Interciencia. 25(5): 254–259.

014 Weaver, P.L.; Bauer, G.P. 2000. Major Meliaceae

in Nicaragua. Gen. Tech. Rep. IITF–10. San

Juan, PR: U.S. Department of Agriculture,

Forest Service, International Institute of Tropical

Forestry. 38 p.
42

Weitz, A.M.; Keller, M.; Linder, E.; Crill, P.M.

1999. Spatial and temporal variability of nitrogen

oxide and methane fluxes from a fertilized tree

plantation in Costa Rica. Journal of Geophysical

Research. 104: 30,097–30,107.

 Wiley, J.; Wunderle, J.; Smith, A. [and others].

2000. Puerto Rican Parrot (Amazona vittata).

In: Snyder, N.; McGowan, P.; Gilardi, J.; Grajal,

A., eds. Parrots: statussurvey and conservation

action plan 2000–2004. Oxford, UK:

International Union for Conservation (IUCN):

117–118.

 Wunderle, J.M., Jr. 1999. Conservation of

Caribbean wildlife diversity: the role of

agroforestry plantations. In: Yocum, C.; Lugo,

A.E., ed. Proceedings of the ninth meeting of

Caribbean foresters. Biodiversity in the

Caribbean: management and benefits; 1998

June 1–5; Santo Domingo, DR. San Juan, PR:

U.S. Department of Agriculture, Forest Service,

International Institute of Tropical Forestry: 115–

120.

 Wunderle, J.M., Jr. 1999. Introductions to

hurricanes and wildlife; migrant birds in the

Caribbean; exotic birds. In: Nellis, D.W.,comp.,

ed. Puerto Rico & Virgin Islandswildlife viewing

guide. Helena, MT: Falcon Press: 22, 27, 41.

015 Wunderle, J.M., Jr. 1999. Pre- and post-hurricane

fruit availability: implications for Puerto Rican

Parrots in the Luquillo Mountains. Caribbean

Journal of Science. 35(3–4): 1-14.

 Wunderle, J.M, Jr.; Latta, S.C. 1999. Birds in

sun and shade coffee plantations in the

Cordillera Central, Dominican Republic:

implications for conservation. El Pitirre. 12(2):

56.

 Wunderle, J.M, Jr.; Latta, S.C. 2000. Winter site

fidelity of Nearctic migrant birds in shade coffee

plantations of different sizes in the Dominican

Republic. Auk. 117: 596–614.

016 Yocum, C.; Lugo, A.E., eds. 1999. Biodiversity

in the Caribbean: management and benefits:

Proceedings of the ninth meeting of Caribbean

foresters; 1998 June 1–5; Santo Domingo, DR.

San Juan, PR: U.S. Department of Agriculture,

Forest Service, International Institute of Tropical

Forestry. 130 p.

B. Other publications still available

 Otras publicaciones disponibles aún

017 Various authors. 1985–1999. IITF Silvics Manual.

SO–ITF–SM 1–88. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry:

sectionally numbered. [complete silvics manual

available in English, has detailed silvicultural

descriptions for 88 species of trees, e.g.,

Swietenia macrophylla, Khaya senegalensis,

Pterocarpus macrocarpus, others].

018 Acevedo-Rodríguez, P. en colaboración con

Woodbury, R.O. 1985. Los bejucos de Puerto

Rico. Gen. Tech. Rep. SO–58. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, Southern Forest Experiment Station,

Institute of Tropical Forestry. 331 p. Vol. 1.

019 Anderson, R.L.; Birdsey, R.A.; Barry, P.J. 1982.

Incidence of damage and cull in PuertoRico’s

timber resource, 1980. Res. Bull. SO–88. New

Orleans: U.S. Department of Agriculture, Forest

Service, Southern Forest Experiment Station,

Institute of Tropical Forestry. 13 p.

020 Arendt, W.J. 1988. Range expansion of the cattle

egret (Bubulcus ibis) in the Greater Caribbean

Basin. Colonial Waterbirds. 11(2): 252–262.

021 Arendt, W.J.; Arendt, A.I. 1988. Aspects of the

breeding of the cattle egret (Bubulcus ibis) in

Monserrat, West Indies, and its impact on nest

vegetation. Colonial Waterbirds. 11(1): 72–84.
43

022 Barres, H. 1964. Rooting media for growing pine

seedlings in hydroponic culture. Res. Note 2.

Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Institute of Tropical

Forestry. 4 p.

023 Bauer, G.P.; Gillespie, A.J.R. 1990. Volume tables

for young plantation-grown hybrid mahogany

(Swietenia macrophylla x S. mahagoni) in the

Luquillo Experimental Forest of Puerto Rico.

Res. Pap. SO–257. New Orleans: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry.8 p.

024 Benstead, J.P.; March, J.G.; Pringle, C.M.;

Scatena, F.N. 1999. Effects of a low-head dam

and water abstraction on migratory tropical

stream biota. Ecological Applications. 9(2):

656–668.

