,6 United States fl Department of 

3/ Agriculture 

Forest Service 

Southern Forest 

Experiment Station 

New Orleans, 

Louisiana 

General Technical Report 

SO-105 

July 1994 

Insects of the Luquillo 

Mountains, Puerto Rico 

Juan A. Torres 
SUMMARY 

In this review of the literature on forest entomology in Puerto 

Rico, emphasis is given to research conducted in the Luquillo 

Experimental Forest (LEF). This review should serve as an introduction to the insects inhabiting the LEF for researchers and as a 

guide for the identification of possible insect pests. There are three 

sections to this review. The first deals with basic insect ecology; the 

second, forest insect pests; and the third, insect attacks on dry 

wood and during wood seasoning. The reference section and appendixes contain information on the systematics and taxonomy of 

different insect orders found in Puerto Rico. 

ACKNOWLEDGMENTS 

I thank the following people for their help in the preparation of 

this work: Maribel Torres, Silverio Medina, and Jose Maldonado. 

The cover drawing of the fungus grower ant Chyphomyrmex 

minutus was drawn by R.A. DeNicola. Figures 1 and 15 were drawn 

by Jesus Gonzhlez. 
CONTENTS 

................................................................ INTRODUCTION 1 

INSECT ECOLOGY .............................................................. 2 

Order Coleoptera (beetles) ..................................................... 2 

Carabidae (ground beetles) ...................................................... 2 

Leiodidae (round fungus beetles) ................................................ 3 

Scarabaeidae (scarab beetles) .................................................... 3 

Scolytidae bark beetles) and Platypodidae (pin-hole borers) ........................ 4 

Order Dipteratflies) ........................................................... 4 

Culicidae ......................................................... (mosquitoes) 4 

Muscidae ........................................................ (muscid flies) 5 

Phoridae (humpbacked flies) .................................................... 5 

Syrphidae (flower flies) ......................................................... 5 

Tipulidae (crane flies) .......................................................... 6 

Order Hymenoptera (ants, bees, and wasps) ................................... 7 

Anthophoridae (carpenter bees. cuckoo bees and digger bees) ...................... 7 

Apidae (honey bees) ............................................................ 7 

Formicjdae (ants) .............................................................. 7 

Vespidae (wasps) .............................................................. 8 

Orderlsoptera (termites) ..................................................... 8 

Order Odonata (dragonflies, damselflies) ...................................... 11 

Order Ortboptera (crickets, grasshoppers, and walking sticks) ............... 12 

Phasmatidae (walking sticks) ................................................... 12 

Order Tricboptera (caddisflies) ................................................ 13 

Faunistic Studies .............................................................. 13 

Bromeliad communities ........................................................ 13 

Seed and fruit consumers ...................................................... 14 

Insect diversity and abundance ................................................. 14 

Herbivores .................................................................... 15 

Jnsectbiomass ................................................................. 17 

Pollinators .................................................................... 17 

Lizard and frog predation on insects ............................................. 17 

Bird predation on insects ....................................................... 18 

INSECTS THAT ATTACK FOREST TREES AND SHRUBS ...................... 19 

Order Coleoptera (beetles) ..................................................... 19 

Bostrichidae (branch and twig borers) ........................................... 19 

Buprestidae (metallic wood-boring beetles) ....................................... 22 

Cerambycidae (long-horned beetles) and Elateridae (click beetles) ................... 22 

Curculionidae (snout beetles) .................................................... 22 

Platypodjdae (pin-hole borers) .................................................. 23 

Scarabaeidae (scarab beetles) ................................................... 23 

Scolytidae (bark and ambrosia beetles) ........................................... 25 

OrderDiptera (flies) .......................................................... 26 

Tephritidae (fruit flies) ......................................................... 26 

Order Hemiptera ....................................................... (bugs) 26 

Pyrrhocoridae (red bugs or stainers) ............................................. 26 

Tingididae (lace bugs) .......................................................... 26 

Order Hornoptera (aphids, scale insects, leafhoppers, and mealybugs) ......... 26 

Aphididae (aphids) ............................................................. 26 

Asterolecaniidae (pit scales) ..................................................... 27 

Cicadellidae (leafhoppers) ....................................................... 27 
Coccidae (soft. wax. and tortoise scales) ......................................... 28 

Diaspididae (armored scales) .................................................... 28 

Margarodidae (giant coccids and ground pearls) .................................. 28 

Pseudococcidae (mealybugs) .................................................... 29 

Order Hymenoptera (ants. bees, sawflies, and wasps) ......................... 29 

hgidae .............................................................. (sawflies) 29 

Formicidae (ants) .............................................................. 29 

Order Lepidoptera (butterflies and moths) .................................... 30 

Cosmopterygidae (cosmopterigid moths) ......................................... 30 

Gelechiidae (gelechiid moths) ................................................... 30 

Gometridae (measuring worms) ................................................ 30 

Hyblaeidae (hyblaeid moths) ..................................................... 30 

Megalopygidae (flannel moths) ................................................... 30 

Noctuidae (noctuid moths) ...................................................... 30 

Psychidae (bagworms) ......................................................... 31 

Pyralidae (snout and grass moths) .............................................. 31 

Order Thysanoptera ................................................... (thrips) 34 

Other ........................................................... Insect Groups 34 

mSECTS THAT ATTACK FOREST PRODUCTS .................................. 35 

Order Coleoptera (beetles) .................................................... 35 

Bostrichidae (branch and twig borers) ........................................... 35 

Lydidae (powderpost beetles) ................................................... 36 

Order Hymenoptera (ants, bees, wasps, and allies) ............................ 36 

Eumenidae (mason and potter wasps) ........................................... 36 

Order Isoptera (termites) ...................................................... 36 

LITERATURE CITED ........................................................... 38 

APPENDIXES ................................................................... 45 

Appendix A - Forest and Plantation Trees and Their Associated 

or Potential Insect Pests ........................................... 46 

Appendix B - List of Important Entomological Studies in the Luquillo Mountains .... 49 

Appendix C - Additional References .............................................. 50 
Insects of the Luquillo Mountains, Puerto Rico 

Juan A. Torres 

There is a generalized belief that Puerto Rico lacks 

problems with forest insects. The reason for this belief 

may be that, unless forests are profoundly disturbed 

through silvicultural practices or natural disasters, 

indigenous insect pests normally do not cause catastrophic or appreciable damage to stands. The initial 

success of exotic tree species does not guarantee that 

they will not eventually face insect or disease problems. 

Time is required for populations of insect pests and 

pathogens to build up and for local pests to adapt to a 

new host (Bakshi 1976). In addition, increases in the 

area of forest cultivation will lead to an increase in 

insect pests associated with the host (El Servicio de 

Extensi6n Agricola 1948, Strong 1979). The cultivation 

of small areas to examine the adaptability of a plant 

species to a new environment cannot test this phenomenom. The rapid growth rates of some tropical tree 

species, as well as the fact that the growth is continuous over much of the year, allow trees to outgrow the 

attacks from most leaf-feeding insects (Hodges and 

McFadden 1987). However, some tropical insects may 

be more difficult to control because they thrive 

throughout the whole year. In tropical areas with a prolonged dry season, the weather can keep insect populations at relatively low levels (Janzen 1973a). 

The problems created by insects in forests often 

require different control approaches than those used in 

agriculture. The use of insecticides can be economically 

prohibitive even when it is a reasonable alternative for 

forest insect pest control. Thus, it is important to 

include entomologists as part of forest management 

teams for evaluating the effects of management practices and natural disturbances on insects and vice 

versa. Management practices developed for agricultural 

systems are labor intensive and incorporate vast quantities of fertilizers and other inputs, which are often 

not economically feasible for use in forest systems. For 

these reasons, many of the inputs that keep a forest in 

a healthy condition must be provided by the organisms 

that comprise the community. 

Insects have several beneficial roles in a forest. Many 

plants require insects for pollination and seed dispersal 

(Beattie 1985). Many vertebrate species that humans 

consider esthetic elements of the biota are completely or 

partially dependent on the presence of insects for their 

nutrition (Garcia 1938, Janzen 1973b, Ross and others 

1982). In bodies of water in the forest, many fish rely on 

insects for much of their food, and aquatic insects 

recycle nutrients in their habitats before the nutrients 

are carried downstream into the ocean (Ross and others 1982). Insects are also important in nutrient recycling in terrestrial systems. Many species of birds 

and other terrestrial predators feed on aquatic insects, 

returning nutrients back to the terrestrial environment. The burrowing activities of invertebrates, including several insect species, help reduce soil erosion by 

facilitating the penetration of water and reducing overland runoff (Anderson 1988). 

In this paper, the literature on forest entomology in 

Puerto Rico is reviewed, giving emphasis to the work 

done in the Luquillo Mountains. The Luquillo Mountains are located in the northeast corner of Puerto Rico 

and mainly include four subtropical life zones: wet forest, rain forest, lower montane wet forest, and lower 

montane rain forest. There are four predominant plant 

associations in the mountains: tabonuco, colorado, 

palm, and dwarf forests. These four plant associations 

are roughly stratified by altitude. The tabonuco forest 

is found below 600 m and occupies nearly 70 percent of 

the Luquillo Mountains. The colorado forest, which 

covers about 17 percent of the mountains, is found 

above 600 m. On peaks and ridges above 750 m (2 

percent of the Luquillo Mountains) the dwarf forest is 

found. The palm forest is interspersed within the colorado and dwarf forests and covers 11 percent of the 

Luquillo Mountains (Brown and others 1983). 

This review is divided in three sections: (1) insect 

ecology, (2) insect pests of forest trees and shrubs, and 

(3) insect pests of forest products. The first section 

covers the research on insects that are not generally 

considered economically important but perform many 

important ecological functions. The section on insect 

Juan A. Torres is an associate professor, Department of Biology, University of Puerto Rico, Bayamon, PR 00959-1919. Research and data 

compilation for thia publication were done by the author while he was conducting postdoctoral research at the Institute of Tropical Forestry, U.S. 

Department of Apiculture, Forest Service, Southern Forest Experiment Station, Rio Piedras, PR 00928-2500. 
pests of forest trees and shrubs deals with insects that 

have created problems in important (or potentially 

important) timber species that grow in the LuquiIlo 

Mountains. Forest insect pests appear to have been 

more common in the first half of this century than at 

present in Puerto Rico. This may be related to the 

widespread use of modern insecticides and the introduction of the frog Bufo marinus to Puerto Rico in the 

5920's (Wolcott 194%). In the last section, insects that 

attack wood during the drying process and insects that 

consume dry wood are discussed. The section contains 

a thorough discussion of termites, which is necessary 

due to the importance of this group of insects and the 

extensive misconceptions about feeding preferences of 

Figure 1. -A ground beetle Antilliscaris megacephalus; actual 2ength 

of the beetle is approximately 35 rnm (adapted from 

Darlington 1970 with permission of "BiotrCpica'>/. 

termites. Appendix A lists forest and plantation trees 

and their associated potential insect pests. Appendix B 

provides a list of contributions that provide a quick 

introduction to the ecology and pest insects in the 

Luquillo Mountains. Appendix C is an annotated bibliography of additional references that deal with the 

systematics of some insect groups. Wolcott (1948a, 

1948b; 1948c, 1948d) and Maldonado and Navarro 

(1967) can be consulted for additional references on 

systematics. 

The radiation experiment referred to in this review 

involved the exposure of a section of the tabonuco forest 

near El Verde Field Station to a 10,000 curie cesium 

source for 92.8 days (Jan. 19 to Apr. 27, 1965). Comparisons with a mechanically pruned area known as 

the north cut center and a little-disturbed south control center were made to determine the effects of radiation. This radiation study was conducted under the 

sponsorship of the U.S. Atomic Energy Commission 

and the results are summarized in the book "A tropical 

rain forest: a study of irradiation and ecology at El Verde, 

Puerto Rico" (U.S. Atomic Energy Commission 1970). 

INSECT ECOLOGY 

Order Coleoptera (beetles) 

Carabidae (ground beetles).-Most species of 

ground beetles (the Carabidae family) hide during the 

day and feed at night. Nearly all are predaceous on 

other insects. Darlington (1970) studied the ground 

beetles, finding a reduction in the number of species at 

high elevations in Puerto Rico and a tendency for wing 

atrophy. Lowland species are usually small and have 

well-developed wings. Darlington reported Antilliscaris 

megacephalus (recorded as Scarites megacephalus) 

from the Luquillo Mountains as the largest Carabidae 

(30 to 35 mm) found in the Antilles (fig. 1). Hlavac 

(1969) presented taxonomic keys for the genus Antilliscaris and compared the Puerto Rican species with 

the ones from Jamaica. There are 8 known montane 

species of carabids in Puerto Rico (3 in the genus 

Antilliscaris) and 17 in Jamaica, an island of similar 

size. In Jamaica, nine of the carabid species belong to 

the genus Colpodes, and there are no Antilliscaris. The 

montane biota of Puerto Rico is unique in not having 

members of the ecologically variable carabid genus 

Colpodes. 

Hlavac (1969) suggested the occurrence of a shift 

from burrowing to crawling on the ground surface in 

the Antilliscaris of Puerto Rico. Short antennae are 

characteristic of diverse burrowing Coleoptera that 

have evolved a pedunculate body form. In A. megacephalus, 50 percent of the antennae extend beyond the 

mandible. If A. megacephalus burrows, the antennae 

would have to be doubled over during this activity. The 

long antennae of this species strongly suggest that it is 
not a burrowing one. Antilliscaris megacephalus is 

doubtless one of the top predators among ground invertebrates (Hlavac 1969). 

Based on larval characters, Antilliscaris is closely 

aligned with genera of the Afrotropical region, 

especially with Prodyscherus, which is endemic to 

Madagascar (Nichols 1986). The abundance of Antilliscaris may have been severely affected by the introduction of predators and omnivores, such as mongooses 

(Herpestes auropurzctatus), feral cats (Felis catus), 

toads (Bufo marinus), the wharf rat (Rattus noruegicus), and the black roof rat (R. rattus) to Puerto 

Rico (Nichols 1988). 

The presence of three species of Antilliscaris and the 

relatively high percentage of endemism (38 percent) in 

the Puerto Rican Scaritinae fauna suggest that Puerto 

Rico is the oldest emergent land mass in the Greater 

Antilles (Nichols 1988). 

The carabid genus Platynus is composed of at least 

66 species in the West Indies. Most of these species are 

restricted to single islands, some to only one mountain 

on an island (Liebherr 1988). There are 25 endemic 

specjes of Platynus in Hispaniola and 12 in the Lesser 

Antilles. Although Puerto Rico has suitable mountain 

and forest habitats for beetles of the genus, no Platynus 

beetles have been found. The small size of Puerto Rico 

and its geographical location at the eastern end of the 

Greater Antilles make the island a comparatively poor 

candidate for receiving water-transported immigrants 

(Liebherr 1988). 

Leiodidae (round fungus beetles). - Round fungus beetles (the Leiodidae family) occur in fungi, under 

bark, in decaying wood, in carrion and in other similar 

habitats (Borror and others 1981). Peck (1970, 1972) 

studied the subfamilies Catopinae and Leiodinae in 

Puerto Rico and Jamaica. Most of the Catopinae beetles occur in carrion. Peck found that members of the 

Catopinae were rare in the Luquillo Forest and suggested that high humidity may be the reason for the 

low numbers of these inseds. In addition, the zoogeography of the family is delineated. 

Scarabaeidae (scarab beetles).- The scarab beetles (the Scarabaeidae family) vary greatly in feeding 

habits. Many feed on dung, decaying plant material, 

carrion, etc.; others feed on live plants. Matthews 

(1965), in his detailed study of the dung beetles of the 

Luquillo Mountains, showed that the species were separated according to altitudinal belts, behavior, and die1 

cycle (figs. 2, 3). He compared the Puerto Rican species 

with the ones from Mauritius (in the Indian Ocean). 

Several patterns were found to be in common, including the distributions of species highly influenced by 

altitude and bird and mollusk excrement as the main 

sources of food for these beetles before humans came. 

Food is now provjded by the excrement of an introduced monkey, Macaca, in Mauritius, and the roof rat, 

Rattus rattus, in Puerto Rico. The parallelisms are the 

hispidurn iunceonurn I I 

111 Ill1 I I I I t I 

0~~3456769101112 

ALTITUDE (HUNDRED METERS) 

Figure 2.-Altiiude distribution of the dung beetles found in the 

Luquillo Mountains of Puerto Rico (adapted from Matthews 1965 with permission of "Transactions of the 

American Entomological Society"). 

result of adaptations by beetles of the same tribe to 

similar tropical oceanic islands of mountainous 

topography. Matthews also deIineated possible historical diet changes for these beetles in Puerto Rico, and, 

for the scarab Canthochilum histeroides, he reported 

altitudinal variation in morphology. The scarab beetles 

are important in nutrient cycling. In Puerto Rico, the 

absence of large dung beetles capable of exploiting large 

masses of excrement in open areas such as cow pastures, results in areas that are temporarily unavailable 

for grazing and provides breeding media for horn fljes. 

Matthews (1966) also wrote a treatise on the taxonomy, zoogeography, ecology, and evolution of the sub- . family Scarabaeinae in the Antilles. The ecogeographic, 

adaptive radiation in the genus Canthochilum, in 

Puerto Rico and Cuba, and other patterns of endemism 

were discussed. Matthews found that the neotropical 

species of the Antillean scarabaeine fauna comprise a 

minority; the majority of the species are possibly of Old 

World origin. The high endemicity of the Greater Antilles Scarabaeinae fauna is a reflection, not of insular 

evolution, but of the survival of an ancient fauna for 

which these islands have become a refuge. The 

restriction of the beetle Oniticellus cubensis to open 

terrain points to the presence of natural savannas 

before the advent of humans to the region. Matthews 

invoked interspecific competition to explain the patterns of distribution on many islands. The highland, 

humid forest belt is not invaded by dung beetles except 

under the pressure of interspecific competition result- 
@ TIMES OF OBSERVED ACTIVITY 

f2j PRESUMED ACTIVITY RANGE 

Figure 3.-Die1 activity cycles of the three most abundant species ofdung beetles in 

the Luquillo Mountains of Puerto Rico (adapted from Maithews 1965 

with permission of "Transactions of the American Entomological 

Society"). 

ing from the invasion of a second species of the same 

genus. One of the species may be displaced into the 

montane zone, while the other remains in the lowland 

belt. 

Scolytidae (bark beetles) and Platypodidae 

(pin-hole borers). - The Scolytidae family contains 

two main groups: the bark beetles, which feed on the 

phloem tissue, and the ambrosia beetles, which bore 

into the wood and cultivate a fungus upon which they 

feed. Some scolytids feed on seeds. Janzen (1972) 

described the interaction between the scolytid beetle 

Coccotrypes carpophagus and the palm Prestoea montuna. Janzen hypothesized that seed predation by the 

beetle is the selective force producing fruiting synchrony in the palm. Nonetheless, Wood (an authority 

on this family of beetles, Janzen 1972) thinks that C, 

carpophagus was introduced to Puerto Rico from the 

Old World around 1900. This implies that the interaction between the beetle and the palm is recent and does 

not represent an example of coevolution. Bannister 

(1970) found that F! montana germination percentages 

in plots where C. carpophagus was abundant djd not 

differ from plots where the beetles were scarce. 

It appears that tropical cyclones can affect the abundance of these organisms because plant phenology 

changes after tropical storms (Bates 1929). When this 

happens, pollinators and seed eaters are stressed. 

Nonetheless, organisms that feed on dead or damaged 

wood like scolytids, platypodids, cerambycids, and termites increase in abundance after storms (Bates 1929, 

Wolcott 1932). 