025 Birdsey, R.A.; Jimenez, D. 1985. The forests of

Toro Negro. Res. Pap. SO–222. New Orleans:

U.S. Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry. 29 p.

026 Birdsey, R.A.; Weaver, P.L. 1987. Forest area

trends in Puerto Rico. Res. Note SO–331.

New Orleans: U.S. Department of Agriculture,

Forest Service, Southern Forest Experiment

Station, Institute of Tropical Forestry. 5 p.

027 Birdsey, R.A.; Weaver, P.L.; Nicholls, C.F. 1986.

The forest resources of St. Vincent, West Indies.

Res. Pap. SO–229. New Orleans: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry. 25 p.

028 Briscoe, C.B. 1962. Crecimiento en diámetro de

los árboles en los cerros de la región caliza seca.

Apuntes Forestales Tropicales ITF-12. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Tropical Forest Research

Center. 2 p.

029 Briscoe, C.B. 1962. Tree diameter growth in the

dry limestone hills. Trop. For. Note ITF–12.

Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Tropical Forest

Research Center. 2 p.

030 Briscoe, C.B.; Nobles, R.W. 1962. Height and

growth of mahogany seedlings. Trop. For. Note

ITF–13. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Tropical Forest

Research Center. 2 p.

031 Chinea, J.D.; Beymer, R.J.; Rivera, C. [and others].

1993. An annotated list of the flora of Bisley

area, Luquillo Experimental Forest, Puerto

Rico, 1987 to 1992. Gen. Tech. Rep. SO–94.

New Orleans: U.S. Department of Agriculture,

Forest Service, Southern Forest Experiment

Station, International Institute of Tropical

Forestry. 12 p.

032 Chudnoff, M.; Goytía, E. 1967. The effect of

incising on drying, treatability, and bending

strength of fence posts. Res. Pap. ITF–5. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Institute of Tropical Forestry.

20 p.

033 Chudnoff, M.; Maldonado, E.D.; Goytía, E. 1966.

Solar drying of tropical hardwoods. Res. Pap.

ITF–2. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Institute of Tropical

Forestry. 26 p.

034 Crow, T.R.; Weaver, P.L. 1977. Tree growth in a

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.

035 Englerth, G.H.; Maldonado, E.D. 1961. Bamboo

for fence posts. Trop. For. Note 6. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, Tropical Forest Research Center. 2 p.

[Disponible en español como Apuntes

Forestales ITF–6].
44

036 Ewel, J.J.; Whitmore, J.L. 1973. The ecological

life zones of Puerto Rico and the United States

Virgin Islands. Res. Pap.ITF–18. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, Institute of Tropical Forestry. 72 p.

037 Figueroa Colón, J.C. 1996. Phytogeographical

trends, centers of high species richness and

endemism, and the question of extinctions in the

native flora of Puerto Rico. In: Figueroa Colón,

J.C., ed. The scientific survey of Puerto Rico

and the Virgin Islands. The annals of the New

York academy of sciences. New York: The

Academy: 89–102. Vol. 776.

038 Figueroa Colón, J.C.; Wadsworth, F.H.; Branham,

S., eds. 1987. Management of the forests of

tropical America: prospects and technologies:

Proceedings of a conference; 1986 September

22-27; San Juan, PR. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry. 469 p.

039 Francis, J.K. 1989. Merchantable volume and

weights of mahoe in Puerto Rican plantations.

Res. Note SO–355. New Orleans: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry. 4 p.

040 Francis, J.K. 1994. English-Portuguese equivalents

of forestry and conservation terms. [Termos

equivalentes em silvicultura e conservaçao

Portugués-Inglés]. Gen. Tech. Rep. SO–109.

New Orleans: U.S. Department of Agriculture,

Forest Service, Southern Forest Experiment

Station, International Institute of Tropical

Forestry. 45 p.

041 Francis, J.K. 1998. Tree species for planting in

forest, rural, and urban areas of Puerto Rico.

Gen. Tech. Rep. GTR–IITF–3. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical

Forestry. 82 p.

042 Francis, J.K. 1999. Especies forestales para plantar

en áreas forestales, rurales y urbanas de Puerto

Rico. Gen. Tech. Rep. IITF–13. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical

Forestry. 88 p.

043 Francis, J.K.; Alemañy, S.; Lioger, H.A.; Proctor,

G.R.1998. The flora of Cañon San Cristóbal,

Puerto Rico. Gen. Tech. Rep. IITF–4. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, International Institute of Tropical

Forestry. 37 p.

044 Francis, J.K.; Gillespie, A.J.R. 1993. Relating gust

speed to tree damage in Hurricane Hugo, 1989.

Journal of Arboriculture. 19(6): 369–373.

045 Francis, J.K.; Liogier, H.A. 1991. Naturalized

exotic tree species in Puerto Rico. Gen. Tech.

Rep. SO–82. New Orleans: U.S. Department

of Agriculture, Forest Service, Southern Forest

Experiment Station, Institute of Tropical

Forestry. 12 p.

046 Francis, J.K.; Parresol, B.R.; de Patino, J.M. 1996.

Probability of damage to sidewalks and curbs

by street trees in the tropics. Journal of

Arboriculture. 22(4): 193–197.

047 Francis, J.K.; Rivera, C.; Figueroa, J. 1994.

Toward a woody plant list for Antigua and

Barbuda: past and present. Gen. Tech. Rep.

SO–GTR–102. New Orleans: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station,

International Institute of Tropical Forestry. 28

p.

048 Francis, J.K.; Rodriguez, A. 1993. Seeds of Puerto

Rican trees and shrubs: second installment. Res.

Note SO–374. New Orleans: U.S. Department

of Agriculture, Forest Service, Southern Forest

Experiment Station, Institute of Tropical

Forestry. 5 p.
45

049 Franco, P.A.; Weaver, P.L.; Eggen MacIntosh, S.

1997. Forest resources of Puerto Rico. 1990.

Res. Bull. SRS–22. Asheville, NC: U.S.

Department of Agriculture, Forest Service,

Southern Research Station. 45 p.

050 Frangi, J.L.; Lugo, A.E. 1991. Hurricane damage

to a flood plain forest in the Luquillo Mountains

of Puerto Rico. Biotropica. 23(4a): 324–335.

051 Fu, S.; Rodríguez Pedraza, C.; Lugo, A.E. 1996.

A twelve-year comparison of stand changes in

a mahogany plantation and a paired natural

forest of similar age. Biotropica. 28(4a): 515–

524.

052 García-Martinó, A.R.; Scatena, F.N.; Warner,

G.S.; Civco, D.L. 1996. Statistical low flow

estimation using GIS analysis in humid montane

regions in Puerto Rico. Water Resources

Bulletin. (Journal of the American Water

Resources Association). 32(94): 413–424.

053 García-Martinó, A.R.; Warner, G.S.; Scatena,

F.N.; Civco, D.L. 1996. Rainfall, runoff and

elevation relationships in the Luquillo Mountains

of Puerto Rico. Caribbean Journal of Science.

32(4): 413–424.

054 Gillespie, A.J.R. 1994. Remote sensing for tropical

forest assessment. [Evaluación de bosques

tropicales utilizando la técnica telesensorial].

Proceedings of a workshop; 1991 April 8–12;

San Juan, PR. Río Piedras, PR and Paris: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry and

UNESCO. 84 p.

055 Jones, S.C.; Nalepa, C.A.; McMahan, E.A.;

Torres, J.A. 1995. Survey and ecological

studies of the termites (Isoptera:

Kalotermitidae) of Mona Island. Florida

Entomologist. 78(2): 305–313.

056 Keller, M.; Clark, D.A.; Clark, D.B. [and others].

1996. If a tree falls in the forest. Science.

273: 201.

057 Kejerfve, B.; de Lacerda, L.D.; Diop, E.H.S., eds.

1997. Mangrove ecosystem studies in Latin

America and Africa. Cosponsored by

UNESCO, International Society for Mangrove

Ecosystems; and U.S. Department of

Agriculture, Forest Service, International

Institute of Tropical Forestry. Paris: UNESCO.

349 p.

058 Kjerfve, B.1998. CARICOMP – Caribbean

Coral Reefs, Seagrass and Mangrove Sites.

Coastal Region and Small Islands Papers 3.

Paris: UNESCO. 345 p.

059 Knowles, O.H.; Parrotta, J.A. 1997. Phenological

observations and tree seed characteristics in an

equatorial moist forest at Trombetas, Para State,

Brazil. In: Lieth, H., comp., ed. Phenology in

seasonal climates I. Leiden, Netherlands:

Backhuys: 67–84.

060 Latta, S.C.; Wunderle, J.M., Jr. 1996. The

composition and foraging ecology of mixed

species flocks in pine forests of Hispaniola. The

Condor. 98: 595–607.

061 Latta, S.C.; Wunderle, J.M., Jr. 1996. Ecological

relationships of two todies in Hispaniola: effects

of habitat and flocking. The Condor. 98: 769–

779.

062 Ledig, F.T.; Whitmore, J.L. 1981. The calculation

of selection differential and selection intensity

to predict gain in a tree improvement program

for plantation grown Honduras pine in Puerto

Rico. Res. Pap. SO–170. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

Institute of Tropical Forestry. 7 p.

063 Liegel, L.H. 1984. Effects of adding magnesium

nitrate before dry-ashing on phosphorus in

Honduras pine foliage. Journal of Agriculture

of the University of Puerto Rico. 68(2): 219–

221.

064 Liegel, L.H. 1984. Results of 5-to 6-year-old

provenance trials of Pinus oocarpa Schiede
46

on eight sites in Puerto Rico. Silvae Genetica.

33(6): 223–230.

065 Liegel, L.H. 1984. Status, growth, and

development of unthinned Honduras pine

plantations in Puerto Rico. Turrialba. 34(3):

313–324.