Schedl (1961), in his treatise of the Scolytidae and 

Platypodidae families, gave information about distribution and hosts for some Puerto Ecan members of the 

two families. Bright (1985) compiled a checklist of the 

West Indies Scolytidae. 

Order Diptera (flies) 

Culicidae (mosquitoes). - Mosquitoes (the Culicidae family) are important because the females are 

blood feeders; many species bite humans, and they 

serve as vectors in the transmission of diseases. In a 

study of the ekts of cesium gamma radiation on the 

mosquito populations at El Verde, Weinbren and 

Weinbren (1970) argued that the mosquito Aedes 
6 0 I 

Aedes rnediavitt~fu~ After much of the research activities ceased, both habL 50 i itats and hosts may have diminished below the level 

P: 30 lGhCDlC7ED CND 

SOUTH CONTROL 

20 

h 2 to 

a 

2 0 

A. taeniorhynchus 

s' lo 

0 

i i 

A. serrolus I 

g 20 i I 

30 

A. aegyptl 8 20- 

I ! 

t RIDIPTION PERIOD ? I 

v 

FMAMJJASONDJ FMAMJ JASONDJ 

1964 1965 

Figure 4. -Monthly capture record of four mosquito species of Aedes 

in El Verde, Puerto Rico, before and after radiation 

(adapted from Weinbren and Weinbren 1970). 

aegypti seemed more prevalent after radiation, although the number captured was small (fig. 4). The 

mosquitoes appeared to belong to a wild strain of this 

species that is generally found in the Old World. Breeding inside dense forests is not documented in the New 

World A. aegypti and is only rarely encountered in the 

wild strains of this species in Africa (Haber and Moore 

1973). 

Weinbren and Weinbren (1970) furnished a list of the 

members of the family Culicidae found in Puerto Rico 

and information on the temporal abundance of other 

mosquito species in the Luquillo Mountains. Because 

there were no replications in the experiment and no 

appropriate statistical analyses, the changes in abundance of these mosquito species could not be directly 

related to the effects of radiation or the opening of the 

canopy. Five years after the study by Weinbren and 

Weinbren, Haber and Moore (1973) conducted a survey 

of the same area and did not find A. aegypti in the 

dense forest. It appears that the increase in numbers of 

A. aegypti at El Verde was not due to the effects of 

radiation, but to the result of the intense human 

activity inside the forest during and after the radiation 

experiment. The traffic of equipment and materials to 

the experimental area increased the breeding area. 

needed to support a breeding population. 

Tulloch (1938) made notes on the mosquitoes associated with bromeliads in the Luquillo Mountains and 

the Maricao State Forest. Information on the temperature and pH of the water where the mosquitoes bred is 

provided. These pH readings, along with those presented in Paz (19741, could serve as a baseline to study 

possible changes in water pH due to atmospheric 

pollution. 

Muscidae (museid flies). - Many muscid flies are 

important pests, among them the house fly, Musca 

domestics, and horn flies, Hwmatobia irritans, which 

are pests of cattle. The larvae of the flies Philornis spp. 

parasitize several bird species in Puerto Rico, including 

the Puerto Rican parrot Amazona uittata (Arendt 

1985a, 1985b, Perez and Collazo 1976, Snyder and others 1987). The incidence of parasitism varies geographically and among bird species. Thrasher nestlings, 

Margarops fuscatus, have been found infested a few 

hours after hatching. Seasonal and annual variations 

occur in the level of parasitism of thrashers. Arendt 

found that 97 percent of nestling mortality in the 

pearly-eyed thrasher was due to parasitism by larval 

Philornis spp. Ninety-six percent of 448 nestlings 

examined from 1979 to 1982, harbored Philornis larvae 

(table I), whereas 31 percent of adult birds harbored the 

larvae. Infested thrasher nestlings were underweight 

and showed retarded tarsus growth as well as delayed 

emergence of the ninth primary pinfeather. The survival of post-fledge juveniles was influenced by parasitism while in the nest (Arendt 1985a, 1985b; Vhazy 

and Arendt 1986). 

Phoridae (humpbacked flies). - Phorid adults are 

abundant in decomposing vegetation. Larvae occur in 

decaying animal or plant matter, in fungi, as internal 

parasites of other insects, and as parasites or commensals in the nests of ants or termites (Borror and others 

1981). Villa and 'J'ownsend (1983) found the larvae of 

the phorid fly, Megaselia scalaris, eating viable eggs of 

the frog Eleutherodactylus coqui. In 1982, the flies infested approximately 3 percent of the frog egg clukhes. 

The fly's larval stage lasts between 34 and 46 hours, 

and the pupal stage lasts a minimum of 242 hours (10 

days). 

Syrphidae (flower flies).-Adult syrphid flies are 

common near flowers and often do a great deal of 

hovering. Many are brightly colored and resemble 

wasps or bees. Syrphid larvae are predaceous on aphids. 

They live in the nests of social insects, decaying wood, 

or polluted aquatic habitats (Borror and others 1981). 

In Puerto Rico, most of the members of this family (fig. 

5) feed on aphids and are beneficial in suppressing 

aphid outbreaks. Life history studies on the subfamily 

Syrphinae have been undertaken on few species. 

Telford (1973) suggested that some Syrphinae species 
Table 1.-Preualence and mean intensity of Philornis spp. larvae on 448 nestling 

pearly-eyed thrashers, 1979-1982 (adapted from Arendt 1985a with permission of The Auk) 

Nestlings Total number Intensity 

Month sampled Prwalence of parasites (range) 

Percent 

January 3 100 

February 25 80 

March 43 81 

April 97 96 

May 115 100 

June 95 100 

July 70 100 

Parasites per bird 

69 23.0 (19-26) 

432 20.6 (0-49) 

1,086 25.3 (0-73) 

4,215 40.9 (0-220) 

4,720 45.4 (6-131) 

6,294 61.7 (3-169) 

2,946 42.1 (5-135) 

-- - - 

Figure 5. -A flower f7y, Baccha clavata (adapted from Wolcott 1948~). 

can be facultative predators or some subsist solely on 

plant food. It is possible that some species feed on 

decomposing plant material as well as on insect prey. 

Tipulidae (crane flies).- Most of the adult tipulids resemble overgrown mosquitoes with extremely 

long legs, which are easily broken off (Borror and others 1981). The larval food of most species consists of 

decayed vegetation. The larvae of some species feed on 

algae and the roots of plants; others are carnivorous 

(Peterson 1982). In Puerto Rico, the various Tipulidae 

species tend to exhibit different patterns of distribution 

(Alexander 1932). Some tend to be isolated at higher 

elevations in the mountains of Puerto Rico, others tend 

to be widespread. The Luquillo Mountains are of 

unusual interest because several crane fly species are 

apparently restricted to this area. The ecology of this 

group is quite variable. Some crane fly larvae are found 

on the wet faces of cliffs at the margins of streams; 

others are aquatic and carnivorous. The latter group 

pupates on land, and the adults occur in swarms close 
to rivers. Lvvae of TipuIidae have been found feeding 

on tabonuco fruits, Dacryodes excelsa, and in fallen 

decomposing wood.) Some crane flies suck the nectar of 

various flowers (Alexander 1932). 

Order Hymenoptera (ants, bees, and wasps) 

Anthophoridae (carpenter bees, cuckoo bees 

and digger bees).- The family Anthophoridae is 

large, diverse, and widely distributed. ?\Jesting females 

may be solitary, but in many genera, they form large 

aggregations (Snelling 1981). 

Bees of the genus Xylocopa include some of the 

largest bees. They construct nests in dead wood. Jackson and Woodbury (1976) published a list of the plants 

used by the carpenter bee, Xylocopa mordax (recorded 

as X. brasilianorum) (fig. 6), for nesting and sources of 

food. In Puerto Rico, this species of bee uses 285 plant 

species for food and is responsible for the pollination of 

25 fruit and seed crop species used by humans. 

Apidae (honey bees). - Eusociality (truly social 

behavior) attains its highest development among the 

bees in the family Apidae. Not all Apidae are social; the 

Euglossinae subfamily includes only solitary or communal species. The Apinae subfamily includes only the 

1Personal observation by the author. 

genus &is and is best known by the common honey 

bee, &is mllifera. This is one of the most important 

animals domesticated by humans; it is valued for polIination and for the production of honey and wax 

(Snelling 1981). 

The number of tree cavities occupied by the honey 

bee A. mellifera is 0.29 colonies per hectare in the 

upper Luquillo Mountains, representing a threat to the 

Puerto Rican parrot, Amazona uittata, by occupying 

potential parrot nest sites. Cavities in palo colorado, 

Cyrilla racemiflora, are the nesting sites most used by 

the Puerto Rican parrot. Apparently, palo colorado nectar and pollen are responsible for poisoning the brood 

of &is mellifera in Florida, U.S.A. (Sanford 1987). 

Most swarms of A. mellifera take place from June to 

September, coinciding with the peak flowering of the 

sierra palm (Prestoea montana) on which A. mellifera 

feeds heavily (Snyder and others 1987). 

Formicidae (ants). - Ants (the Formicidae family) 

are numerically the most abundant of the social 

insects. They occur nearly everywhere except on Antarctica, in the extreme north, and on a few oceanic 

islands (Snelling 1981). Janzen (1973a) analyzed the 

interaction between ants and yagrumo hembra, 

Cecropia peltata, in the Caribbean. He postulated that 

a reduction in herbivores and interspecific competition 

from trees and vines, in insular habitats, allows the 

Figure 6. -The carpenter bee Xylocopa mordax (ada9ted from Wolcott 1948d) 
plant to survive without A-teca ants. Fonetheless, 

Cecropia leaves are not defenseless without ants; they 

are physically t~ugh, have dense trichomes, md contain alkaloids and phenolic compounds (Putz and 

Holbrook 1988). Schupp (1986) found that ant-free 

trees suffered more destruction by herbivores in 

Ecuador, but the leaves were not completely destroyed 

and the saplings continued their growth. In Malaysia, 

Cecropia trees without ants suffered less from vines 

than did other pioneer species (Putz and Holbrook 

1988). De Andrade and Carauta (1982) cast further 

doubts on the importance of Azteca ants for the survival of Cecropia. De Andrade and Carauta found 

severe damage caused by insects on these plants. Ants 

attacked only the insects that competed with them for 

the plant resources while ignoring the insects that fed 

in the adaxial surface of the leaves (de Andrade 1984, 

de Andrade and Carauta 1982). 

The level of protection that Cecropia trees receive 

from Azteca ants depends on the ant species that inhabit the tree. Members of the Azteca alfari group 

usually do not provide good protection. Also, 

aggressiveness varies through the ranges of the Azteca 

species (Longino 1991). 

Lavigne (1970a, 1977) presented information on 

nesting sites, colony size, die1 activity, feeding relations, 

and predators on ants at the Luquillo Mountains. On a 

numerical basis, ants may constitute more than a third 

of the diet of many species of frogs and lizards inhabiting the Luquillo Mountains. Lavigne (1970a) developed 

a taxonomic key to identify ants, placing emphasis on 

the antennae and mandibular dentition to aid in the 

identification of isolated heads as typically found in the 

stomachs of predators. Torres and Canals (1983) 

reported on the nesting sites of ants in the different 

forest associations and found a reduction in species 

diversity along an altitudinal gradient at the Luquillo 

Mountains. Hernhdez (1986), working in the dwarf 

forest, reported four ant species and other insects that 

participated in the robbing of nectar from roble de 

sierra, 'lhbebuia rigida. 

Vespidae (wasps). - The family Vespjdae is virtually cosmopolitan, present on all continental land 

masses, except Antarctica, and is found on distant 

oceanic islands. The genus Polistes is worldwide in distribution (Snelling 1981). Nevling (1971) found that the 

most conspicuous insect in the dwarf forest was the 

paper wasp I? crinitus. Many wasps hovering above the 

canopy were observed after the forest became illuminated, but they were not involved in the pollination of 

vegetation. Nests of this wasp are located in open areas, 

not usually found under the closed canopy of the forest. 

Polistes crinitus has been observed in enormous 

numbers on El Yunque Rock (Wolcott 1948d). The 

wasps were so active and abundant that one could not 

safely approach them. Nothing was observed in the 

surrounding vegetation on which they might have been 

feeding. One suggested reason was that air currents 

had carried the wasps up the mountain and concentrated them, despite their efforts to fly away Wolcott 

194811). Another reason could be that these wasps were 

part of a mating swarm. In Florida (U.S.A.), thousands 

of paper wasps are attracted to tall structures such as 

the space shuttle launch pads, where they have been 

observed mating (Landolt and Reed 1990). 

Order Isoptera (termites) 

Termites (the order Isoptera) play a major part in 

decomposing dead trees in the Tropics. 'Ib help digestion, termites have micro-organisms in the hind gut. In 

the more primitive families, obligate flagellate protozoa 

digest intracellularly, but in the Brmitidae, bacteria 

help in the digestion of cellulose. Some members of the 

Termitidae produce their own enzymes to digest eellulose. Most wood eaters leave living wood alone, and 

many make a distinction between dry wood and wet 

wood (Brian 1983, Wilson 1971). 

Martorell (1941) and Martorell and Garcia (1971) 

published a list of the trees infested by the termite 

Nasutitermes costalis (fig.7). This species is common at 

low altitudes; it has not been observed above 933 m. 

Nasutitermes costalis does not feed on live trees and is 

highly abundant at Guhica State Forest and in mangrove areas. Martorell (1941, 1945a, 1976) reported 

food habits and altitudinal distribution of other termite species. Martorell (1976) reported the termite 

Glyptotermes pubescens on the high altitudes of the 

island attacking palo colorado, Cyrilla racemiflora, and 

nuez moscada, Ocotea moschata. 

The first record of the termite G. liberatus for the 

island was made by Martorell (1973) in the Luquillo 

Mountains and Maricao. Snyder (1956) developed keys 

for termite workers and alates from the West Indies. He 

suggested that many termite species in the West Indies 

have a limited distribution. The tropicosmopolitan drywood termite (Cryptotermes brevis) and N. costalis 

readily become established on any island in which 

environmental conditions are suitable (Snyder 1956). 

McMahan (1968b, 1970~) studied the effects of 

cesium 137 gamma radiation on termites and found a 

decrease in both nests and tunnel occupancy in the 

irradiated area (fig. 8). In 1968, 3 years after the 

removal of the gamma source from the radiation tenter, twice as many nests were abandoned by N. costalis 

in the radiation center as compared to the south control 

center, and N. costalis had established a new nest in 

the irradiated area (McMahan 1969). Four years after 

the radiation exposure, tunnel occupancy in the radiation center was comparable to that in the control tenters (table 2). In the radiation center, there was a 

marked increase in tunnels occupied by N. costalis and 

only a slight increase in tunnel occupancy by the termite Parvitermes discolor. In the south control center, 
SOLDIER 

Figure 7. -Worker and soldier of the termite Nasutitermes costalis (adapted frvm 

WoIcoti 1948a) 

there was an increase in tunnel occupancy by N, costalis 

but a decrease by I? discolor. This was correlated with the 

establishment of two new N. costalis nests in this area. 

McMahan (1970a, 1970b) also found behavioral differences (polyethism) in expansion and repairs of nests 

within the worker caste of N. costalis in the irradiated 

and control nests. Nonetheless, the average number of 

termites were slightly lower at disturbed surfaces in 

the irradiated area compared to the control. 

Termites exposed to a cobalt 60 source in the laboratory were more sensitive to radiation than expected 

when compared to other insects of their size (McMahan 

1968a). McMahan discovered significant differences in 

radiation sensitivity between N. costalis and I? discolor, but not between N. costalis and N. nigriceps. In 

addition, workers of all species were more radiosensitive than soldiers (fig. 9). 

Weigert (1970a) conducted a study on the energetics 

of N. costalis, finding that the role of this termite was 

relatively minor in the total decomposer energy budget 

in an El Verde tabonuco forest. The food supply of this 

species is restricted to dead wood not yet in an 

advanced state of decay. Although thoroughly rotted 

wood is attractive to some termite species, it is avoided 

by others, possibly because of its low carbohydrate content (La Fage and Nutting 1978). Weigert concluded 

that the number of termites in an N. costalis colony 

was correlated with the surface area of the arboreal 

nest, possibly due to the necessity for gas exchange to 

occur throughout the nest surface. Although Weigert 

was impressed by the small amount of dead wood consumed by N. costalis, it should be recognized that I? 

discolor is quite abundant in the forest and may be 

more important than N. costalis in the consumption of 

dead wood. Paruitermes discolor can be found in logs 

and branches in advanced stages of decay and has a 

greater altitudinal distribution than N. costalis, which 

is found mostly in the foothills of the forest. Since P. 

discolor does not build arboreal nests, its presence is 

more difficult to detect. In addition, coleopterans (e.g., 

cerambycids) are very important in dead wood consumption in the Luquillo Mountains.] 
Por vif ermes disco/or 

S 15 

METERS FROM MIDPOINT 

t. 

Z 

Figure 8. -Histograms of tunnel-bearing trees ai increasing distances from midpoints of radiation 

and control centers 18 months afier radiation (adapted from McMahan 1970~). 

- CONTROL CENTER 

- a Abandoned 

Table 2.- nnnel survey daia from Nasutitermes costalis in the three centers (adapted from 

McMahan 1970~) 

Irradiated area South control North cut center 

Trees with tunnel^ Trees with Tunnels Trees with Tunnels 

Year tunnels occupied tunnels occupied tunnels occupied 

------- -- Number --------- --------- Number -------- - --------- Number --------- 

1966 90 3 92 22 29* 0 

1967 102 1 108 2 1 78 7 

1968 93 15 136 23 82 9 

1969 89 39 120 44 82 10 

*Survey to 20-m radius only. 
- - - Porvitermes discolor, soldiet 

--- P. discolor, workel 

------ Nosutitermes costolis, soldier 

N. costolis, worker 

DAYS AFTER TREATMENT 

Figure 9. -Soldier and worker survival rates for the termites, Parvitermes 

discolor and Nasutitermes costalis, afler radiation treatments. 

Percentages exceed 100 because all data were corrected for control 

mortality (adapted from McMahan 1968b). 

Order Odonata (dragonflies and damselflies) 

Members of the order Odonata are relatively large 

insects that spend a large part of their time in flight. 

The nymphs are aquatic, and the adults are generally 

found near water (Borror and others). Garcia (1938), in 

the appendix to his ecological survey of the freshwater 

insects of Puerto Rico, listed the aquatic species by 

order, with the exception of the Odonata, which he 

covered extensively. The work of Garcia is rich in information about the Odonata in Puerto Rico. Garcia found 

that species are distributed according to altitude (table 

3), and he summarized the information on the rate of 

predation by insectivorous birds on insects of the 

Odonata. In some insectivorous bird species, as much 

as 84 percent of their stomach contents consisted of 

nymphs and adults of the Odonata. Fish are also heavy 

predators of the Odonata, but data do not exist for fish 

predation on the Odonata in Puerto Rico. There is 

interspecific predation among odonate species (among 

adults and nymphs). Cannibalism is practiced by several species, which could explain the fact that few 

nymphs reach the adult stage. 

Odonates in their adult stage are beneficial because 

they destroy many harmful insects such as mosquitoes. Nymphs could also be beneficial, but their contribution is reduced by the fad that they may destroy 

young fish. Dragonfly nymphs, together with the predaceous water beetle Megadytes gigantea (Dytiscidae) 

(fig. lo), were found to be the worst enemies of tadpoles 

of the introduced toad, Bufo marinus (Wolcott 1948b). 