066 Liegel, L.H. 1990. Didymopanax morototoni

(Aubl.) Decne. & Planch. Yagrumo macho.

Araliaceae. Ginseng family. In: Burns, R.M.;

Honkala, B.H., comps., eds. Silvics of North

America: Hardwoods. Agric. Handb. 654.

Washington, DC: U.S. Department of

Agriculture: 288–293. Vol. 2.

067 Liegel, L.H., comp. 1991. Growth and site

relationships of Pinus caribaea across the

Caribbean Basin. Gen. Tech. Rep. SO–83. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Southern Forest Experiment

Station, Institute of Tropical Forestry. in

cooperation with: The University of Puerto

Rico. 70 p.

068 Liegel, L.H.; Stead, J.W. 1990. Cordia alliodora

(Ruiz & Pav.) Oken. Laurel, Capá Prieto.

Boraginaceae. Borage family. In: Burns, R.M.;

Honkala, B.H., comps., eds. Silvics of North

America: Hardwoods. Agric. Handb. 654.

Washington, DC: U.S. Department of

Agriculture: 270–277. Vol. 2.

069 Liegel, L.H.; Venator, C.R. 1987. A technical guide

for forest nursery management in theCaribbean

and Latin America. Gen. Tech. Rep. SO–67.

New Orleans: U.S. Department of Agriculture,

Forest Service, Southern Forest Experiment

Station, Institute of Tropical Forestry. 156 p.

070 Little, E.L., Jr.; Woodbury, R.O.; Wadsworth, F.H.

1976. Flora of Virgin Gorda (British Virgin

Islands). Res. Pap. ITF–21. Río Piedras, PR:

U.S. Department of Agriculture, Forest Service,

Institute of Tropical Forestry. 36 p.

071 Lodge, D.J.; Scatena, F.N.; Asbury, C.E.;

Sanchez, M.J. 1991. Fine litterfall and related

nutrient inputs resulting from Hurricane Hugo in

subtropical wet and lower montane rain forests

of Puerto Rico. Biotropica. 23(4a): 336–342.

072 Lugo, A.E. 1988. Forest lands in Puerto Rico and

the Caribbean 500 years after their discovery

by Cristoforo Colombo. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station.16 p. [slide

presentation with references and appendix]

073 Lugo, A.E. 1992. Comparison of tropical tree

plantations with secondary forests of similar age.

Ecological Monographs. 62(1): 1–41.

074 Lugo, A.E. 1992. Tree plantations for rehabilitating

damaged forest lands in the tropics. In: Wali

Mohan, K., ed. Ecosystem Rehabilitation:

Ecosystem Analysis and Synthesis.The Hague,

The Netherlands: SPB Academic Publishing:

247–255. Vol. 2.

075 Lugo, A.E. 1994. Preservation of primary forests

in the Luquillo Mountains, Puerto Rico.

Conservation Biology. 8(4): 1122–1131.

077 Lugo, A.E. 1996. Ninety years of plant ecology

research in Puerto Rico. In: Figueroa, J.C., ed.

The Scientific Survey of Puerto Rico and the

Virgin Islands. Annals of the New York

Academy of Sciences. New York: The

Academy: 73–88. Vol. 776.

078 Lugo, A.E. 1997. Old-growth mangrove forests

in the United States. Conservation Biology.

11(1): 11–20.

079 Lugo, A.E. 1999. Point-counterpoints on the

conservation of big-leaf mahogany. Gen. Tech.

Rep. WO–64. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

21 p.
47

080 Lugo, A.E.; Brown, S. 1998. Los trópicos pueden

sobrevivir. Tierramérica. 4(1): 6–7.

081 Lugo, A.E.; Ford, L.B., eds. 1987. Forest

recreation in the Caribbean Islands.

Proceedings of the third meeting of Caribbean

foresters; 1986 May 19–23; Guadeloupe,

F.W.I. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Southern Forest

Experiment Station, Institute of Tropical

Forestry and the Caribbean National Forest.

95 p.

082 Lugo, A.E.; Frangi, J.L. 1993. Fruit fall in the

Luquillo Experimental Forest, Puerto Rico.

Biotropica. 25(1): 73–84.

083 Lugo, A.E.; García, A.R. 1996. Cartilla del agua

para Puerto Rico. Acta Científica. 10(1–3).

89 p.

084 Lugo, A.E.; Mastrantonio, L. 1999. Institute of

Tropical Forestry: the first fifty years. Gen. Tech.

Rep. IITF–GTR–7. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

56 p.

085 Lugo, A.E.; Ramos, O.; Molina, S.[and others].

1996. A fifty-three year record of land use

change in the Guanica Forest Biosphere Reserve

and its vicinity. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry with

Fundación Puertorriqueña de Conservación.13

p. [photographs and maps{maps by Velez

Rodriguez, L.L.}]

086 Lugo, A.E.; Scatena, F.N. 1996. Background and

catastrophic tree mortality in tropical moist, wet,

and rain forests. Biotropica. 28(4a): 585–599.

087 Maldonado, E.D. 1961. Peladora de postes de

cadena ajustada [A tight chain post peeler].

Apuntes Forestales Tropicales ITF–8. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Tropical Forest Research

Center. 5 p.

088 Maldonado, E.D. 1962. Radiación solar para

secar caoba en Puerto Rico. Apuntes

Forestales Tropicales ITF–14. Río Piedras, PR:

U.S. Department of Agriculture, Forest Service,

Tropical Forest Research Center. 5 p.

089 Maldonado, E.D. 1962. Solar radiation used to

dry mahogany lumber in Puerto Rico. Trop. For.

Note ITF–14. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

Tropical Forest Research Center. 5 p.

090 Mastrantonio, J.L.; Francis, J.K. 1997. A student

guide to tropical forest conservation. Gen. Tech.

Rep. GTR–SO–123. Asheville, NC: U.S.

Department of Agriculture, Forest Service,

Southern Research Station; Río Piedras, PR:

International Institute of Tropical Forestry.

20 p.

091 Molina, S.; Alemañy, S. 1997. Species codes for

the trees of Puerto Rico. Gen. Tech. Rep. GTR–

SO–122. Asheville, NC and Río Piedras, PR:

U.S. Department of Agriculture, Forest Service,

Southern Research Station and International

Institute of Tropical Forestry. 67 p.

092 Nobles, R.W.; Briscoe, C.B. 1966. Height growth

of mahogany seedlings. Res. Note ITF–10. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Institute of Tropical Forestry.

4 p.

093 Parrotta, J.A. 1993. Assisted recovery of degraded

tropical lands: plantation forests and ecosystem

stability. In: Paoletti, M.G.; Foissner, W.;

Coleman, D., eds. Soil Biota,Nutrient Cycling,

and Farming Systems. Boca Raton, FL: Lewis

Publishers: 169–182.

094 Parrotta, J.A. 1999. Tropical restoration for the

new millennium. Fourth annual Puerto Rico

forestry conference; 1999 May 23–28; San

Juan, PR. Río Piedras, PR: U.S. Department
48

of Agriculture, Forest Service, International

Institute of Tropical Forestry. 139 p. [abstracts].

095 Parrotta, J.A.; Baker, D.D.; Fried, M. 1994.

Estimation of nitrogen fixation in Leucaena

leucocephala using 15N-enrichment

methodologies. In: Sprent, J.I.; McKey, D., eds.

Advances in legume systematics 5: the nitrogen

factor. Kew, London, UK: Royal Botanic

Gardens: 75–82.

096 Parrotta, J.A.; Baker, D.D.; Fried, M. 1996.

Changes in nitrogen fixation in maturing stand

of Casuarina equisetifolia and Leucaena

leucocephala. Canadian Journal of Forest

Research. 26: 1684–1691.

097 Parrotta, J.A.; Francis, J.K.; de Almeida, R.R.

1995. Trees of the Tapajos:a photographic field

guide. Gen. Tech. Rep. IITF–1. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical

Forestry. 370 p.

098 Parrotta, J.A.; Knowles, O.H.; Wunderle, J.M.,

Jr. 1997. Development of floristic diversity in

10-year-old restoration forests on a bauxite

mined site in Amazonia. Forest Ecology and

Management. 99(1–2): 21–42.

099 Parrotta, J.A.; Lodge, D.J. 1991. Fine root

dynamics in a subtropical wet forest following

hurricane disturbance in Puerto Rico.

Biotropica. 23(4a): 343–347.

100 Parrotta, J.A.; Turnbull, J.W.; Jones, N. 1997.

Catalyzing native forest regeneration on

degraded tropical lands. Forest Ecology and

Management. 99(1–2): 1–7.

101 Reardon, R.C.; Edwards, W.G.; Meyerdirk, D.

1998. Pink hibiscus mealybug Maconellicoccus

hirsutus (Green). [La cochinilla rosada del

Hibisco Maconellicoccus hirsutus (Green)].

State and Private Forestry HT-1. Morgantown,

WV and Río Piedras, PR: U.S. Department of

Agriculture, Animal and Plant Health Inspection

Service, U.S. Department of Agriculture, Forest

Service, International Institute of Tropical

Forestry and Forest Health Technology

Enterprise Team. 14 p. [in English and Spanish].

102 Reyes, G.; Brown, S.; Chapman, J.; Lugo, A.E.

1992. Wood densities of tropical tree species.

Gen. Tech. Rep. SO–88. New Orleans: U.S.

Department of Agriculture, Forest Service,

Southern Forest Experiment Station, Institute

of Tropical Forestry. 15 p.

103 Sánchez, M.J.; Lopez, E.; Lugo, A.E. 1997.

Chemical and physical analyses of selected

plants and soils from Puerto Rico (1981–1990).

Res. Note IITF–RN–1. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

112 p.

104 Scatena, F.N. 1995. Relative scales of time and

effectiveness of watershed processes in a

tropical montane rain forest of Puerto Rico.