Adult bullfrogs, Rana catesbiana, feed on ants, cockroaches, spiders, millipides, snails,-and aquatic insects, 

such as dragonfly nymphs, the electric light bug, 

Belostoma boscii, and the dytiscid beetle, Megadytes 

gigantea. These aquatic insects are in turn predaceous 
ADULT MALE 

but did not identify the insect species. Silander (1979) 

found beetle larvae (Coleoptera: Nitidulidae) feeding on 

the seeds of yagrumo hembra, Cecropia peltata. Beetles 

(Nitidulidae) and Lepidoptera larvae eat the mesocarp 

of granadillo, Buchenwia capitaia, fruits at EI Verde 

(Sastre 1979). 

Insect Divemity and Abundance. - Drewry (1970) 

furnished a list of 1,200 insect species collected at El 

Verde, many of which are deposited in the collection at 

El Vexde Field Station. This information, along with 

the findings of Allan and others (1973) on foliage 

arthropods in second-growth vegetation, shows the 

depauperate nature of the Puerto Rican insect fauna. 

Mosquitoes and other blood- sucking Diptera are scarce 

in the Luquillo Mountains despite the prevalence of 

water (Snyder and others 1987). Martorell (1945a) presented several reasons that explain the depauperate 

Puerto Rim insect fauna: (1) Puerto Rico's position in 

relation to the other islands of the Greater and Lesser 

Antilles restricts the arrival of colonizers (Puerto Rico 

is farthest east of the large islands in the Greater 

Antilles and farthest north of the islands of the Lesser 

Antilles), (2) the Antiliean trade winds coming from 

the east or northeast, where land masses are almost 

nonexistent, do not help in the distribution of insects in 

a west to east direction, and (3) the destruction of 

lowland forests and the partial deforestation in the 

mountains might have contributed to the local extinction of some insect species. Estrada and others (1982) 

contributed a list of the flora and fauna near El Verde 

Field Station. For the invertebrates, information is 

presented on the strata where they occur (e.g., soil, 

11. -Larvae aqd adult of the common caddisfly Chimarrha 

albomaculata (adapted from Wokoti 1948~). litter, understory, and canopy). The list has'information on major components of food webs based on abundance or size of the organisms. 

lrious groups. He postulated that the negative asso3tions between mosquitoes and ostracods, copepods, 

id nauplii were the result of competition (fig. 12). 

aguire used glass jars to simulate microhabitats for 

patic organisms and found that cesium gamma radiion had no appreciable effects on these communities. 

3out a year after termination of irradiation, autowed glass bottles filled with water from the nearby 

~nadora River were left in the field to determine the 

;te of invertebrate colonization. The rate of colonizaIn was remarkably rapid. Populations were estab- 

;hed within 2 weeks in these autoclaved glass bottles. 

Seed and Fruit Consumers. - Lavigne (1970b) 

iblished a list of insect species that use fruit and 

eds as food or breeding grounds (table 4). The princi11 users of these resources were members of the 

,ders Diptera, Coleoptera, and Hymenoptera (Foricidae). The study was preliminary in design, and 

ivigne indicated that the data were insufficient to 

ttermine if there was resource specialization. Nieves 

979) found heavy attack by insects on the fruits and 

eds of yagrumo macho, Didympanax morototoni, 

Snyder and others (1987) used sticky-paper strips to 

monitor the abundance of insects in the Luquillo 

Mountains. Insect abundance in disturbed forests was 

double that of virgin forests; and abundance was lowest 

in the palo colorado and dwarf forests. More insects 

were trapped in the sierra palm forest than in the 

tabonuco, although this could be an artifact due to the 

deficient understory growth in the sierra palm forest. 

Diptera comprised 63 percent, Homoptera, 9 percent, 

Coleoptera, 9 percent, Lepidoptera, 5 percent, and 

other insect orders, 16 percent of the insects captured 

in aU forest types. Because a relative method of abundance estimation was used (Southwood 1971), results 

cannot be used to estimate the absolute density of 

insects in the different forest types. Snyder and others 

(1987) found seasonal changes in insect abundance. 

Numbers began to increase in February (during the 

dry season), they peaked in May, at the start of the 

rainy season, and decreased thereafter. The insect 

abundance peak in late spring and summer was correlated with the period of reproduction of many insectivorous birds. The pattern of bird abundance and 
Hor~aclicoid copfpod + -4 ~stracod (long thin shape),-A+ -+ 

I c.4)- 

Flat worm * 

(E 2 4 STANDARD ERRORS 

I==: 2 3 STANDARD ERRORS 

POSITIVE ASSOCIATION - 2 2 STANDARD ERRORS 

I-- !: 1.5 TO 1.9 STANDARD ERRORS 

Figure 12.- Patterns of fwnal associations in aquatic biotas of bromeliads (adapted from Maguire 1970). 

diversity parallels the trends exhibited by the flora and 

insects. The major factor in the reduction of bird species with increasing altitude may be the reduction in 

insect prey. Though the data on insect abundance in 

this study cannot be considered definitive due to the 

limitations in the sampling methods, the study suggests interesting patterns that deserve to be studied 

with more precise sampling techniques. 

McMahan and Sollins (1970) found no cesium radiation effects on microarthropods at El Verde, 2 years 

after the irradiation experiment. However, the study 

lacks replication of the experimental units, and the 

species were not identified. The study does not present 

evidence of the adequacy of the sampling scheme, in 

order to compare the different sites, and lacks the necessary statistical analyses. Although the results indicate that some groups differed in abundance among the 

sites, the above limitations cast doubts on the results of 

the study. Weigert (1970b) found neither radiation 

effects on soil and litter microarthropods 1 year after 

the irradiation, nor a relation between litter standing 

crop and microarthropod abundance. This study also 

lacks experimental replications. 

Drewry (1969) discussed the pitfalls of measuring 

and comparing insect diversity among sites. Linear 

methods, such as William's log series or an incorrect 

assumed constancy of Brillouin's H with increasing 

sample size, .can be misleading. Plotting a diversity 

index curve can reveal nonlinearities, and the composite ratio provides a view of total sample composition, giving a foundation to whichever diversity index is 

chosen. 

Herbivores.-Odum and Ruiz (1970) did not find 

significant changes in the level of herbivory in an irradiated area at El Verde. This was contrary to the findings at Brookhaven, where an epidemic of aphids 

occurred after exposure to radiation. One of the drawbacks of the El Verde study is that the authors did not 

distinguish between insect herbivory and herbivory 

caused by other herbivores, such as snails. 

Fifteen species of Lepidoptera occurred in large 

numbers 7 months after the passage of Hurricane 

Hugo over the Luquillo Mountains. Spodoptera 

eridania (Lepidoptera: Noctuidae) was the most common of the larvae and fed on 56 plant species belonging 

to 31 families. All the Lepidoptera fed on early successional vegetation. The outbreaks appeared to be based 

on the flush of new foliage that developed in the 

Luquillo Mountains after the passage of the hurricane. 

The end of the S. eridania outbreak was concurrent 

with the consumption of its preferred host plants and 

to an apparent increment in parasitism by ichneumonids (Torres 1992). 

A factor that reduces the metabolic production of 

plants in the dwarf forest is damage to the foliage 

(Howard 1969). Howard found that many plants pre- 
nble 4.-lnsects reared from or associated with the tree fruits found in the forest litter, El Verde Field Station, Puerto Rico (Lwigne 1970bj 

Insects 

Tree species Diptera Coleoptera Other orders 

Tipulidae 

Limonia willistoniana* 

Scarabaeidae 

Canthochilurn histemides 

Limonia domestics* 

Ceratopoponidae Endomychidae 

Forczpomyia genualis' Endomychid Bt 

Sciaridae Staphylinidae 

Sciarid B* Staphylinid C 

Sciarid E* Curculionidae 

Stratiomyidae Curculionid C 

Nothornyia nigra* 

Chironomidae 

Chironomid DD 

Chironomid 00 

Psychodidae (undet.) 

Phoridae (undet.) 

Sloanea berteriana Chironomidae 

Chironomid KK* 

Psychodidae (undet.) 

Byrsonima coriacea Drosophilidae (undet.) 

.nclkara bidentata Drosophilidae (undet.)* 

Genipa americana 

Prestoea mntana 

Dacryodes ercelsa Drosophilidae (undet.)" 

Psychodidae (undet.1' 

Chironomidae 

Chironomid KK* 

Eugenia stahlii 

Inga fagifolia Tipulidae 

Limnin willistoniana* 

Chironomidae (undet.)* 

Inga Vera Phoridae 

Phorid Q* 

Nitidulidae 

Nitidulid A 

Nitidulidae 

Nitulid A 

Nitidulidae 

Nitulid A 

Nitidulidae 

Nitulid A 

Staphylhidae 

Staphylinid G 

Staphylinid N 

Scolytidae 

Scolytid F* 

Nitidulidae 

Nitulid A 

Endomychidae 

Endomychid B 

Nitiduiidae 

Nitidulid B 

Hymenoptera 

Formicidae 

Paratrechina sp. (EE) 

Formicid Z 

Hymenoptera 

Formicidae 

Cyphomyrmex minutus 

Hymenoptera 

Formicidae 

Mycocepurus smithi 

Hymenoptera 

Formicidae 

Pkidde moerens 

Soknopsis sp. (JJ) 

Formicid Z 

Hymenoptera 

Formicidae 

Soknopsis ap. (JJ) 

Pheidole moerens 

Orthoptera (undet.)* 

Hymenoptera 

Formicidae 

Pheidole moerens 

Soknopsis sp. (JJ) 

Formicid Z 

Lepidoptera 

Micrulepidoptera (undet.)' 
nble 4.-Insects reared from or associaied with the tree fruits found in the forest litter, El Verde Field Station, Puerto Rico (Lauigne 1970b)- 

(Continued) 

- 

'l'ree species Diptera Coleoptera Other orders 

Guarea guidonia 

Cecidomyiidae 

Cecidomyiid F* 

Lonchaeidae 

Lonchaea sp. (B)* 

Drosophilidae (undet.)* 

Ceratopogonidae (undet.)* 

Mycetophilidae 

Boktina sp. (BY 

Sciaridae (undet)* 

Drosophilidae (undet.) 

*Reared from fruit. 

tLetters represent different species and relate the species tn previour 

sent in the dwarf forest were consumed by insects. In 

six species, the leaf area was reduced by over 16 percent 

(table 5). Succulent young leaves were preferentially 

consumed by insects. Howard (1969) provided common 

names of insects that consumed the plants during the 

study, as well as information about their appearance. 

The author could not correlate leaf texture, liquid content, aromatic constituents, or pH with the degree of 

consumption' by insects. 

Assuming that the only herbivores present were 

insects, Benedid (1976) studied herbivory rates in four 

forests, including the dwarf forest at the Luquillo 

Mountains, and correlated herbivory rates with some 

physical and chemical properties of the leaves. She 

found that plant species with leaves that were tough, 

thick, dry, or highly pubescent were avoided (table 6). 

Many species that had leaves with palatable textures 

were not consumed. These species were either pubescent or members of families that produce aromatic oils 

or calcium carbonate concretions. Most dwarf forest 

leaves do not possess any apparent defenses, although 

many are thick, which may reduce palatability. Several 

dwarf forest plant species belonging to aromatic-oil producing families were among the species with high herbivory rates. Probably, volatile oils do not reduce 

herbivory in the dwarf forest due to the high moisture 

content of the leaves and the constant washing by rainfall. Contrary to the findings of Howard, Benedict considered that the herbivory rate was relatively low in the 

dwarf forest, and she attributed this to unfavorable 

weather conditions for insects, rather than to the lack 

Staphylinidae 

Staphylinid C 

Staphylinid G 

Staphylinid K 

Staphylinid L 

Staphylinid M 

Endomychidae 

Endomychid B 

Scarabaeidae 

Canthochilum borinquensis 

Nitidulidae 

Nitidulid A 

Hymenoptera 

Formicidtie 

Formicid Z 

Myrmelachisto ramulorum 

Pheidole sp. (HH) 

Hymenoptera 

Formicidae 

Formicid Z 

Solenopsis sp. (GG) 

Sokmpsis sp. (JJ) 

Sokmpsis corticalis 

Pheidde moemns 

; diversity studies conducted at El Verde Field Station. 

of palatable food. The low herbivore pressure, due to 

environmental limits on insect populations, may 

explain the apparent palatability of leaves in the dwarf 

forest. Vines (e.g., Marcgrauia sintenisii) in the dwari 

forest had some of the highest herbivory rates of the 

species studied and tended to be low to intermediate in 

toughness, thickness, and dryness (Benedid 1976). 

Insect Biomass. -0dum and others (1970) presented estimates of biomass from different anima 

groups in a tabonuco, Dacryodes excelsa, forest i~: 

Puerto Rico. Although organisms were not sample 

with accuracy, the animal (including the insect) con. 

sumer biomass was small, relative to the great mass o: 

live plant material. 

Pollinators. - ln a study of the flowering cycle i~ 

the dwarf forest, Nevling (1971) found an apparent lac1 

of insect pollinators. Only one definite pollinator, i 

species of bee fly (Bombyliidae), was observed pollinat 

ing the sierra palm, Prestoea rnontana. The lack o 

pollinators could be one of the reasons for the lov 

percentage of fruit set in the dwarf forest. This lack o 

pollinators may be due to the high frequency of rainf'al 

and high humidity, which dilutes nectar concentratioi 

and decreases flower attractiveness. Also, where fruit 

set occurred, the fruits were subject to insect and bin 

predation to the extent that few plant species brough 

fruit to maturation (Nevling 1971). 

Lizard and Frog Predation on Insects.- Liste 

(1981) showed that arthropod densities within th 

Luquillo Mountains remained stable between the sun 

mer and winter, in the lower strata of the forest, bu 
%ble 5.- Plants most susceptible to insect damage in the dwarfforest (adapted fmm 

Howard 1969 with permission of the "Journal of the Arnold Arboretum," 

copyright President and Fellows of Harvard College) 

Tree speciee 

Clusia clusioides 

Haenianthus salicifolius 

Wallenin yunquensis 

Eugenia borinqunsis 

Miconk foveolaia 

Miconia pachyphylh 

Cybianthus sintenisii 

Hornemannia racemosa 

Rqiania cordata 

Pe~emmia hernandiifolia 

Reduction in 

Damage surface am 

Leaf water 

pH* content 

Percent 

3.9-4.3 66 

5.14.2 62 

3.5-4.2 75 

4.8 30 

3.9-4.0 71 

3.7-4.3 60 

4.1-4.5 84 

3.9-5.3 67 

4.9 ... 4.6-5.1 ... 

*The pH of liquid in leaves. 

'Blank fields indicate no data available. 

Qic-v3reased during the winter, at the higher strata. The 

~3~xonomic composition of samples did not differ 

between the seasons. Lister displayed the taxonomic 

distribution of insect prey biomass for three lizard 

species. Members of the Orthoptera, Coleoptera, and 

Hymenoptera (Formicidae) were the most important 

prey items for these lizards. Rivero and others (1963), 

in a preliminary study of the food habits of the frog 

Eleutherodactylus karlschmidti (the most aquatic of 

the Eleutherodactylus) in the Luquillo Mountains, 

found that most of the food eaten by these frogs was 

terrestrial. About 90 percent of the food was composed 

of insects. Dipterans (many aquatic larvae) comprised 

44 percent of the insects. 'R-ichopterans and hymenopterans each comprised about 15 percent of the insects 

consumed. Most of the hymenopterans were ants. 

Lavigne and Drewry (1970) studied the feeding habits 

of frogs and lizards at El Verde, listing the prey items 

consumed by order or class (tables 7, 8). Members of 

the Hymenoptera and Diptera were among the dominant food items. Lavigne and Drewry (1970) also developed a classification of prey types to be used as 

indicators of habitat type. Cintr6n (1970) found a high 

consumption of coleopterans and hymenopterans 

(mainly ants) by tree frogs of the genus Eleutherodactylus in palm (Prestoea montana) forests (fig. 13). 

Bird Predation on Insects. - Wetmore (1916) studied bird predation on insects throughout Puerto Rico. 

The following are highlights of his observations by bird 

species: 

Puerto Rican woodpecker, Melanerpes portoricensis. 

Larvae and adults of wood borers (mainly cerambycids) form 24 percent of this bird's diet. The woodpecker uncovers and drags these larvae from beneath 

the bark, where they are protected from other birds. 

Ants form 10 percent of the diet. La hormiguilla, 

Myrmelachista ramulorum, is one of the preferred 

ants. 

Puerto Rican tody, Ilbdus micanus. Diptera formed 

30 percent, coleopterans, 23 percent, and homopterans, 9 percent of the animal food items consumed 

by this species. Many insects eaten by this bird are 

injurious in coffee plantations. 

Puerto Rican screech-owl, Otus nudipes. This owl 

feeds largely on May beetles (Phyllophaga) and weevils (e.g., Diaprepres) that damage crops and forest 

trees. The owl, a night feeder, is highly adapted to eat 

the nocturnal May beetles. 

Puerto Rican flycatcher, Myiarchus antillarum. Caterpillars and fragments of adult lepidopterans comprise 25 percent, and hymenopterans, 11 percent of 

the animal food eaten by this species. 

Adelaide's warbler, Dendroica adelaidae. This warbler feeds mostly on animal matter and consumes 

enormous numbers of small homopterans. Lantern 

flies (Fulgoridae) appear to be a favorite food item of 

all warblers in Puerto Rico. 

Black and white warbler, Mniotilta varia. Coleopterans constitute the highest percentage of this bird's 

diet. Engraver beetles (Platypus sp.) were also taken 

by this species and by other warblers. 

Bananaquit, Coereba flaveola. This bird feeds mostly 

on small caterpillars and lantern flies (Fulgoridae). 

Greater Antillean grackle, Quiscalus niger. Lepidopteran~ and coleopterans (may beetles and weevils) are 

important items in the diet of this species. 

* Puerto Rican tanager, Nesospingus speculiferus. Caterpillars constituted the large bulk of the animal food 

consumed by this bird. 
Table 6.-Leaf area indices, leaf characteristics, and herbivory raies for ihe irees in the dwarf forest (adapted fmrn Benedict 1976) 

Moisture Toughnesspercent thicknessLeaf area of wet dryness Weight/- Herbivory Browning 

Tree species index Toughness Thickness weight index ratio rate rate 

kgscrn - 2 rnrn 

Alsophila bryophila 

(Cyatheaceae) 0.42 ,.. * 0.32 78 n.d. 10 0.57 0.14 

Calycogonium squarnulosum 

(Melastomataceae) 0.60 4.47 0.33 79 25 9 0.42 0.47 

Calyptmnthes krugii 

(Myrtaceaet) 0.05 9.91 0.44 64 125 20 0.00 7.89 

Eugenia borinquensis 

(MYT~~~) 0.25 9.70 0.91 67 235 18 1.04 0.69 

Gwrnanin berteroniana 

(Bromeliaceae) 0.13 12.77 0.27 86 39 7 0.02 0.54 

Miconin pachyphylla 

(Melastomataceae) 0.12 ... 0.42 66 n.d. 11 0.35 1.34 

Ocoiea spathulnta 

(Lauraceaet) 0.15 7.86 0.44 67 91 13 0.22 2.35 

'Ihbebuia rigida 

(Bignoniaceae) 

Total 2.60 

Weighted average 7.02 0.40 76 75 12 0.53 1.00 

'Blank field indicates no data available. 

tFamily that characteristically produces aromatic oils or cystoliths. 