Natural and anthropogenic influencesin fluvial

geomorphology. Geophysical monograph 89.

Washington, DC: American Geophysical Union:

103–111.

105 Scatena, F.N. 1998. An assessment of climate in

the Luquillo Mountains of Puerto Rico. In:

Segarra García, R.I., comp., ed. Proceedings:

Tropical hydrology and Caribbean water

resources: Third international symposium on

tropical hydrology and fifth Caribbean Islands

water resources congress; 1998 July 12–16;

San Juan, PR. Herndon, VA: American Water

Resources Association: 193–198.

106 Scatena, F.N.; Larsen, M.C. 1991. Physical

aspects of Hurricane Hugo in Puerto Rico.

Biotropica. 23(4a): 317–323.

107 Scatena, F.N.; Moya, S.; Estrada, C.; Chinea, J.D.

1996. The first five years in the reorganization

of aboveground biomass and nutrient use

following Hurricane Hugo in the Bisley

Experimental Watersheds, Luquillo
49

Experimental Forest, Puerto Rico. Biotropica.

28(4a): 424–440.

108 Schubert, T.H.; Zambrana, J. 1978. Honduras or

big-leaf mahogany. Urban Forestry Bulletin,

Caribbean Area. U.S. Department of

Agriculture, Forest Service, Southeastern Area.

2 p.

109 Schubert, T.H.; Zambrana, J. 1978. West Indies

or small-leaf mahogany. Urban Forestry Bulletin,

Caribbean Area. U.S. Department of

Agriculture, Forest Service, Southeastern Area.

2 p.

110 Silander, S.R.; Lugo, A.E. 1990. Cecropia peltata

L. Yagrumo hembra, Trumpet-Tree. Moraceae.

Mulberry family. In: Burns, R.M.; Honkala,

B.H., comps., eds. Silvics of North America:

Hardwoods. Agric. Handb. 654. Washington,

DC: U.S. Department of Agriculture: 244–249.

Vol. 2.

111 Torres, J.A. 1992. Lepidoptera outbreaks in

response to successional changes after the

passage of Hurricane Hugo in Puerto Rico.

Journal of Tropical Ecology. 8: 285–298.

112 Torres, J.A. 1994. Insects of the Luquillo

Mountains, Puerto Rico. SO–GTR–105. New

Orleans: U.S. Department of Agriculture, Forest

Service, Southern Forest Experiment Station.

53 p.

113 Torres, J.A. 1994. Wood decomposition of

Cyrilla racemiflora in a tropical montane

forest. Biotropica. 26(2): 124–140.

114 Tosi, J.A., Jr.; Velez-Rodriguez, L.L. 1983.

Provisional ecological map of the Republic of

Brazil. Washington, DC: U.S. Department of

Agriculture, Forest Service, Southern Forest

Experiment Station, Institute of Tropical

Forestry. in cooperation with: Center for Energy

and Environment Research, University of Puerto

Rico and U.S. Department of Energy.16 p.

115 Veldkamp, E.; Keller, M. 1997. Nitrogen oxide

emissions from a banana plantation in the humid

tropics. Journal of Geophysical Research.

102(D13): 15,889–15,898.

116 Veldkamp, E.; Keller, M.; Núñez, M. 1998. Effects

of pasture management on N2

0 from NO

emissions from soils in the humid tropics of

Costa Rica. Global Biogeochemical Cycles.

12(1): 71–79.

117 Venator, C.R. 1972. Effect of gibberelic acid on

germination of low-vigor Honduras pine seeds.

Forest Science. 18(4): 331.

118 Venator, C.R. 1976. A mutant Pinus caribaea

var. hondurensis seedling incapable of

developing normal secondary needles.

Turrialba. 26(1): 98–99.

119 Venator, C.R.; Muñoz, J.E.; Barros, N.F. 1977.

Root immersion in water: a promising method

for successful bare-root planting of Honduras

pine. Turrialba. 27(3): 287–291.

120 Wadsworth, F.H. 1997. Forest production for

tropical America. Agric. Handb. 271.

Washington, DC: U.S. Department of

Agriculture. 563 p.

121 Wadsworth, F.H. 1999. Montane forest

management in the insular Caribbean. Gen.

Tech. Rep. IITF–GTR–8. Río Piedras, PR:

U.S. Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

29 p.

122 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, Institute of Tropical Forestry. 24 p.

123 Weaver, P.L. 1990. Calophyllum calaba L.

Maria, Santa Maria. Guttiferae. Mangosteen

family. In: Burns, R.M.; Honkala, B.H., eds.
50

Silvics of North America: Hardwoods. Agric.

Handb.654: Washington, DC: U.S. Department

of Agriculture: 172–178. Vol. 2.

124 Weaver, P.L. 1990. Manilkara bidentata (A.

DC.) Chev. Ausubo, Balata. Sapotaceae.

Sapodilla family. In: Burns, R.M.; Honkala,

B.H., eds. Silvics of North America:

Hardwoods. Agric. Handb. 654. Washington,

DC: U.S. Department of Agriculture: 455–460.

Vol. 2.

125 Weaver, P.L. 1990. Tabebuia heterophylla (DC.)

Britton. Roble blanco, White-cedar.

Bignoniaceae. In: Burns, R.M.; Honkala, B.H.,

eds. Silvics of North America: Hardwoods.

Agric.Handb. 654. Washington, DC: U.S.

Department of Agriculture: 778–783. Vol. 2.

126 Weaver. P.L. 1994. Baño de Oro Natural Area

Luquillo Mountains, Puerto Rico. Gen. Tech.

Rep. SO–111. New Orleans: U.S. Department

of Agriculture, Forest Service, Southern Forest

Experiment Station. 55 p.

127 Weaver, P.L. 1994. Effects of Hurricane Hugo on

trees in the Cinnamon Bay watershed, St. Johns,

U.S. Virgin Islands. Caribbean Journal of

Science. 30(3–4): 255–261.

128 Weaver, P.L. 1996. Forest productivity in the

Cinnamon Bay watershed, St. Johns, U.S.

Virgin Islands. Caribbean Journal of Science.

32(1): 89–98.

129 Weaver, P.L. 1998. The effects of environmental

gradients on hurricane impact in the Cinnamon

Bay watershed, St. John, U.S. Virgin Islands.

In: Dallmeier, F.; Comiskey, J.A., comps., eds.

Forest biodiversity in North, Central and South

America, and the Caribbean: research and

monitoring. Man and the biosphere series. Paris:

UNESCO and the Parthenon Publishing Group:

333–346. Vol. 21.

130 Weaver, P.L. 1998. Hurricane effects and longterm recovery in a subtropical rain forest. In:

Dallmeier, F.; Comiskey, J.A., comps., eds.

Forest biodiversity in North Central and South

America, and Caribbean: research and

monitoring. Man and the biosphere series. Paris:

UNESCO: 249–270. Vol. 21.

131 Weaver, P.L. 1999. Impacts of Hurricane Hugo

on the dwarf cloud forests of Puerto Rico’s

Luquillo Mountains. Caribbean Journal of

Science. 35(1–2): 101–111.

132 Weaver, P.L.; Birdsey, R.A.; Nicholls, C.F. 1998.

Los recursos forestales de San Vicente,Indias

Occidentales. Res. Pap. SO–244. New

Orleans: U.S. Department of Agriculture, Forest

Service, Southern Forest Experiment Station.

27 p.

133 Weaver, P.L.; Medina, R.R.; Coll Rivera, J.L.

1999. Las Cabezas de San Juan: a natural

treasure in northeastern Puerto Rico. Gen. Tech.

Rep. IITF–6. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

27 p.

134 Weaver, P.L.; Ramirez, J.L.; Coll Rivera, J.L.

1999. Las Cabezas de San Juan Nature

Reserve (El Faro). Gen. Tech. Rep. IITF–5.

Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, International

Institute of Tropical Forestry. 58 p.

135 Weaver, P.L.; Sabido, O.A. 1997. Mahogany in

Belize: a historical perspective. Gen. Tech. Rep.

GTR–IITF–2. Asheville, NC: U.S. Department

of Agriculture, Forest Service, International

Institute of Tropical Forestry. 31 p.

136 Whitmore, J.L. 1972. Pinus merkusii unsuitable

for plantations in Puerto Rico. Turrialba. 22(3):

351–353.

137 Whitmore, J.L. 1978. Bibliography on Eucalyptus

deglupta Bl. Res. Note ITF–17. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, Institute of Tropical Forestry. 18 p.
51

138 Whitmore, J.L. 1978. Cedrela provenance trial in

Puerto Rico and St. Croix: establishmentphase.

Res. Note ITF–16. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

Institute of Tropical Forestry. 11 p.

139 Whitmore, J.L.; Hinojosa, G. 1977. Mahogany

(Swietenia) hybrids. Res. Pap.ITF–23. Río

Piedras, PR: U.S. Department of Agriculture,

Forest Service, Institute of Tropical Forestry.

8 p.

140 Whitmore, J.L.; Liegel, L.H. 1980. Spacing trial

of Pinus caribaea var. hondurensis. Res. Pap.

SO–162. New Orleans: U.S. Department of

Agriculture, Forest Service, Southern Forest

Experiment Station, Institute of Tropical

Forestry. 9 p.

141 Wickel, A.J. 1997. Rainfall interception modeling

for two tropical forest types in the Luquillo

Experimental Forest, Puerto Rico. In:

Hydrological modeling in humid tropical island

setting: with special reference to the Luquillo

Experimental Forest, Puerto Rico. Working

paper No. 2. Amsterdam, The Netherlands:

Vrije Universiteit, Faculty of Earth Sciences.

42 p. + appendix.

142 Wiley, J.W.; Wunderle, J.M., Jr. 1993. The effects

of hurricanes on birds, with special reference

to Caribbean islands. Bird Conservation

International. 3: 319–349.