INSECTS THAT ATTACK FOREST 

TREES AND SHRUBS 

In this section, insects that attack vines, weeds, and 

grasses are excluded; those insects are reviewed by 

Martorell (1976). Some insects presented in Martorell's 

work (e.g., many homopterans) are not included here if 

there is no indication of the damage these insects can 

cause to trees. Several of Martorell's records pertain to 

insects that have been collected only from the Luquillo 

Mountains. Insects that attacked plants or shrubs in 

other areas of Puerto Rico are included here because 

those plants can also occur in the Luquillo Mountains. 

Order Coleoptera (beetles) 

Bostrichidae (branch and twig borere). - Most 

of the species of the Bostrichidae family attack wood, 

either living trees or seasoned lumber. These beetles, in 

rare instances, can cause damage to older dwellings 

and furniture (Dillon and Dillon 1972). Sein (1931) 

reported an outbreak of the boatrychid beetle, Apate 

monacha (fig. 14) in coffee, Coffea arabica, plantations 

and pigeon peas, Cajanus cajan. This wood borer 

attacked live trees of several important timber species 

like cedar. Sein suggested ideas for controlling the 

insect. The bostrychid beetle also attacked many agri- 
Bble 7.-Feeding habits of adult frogs by ser, based on stomach conteni, in the Lquillo Mountains (Lavigne and Drewry 1970) 

Eleutherodactylus Leptodactylus 

coqui portoricensis richmndi wightmanae eneidne antillensis brittoni locustus albilabris 

dPd Pd Pd 0 d dPdPdQ 9 

Number of frogs examined 

%on 17 12 6 88511 2 5 1376331 3 

___________________-----------------.--------------------------- Number of organisms/fmg ............................... -------------------------------- 

Araneida 0.4 0.6 0.3 0.5 0.6 0.4 0.2 0.5 0.2 0.6 0.3 0.5 0.6 1.3 1.0 0.3 

Acarina 0.6 0.3 ... ... ... ...* 3.1 0.4 0.4 2.0 0.2 0.5 0.4 1.6 3.6 5.0 

Chilopods 0.05 ...... ... ... ... ......... ......... ...... 0.2 ... Diplopoda 0.1 0.15 ... ... ......... ......... ... 0.1 0.3 0.4 0.5 0.3 

Chelonethida 0.05 ...... ...... ... 0.2 1.0 ... ......... ......... ... Isopoda ... ...... 0.1 0.3 0.2 ... ... ... ... 0.1 0.15 ......... ... Mollusca ... ...... 0.1 0.1 0.2 0.1 ... ... 0.07 0.25 ... ......... ... Collembola ... ...... ...... ...... .., 0.4 ...... 0.15 0.3 1.3 3.0 ... Orthoptera 0.5 0.9 ... ... ... ... ... ... 0.4 0.4 0.1 0.2 0.2 0.1 0.3 ... Psocoptera 0.05 0.25 ... 0.1 ... ... 0.1 ... 0.2 0.07 ...... ...... 1.0 ... Coleoptera 0.2 0.25 0.5 0.5 0.7 1.8 0.9 ... ... 0.15 0.3 ...... 0.3 ... ... Homoptera 0.6 0.5 0.3 0.4 0.1 ... 0.6 1.5 0.4 0.6 0.4 0.8 ......... ... FIemiptera 0.05 0.15 ... 0.2 ... ... 0.2 ... ... 0.15 0.3 0.25 0.3 ...... 0.3 

Diptera 0.7 4.1 1.3 1.1 1.0 0.4 0.6 ... 1.8 1.8 3.0 1.15 4.6 3.3 3.0 ... Hymenoptera 7.6 6.8 3.5 5.1 1.0 1.0 10.0 11.0 3.0 1.8 6.7 1.8 2.0 3.3 4.0 3.0 

Lepidoptera 0.5 0.5 0.6 0.6 ... ... 0.2 ... ... 1.5 4.6 0.7 1.3 0.3 ... ... Isoptera ... ...... ...... ... 1.3 0.5 ... .................. ... Thysanoptera ... ...... ...... ...... ... ... ......... 0.3 ...... ... - 

*Blank fields indicate no data available. 

Table &-Feeding hobits oflizards, based on organisms collected from lizard stomachs in the Luquillo Mountains (adapted from Lavigne and 

Drewry 1970) 

- - - 

gundlachi stratulus ewrmanni gundlochi ewrmanni 

8 9 d 9 6 juvenile juvenile 

Number of lizards examined 

Number of organismsllizard 

1.0 0.6 

4.5 0.3 

Araneida 

Acarina 

Chilopoda 

Diplopoda 

CheIonethida 

Isopoda 

MoIIusca 

Collembola 

Orthoptera 

Psocopt4?m 

Cofeoptera 

Homoptera 

Hemiptera 

Diptera 

Hymenoptera 

Lepidoptera 

Isoptere 

Thysanoptera 

'Richoptera 

- 

*3lank fields indicate no data available. 
COLEOPTERA 

LARVAE 

DIPTERA 

(CULIC~DAE) 

COLEOPTERA 

HYMENOPTERA 

ICHNEUMONOIDEA 

ISOPTERA 

OLIGOCHAET A 

OTHER DIPTERA 

HYMENOPTERA 

FORMICIDAE 

CHILOPODA 

0 20 40 60 0 20 40 60 0 20 40 60 

STOMACH CONTENTS (PERCENT) 

Figure 13. -Stomach contents of three Eleutherodadylus species of tree frogs trapped in the Luquillo 

Mountains of Puerto Rico (adapted from Cintr6n 1970). 

Imm 

H 

Figure 14. -A wood/twig boring beetle, Apate monacha (adapted from Wolcott 1948b). 
cultural trees including grapefruit, Citrus paradisi; 

orange, Citrus sinensis; avocado, Persea americana; 

and others. Apate monacha attacked Dominican 

mahogany, Swietenia mahogani, near Guayanilla in 

1937. The outbreak was related to the presence of dead 

wood remaining after the cutting of inferior species in 

the mahogany plantation. Dead wood served as breeding grounds for the borer. Some trees were so damaged 

that they died a few months after infestation (Martorell 

1939). During this outbreak, the infestation spread to 

adjacent 4-year-old Australian pines, Casuarina equisetifolia, and to scar trees, Bucida buceras. Apate monacha breeds in dead wood and, when abundant, can 

attack every tree in the vicinity, whether alive or dead. 

Attacks by A. monacha have been reported on the following trees; achiote, Bixa orellana; flamboyhn, 

Delonix regia, guaba, Inga vera; palo de hueso, Chionanthus domingensis; alilaila, Melia azedarach; dry 

post of hueso, Picramnia pentandra; tamarindo, illzmarindus indica; pomarrosa, Syzygium jambos; granada, Punica granatum; and algoddn, Gossypium 

barbadense (Martorell 1945a, Wolcott 1948b). 

Buprestidae (metallic md-boring beetlea). - 

Many adult buprestids are attracted to dead or dying 

trees, logs, or slash. Most buprestid larvae bore under 

bark or in wood (Borror and others 1981). In Puerto 

Rico, buprestids seem to be more abundant in lowland 

habitats, such as dry forests and mangrove areas. 

Wolcott (1948b) reported the introduction of the 

buprestid Polycesta depressa to Puerto Rico as a result 

of the unrestricted lumber traffic from Hispaniola. The 

larvae attack West Indian mahogany (Swietenia 

mahogani) and other trees of lesser economic value, 

feeding on the cambium, when trees are young, and on 

the heartwood, when they approach maturity. 

Cerambycidae (Long-horned beetles) and Elateridae (click beetles). - The brightly colored adult 

cerambycids generally feed on flowers. Many are nocturnal (Borror and others 1981). Most cerambycid larvae are wood borers. Wolcott and Martorell (1942) 

reported the presence of the cerambycid Lagochirus 

araneiformis and the elaterid Chalcolepidius silbermanni (fig. 15) in Puerto Rico. The larvae of elaterids 

are mostly predaceous on other insect larvae, and C. 

silbermanni appears to be a predator of the cerambycid. Wolcott and Martorell (1942) hypothesized that 

these beetles arrived in wood shipped from Santo 

Domingo to Puerto Rico. Mahogany planks and 

unbarked railroad ties were observed to be heavily 

infested with cerambycid larvae upon arrival to Puerto 

Rico. It is possible that the wood and sugarcane trades 

were primary routes for the introduction of many 

insect species to Puerto Rico. 

Cerambycids, along with buprestids and scolytids, 

were attracted to guanacaste trees, Enterolobium 

cyclocarpum, after the trees were infested by a fungus 

that caused an unidentified disease. Weakened limbs 

may fall suddenly in the absence of wind or external 

force, due to their own weight (Francis 198833). 

Curculionidae (snout beetlea). - Beetles that have 

a well-defined beak compose this large family. Most 

snout beetles are plant feeders (Dillon and Dillon 1972). 

They may be found on flowers, leaves, fruits, or roots. 

nYo species of vaquitas, Diaprepes abbreuiatus (fig. 16) 

and Compsus maricao, attacked the seedlings of 

almendrbn, Prunus occidentalis, in nurseries, often 

defoliating the young plants CWoIcott 1956). 

The weevil D. abbrevidus is one of the major causes 

of defoliation in Puerto Rico. Martorell (1945b) listed 

44 plant species attacked by this insect. Among the 

trees attacked by this weevil are the following: acacia 

amarilla, Albizia lebbeck; gongoli, Iler sidemxyloicles; 

moca, Andira inermis; and cedro, Cedrela odorata. 

Diaprepes abbreviatus partially defoliated mahogany 

seedlings and fed on their roots at the Catalina Nursery 

(USDA FS 1981). The larvae caused about 25 percent 

Figure 15.-An elaterid beetle, Chalcolepidius silbermanni. 
Figure 16. -A snout beetle, Diaprepes abbreviatus (adqted from W'da~tt 1948b). 

mortality in transplanted stocks from nurseries at 

Dorado and Monterrey (USDA FS 1976). Outbreaks of 

this weevil resulted in intensive defoliation of ceiba, 

Ceiba pentandra, at different localities in Puerto Rico. 

Diaprepes abbreviatus is one of the favorite foods of 

insectivorous birds in Puerto Rico (Martorell 1945b). 

A weevil larva (subfamily Cryptorhynchinae) was 

reported to attack the bark of eucalyptus trees 

(Eucalyptus deglupta and E. decaisneana) in a trial 

experimental planting on the Rio Abajo Forest. Larval 

girdling and mining often result in windthrow at the 

point of attack (USDA FS 1976). Larvae of the weevil, 

Pseudomopsis cucubam, were reported to feed on the 

seeds of ortegcin, Coccoloba swartzii, but the extent of 

seed damage is not known Wolcott 1948b). 

Platypod idae (p in-hole borers). - The platypodids (fig. 17) are wood-borers. Their larvae feed on 

fungi cultivated in their galleries. Platypuspunctulaius 

appears to be the most abundant member of the 

Platypodidae in Puerto Rico. It was found boring into 

broken limbs of moca, Andira inermis; diseased 

coconut palms, Cocos nucifera; logs of tabonuco, 

Dacryodes excelsa; and almhcigo, Bursera sirnaruba. 

Platypus compositus was collected in a tabonuco log at 

Toro Negro. The platypodid Platypus ratzebui.gi was 

found under the bark of guamb, Inga fagifolia, and in 

logs of guaba, Inga vera, in Lares and Ciales (Martorell 

1976). Platypus ratzeburgi was also found recently 

attacking coffee, Coffea ezcelsa, at Castafier (Gallard0 

1987). Another platypodid, Platypus aisus, has been 

reported in logs of Inga vera at Aibonito (Martorell 

1976). 

Scarabaeidae (scarab beetles). - Individual feeding habits indicate that there are two distinct groups in 

the Scarabaeidae family. In one group, both larvae and 

adults feed on dung, carrion, and skin fixthem. The 

second group feeds on roots, plant juices, and decaying 

wood, as larvae, and on leaves, flowers, and pollen, as 

adults. The latter group includes insects of great economic importance (Dillon and Dillon 1972). Larvae of 

the coconut rhinocerus beetle (fig. 181, Strategus 

oblongus, feed in rotting wood of coconut palms, Cocos 

nucifera; guaba, Inga vera; guamii, Inga fagifolia; 

chinas, Citrus sinensis; tulipan africano, Spathodea 

campanulaia; jobo, Spondias dulcis; and pdma de 

sombrero, Sabal causiarum. In coconut palms, the 

result of larval infestation is nearly complete pulverization of the interior of the wood (Plank 1948b). Strategus oblongus is one of the largest beetles found in 

Puerto Rico. The adult beetle feeds mostly on the ger 

mind tissue and meristem of young coconut palms. 

Adults also feed on young palma real, Roystonea borinquena, and on some palms of the genera Phoenix and 

Pritchardia. 

Approximately 2 years after a major hurricane, 

reports of extensive damage to replanted and naturally 
F'igure 17.-A pin-hole borer, Platypus parallelus (adapted from Equihua and Atkinson 

1987 with permission of "Fdin Entomolbgica Maicam"). 

ADULT 

LARVA 

w 

I cm 

Figure 18.-Adult arid larva of the coconut rhirwcerus beetle Strategus oblongus (adapted from Plank 19486, Wolcott 

1948b). 
regenerated palms appeared (Plank 1948b). Hurricanes 

uprooted or broke off many coconut palms. In subsequent years, the decay of this wood increased the food 

supply for the larvae and ultimately resulted in a population explosion of the adult beetles. The damage to 

coconut plantations caused by this beetle was so large 

that in 1935, as part of a clean-up campaign, a bounty 

of one cent was paid for each beetle, and one-half cent 

was paid for each larva, by the superintendent of the 

Puerto Rico Emergency Relief Administration. After 3 

days of this campaign, 2,405 beetles were collected in 1 

large grove near the town of Rio Grande, where some 

22,000 palms had been blown down by hurricanes 

(Plank 1948b). 

Adults of the May beetle, Phyllophaga portoricensis 

(fig. 19), defoliated large trees of yagrumo macho, 

Didymopanax rnorototoni, at Rio Blanco, Naguabo, 

from 1940 to 1941 (Martorell 1945a). The adults of the 

May beetle l? vandinei fed on the foliage of palma real, 

Rqystonea borinquena, causing considerable damage at 

Isabela in 1941. These beetles feed voraciously in caoba 

hondureiia, Swietenia rnacrophylla (Martorell 1945a). 

Scolytidae (bark and ambrosia beetles). Adults 

and larvae of the Scolytidae family live in timber and 

beneath the bark of trees. Usually, dead or dying trees 

are attacked, but some species live in apparently 

healthy trees (Dillon and Dillon 1972). Boring holes 

caused by the scolytid beetle, Xylosandrus compactus, 

appear to provide infection courts for a fungus that 

causes dieback in mahogany (Swietenia macmphylla). 

Generally, beetle incidence at Catalina, Rio Grande, 

was less than 5 percent, but areas with a beetle incidence of 50 percent have been reported (USDA FS 

1982). The susceptibility of mahogany to these beetles 

was not related to the tree vigor (USDA FS 1984). 

Xylosandrus compactus was also reported attacking 

orchid pseudobulbs, Caitleya spp., in a greenhouse at 

Barranquitas (Martorell and Medina 1974). 

Guaba, Inga uera, and guarn8, I. fagifolia, are two of 

the most important shade trees in coffee groves. The 

scolytid Xyleborus affinis was reported to attack live I. 

vera and I. fagifolia trees, killing many. Infested trees 

recovered only occasionally and, then, only from very 

light attacks (Porto Rico Agricultural Experiment Station 1915, Wolcott 1948b). Martorell (1945a) found X. 

afinis attacking I. fagifolia, the dry fruits of guava, 

Psidium guajava, and the stalks of sugarcane, Saccharum officinarurn Wolcott 1948b). 

The scolytids Xykborus afinis and X. dvulus killed 

apparently healthy trees of acacia amarilla, Albizia lebbeck, in Rio Piedras in 1945. Xyleborus afinis was 

collected in decaying coconut palms, Cocos nucifera; 

and X. voluulus was collected from under the dead bark 

of granadillo, Buchenauia capitata, and in logs of 

almkigo, Bursera simaruba, Wolcott 1948b). Another 

scolytid, X. ferrugineus, was collected from coconut 

Figure 19.- The Puerto Rican May beetle Phyllophaga portoricensis (adapted from Wolcott 1948b). 
palms; the bark of moca, Andira inermis; dead 

alm6cigo trees; and dead bucayo trees, Erythrinapoeppigiana (Martorell 1976). The beetle X. xylographus 

was seen on mango, Mangifera indica (Bright 1985, 

Martorell 1945a). 

Members of the scolytid genus Hypothenemus have 

been collected from several hosts. Hypothenemus 

obscurus was reared from a dead branch of flamboytin, 

Delonix regia, and from fruits of almendra, Terminalia 

caiappa. Hypothenemus obscurus was apparently common in pods of algarroba, Hymenaea courbaril, and 

tamarindo, 'Ihmarindus indica, in several localities in 

Puerto Rico. Hypothenemus trinitatis was collected in 

dead branches of guaraguao, Guarea guidonia (Martorell 1945a). Hypothenemus sp. (near eruditus) was 

collected in seed pods of maga, Thespesia grandiflora; 

H. crudiae, H. obscurus, and H. seriatus were collected 

in the dry fruits of guava, Psidium guajava. Hypothenemus eruditus was collected in pods of emajagiiilla, 

Thespesia populnea; in a dead pole of maricao, Byrsonima coriacea; and in dry twigs of masa, 

Tetragastris balsamifera (Bright 1985; Martorell 

1945a, 1976). Hypothenemus brunneus was recorded in 

the seed balls of the mangrove, Conocarpus erectus, 

and Hypothenemus sp. (near hampei) was found in 

mummied or culled berries of Liberian coffee, Coffea 

dewevrei, in 1941 (Wolcott 1948b). 

The scolytid Arnbrosiodmus lecontei was abundant 

in dead branches of Spanish cedar, Cedrela odorata, at 

El Verde in 1940 (Martorell 1945a); it was also found 

attacking logs of tabonuco, Dacryodes accelsa, and dead 

twigs of aceitillo, Zanthoxylum flavum. Another 

scolytid, Monarthrum sp., was colleded in tabonuco 

logs (Martorell 1976). Plank (1950a) reported that the 

scolytid Poecilips sp. attacked the seeds of mamey, 

Mammea americana. 

Smith (1966) reported that after the termination of 

cesium irradiation at El Verde, bark beetles were very 

active in severely damaged trees, but not in moderately 

damaged ones. He found that bark beetle activity was 

delayed in sabinbn, Croton poecilanthus; ausubo, 

Manilkara bidentata; and palo colorado, Cyrilla 

racemiflora; a fad suggesting that tree species of more 

hydric climatic preference were slower to read to radia- ' 

tion. One problem with Smith's analysis is that he did 

not present information on bark beetle attacks outside 

the irradiated area for control purposes. The tree species early attacked by bark beetles (Inga spp., 

Didymopanax morototoni, Buchenavia capitata, and 

Dacryodes excelsa) at El Verde are highly susceptible to 

bark beetles. Smith did not take into account tree susceptibility when assessing radiation effects. 

Order Diptera (flies) 

Tkphritidae (fruit flies). - The larvae of many 

tephritids feed on plants, and some are serious pests. 

The adults are found on flowers or vegetation (Borror 

and others 1981). An outbreak of the fruitfly Anastrepha suspensa (fig. 20) occurred on the fruits of 

pomarrosa, Syzygium jambos, at Maricao in 1917 (Martorell 1945a). Anastrepha suspensa also causes considerable damage to guava fruits, Psidium guajwa, and 

jobo, Spondias dulcis. The fruits of almendrbn, 

Prunus occidentalis, are edible but are heavily infested 

by A. suspensa CWolcott 1956). Another fruitfly, Anastrepha obliqua, is an important fruit pest of mango, 

Mangifera indica, in Puerto Rico (Martorell 1976). 

Order Herniptera (bugs) 

Pyrrhocoridae (red bugs or stainers). - Members 

of the Pyrrhocoridae family are elongate-oval bugs that 

are usually brightly marked with red or brown and 

black (Borror and others 1981). The cotton-stainers 

Dysdercus andreae and D. neglectus were abundant on 

maga, Thespesia grandiflora, where they were found 

feeding on its seeds (Martorell 1976). Dysdercus 

andreae also feeds on the seeds of ceiba, Ceiba pentandra, and cotton, Gossypium spp. Wolcott 1948b). 

Tingididae (lace bugs). - The lacelike appearance 

of lace bugs (the Tingididae family) is found only in the 

adults; nymphs are usually spiny. Lace bugs are plant 

feeders; their feeding causes a yellow spotting of the 

leaves, with continued feeding, the leaves become 

entirely brown and fall off (Borror and others 1981). A 

heavy infestation of cap6 prieto, Cordia alliodora, by 

the tingid Dictyla monotropidia occurred in several 

parts of Puerto Rico in 1940. The attack caused a 

chlorotic condition on the leaves and subsequent 

defoliation. Many trees were completely defoliated due 

to the intense attack and the high abundance of 

nymphs and adults on the leaves CMartorell 1940b). 

Another lace bug, Dictyla momtropidia, fed on mago, 

Hernandia sonora, along Highway 191 near Mameyea 

in 1940. Chlorosis of the tree$ was reported there (Martorell 1946a). 

Order Homoptera (aphids, scale insects, 

leafhoppers, and mealybugs) 

Aphididae (uphi&).-Aphids are found in large 

numbers sucking sap from the stems or leaves of 

plants. They excrete honeydew, which may be produced 

in sufficient quantities to cause the surface of objeds 

beneath them to become sticky (Borror and others 

1981). The honeydew is a favorite food of many ant 

species. Smith and others (1963, 1971) discussed the 

Puerto Rican Aphididae, providing information on the 

Luquillo Mountains, but did not provide sufficient 

quantitative data on damage to trees. The foliage of 

madre de cacao (Gliricidia sepium), a popular ornamental or hedge plant that was naturalized in Puerto 

Rico and is common in dry regions and lower mountain 
Figure 20.-A fruit fly, htrepha suspensa (adapted from Wdcott 194&). 

I rnrn 

H 

regions, is often attacked by the aphid Aphis craccivora. The aphid's honeydew attracts ants and promotes the growth of fungus over the leaves (Little and 

Wadsworth 1964). 

hterolecaniidae (pit sealea). -Members of the 

Asterolecaniidae family are called pit scales because 

many of them produce gall-like pits in the bark of trees 

(Borror and others 1981). The pustule scale, 

Asterolecaniurnpustulans, was responsible for the near 

eradication of the silver oak, Grevillea robusta, in 

Puerto Rico (Martorell 1943). The scale also killed 

many trees of guara, Cupania americana, and yellow 

cassia, Cassia siamea, which is a good species for firewood. Wolcott (1940~) found the scale attacking maga, 

Thespesia grandifbra; its infested leaves and branches 

turned brown and died. Sometimes as many as half of 

the lateral branches were infested and killed. Wolcott 

suggested the eradication of the introduced tree, C. 

siamea, to reduce the infestation on maga. The pustule 

scale was very common on several exotic trees. 

Asterolecanium pustulans is susceptible to attack by 

three species of parasitic wasps that could control the 

scale in maga if some of the alternative host trees were 

eliminated (Wolcott 1940~). Wolcott's suggestion was 

not taken into consideration since the introduction of 

the ladybird beetle, Chilocorus cactiJ for the control of 

the bamboo scale resulted in the control of the pustule 

scale (Wolcott 1953). The ladybird beetle preferred A. 

1 

pustulans over the bamboo scales A. bambusae and A. 

miliaris. That is an example of fortuitous biological 

control of a pest. 

The scales Astemlecanium barnbusae and A. miliaris 

feed on bamboo. In many countries, these species are 

confined to the common bamboo, Bambusa vulgaris. 

In Puerto Rico, these scales also attack most of the 

introduced species of bamboo. These scales attack B. 

vulgaris from the lowlands to altitudes up to 600 m in 

the Luquillo Mountains (Martorell 1976). Several apecies of predatory beetle were introduced from the 

Guianas and Trinidad in an attempt to control the scale 

(Bartlett 1938). White (1948) reported that the beetles 

achieved good control of the scales. The control of A. 

bambusae by the ladybird beetle, Cladis nitidula, is 

another example of effective biological control. 

Cicadell idae (leafhoppers). - The Cicadellidae 

family is the largest in the order Hemiptera. Leafhoppers are often destructive to certain plants, not only by 

damage caused by their feeding, but also because they 

transmit plant diseases (Ross and others 1982). 

Wood of the roble blanco, lbbebuia hetemphylla, has 

many uses, and the tree is also planted as an ornamental. During long dry periods, the leafhopper Rabela 

tabebuiae feeds on roble blanco, defoliating it or causing 

its leaves to turn yellow (Little and Wadsworth 1964, 

Martor& 1976). Another leafhopper, DikrelZu cedrelae, 

became so abundant on the cedar (Cedrela spp.) leaves 
that premature yellowing and shedding occurred 

(DeLeon 1941, Wolcott 1948a). The damage can affect 

the annual growth rate of cedar trees. These leafhoppers, together with the mahogany shoot borer, Hypsipyla grandella (Lepidoptera-Pyralidae), which causes 

low branching, prevented the cedars from rapidly developing marketable lumber in Puerto Rico (Wolcott 

1948a). The leafhopper Omgalebra cordiae caused yellowing of foliage and subsequent defoliation on cap6 

prieto, Cordia alliodora, in Cayey and San Lorenzo 

between 1940 and 1947 (Martorell 1976). 

Coccidae (soft, wax, and tortoise scales).- The 

females in the Coccidae family are elongate-oval, usually convex, but sometimes flattened, with a hard 

smooth exoskeleton or covered with wax. The males 

may be winged or wingless (Borror and others 1981). 

Soft scales are pests of ornamentals throughout most 

of the world, although they are more abundant in tropical and subtropical areas. 

A heavy infestation of the black scale Saissetia oleae 

on twigs and smaller branches resulted in terminal 

death in moca, Andira imrmis, in Patillas in 1940 

(Martorell 1976). Heavy infestations of the greenishbrown scale, Pulvinaria psiddii, on trunks and 

branches of 4 to 5-year-old Spanish cedar, Cedrela 

odorata, were observed along the Cayey-Salinas road in 

1941 (Martorell 1976). The present scarcity of the 

greenish-brown scale on cedar is due to the introduction of the ladybeetle, Cryptolaemus montmuzieri, into 

Puerto Rico for the control of mealybugs on sugarcane 

Wolcott 1948a). 

The coccoid, Octaspidiotus araucariae, attacked an 

Araucaria cunninghamii plantation in Carite (Francis 

1988a). The twigs soon blackened and fell off, leaving 

the plantation nearly defoliated. If A. cunninghamii is 

attacked by 0. araucarim in all parts of Puerto Rico, it 

will be impractical to grow A. cunninghamii on the 

island. 

Diaspididae (armored scales). Diaspididae is the 

largest family of scale insects. The actual insect has a 

delicate oval body hidden beneath a scale. The appendages are reduced, the body, at maturity, is a little more 

than an egg sac (Ross and others 1982). An intense 

attack of the white peach scale, Pseudaulacaspis pentagona, occurred on ash, Fraxinus uhdei, resulting in 

the destruction of 44 percent of the introduced trees at 

Toro Negro plantations (Martorell 1940a, 1943, 1976). 

Quina, Cinchona spp., (which is used in the preparation of quinine) plantings near Maricao also suffered 

heavy losses (Martorell 1943, Plank and Winters 1949). 

Several trees of almendrtin, Prunus occidentalis, were 

attacked by P. pentagona in a nursery in the Doiia 

Juana Forest (Wolcott 1956), and a heavy scale infestation was observed on a cabfi tree, 'lFemu lumarckianun, in the Luquillo Mountains in 1940 (Martorell 1976). 

Pseudaulacaspis pentagona deformed and killed many 

mahoe trees, Hibiscus elatus, in the Toro Negro Forest 

(nopical Forest Experiment Station 1954). 

The white peach scale attacked toon, 'lbona ciliata, 

seedlings in Corozal and Vega Alta in 1972 (Whitmore 

and Medina 19'74). The scale affects tree vigor and 

form. There are several parasitic wasps (Eulophidae) 

that attack this scale in Puerto Rico (Wolcott 1948d). 

Toon is a promising tree for replacement of cedar, 

Cedrela odorata, one of the most important timber 

trees native to the neotropics. Cedar encountered problems in Puerto Rico related to site requirements and 

attacks by the shoot borer, Hypsiwla grandella (Lepidoptera: Pyralidae). 

The black thread scale, Ischnaspis longirostris, 

attacked guaraguao, Guarea guidonia, quite severely at 

El Verde. It also caused chlorosis followed by defoliation 

on caoba hondureiia, Swietenia macmphylla, in 1940 

and 1942 (Martorell 1940b, 1976). 

Margarodidae (giant coccids and ground 

pearls). -The Margarodidae family includes some of 

the largest species in the superfamily Coccoidea. Some 

species may reach a length of about 25 mm. The name 

ground pearls was given because of the pearllike 

appearance of the cysts of females in the genus Margarodes; those insects live in the roots of plants (Borror 

and others 1981). The cottony cushion scale Iceryapurchasi (fig. 21) attacks the needles of the Australian 

pine, Casuarina equisetifolia, and is also a pest of 

citrus in Puerto Rico. The scale is controlled by the 

Figure 21. -The cottony cushion scale Iwrya pwchasii. 
imported ladybird beetle, Vedalia cardinalis CWolcott 

1943b, 1960). In some humid citrus groves, the fungus 

Spicaria javanica, caused complete eradication of the 

scale CWolcott 1951); that was an excellent example of 

biological control of an insect pest in Puerto Rico. 

Pseudococcidae (mealybugs). - The Pseudococcidae family is cosmopolitan in distribution. The 

mealybugs make no scale but secrete waxy filaments 

that are especially noticeable about the body margin 

(ROSS and others 1982). The mealybug, Nipaecoccus 

nipae, is the most common member of the genus found 

in Puerto Rico. It caused damage to 32 species of trees 

and shrubs, many found in the Luquillo Mountains 

(Martorell 1945b). It is found from the lowest elevations 

in Puerto Rico up to around 830 m, but rarely at higher 

elevations. The insect is controlled in Puerto Rico by a 

group of parasites and predators, among them, ladybird 

beetles and entomogenous fungi (Martorell 1945b, 

1976). 

Order Hymenoptera (ants, bees, sawflies, and 

wasps) 

Argidae (sawflies). -Most of the argids are black 

or a dark color. The larvae feed mainly on trees. The 

sawfly, Schizocerina krugii (fig. 22), is the only species 

of this family found in Puerto Rico. The larvae of S. 

krugii cause intensive defoliation on seagrape, Coccoloba uvifera, which has been introduced to the 

Luquillo Mountains (Little and Woodbury 1976). Schizocerina krugii is also considered a pest of moral6n, C. 

pubescens, and other Coccoloba species (Martorell 

1976). 

Formicidae (ants). -The use of seeds to supplement a diet based on insect prey is common in ants of 

the subfamily Myrmicinae. Many of the seeds that the 

ants transport are not eaten; thus the ants contribute 

to the dispersal of plants (Brian 1983). Natural 

regeneration of the Australian pine Casuarinu equiI ADULTS 

I J 

Figure 22.-Adults, larvae. cocoons, and eggs of the tenthredinid wasp SchFzmrina krugii (adapted from Martorell 194561 
setifolia is rare in Puerto Rico due to seed consumption 

by ants (Little and Wadsworth 1964). The introduction 

of guam6 venezolano, Inga quaiernclta, for use as shade 

in coffee plantations, failed because the tree is susceptible to the ant Myrmelachista ramulorum, a pest of 

coffee. Nonetheless, the tree survives well in several 

areas in Puerto Rico (Little and Wadsworth 1964). 

Myrmelachista ramulorum forms irregular tunnels 

in coffee trees. In some parts of the tunnels, the ants 

tend mealybugs or scale insects, which suck sap from 

the trees and reduce the production of coffee berries 

(Wolcott 1933, 1951). Tunneled coffee trees are easily 

broken by wind and storms. This ant is highly abundant in the canopy of Puerto Rican forests and can be 

responsible for the death of small branches. Myrmelachista ramulorum infestations on pomarrosa, Syzygium jambos, caused the death of more than 50 

percent of the tree terminals at Lares in 1940 (Martorell 1976). 

Order Lepidoptera (butterflies and moths) 

Cosmopterygidae (cosmopterigid moths).- 

Most of the members of the Cosmopterygidae family 

are leaf miners in their larval stage. Severe infestations 

of the palmetto leaf-miner, Homaledra sabalella, 

occurred in palma de sierra, Prestoea montana, in the 

Luquillo Mountains, Toro Negro, and Cayey in 1940 

(Martorell 1945a). The caterpillars of H. sabalella are 

gregarious and feed on the underside of palm leaves. 

This moth also feeds on coconut palms and ornamental 

palms. The moth occurs in all parts of the island 

(Wolcott 1948~). Parasitic wasps (Chalcididae) are 

responsible for the control of the caterpillars of H. 

sabalella in the field (Martorell 1945b). 

Gelechiidae (gelechiid moths). -Members of the 

Gelechiidae family may mine, roll, or web together 

leaves of several plants. Some species produce galls, 

others are pests in stored seeds, cereals, and tubers 

(Peterson 1984). 

Maga, Thespesia grandiflora, was formerly planted 

for timber throughout Puerto Rico and for esthetics 

along roadsides. Its wood is superior to mahogany in 

many aspects (Wolcott 1940a). Since maga was an alternative host to the pink bollworm, Pectinophra gossypiella, its planting was discontinued in the cotton 

regions (Little and Wadsworth 1964). Pectinophra gossypiella feeds on the pods or seeds of maga and emajagiiilla, T. populnea. 

Geometridae (measuring worms). - The larvae 

of geometers have two or three pairs of prolegs at the 

posterior end of the body and none in the middle. The 

locomotion is accomplished by placing the posterior end 

of the body close to the thoraxic legs and then moving 

the anterior end, thus progressing in a looping fashion 

(Borror and others 1981). The larvae feed chiefly on 

foliage (Peterson 1984). Larvae of the moth Semiothisa 

diffusata (recorded as Macaria diffusata) defoliated 

flamboyh, Delonix regia, trees in Guhica in 1916 

(Martorell 1976). 

Hyb2aeidae lhyblaoid moths). - The Hyblaeidae is 

an Old World tropical family. The leafroller, Hyblaea 

puera, is an introduced species in Puerto Itico. The 

caterpillars of H. puera do considerable damage to teak, 

Tectona grandis, in the Orient (Forbes 1930). Hyblaea 

puera heavily infested capd blanco, Petitia domingensis, in a USDA Forest Service nursery at Cayey in 

1937. Subsequently, only small outbreaks have 

occurred (Martorell 1939). 

Megalopygidae (flannel moths). - Moths of the 

Megalopygidae family have a coat of scales mixed with 

fine curly hairs, which give the moth a somewhat 

woolly appearance (Borror and others 1981). The larvae 

have stinging spines under the hairs and can cause 

severe irritation to humans. The family is mainly 

found in the New World, in the Tropics and warm 

temperate regions (Forbes 1930). 

The larvae of la plumilla, Megalqyge krugii (fig. 23), 

caused considerable damage and defoliated several marim trees, Byrsonima coriacea, at Aguas Buenas in 

1940. An outbreak of caterpillars occurred on moca, 

Andira inermis, causing complete defoliation at Poncx! 

(Martorell 1945a). Guamd, Inga fagifolia, one of the 

most important shade trees in coffee groves, was 

attacked by the larvae of M. krugii. Caterpillars were 

very abundant, causing considerable damage to leaves 

(Porto Rico Agricultural Experiment Station 1915). 

Megalopyge krugii defoliated flamboyh trees, Delonix 

regia, at Ponce in 1940 (Martorell 1945a). 

Noctuidae (noctuid moths). - Noctuidae is the 

largest family in the order Lepidoptera. The larvae are 

usually leaf feeders. Some are stem or root borers, and 

others gnaw in fruits (Ross and others 1982). 

The flarnboyan, Delonix regia, widely planted as an 

ornamental and found in some island forests, is 

attacked occasionally by the caterpillars of the moth 

Melipotis acontioides (recorded as Lyncestis acontioides). nees can be completely defoliated (Little and 

Wadsworth 1964) and damage can be extensive 

(Wolcott 1945b). This moth is usually a lowland insect 

and has not been observed at middle elevations. Caterpillars of another moth, Spodoptera frugiperda, 

attacked small seedlings of Eucalyptus robusta in 

USDA Forest Service nurseries at Rio Piedras (Martorell 1976). The caterpillars were kept under control by 

parasitic larvaevorid flies, parasitic wasps, and 

entomogenous fungi. Caterpillars were also attacked by 

a carabid beetle and an assassin bug. Insectivorous 

birds and some lizards preyed on the caterpillars or the 

moths. Caterpillars of the moth Diphthera festiva 

(recorded as Noropsis hieroglyphics) bored in .the 

trunks of Casuarina equisetifolia causing severe 

damage to young trees at Guhica in 1923 (Martorell 

1976). 
Figure 23.-Adults and cocoons of la plumilla Megalopyge krugii (adapted from Wolcott 1948~). 

Psychidue (bagworms). - The adult males of the 

Psychidae moths have wings, but the females are wingless, legless, and usually never leave the bag in which 

they pupated. The caterpillars of Oiketicus kirbyi 

almost defoliated trees of guano, Ochroma lagopus, at 

Catalina in the Luquillo Mountains in 1940. Also, caterpillars of 0. kirbyi caused defoliation of several 

almendra, Termimlia catuppa, trees at La Muda in 

1938 CMarttorell1945a). 

Pyralidae (snout and grass moths). - The Pyralidae is one of the most economically important families 

in the order Lepidoptera. The larvae, some being 

saprophagous and a few, predaceous, exhibit a broad 

range of habits such as feeding on leaves, fruits, 

flowers, and boring in stems (Ross and others 1982). 

The shoot borer, Hypsipyla grandella, does only 

minor damage in native forests, but causes severe 

damage in mahogany plantations (Hodges and McFadden 1987). HypsiM grandella larvae bore into stem 

tissues and hollow out the shoots. This results in retardation of stem growth, profuse branching, and possible 

shoot deformities. After continual attack, the trees 

either die or become so deformed that the possibility of 

reaching merchantability is minimal (Gara and others 

1973). The ichneumonid wasp, Calliephialtes ferrugineus, attacks the shoot borer (Martorell 1945a, 

1945b; Wolcott 1948d). 