143 Woodbury, R.O.; Little, Jr., E.L. 1976. Flora of

Buck Island Reef National Monument (U.S.

Virgin Islands). Res. Pap. ITF–19. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, Institute of Tropical Forestry. 27 p.

144 Wunderle, J.M., Jr. 1994. Census methods for

Caribbean land birds. Gen. Tech. Rep. SO–

GTR–98. New Orleans: U.S. Department of

Agriculture, Forest Service, Southern Forest

Experiment Station. 26 p.

145 Wunderle, J.M., Jr. 1995. Métodos para contar

aves terrestres del Caribe. Gen. Tech. Rep.

SO–100. New Orleans: U.S. Department of

Agriculture, Forest Service, Southern Forest

Experiment Station. 28 p.

146 Wunderle, J.M., Jr. 1995. Population

characteristics of Black-throated Blue Warblers

wintering in three sites on Puerto Rico. The Auk.

112(4): 931–946.

147 Wunderle, J.M., Jr. 1995. Responses of bird

populations in a Puerto Rican forest to Hurricane

Hugo: the first 18 months. The Condor. 97:

879–896.

148 Wunderle, J.M., Jr. 1997. The role of animal seed

dispersal in accelerating native forest

regeneration on degraded tropical lands. Forest

Ecology and Management. 99(1–2): 223–235.

149 Wunderle, J.M., Jr. 1999. Avian distribution in

Dominican shade coffee plantations: area and

habitat relationships. Journal of Field

Ornithology. 70(1): 58–70.

150 Wunderle, J.M, Jr.; Cortes, R.A.; Carromero, W.

1992. Song characteristics and variation in a

population of bananaquits on Puerto Rico. The

Condor. 94: 680–691.

151 Wunderle, J.M., Jr.; Latta, S.C. 1996. Avian

abundance in sun and shade coffee plantations

and remnant pine forest in the Cordillera Central,

Dominican Republic. Ornitología Neotropical.

7: 19–34.

152 Wunderle, J.M., Jr.; Latta, S.C. 1998. Avian

resource use in dominican shade coffee

plantations. Wilson Bulletin. 110(2): 271–281.

153 Wunderle, J.M., Jr.; Lodge, D.J.; Waide, R.B.

1992. Short-term effects of Hurricane Gilbert

on terrestrial bird populations on Jamaica. The

Auk. 109(1): 148–166.
52

154 Wunderle, J.M., Jr.; Waide, R.B. 1993.

Distribution of overwintering Nearctic migrants

in the Bahamas and Greater Antilles. The

Condor. 95: 904–933.

155 Yocum, C.; Lugo, A.E., eds. 1995. The economics

of Caribbean forestry: Proceedings of the

seventh meeting of Caribbean foresters at

Jamaica; 1994 June 13–17; Kingston, Jam.

Tech. Proceedings R8–TP 23. Atlanta: U.S.

Department of Agriculture, Forest Service,

Southern Region and Río Piedras, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry. in

cooperation with the University of Puerto Rico.

144 p.

156 Yocum, C.; Lugo, A.E., eds. 1997. Protected

areas management: Proceedings of the eighth

meeting of Caribbean foresters at Grenada;

1996 June 3–7; St. Georges, GJ. Río Piedras,

PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical

Forestry. 144 p.

157 Zepeda, G.; Lugo, A.E., eds. 1994. Towards

sustainable forest resource management in the

Caribbean. Proceedings of the sixth meeting of

Caribbean foresters at Martinique; 1994 July

20-24; Martinique. San Juan, PR: U.S.

Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

55 p.
53

If you would like to receive hardcopies of available publications, please contact Gisel Reyes at

the mailing or e-mail address below. Please specify the numbers you want to receive and the

Annual Letter year, i.e. 199-2000. All publications are sent while supplies last.

Si desea recibir separatas de las publicaciones disponibles, favor de comunicarse con Gisel

Reyes a la dirección postal o electrónica indicada abajo. Por favor, indique los números

deseados y el año del “Annual Letter”, i.e. 1999-2000. Todas las publicaciones se enviarán

hasta agotarse las cantidades impresas.

USDA Forest Service – Intl. Institute of Tropical Forestry

Attn: Annual Letter 1999-2000 publications

Box 25,000

Rio Piedras, Puerto Rico 00928-5000

greyes@fs.fed.us
54

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.

The USDA prohibits discrimination in all its programs and activities

on the basis of race, color, national origin, sex, religion, age,

disability, political beliefs, sexual orientation, or marital or family

status. (Not all prohibited bases apply to all programs.) Persons

with disabilities who require alternative means for communication of

program information (Braille, large print, audiotape, etc.) should

contact USDA's TARGET Center at (202) 720-2600 (voice and TDD).

To file a complaint of discrimination, write USDA, Director, Office of

Civil Rights, Room 326-W, Whitten Building, 1400 Independence

Avenue, SW, Washington, D.C. 20250-9410 or call (202) 720-5964

(voice and TDD). USDA is an equal opportunity provider and

employer.“