In Puerto Rico, Dominican mahogany (Swietenia 

mahogani) is seldom attacked, but Honduras 

mahogany, S. macrophylla, is highly susceptible to H. 

grandella. Weaver and Bauer (1986) reported infestation rates as high as 57 percent (table 9) in areas using 

line plantings, a silvicultural technique in which most 

of the native forest is left intact. Mahogany trees in 

plantations or dense line plantings present ideal conditions for maintaining large borer populations. Planting 

mahogany alternately with other species in line planting~ in Africa has provided some control of the shoot 

borer (Roberts 1968). Attacks are either reduced or 

nonexistent in shaded plantations in some instances, 

often due to factors such as low seedling density (Whitmore 1978b). Weaver and Bauer (1986) suggested that 
Table 9.-Percent of shoot borer damage to suruiuing trees in line 

plantings established in 1974 and 1980 in Pwrto Rico 

(adapted from Weaver and Bauer 1986 with permission of 

lLrrialba) 

- 

ks ks 

Year damaged measured 

Percent Number 

1974 57 214 

1979 11 672 

1980 17 643 

the presence of surrounding shade helps in the control 

of weeds and probably assists the mahogany tree in the 

development of a single shoot, even if infested by the 

moth. Weaver and Bauer (1986) also suggested that 

chemical control of the borer is most easily accomplished when seedlings are kept in the nursery until 

they reach at least 1 meter in height. This will reduce 

weeding costs and the time available for infestation to 

take place. Mahogany grown on open sites that were 

extensively site prepared suffered severe attacks by the 

borer (USDA FS 1976). Gara and others (1973) found 

that the attacked mahogany trees had new flushes of 

growth. The most serious attack occurred on trees with 

a high new leaf to total leaf ratio. This indicates that 

faster growing trees that; produce abundant new leaves 

are more subject to attack than moderate- to slowgrowing trees. The fad that shaded trees are slower 

growing than open grown trees may explain the conclusions of silvicultural studies that recommend growing 

mahogany under shade to prevent attacks by H. 

grandella. 

Toon, Tbona ciliaia, (an Australian Meliaceae) is not 

attacked by H. grandella in Latin America, but it is 

heavily attacked in its native habitat by the moth H. 

robusta. It appears that the family Meliaceae has 

evolved the capacity to produce poisonous substances 

to deter Hypsipyla. The shoot borers have coevolved 

with each particular tree species, adapting to the secondary substances to the point that the toxins have 

become an attractant to the insect. Due to chemical 

differences between the compounds produced by different Meliaceae species, trees introduced into new geographical areas will be immune to the native shoot 

borer for an unknown period (Gara and others 1973). 

Contrary to the situation in Puerto Rico, in Haiti the 

shoot borer prefers S. mahogani, which is native, and 

refuses S. macrophylla. This suggests that the borer 

and S. mahogani coevolved in Haiti and supports the 

conclusion that S. rnahogani is not native to Puerto 

Rico. Apparently, the shoot borer has not evolved a 

taste for S. mahogani in some 200 years since the tree 

was introduced to Puerto Rico (Whitmore and Hinojosa 1977). Recently S. macrophylla has been attacked 

by H. robusta in Timur (Ngatiman 1989). 

Spanish cedar, Cedrela odorata, a member of the 

mahogany family, is also attacked by H. grandella in 

Puerto Rico. Trees growing in the open were attacked 

more often than trees growing under shade. Whitmore 

(1978a) found a weak relationship between vertical 

growth and borer attack (the borer tends to attack trees 

that grow fast and have fresh, green growth). Lamb 

(1968) found that heavy resin flow in vigorous trees 

discourages the insect. 

The roble leaf-webber, Eulepte concordalis, is considered a destructive foliage pest of roble rosado, llhbebuia 

heterophylla, and roble de sierra, T. rigida. This moth 

occurs from the lowlands to elevations of 800 m. About 

50 percent of the trees in a large group were attacked 

by the larvae of this species along the Mount Britton 

trail in the Luquillo Mountains in 1940. The larva of E. 

concordalis is a leaf-webber that eats the leaf epidermis 

of Tabebuia and higiiera, Crescentia cujete. After 

attack, trees exhibit skeletonized, brownish leaves that 

are webbed together in bunches. Eulepte concordalis is 

parasitized by the tachinid Sturmia albincisa, the 

chalcid Brachynteria incerta, and the braconid Bracon 

cushmani (Martorell 1940~). The caterpilIars of the 

moth Epicorsia oedipodalis (recorded as Pyrausta 

cerata) fed on p6ndula de sierra, Citharexylum 

caudatum, in the Luquillo Mountains, but there was no 

indication of the damage caused by this species. Epicorsia oedipodalis caterpillars caused heavy defoliation on 

p6ndula colorada, Citharexylum fruticosum, particularly in the middle elevations of Puerto Rico. The caterpillars are parasitized by the ichneumonid wasp, 

Eiphosoma insularis. Another moth, Pachymorphus 

subductellus, is a common twig borer of %buia heterophylla. It also attacked Wbuia rigida. The caterpillars of Tetralopha scabridella caused heavy 

defoliation to guaba, Inga Vera, during the insect's 

peak of abundance (Martorell 1945a, 1945b, 

1976). 

The caterpillars of Agathodes &signalis (fig. 24) are 

leafrollers and twig borers in bucayo, Erythrina spp. 

This moth is a voracious pest that causes heavy defoliation and destruction of new shoots, year after year. The 

caterpillars also bore into guaba, Inga uera. The caterpillars of Erastia meticulosalis (fig. 25) are twig and 

pod borers of Erythrina fusca. They infested 90 percent of the trees in an experimental planting at Rio 

Piedras in 1921. Terastia meticulosalis attacked other 

Erythrina species and cap6 blanco, Petitia domingensis 

(Martorell 1945b, 1976). 

The moth Phostria originalis attacked moca, Andira 

imrmis, in several towns in February and March of 

1940 (Martorell 1976). This insect is a webworm and is 

found in clusters of 80 to 100 larvae at the end of 

branches. Its larvae can completely defoliate trees 

(Martorell 1939). Another moth, Blepharomastix 

ebulealis, has smd, but very voracious, caterpillars 

that feed on camasey peludo, Heterotrichum cymosum. 
t I cm -I 

* 

Figure 24. -A pynJid moth, Agathodes designah ~~~ 

I cm I 

* 

Figure 25.-A pymlid moth, Terastia meticulosalis (adapted fmm Wdcott 19481:). 
Figure 26.-Adult of the Cecropia leaf-dkr Syllepte silicalis (adapted from Wobtt 1948~). 

Figure 27.-AduU of the thrip Selenothrips rubrocindus. 

Blepharomastix ebulealis infested many trees but most 

of the caterpillars were parasitized by larvaworid flies 

(Martorell 1945b). 

The caterpillars of the yagrumo leafroller, Syllepte 

silicalis (fig. 26), fed on the buds of terminal branches 

of Cecropiapeltata, along the Mount Britton trail in the 

Luquillo Mountains, causing considerable damage. 

There are four additional species of Lepidoptera that 

feed on C. peltata in Puerto Rico (Martorell 1976). 

Order Thysanoptera (thrips) 

Most thysanopterans are between 2 and 3 mm long. 

Thrips occur commonly in flowers. Many species are 

destructive to plants and are found in the leaves of the 

infested hosts (Ross and others 1982). The thrips, 

Selenothrips rubrocinctus (fig. 271, attack kadam trees, 

Anthcephalus chinensis, reducing their leaf area, but 

do not kill them (Lugo and Figueroa 1984). Thrips 

attack old leaves, but damage does not appear visible 

until the leaves are nearly ready to drop (USDA FS 

1976). Heavy feeding on leaves caused premature leaf 

drop and, ultimately, complete defoliation and death of 

trees (USDA FS 1972). A growth study is needed to 

measure any effects of infestations by thrips on tree 

height and diameter over time. If damage is significant, 

possible solutions may include growing kadam in mixed 

stands or selection of resistant clones. Clone selection 

in cocoa, Theobroma cacao, was successful in reducing 

damage by the same species of thrips in Trinidad 

(USDA FS 1976). This thrip also attacks almendra, 

Z'krminalia catuppa, causing a silvery appearance in 

the leaves (Martorell 1976). 

Other Insect Groups 

El Servicio de Extensicin Agricola (1948) listed the 

common names of insects that attack laurel sabino, 

Magnolia splendens; guaba, Inga vera; pines; 

eucalyptus; etc. Several suggestions for dealing with 

these insect pests are available. 
Plank (1950a) studied the insecticidal properties of 

48 species of plants in 23 families. Thirteen local, economically important species of insects belonging to four 

orders were tested. Mamey, Mammea americana, 

exhibited the greatest insecticidal potential of the 

plants examined. The half-ripe fruits, seeds, and leaves 

showed insecticidal value, either as an internal poison, 

as a contact poison, or both. The active chemical in the 

mature seeds (the most toxic part of the plants) was 

mameyin, a substance that is similar in composition 

and has similar effects to those of pyrethrins. One of 

the earliest uses of mamey in the Caribbean was in the 

control of fleas that attack humans and domestic animals. Mamey leaves were used successfully for many 

years in Puerto Rico to wrap the trunks of newly set 

garden plants for the prevention of insect attack (Plank 

1950a). 

INSECTS !ITEAT ATTACK FOREST 

PRODUCTS 

Order Coleoptera (beetles) 

Bostrichidae (branch and twig borere). - In 

Puerto Rico, harvested bamboo (one of the most commonly used plants in the Tropics) is attacked by the 

escarabajo del bambil seco, Dinoderus minutus, a 

member of the Bostrichidae. Few beetles are found in 

the field, but high populations may be found under 

storage conditions White 1948). Dead or dying culms 

left in the field are rarely attacked (Plank 1948a). 

Severe infestations weaken and mar the stored culms 

to the extent that they are worthless. Bamboo culms 

that do not become infested within 1 or 2 months are 

usually not attacked later. 

The adults of D. minutus appear to be crepuscular, 

and they attack dry vegetable products like corn, Zea 

mays. A braconid parasite, Doryctes jarvus; a 

pteromalid parasitic wasp, Pmamotura aguila; and a 

reduviid predator, Peregrinator biannulipes, proved to 

be ineffective in reducing the beetle populations before 

considerable damage had occurred (Plank 1939,1948a). 

Plank (1950b) found that infestations were correlated 

more with starch concentration (table 10) than moisture content or specific gravity of the wood (table 11). 

Susceptibility to the beetles varies along the length of 

the culm. The distal ends of culms are more resistant 

than basal parts. The susceptibility to the attack by 

the beetle differed among species and varieties of bamboo. In addition, susceptibility was correlated with the 

age of the bamboo at harvest. Plank (1950b) proposed 

ways to protect the wood from the beetle. White and 

others (1946) studied the relation between curing and 

durability in bamboo, Banbusa tuldoides, finding that 

even the best curing treatment resulted in 66 percent 

of infestation by the beetle. 

Boone and others (1969) discussed chemical control 

of the bostrichid Tetrapriocera longicornis during airdrying of fenceposts. Fenceposts require drying to 

reach a moisture content compatible with a preservation treatment. During the drying interval, posts are 

attacked by the bostrichid T. longicornis. This 

bostrichid can also attack freshly felled wood. 

10. -Susceptibility of I-yead culm of various species and 

varieties of bamboo to attack by the bamboo powderpost 

beetle compared with Bambusa vulgaris and the corresponding iodine-starch score at time of erposure to infestation (adapted fforn Plunk 19SOb) 

A=rage 

iodine-atarch 

Bamboo species Susceptibility score 

Bambusa vulgMis 

B. vulgar% vittato 

B. pdymovha 

Dendmalamus strictus 

(large-leafed var.) 

B. tuldoides 

D. membmnaceous 

D. giganteus 

B. longispiculata 

B. tuldO 

B. tuldO* 

Siriadamu oldhami 

B. tQtilis 

Percent Number 

'Variety P.I. No. 74413. 

Table 11. -Moisture content and specific pity of wood in 1-yeadd 

culms of various species and varieties of bamboo at the 

time of erposure to infestation, with comsponding number of powderpost beetk attacks (adapted fmm Plank 

195Ob) 

Average* Aventge A.clerage attacks 

moiatura specific m6OW 

 amb boo species content iimitv ~ieces 

Bambusa tdlis 

B. longispiculatn 

Dendrocalarnus 

membranuceous 

B. vulgaris vittata 

B. vulgaris 

B. tulda 

D. strictus 

(large-leafed m.1 

B. tuMoides 

B. polymorpha 

B. tuldat 

Sinocalamus oldhumi 

D. gigantew 

Percent glcma Number 

- - -- 

*Because moisture content is expressed as a percentage of the 

weight of the ovendry wood, it is possible for this percentege to be 

100 or greater. 

+Variety P.I. 74413. 
Lyctidae (powderpost beetles). - Beetles of the 

Lyctidae family attack seasoned timber (preferring a 

moisture content of 10 to 18 percent), the pith of vines, 

and the dried roots of herbaceous plants (Dillon and 

Dillon 1972). The lyctid, IZ-ogoxylum aequale, was 

reported to attack the sapwood but to spare the heartwood of furniture made of West Indies mahogany, 

Swietenia mahogani (Wolcott 1948b). 

Two species of the Lyctidae family were reported to 

infest cut culms of bamboo, but the damage done was 

relatively minor (Plank 1948a). 

Order Hymenoptera (ants, bees, wasps, and 

allies) 

Eumenidae (mason and potter wasps).- Wasps 

of the genus Zethus are frequently attracted to flowers. 

They use old ins& burrows in twigs or wood for nesting (Bohart and Stange 1965). Larvae of the wasps feed 

on caterpillars deposited in the nest by the reproductive 

female. These wasps do not use mud to line the nest 

tunnels (Bohart and Stange 1965, Wolcott 1948d). In 

Puerto Rico, the eumenid wasp, Zethus ruffnodis, 

apparently causes damage by burrowing in fenceposts 

made of different kinds of woods (Martorell 1941). 

Order Isoptera (termites) 

There are 17 termite species in Puerto Rico, and 4 

are economically important. These are Nasutitermes 

costalis, the dry-wood termite, Cryptotermes brevis, 

and the subterranean termites, Heterotermes tenuis 

and H. convexinotatus (Wolcott 1921, 1939, 1946~). 

Nasutitermes costalis builds arboreal nests that are 

easily seen and destroyed, either mechanically or chemically. Nasutitermes costalis requires subterranean 

sources of water in addition to water obtained from 

dead branches. The termite will perish if the ground 

water supply fails (Wolcott 1954). It is mainly a rural 

dweller that can destroy fenceposts and other wooden 

structures. 

The subterranean termites (Heterotermes spp.) may 

occur anywhere in Puerto Rico, and the geographical 

area of infestation appears to be increasing. The historical concentration of subterranean termites was in the 

cities. These species do not build visible nests, and for 

that reason they are difficult to control. Species of 

Heterotermes feed on both topwood and roots and can 

penetrate concrete houses through cracks in the floor 

or through pieces of wood that contact the soil Wolcott 

1939, 1954). Heterotermes tenuis was considered the 

most destructive of the subterranean termites in much 

of the New World Tropics. In Trinidad, H. tenuis was 

reported to attack all dead wood in contact with the soil 

within a few months (Brooks and others 1941). 

The dry-wood termite, C. brevis, is able to obtain all 

required moisture from dry wood. Cryptotermes brevis 

does not appear to live in the ground at any point of its 

existence, and it usually does not live outside buildings. 

Only the heartwood of West Indies mahogany, 

Swietenia mahogani, is resistant to C. bmis; but even 

this wood may be eaten to a slight extent (Wolcott 

1940b). Philippine mahogany, Shorea negrosensis; 

African mahogany, Khaya ivorensis; and even Honduras mahogany, Swietenia macrophylLa, are damaged 

by C. brevis (Wolcott 1954). lkrmite resistance of S. 

macropylla heartwood is comparable to the resistance 

of S. mahogani sapwood, which is relatively low 

(Wolcott 1948e, 1950). 

The defenses of wood against termites involve toxic 

or repellent chemicals, adverse physical characteristics, 

moisture content, nutrient imbalance or lack of specific 

growth factors needed by the insects, and microbial 

associations present in the wood (La Fage and Nutting 

1978, Waller 1988). These factors vary inter- and intraspecifically (e.g., a log may contain palatable wood near 

unacceptable wood). In addition, the termites may 

respond to these defenses in a hierarchical fashion such 

that wood may be accepted or rejected depending on 

other available food. Woods detrimental to protozoans 

living in the hind gut of termites can be eaten if beneficial woods are also available. Workers that have lost 

their protozoans can be reinoculated by their nestmates (Waller 1988). Age, degree of seasoning, fungal 

damage, weathering, and even peculiarities of individual trees also affect termite attack (Brooks and 

others 1941). 

The extractable organic compounds deposited in the 

heartwood during its transformation from the sapwood generally are more important contributors to the 

natural resistance of wood than are inorganic constituents or physical factors, such as wood density and 

degree of cellulose crystallinity. Due to the mechanical 

nature of termite attack on wood, density probably 

plays some initial protecting role, since harder woods 

are more difficult to fragment by termite mandibles. 

For woods with low density, antitermitic substances 

must constitute the major line of defense (Bultman and 

Southwell 1976). The natural resistance of wood to 

termites tends to decrease with time. The following 

mechanisms have been proposed to explain this tendency: oxidative enzymes of host origin detoxify the 

protectant, auto-oxidation of the protectant, microbial 

or fungal degradation, or leaching of the protectant 

over the years (Bultman and Southwell 1976). 

Wolcott's work (1921, 1924, 1939, 1940c, 1943a, 

1945a, 1946a, 1946b, 1946c, l947,1948e,1950) is often 

cited in publications dealing with wood resistance to 

termites (e.g., Little and Wasdworth 1964, Longwood 

1971, Weaver 1988). Nonetheless, his work deals mostly 

with the dry-wood termite, C. brevis. There is some 

discussion by Wolcott on the habits of N. costalis and 

Heterotermes spp., but his research on wood resistance 

to termite attack applies only to C. brevis. 
Wolcott's (1943a, 1945a, 1946c, 1947) experiments on 

termite repellents have several drawbacks. Among 

them: lack of replications (sample size is one), use of 

several treatments in the same experimental petri 

dishes, and use of the same termites in subsequent 

experiments. The use of several experimental treatments in the same petri dish could create interactions 

that could affect the results of the experiments. In the 

case of use of termites from previous tests, the results 

could be affected by previous exposure of the termites 

to chemical substances. McMahan (1966) found that 

conditioning in C. breuis affects food selection. Rrmites preferred poplar, on which they had been previously fed, instead of maple, to which they were 

unaccustomed, though maple was preferred over poplar 

when the termites had not been conditioned. 

Wolcott (1924,1940b, 1946b, 1948e, 1950) elaborated 

lists of the comparative resistance of several woods to 

the attack of the dry-wood termite, C. breuis. The information on these lists can only be used in a qualitative 

manner. Because of sample size, none of Wolcott's 

papers presented estimates of mean resistance and 

variance to termite attack. In many of his experiments, 

the results on wood preference by C. breuis were contradictory (see Wolcott 1924). Wolcott (1940b) tried to 

explain the inconsistency of his experimental results 

and recognized that "the positions assigned in the list 

for each wood are relative only to some of the other 

woods, and not absolute for each of the other woods." 

The contradictions were often easily dismissed as can 

be seen in the following explanation: "While one cannot 

be sure that all of the samples used in either of these 

series of tests were typical of the species, the error due 

to this fador is naturally less in woods of most uniform 

composition." Also, wood samples used in his experiments were of straight grain, without knotholes or 

other imperfections, to avoid usual or exceptional 

points of attack by the termites (Wolcott 1940b). The 

presence of these factors could facilitate termite attack 

under normal circumstances, affecting the ranking of 

woods in terms of resistance to the dry-wood termite. 

These fadors have to be taken into account in practical 

situations to assess the resistance of different woods to 

termite attack. 

In order to evaluate Wolcott's experiments and others 

dealing with wood preferences of termites, it is important to recognize other factors that could affiect their 

food selection. Fungi may attract termites and stimulate wood consumption. Decay caused by some fungi 

can improve the nutritional conditions of wood for the 

termites. Also, some fungi can break down inhibitory 

compounds in the wood, rendering it suitable as food 

for termites (Becker 1975, Carter 1975, Wood 1978). 

For example, the termite Coptoterms niger lives in the 

heartwood of standing pino hondurefio, Pinus caribaea, only if the heartwood has been attacked by the 

fungus Lentinus pallidus. The resins and turpentine, 

repellent and toxjc to C. niger, are broken down by L. 

pallidus (Williams 1965). The improvement in nutritional value of the wood after fungal attack is probably 

based on the increase in protein, vitamins, and other 

substances. Some termite species survive better in 

wood decayed by fungi than in undecayed wood (Ekker 

1975, Breznak 1982, Carter 1975). Nonetheless, some 

fungi have harmful effects (repellent or toxic) on termites (Becker 1975). Sands (1969) found that adaptation of the termites to special fungi in their diet can 

influence their response in comparative nutrition tests. 

The importance of choice tests in termite wood preference studies is demonstrated by Waller (1988). Waller 

found that less-preferred woods were eaten in large 

amounts in nonchoice tests, and the woods appeared 

favorable for the termites and their protozoans symbiont~. Food selection is affected by the availability of 

the various food sources, so that in the absence of the 

preferred food alternatives, other sources may be 

exploited (Lepage 1974, Wood 1978). Lastly, it is impo1c 

tant to have appropriate controls in feeding preference 

studies. Usually, samples from the colonized wood are 

used as a control (Carter 1975). 

Brooks and others (1941) studied the resistance of 

untreated wood to termites in Trinidad. Heteroternes 

tenuis and N. costalis, two of the most damaging te~ 

mites according to their experiments, are present in 

Puerto Rico. Ten-year old teak, Tkctona grandis, which 

is supposedly very resistant to termite attack, was 

easily attacked. Under their experimental conditions 

(favorable to fungi that attack wet wood), fungi attack 

was responsible for more damage than termite attack, 

although very large populations of Hetemtermes spp. 

developed during the fxst 6 months of the experiments. 

They did not find that termite attack was favored by 

fungal attack or that penetration of fungi was greatly 

accelerated by the inroads of termites. 

Marchan (1946) gave information on the concentration of lignin in wood from different species and the 

degree of susceptibility to the attack by the dry-wood 

termite, C. breuis. Although Marchan did not conduct 

any statistical analyses on the data, it appears that the 

species most resistant to the dry-wood termite tend to 

exhibit high lignin content. However, Wolcott (1940b) 

did not think that lignin is a predictor of wood 

resistance to C. braris attack 

The preservative treatments and service life offenceposts of several species are discussed in the publications of Angler15 (1963), Chudnoff and Goytia (1971, 

1972), Englerth (19601, Maldonado (1962), Ortiz (1963), 

and Tropical Forest Experiment Station (1952, 1953, 

1958). Density was not a predictor of fencepost service 

life (Englerth 1960). Average service life of untreated 

fenceposts is about 1.5 years. The longer life for some 

species can be attributed to the presence of a large 

volume of heartwood (Chudnoff and Goytia 1972). 

Additional information on timber durabilities can be 

found in Longwood (1971). 
Fungi and termites are the principal damaging 

agents of fenceposts. Decay fungi were responsible for 

most of the deterioration of posts (Englerth 1960). 

There is a tendency for termites to be more important 

as fencepost-damaging agents in the dry forests and for 

fungi to be more important in highland wet forests 

(Chudnoff and Goytia 1972). Bultman and Southwell 

(1976) also found that in Panamanian humid forests, 

fungi were more important agents of decay than termites. Nonetheless, Bultman and Southwell found that 

wood attacked by termites decomposed faster than 

wood attacked by fungi. 

Termites of the genus Nasutitermes, such as N. corniger, fur nitrogen at a rate that, in theory, could provide 

all the nitrogen for one to two doublings of the colony 

per year. This could explain the ability of some termite 

species to live in wood with low amounts of nitrogen. 

Termites are also very efficient at recycling nutrients, 

which can also explain these responses. They eat dead 

and unhealthy nestmates and, with the help of symbiotic bacteria, can recycle uric acid to obtain nitrogen 

(Breznak 1982). Termites are so efficient at recycling 

that some species do not even discard their feces; they 

use them for the construction of nests and tunnels 

(Wood 1976). The ecological implications of these findings are yet to be explored in the Luquillo Mountains. 

LITERATURE CITED 

Alexander, C.P. 1932. The crane-flies of Puerto Rico 

(Diptera). Journal of the Department of Agriculture 

of Puerto Rico. 16(4): 347-387. 

Allan, J.D.; Barnthouse, L.W.; Prestbye, R.A.; Strong, 

D.R. 1973. On foliage arthropod communities of 

Puerto Rican second growth vegetation. Ecology. 

54(3): 628-4532. 

Anderson, J.M. 1988. Invertebrate-mediated transport 

processes in soils. Agriculture, Ecosystems, and 

Environment. 24: 5-19. 

Angld, J.I. 1963. Los espeques duran m&...mucho m& 

con pentaclorofenol: Fence posts last longer with 

pentachlorophenol. Revista del Caf6. 18(4): 37-39. 

Arendt, W.J. 1985a. Philornis ectoparasitism of pearlyeyed thrashers. 1: Impacts on growth and development of nestIings. Auk. 102: 270-280. 

Arendt, W.J. 1985b. Philornis ectoparasitism of pearlyeyed thrasher. 2: Effects on adults and reproduction. 

A&. 102: 281-292. 

Bakshi, B.K. 1976. Dealing with pests and diseases in 

tropical forests. Unasylva. 1: 21-24. 

Bannister, B.A. 1970. Ecological life cycle of Euterpe 

globosa Gaertn. In: Odum, H.T.; Pigeon, R.F., eds. A 

tropical rain forest. Washington, DC: U.S. Atomic 

Energy Commission: B-299-314. 

Bartlett, K.A. 1938. A search in the Guianas and Rinidad for predatory beetles of the bamboo scales. 

Journal of Agriculture of the University of Puerto 

Rico. 22(4): 493495. 

Bates, C.Z. 1929. Efectos del huradn del 13 de septiembre de 1928 en distintos iirboles: Effects of the 

September 13, 1928 hurricane on different trees. 

Revista de Agricultura de Puerto Rim. 23(3): 113-228. 

Beattie, A.J. 1985. The evolutionary ecology of antplant mutualjsm. Cambridge, UK: Cambridge University Press, 182 p. 

Becker, G. 1975. Termites and fungi. In: Becker, G.; 

Liese, W., eds. Organismen und holz. Berlin: Duncker and Humbolt: 465-478. 

Benedjct, F.F. 19 76. Herbivory rates and leaf properties 

in four forests in Puerto Rico and Florida. Gainesville, FL: University of Florida. 78 p. M.S. thesis. 

Bohart, R.M.; Stange, LA. 1965. A revision of the 

genus Zethus Fabricius in the Western Hemisphere. 

University of California Publications in Entomology. 

Berkeley, CA: University of California Press: 12.147. 

Boone, R.S.; Chudnoff, M.; Goytia, E. 1969. Chemical 

controI of Bostrichidae during air drying of fence 

posts. Res. Pap. ITF-8. Rio Piedras, PR: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station, Institute of Tropical 

Forestry. 8 p. 

Borror, D.J.; De long, D.M.; 'IYiplehorn, C.A. 1981. An 

introduction to the study of insects. Philadelphia: 

Saunders College Publishing. 827 p. 

Breznak, J.A. 1982. Intestinal microbiota of termites 

and other xylophagous insects. Annual Review of 

Microbiology. 36: 323-343. 

Brian, M.V. 1983. Social insects: ecology and 

behavioural biology. London: Chapman and Hall. 

377 p. 

Bright, D.E. 1985. Studies on West Indian Scolytidae 

(Coleoptera). Entomologische Arboretum Museum 

George Frey. 34: 169-187. 

Brooks, R.L. ; Adamson, A.M.; Baker, R.E.D.; Crowdy, 

S.H. 1941. Durability tests on untreated timbers in 

lkinidad. Caribbean Forester. 2(3): 101-ll9. 

Brown, S.; Lugo, A.E.; Silander, S.; Liegel, L. 1983. Research history and opportunities in the Luquillo 

Experimental Forest. Gen. lkh. Rep. SO-44. New 

Orleans, LA: U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station. 128 p. 

Bultman, J.D.; Southwell, C.R. 1976. Natural 

resistance of tropical American woods to termstrial 

wood-destroying organisms. Biotrcipica. 8(2): 71-95. 

Carter, F.L. 1975. Responses of subterranean termites 

to wood extractives. In: Becker, G.; Liese, W., eds. 

Organismen und holz. Berlin: Duncker and Humbolt: 357-364. 

Chudnoff, M.; Goytia, E. 1971. Treatability of Puerto 

Rican woods. Res. Pap. ITF-11. Rio Piedras, PR: U.S. 

Department of Agriculture, Forest Service, Southern Forest Experiment Station, Institute of hpical 

Forestry. 8 p. 
Chudnoff, M.; Goytia, E. 1972. Preservative treatments 

and service life of fence posts in Puerto Rico. Res. 

Pap. ITF-12. Rio Piedras, PR: U.S. Department of 

Agriculture, Forest Service, Southern Forest Experiment Station, Institute of %pica1 Forestry. 28 p. 

Cintrbn, G. 1970. Niche separation of tree frogs in the 

Luquillo Forest. In: Odum, H.T.; Pigeon, R.F., eds. A 

tropical rain forest. Washington, DC: U.S. Atomic 

Energy Commission: E-51-53. 

Darlington, P.J. 1970. Carabidae on tropical islands, 

especially the West Indies. Biotrcipica. 2(1): 7-15. 

de Andrade, J.C. 1984. Desempenho e distribuiciio de 

Azteca alfari (Formicidae) em Cecropia lyratiloba 

var. nana (Cecropiaceae) na restinga do recreio dos 

bandeirantes, Rio de Janeiro-RJ: Performance and 

distribution of Azteca alfari (Formicidae) in 

Cecropia lyratiloba var. nana (Cecropiaceae). Rio de 

Janeiro-RJ, Atas Society Biologia. 24: 11-15. 

de Andrade, J.C.; Carauta, J.P. 1982. The CecropiaAzteca association: a case of mutualism? Biotr6pica. 

14(1): 15. 

Debon, D. 1941. Some observations on forest entomology in Puerto Rico. Caribbean Forester. 2(4): 160- 

163. 

Dillon, E.S.; Dillon, L.S. 1972. A manual of common 

beetles of eastern North America. New York: Dover 

Publications. 894 p. 

Drewry, G.E. 1969. Animal diversity. In: Jordan, C.F.; 

Drewry, G.E., eds. The rain forest project. Annual 

Rep. 129. Rio Piedras, PR: Puerto Rico Nuclear Center: 107-121. 

Drewry, G.E. 1970. A list of insects from El Verde, 

Puerto Rico. In: Odum, H.T.; Pigeon, R.F., eds. A 

tropical rain forest. Washington, DC: U.S. Atomic 

Energy Commission: E-129-150. 

El Servicio de Extensibn Agricola. 1948. Silvicultor 

hace observaciones sobre las plagas que atacan 10s 

bosques: Forester makes observations about pests 

that attack forests. Revista de Agricultura de Puerto 

Rico. 39(2): 185-186. 

Englerth, G.H. 1960. The service life of untreated 

posts in Puerto Rico after one year in test. l'kop. For. 

Notes 5. Rio Piedras, PR: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Stetion, Institute of Tropical Forestry. 2 p. 

Estrada, A.; Garrison, R.W.; Reagan, D.P. [and others]. 

1982. Flora and fauna of the El Verde Field Station. 

CEER-T-159. Rio Piedras, PR: Center for Energy 

and Environment Research. 18 p. 

Federal Experiment Station. 1950. Report of Federal 

Experiment Station in Puerto Rico. Mayagiiez, PR: 

University of Puerto Rico, Agricultural Experiment 

Station. 30 p. 

Flint, O.S. 1964. The caddisflies (Trichoptera) of 

Puerto Rico. %h. Pap. 40. Rio Piedras, PR: University of Puerto Rico, Agricultural Experiment Station: 5-80. 

Forbes, W.T.M. 1930. Insects of Puerto Rico and the 

Virgin Islands: Heterocera or moths (excepting the 

Noctuidae, Geometridae and Pyralidae). New York 

Academy of Sciences Scientific Survey of Puerto 

Rico and the Virgin Islands. 12(1): 4-171. 

Francis, J.K. 1988a. Araucariaceae in Puerto Rico. %- 

rialba. 38(3): 202-207. 

Francis, J.K. 1988b. Entemlobium qclocarpum (Jacq.) 

Griseb. SO-ITF-SM-15. New Orleans, LA: U.S. 

Department of Agriculture, Forest Service, Southern Forest Experiment Station. 4 p. 

Gallardo, F. 1987. Plaiypus ratzeburgi Chapuis (Coleoptera: Platypodidae): A new pest attacking coffee. 

Journal of Agriculture of the University of Puerto 

Rico. 71(3): 335-336. 

Gara, R.I.; Man, G.G.; Wiikins, R.M.; Whitmore, J.L. 

1973. Flight and host selection behaviour of the 

mahogany shoot borer, HypsiA grandella Zeller 

(Lepid., Phycitidae). Zeitschrift fiir Angewandte 

Entomologie. 72(3): 259-266. 

Garcia, J. 1938. An ecological survey of the fresh water 

insects of Puerto Rico. Journal of Agriculture of the 

University of Puerto Rim. 22(1): 43-97. 

Haber, W.A.; Moore, C.G. 1973. Ads aegypti in the 

Puerto Rican rain forest: results of a one-year survey. Mosquito News. 33(4): 576378. 

Hernhdez, E. 1986. Pollination and nectar robbing of 

llzbebuia rig& (Bignoniaceae) at an elfm forest in 

the LuquiIlo Mountains of Puerto Rim. Rfo Piedraa, 

PR: University of Puerto Rico. 77 p. M.S. thesis. 

Hlavac, T.F. 1969. A review of the species of Scarites 

(Antilliscaris), (Coleoptera: Carabidae) with notes 

on their morphology and evolution. Psyche. 76(1): 1- 

17. 

Hodges, C.S.; McFadden, M.W. 1987. Insects and diseases affecting forest plantations in tropical America. In: Figueroa, J.C.; Wadsworth, F.W.; Branham, 

S., eds. Management of the forests of tropical Am* 

ica. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment 

Station: 365-4376. 

Howard, R.A. 1969. The ecology of an elfin forest in 

Puerto Rim. 8: Studies of stem growth and form and 

of leaf structure. Journal of the Arnold Arboretum. 

50: 225-262. 

Jackson, G.C.; Woodbury, R.O. 1976. Host plants of the 

carpenter bee, Xybcopa brasilianorum L. (Hymenoptera: Apoidea), in Puerto Rico. Journal of Agriculture of the University of Puerto Ria. 60(4): 639460. 

Janzen, D.H. 1972. Association of a rainforest palm 

and seed-eating beetles in Puerto Rim. Ecology. 53 

(2): 258-261. 

Janzen, D.H. 1973a. Dissolution of mutualism between 

Cecropia and Azteca ants. Biotr6pica. 5(1): 15-28. 

Janzen, D.H. 1973b. Sweep samples of tropical foliage 

insects: effects of seasons, vegetation types, elevation, 

time of day, and insularity. Ecology. !54(3): 687-701. 
La Fage, J.P.; Nutting, W.L. 1978. Nutrient dynamics 

of termites. In: Brian, M.V., ed. Production ecology 

of ants and termites. Cambridge, UK: Cambridge 

University Press: 165-232. 

Lamb, A.F. 1968. Fast growing timber trees of the 

lowland tropics. 2: Cedrela odorata. Oxford, UK: 

Commonwealth Forest Institute. 46 p. 

Landolt, P.J.; Reed, H.C. 1990. Looking for love in all 

the wrong places. Agricultural Research. 38: 14-16. 

Lavigne, R.J. 1970a. The ecology of ants of Luquillo 

Forest in the vicinity of El Verde Field Station. In: 

Clements, R.G.; Drewry, G.E.; Lavigne, R.J., eds. 

The rain forest project. Annual Rep. 147. Rio 

Piedras, PR: Puerto Rico Nuclear Center: 74-32. 

Lavigne, R.J. 1970b. The role of insects in the food web 

of the tropical wet forest. In: Clements, R.G.; 

Drewry, G.E.; Lavigne, R.J., eds. The rain forest 

project. Annual Rep. 147. Rio Piedras, PR: Puerto 

Rico Nuclear Center: 83-85. 

Lavigne, R.J. 1977. Notes on the ants of Luquillo Forest, Puerto Rico (Hymenoptera: Formicidae). Proceedings of the Entomological Society of 

Washington. 79(2): 216-237. 

Lavigne, R.J.; Drewry, G. 1970. Feeding behavior of the 

frogs and lizards in the tropical wet forest. In: Clements, R.G.; Drewry, G.E.; Lavigne, R.J., eds. The rain 

forest project. Annual Rep. 147. Rio Piedras, PR: 

Puerto Rico Nuclear Center: 64-72. 

Lepage, M. 1974. Recherches 6cologiques sur une savane sahelienne du Ferlo septentrional, SCnCgal: donnCes prhliminaires sur l'hcologie des termites: 

Eco1ogic.l research south of a sahel savanna, Senegal: preliminary ecological studies of the termites. 

Terre et la vie. 26: 388-409. 

Liebherr, J.K. 1988. Biogeographic patterns of West 

Indian Platynus carabid beetles (Coleoptera). In: 

Liebherr, J.K., ed. Zoogeography of Caribbean insects. 

Ithaca, NY: Corn& University Press: 121-152. 

Lister, B.C. 1981. Seasonal niche relationships of rain 

forest moles. Ecology. 62(6): 1,548-1,560. 

Little, E.L., Jr.; Wadsworth, F.H. 1964. Common trees 

of Puerto Rico and the Virgin Islands. Agric. Handb. 

249. Washington, DC: U.S. Department of Agriculture. 548 p. 

Little, E.L., Jr.; Woodbury, R.O. 1976. ks ofthe 

Caribbean National Forest, Puerto Rico. Res. Pap. 

ITF-20. Rio Piedras, PR: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station, Institute of 'lkopical Forestry. 27 p. 

Longino, J.T. 1991. Azteca ants in Cecropia trees: taxonomy, colony structure, and behaviour. In: Huxley, 

C.R.; Cutler, D.F., eds. Ant-plant interactions. 

Oxford, UK: Oxford University Press: 271-288. 

Longwood, F.R. 1971. Present and potential commercial timbers of the Caribbean. Agric. Handb. 207. 

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

168 p. 

Lugo, A.E.; Figueroa, J. 1984. Anthocephalus chinensis 

(Lam.) A. Rich. ex Walp. SO-ITF-SM-1. New 

Orleans, LA: U.S. Department of Agriculture, Forest 

Service, Southern Forest Experiment Station. 6 p. 

Maguire, B. 1970. Aquatic communities in bromeliad 

leaf axils and the influence of radiation. In: Odum, 

H.T.; Pigeon, R.F., eds. A tropical rain forest. Washington, DC: U.S. Atomic Energy Commission: 

E-95-102. 

Maldonado, E.D. 1962. Durabilidad de postes tratados 

para su finca: Durability of farm treated posts. 

Revista de Agricultura de Puerto Rico. 49(1): 165- 

168. 

Maldonado, J.; Navarro, CA. 1967. Additions and corrections to Wolcott's "Insects of Puerto Rico." 

Caribbean Journal of Science. 7: 45-64. 

Marchan, F.J. 1946. The lignin, ash, and protein content of some neotropical woods. Caribbean Forester. 

7(2): 135-138. 

Martorell, L.F. 1939. Some notes on forest 

entomology - part 1. Caribbean Forester. l(1): 25- 

26. 

Martorell, L.F. 1940a. Some notes on forest 

entomology-part 2. Caribbean Forester. l(2): 31- 

32. 

Martorell, L.F. 1940b. Some notes on forest 

entomology -part 3. Caribbean Forester. l(3): 23- 

24. 

Martorell, L.F. 1940c. Notes on the biology of Mesocondyla concordalis Hiibner and its parasites. Caribbean Forester. 20): 18-19. 

Martorell, L.F. 1941. Some notes on forest 

entomology-part 4. Caribbean Forester. 2(2): 80- 

82. 

Martorell, L.F. 1943. Forests and forest entomology. 

Caribbean Forester. 4(3): 132-134. 

Martorell, L.F. 1945a. A survey of the forest insects of 

Puerto Rim -part 1. Journal of Agriculture of the 

University of Puerto Rim. 29(3): 69-354. 

Martorell, L.F. 1945b. A survey of the forest insects of 

Puerto Rico -part 2. Journal of Agriculture of the 

University of Puerto Rico. 29(4): 355-608. 

Martorell, L.F. 1973. Glyptoterrnes liberatus (Snyder) 

(Isoptera: Kalotermitidae): a new termite record for 

Puerto Rim. Journal of Agriculture of the University of Puerto Rico. 57(4): 355-356. 

Martorell, L.F. 1976. Annotated food catalog of the 

insects of Puerto Rico. Rio Piedras, PR: University 

of Puerto Rico, Agricultural Experiment Station. 

303 p. 

Martorell, L.F.; Garcia, J.C. 1971. New locality and host 

plant record for two species of Nasutitermes (Isoptera: Termitidae). Journal of Agriculture of the University of Puerto Rico. 40(1): 135-137. 

Martorell, L.F.; Medina, S. 1974. Orchid pests in 

Puerto Rico. Journal of Agriculture of the University of Puerto Rico. 58(1): 105-124. 
PR: U.S. Department of Agriculture, Federal 

Experiment Station. 17 p. 

Plank, H.K. 1950b. Studies of factors influencing 

attack and control of the bamboo powder-post beetle. 

Bull. 48. Mayagiiez, PR: U.S. Department of Agriculture, Federal Experiment Station. 39 p. 

Plank, H.K.; Winters, H.F. 1949. Insect and other animal pests of Cinchona and their control in Puerto 

Rico. Bull. 46. Mayagiiez, PR: U.S. Department of 

Agriculture, Federal Experiment Station. 16 p. 

Porto Rico Agricultural Experiment Station. 1915. 

Report of the Porto Rico Agricultural Experiment 

Station, 1914. Mayagiiez, PR: U.S. Department of 

Agriculture, Federal Experiment Station. 35 p. 

Putz, F.E.; Holbrook, N.M. 1988. Further observations 

on the dissolution of mutualism between Cecropia 

and its ants: the Malaysian case. Oikos. 53: 121-125. 

Rivero, J.A.; Maldonado, J.; Mayorga, H. 1963. On the 

habits and food of Eleuthemdactylus karlschmidti 

Grant. Caribbean Journal of Science. 30): 25-27. 

Roberts, H. 1968. An outline of the biology of l-lypsipyh 

robusta Moore, the shoot borer of the Meliaceae 

(mahoganies) of Nigeria, together with brief comments on two stern borers and one other lepidopteran 

fruit borer also found in Nigerian Meliceae. The 

Commonwealth Forestry Review. 47(3): 225-232. 

Ross, H.H.; Ross, CA.; Ross, J.R.P. 1982. Entomology. 

New York: John Wiley and Sons. 666 p. 

Sands, W.A. 1969. The association of termites and 

fungi. In: Krishna, K.; Weesner, F.M., eds. Biology 

of termites. New York: Academic Press: 495-524. 

Vol. 1. 

Sanford, M.T. 1987. Florida bee botany [Circular 6861. 

Gainesville, FL: University of Florida, Florida 

Cooperative Extension Service. 15 p. 

Sastre, 1.1979. Ecological life cycle of Buchenauia capit& CVahl.1 Eichl., a late secondary succesional species in the rain forest of Puerto Rico. Knoxville, TN: 

University of Tennessee. 45 p. M.S. thesis. 

Schedl, K.E. 1961. Scolytidae und Platypodidae 

Mrikas. Revista de Entomologia de Mo~bique. 

4(2): 235-742. 

Schupp, E.W. 1986. Azteca protection on Cecropia: ant 

occupation benefits juvenile trees. Oecologia. 70: 

379-385. 

Sein, F. 1931. Informe sobre el brote del Apate francisca 

en Lares: Report of an outbreak of hate francisca 

in Lares. El Agricultor Puertorriqueiio. ll(7): 24. 

Silander, S.R. 1979. A study of the ecological life history of Cecropia peltata L., an early secondary succesional species in the rain forest of Puerto Rico. 

Knoxville, TN: University of 'knnessee. 94 p. M.S. 

thesis. 

Smith, C.F.; Martorell, L.F.; PBrez, M.E. 1963. 

Aphididae of Puerto Rico. Tech. Pap. 37. Rio Piedras, 

PR: University of Puerto Rico, Agricultural Experiment Station. 121 p. 

Smith, C.F.; Martorell, L.F.; Pkrez, M.E.; Medina, S. 

1971. Additions and corrections to the Aphididae of 

Puerto Rico. Journal of Agriculture of the University of Puerto Rico. 55(2): 192-258. 

Smith, R.F. 1966. Vegetation structure of a Puerto 

Rican rain forest. Athens, GA: University of Georgia. 173 p. Ph.D. thesis. 

Snelling, R.R. 1981. Systematics of social Hymenoptera. In: Hermann, H., ed. Social insects, part 2. 

New York: Academic Press: 369-458. 

Snyder, N.F.; Wdey, J.W.; Kepler, C.B. 1987. The parrots of Luquillo: natural history and conservation of 

the Puerto Rican parrot. Los Angeles: Western 

Foundation of Vertebrate Zoology. 384 p. 

Snyder, T.E. 1956. Termites of the West Indies, 

the Bahamas, and Bermuda. Journal of Agriculture 

of the University of Puerto Rico. 40(3): 

189-202. 

Southwood, T.R.E. 1971. Ecological methods. London: 

Chapman and Hall. 391 p. 

Strong, D.R. 1979. Biogeographic dynamics of insecthost plant communities. Annual Review of Entomology. 24: 89-119. 

Telford, H.S. 1973. The Syrphidae of Puerto Rico. Journal of Agriculture of the University of Puerto Rico. 

57(3): 217-246. 

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. 

Torres, J.A.; Canals, M. 1983. Components of ant diversity and other miscellaneous notes on ants. Boletin 

Cientifico del Sur. lO(1) : 38-43. 

Tropical Forest Experiment Station. 1952. Twelveth 

annual report. New Orleans, LA: U.S. Department 

of Agriculture, Forest Service, Southern Forest 

Experiment Station: 19-21. 

'Ik.opical Forest Experiment Station. 1953. Thirteenth 

annual report. New Orleans, LA: U.S. Department 

of Agriculture, Forest Service, Southern Forest 

Experiment Station: 30-33. 

Tropical Forest Experiment Station. 1954. Fourteenth 

annual report. Caribbean Forester. 15: 1-13. 

Tropical Forest Experiment Station. 1958. Eighteenth 

annual report. New Orleans, LA: U.S. Department 

of Agriculture, Forest Service, Southern Forest 

Experiment Station: 19-20. 

Tulloch, G.S. 1938. Ecological notes on mosquitoes 

associated with bromeliads. Journal of Agriculture 

of the University of Puerto Rico. 22(4): 499-501. 

Uhazy, L.S.; Arendt, W.J. 1986. Pathogenesis associated with philornid myiasis (Diptera: Muscidae) on 

nestling pearly-eyed thrashers (Aves: Mimidae) in 

the Luquillo Rain Forest, Puerto Rico. Journal Wildlife Diseases. 22(2): 224-237. 

U.S. Atomic Energy Commission. 1970. A tropical rain 

forest: a study of irradiation and ecology at El Verde, 
Matthews, E.G. 1965. The taxonomy, geographical distribution, and feeding habits of the canthonines of 

Puerto Rico (Coleoptera: Scarabaeidae). Transactions of the American Entomological Society. 91: 

431-465. 

Matthews, E.G. 1966. A taxonomic and zoogeographic 

survey of the Scarabaeinae of the Antilles (Coleoptera: Scarabaeidae). Memoirs of the American 

Entomological Society. 21: 1-134. 

McMahan, E.A. 1966. Studies of termite wood-feeding 

preferences. In: LaPlante, A.A., ed. Proceedings of 

the Hawaiian Entomological Society; 1966 November 30; Honolulu, HI. I-lonolulu, HI: Hawaiian 

Entomological Society; 19(2): 239-250. 

McMahan, E.A. 1968a. Effects of ionizing radjation on 

three neotropical termites species (Isoptera, Termitidae). In: Kline, J.R.; Jordan, C.J.; Drewry, G.E., 

eds. The rain forest project. Annual Rep. 119. Rio 

Piedras, PR: Puerto Rico Nuclear Center: 124-140. 

McMahan, EA. 1968b. Termite nests and tunnels in 

the radiation center at El Verde. In: Kline, J.R.; 

Jordan, C.F.; Drewry, G.E., eds. The rain forest project. Annual Rep. 119. Rio Piedras, PR: Puerto Rico 

Nuclear Center: 79-81. 

McMahan, E.A. 1969. Termites at El Verde. In: Jordan, 

C.F.; Drewry, G.E., eds. The rain forest project. 

Annual Rep. 129. Rio Piedras, PR: Puerto Rico 

Nuclear Center: 142-144. 

McMahan, E.A. 1970a. Polyethism in workers of 

Nasutitermes costalis (Holmgren). In: Clements, 

R.G.; Drewqy, G.E.; Lavigne, R.J., eds. The rain forest project. Annual Rep. 147. Rio Piedras, PR: 

Puerto Rico Nuclear Center: 118-126. 

McMahan, EA. 1970b. Radiation and the termites at 

El Verde. In: Odum, H.T.; Pigeon, R.F., eds. A tropical rain forest. Washington, DC: U.S. Atomic Energy 

Commission: E-105-122. 

McMahan, EA. 1970c. The termites at El Verde: 1969 

survey. In: Clements, R.G.; Drewry, G.E.; Lavigne, 

R.J, eds. The rain forest project. Annual Rep. 147. Rio 

Piedras, PR: Puerto Rico Nuclear Center: 111-ll7. 

McMahan, E.A.; Sollins, N.F. 1970. Diversity of microarthropods after irradiation. In: Odum, H.T.; 

Pigeon, R.F., eds. A tropical rain forest. Washington, 

DC: U.S. Atomic Energy Commission: E-151-158. 

Nwling, L.I. 1971. The ecology of an elfin forest in 

Puerto Rico. 16: The flowering cycle and an interpretation of its seasonality. Journal of the Arnold 

Arboretum. 52: 586-613. 

Ngatiman. 1989. Serangan hama penggerek pucuk 

Hypsipyla robusta Moore pada tanaman muda 

mahoni (Swietenia macmphylla King) di Pt Lestari, 

Batu Ampar kdimantan Timur: Attack of Hypsimla robusta Moore bud bore pest on mahoni plantation (Swietenia macrophylla King) at the Pt Kana 

Lestari, Batu Arnpar Kalimatan Timur. Wanatrop, 

4: 18-30. 

Nichols, S.W. 1986. Descriptions of larvae of Puerto 

Rican species of Antilliscaris Blinninger and notes 

about relationships and classification ofAntilliscaris 

(Coleoptera: Carabidae: Scaratini: Scaritina). Coleopterist Bulletin. 40: 301-311. 

Nichols, S.W. 1988. Kaleidoscopic biogeography of West 

Indian Scaritinae (Coleoptera: Carabidae). In: 

Liebherr, J.K., ed. Zoogeography of Caribbean 

insects. Ithaca, NY: Cornell University Press: 71- 

120. 

Nieves, L.O. 1979. Ecological life history study of 

Didymopanax momtotoni. Rio Piedras, PR: University of Puerto Rico. 85 p. M.S. thesis. 

Odum, H.T.; Abbott, W.; Selander, R.K. [and others]. 

1970. Estimates of chlorophyll and biomass of the 

tabonuco forest of Puerto Rim. In: Odum, H.T.; 

Pigeon, R.F., eds. A tropical rain forest. Washington, 

DC: U.S. Atomic Energy Commission: 1-1-19. 

Odum, H.T.; Ruiz, J. 1970. Holes in leave8 and the 

grazing control mechanism. In: Odum, H.T.; Pigeon, 

R.F., eds. A tropical rain forest. Washington, DC: 

U.S. Atomic Energy Commission: 1-69-80. 

Ortiz, V. R. 1963. Preservation of Puerto Rican fence 

posts treated by pressure methods. Caribbean FOP 

ester. 24(2): 91-93. 

Paz, N. 1974. Estudio bio-ecol6gico de la fauna de las 

bromelias Guzmania berteronia~uz y Vriesia sientenisii en el Bosque Pluvial de Luquillo (El Yunque), 

Puerto Rico. Rio Piedras, PR University of Puerto 

Rim. 99 p. M.S. thesis. 

Peck, S.B. 1970. The Catopinae (Coleoptera: Leiodidae) 

of Puerto Rico. Psyche. 77: 237-242. 

Peck, S.B. 1972. Leiodinae and Catopinae (Coleoptera: 

Leiodidae) from Jamaica and Puerto Rim. Psyche. 

79: 4957. 

Pkrez, R.A.; Collazo, J. 1976. Aves en Puerto Rico parasitadas por la larva de la mosca del g6nero Neomusca: Pueto Rican bird species parasitized by fly 

larvae of the genus Neomusca. Boletin Cientifico del 

Sur. 4(2): 50-51. 

Peterson, A. 1982. Larvae of insects, part 2. Columbus, 

OH: Ohio State University. 416 p. 

Peterson, A. 1984. Larvae of insects, part 1. Columbus, 

OH: Ohio State University. 315 p. 

Plank, H .K. 1939. Peregrinator bianulipes Montr., a 

predator of the bamboo powderpost beetle in Puerto 

Rico. Journal of Economic Entomology. 32(1): 151. 

Plank, H.K. 1948a. Biology of the bamboo powder-post 

beetle in Puerto Rim. Bull. 44. Mayagiiez, PR: U.S. 

Department of Agriculture, Federal Experiment 

Station. 29 p. 

Plank, H.K. 1948b. Life history, habits, and control of 

the coconut rhinoceros beetle in Puerto Rim. Bull. 

45. Mayagiiez, PR: U.S. Department of Agriculture, 

Federal Experiment Station. 35 p. 

Plank, H.K. 1950a. Insecticidal properties of some 

plants growing in Puerto Rico. Bull. 49. Maymez, 
Wilson, E.O. 1971. The insect societies. Cambridge, 

MA: Harvard University Press. 548 p. 

Wolcott, G.N. 1921. Los comejenes de Puerto Rico: The 

termites of Puerto Rico [Circular 441. Rio Piedras, 

PR: Agricultural Experiment Station. 14 p. 

Wolcott, G.N. 1924. The comparative resistance of 

woods to the attack of the termite C~yptotermes 

brevis Walker. Bull. 33. Rio Piedras, PR: Insular 

Experiment Station. 15 p. 

Wolcott, G.N. 1932. The effect of the hurricane of San 

Cyprih on insects in Puerto Rico. The Insect Pest 

Survey Bulletin. 12(1): 409-410. 

Wolcott, G.N. 1933. The changed status of some insect 

pests in Puerto Rico. The Journal of the Department of Agriculture of Puerto Rico. 17: 265-270. 

Wolcott, G.N. 1939. Comejhn y polilla: 'krmites. Boletin 

48. Rio Piedras, PR: Estacicin Experimental Agricola. 26 p. 

Wolcott, G.N. 1940a. The entomologist looks at 

"maga". Caribbean Forester. l(2): 29-30. 

Wolcott, G.N. 1940b. A list of woods arranged according to their resistance to the attack of the "polilla," 

the dry-wood termite of the West Indies, Cryptotermes brevis Walker. Caribbean Forester. l(4): 

1-10. 

Wolcott, G.N. 1940~. An outbreak of the scale insect, 

Asterolecanium pustulans Cockerell on "maga", 

Montezuma speciosissima. Caribbean Forester. 2(1): 

6-7. 

Wolcott, G.N. 1943a. How to make wood unpalatable to 

the West Indian dry wood termite, Cryptotermes 

bmis Walker. 1: With inorganic compounds. Caribbean Forester. 4(4): 145-157. 

Wolcott, G.N. 1943b. Lady-beetles don't behave. Caribbean Forester. 4(2): 81-32. 

Wolcott, G.N. 1945a. How to make wood unpalatable to 

the West Indian dry-wood termite, Cryptotermes 

brevis Walker-part 3: Caribbean Forester. 6(4): 

245-257. 

Wolcott, G.N. 1945b. Trees for roadside planting in 

Puerto Rico. Caribbean Forester. 6(3): 115-121. 

Wolcott, G.N. 1946a. Factors in the natural resistance 

of woods to termite attack. Caribbean Forester. 7(2): 

121-134. 

Wolcott, G.N. 1946b. A Iist of woods arranged according to their resistance to the attack of the West 

Indian dry-wood termite, Cryptotermes brevis 

Walker). Caribbean Forester. 7(4): 329-334. 

Wolcott, G.N. 1946c. What to do about "polilla." Bull. 

68. Rio Piedras, PR: Agricultural Experiment Station. 29 p. 

Wolcott, G.N. 1947. Termite repellents: a summary of 

laboratory tests. Bull. 73. Rio Piedras, PR: Agricultural Experiment Station. 17 p. 

Wolcott, G.N. 1948a. The insects of Puerto Rico-part 

1. Journal of Agriculture of the University of Puerto 

Rico. 32(1): 1-224. 

Wolcott, G.N. 1948b. The insects of Puerto Rico (Coleoptera). Journal of Agriculture of the University of 

Puerto Rico. 32(2): 225-416. 

Wolcott, G.N. 1948~. The insects of Puerto Rico (Diptera, Siphonaptera, and Lepidoptera). Journal of 

Agriculture of the University of Puerto Rico. 32(3): 

417-748. 

Wolcott, G.N. 1948d. The insects of Puerto Rico 

(Hymenoptera). Journal of Agriculture of the University of Puerto Rico. 32(4): 749-975. 

Wolcott, G.N. 1948e. The resistance to dry-wood termite attack of some Central American woods. Caribbean Forester. 9(1): 53-56. 

Wolcott, G.N. 1950. An index to the termite-resistance 

of woods. Bull. 85. Rio Piedras, PR: Agricultural 

Experiment Station. 25 p. 

Wolcott, G.N. 1951. The present status of economic 

entomology in Puerto Rico. Bull. 99. Rio Piedras, 

PR: Agricultural Experiment Station. 21 p. 

Wblcott, G.N. 1953. Biological control of pustule scale 

in Puerto Rico. Journal of Agriculture of the University of Puerto Rico. 37(3): 228-233. 

Wolcott, G.N. 1954. Brmite damage and control as 

factors in the utilization of timber in the Caribbean 

area. Journal of Agriculture of the University of 

Puerto Rico. 38(2): 115-122. 

Wolcott, G.N. 1956. The insects of "almendrbn" 

Prunus occidentalis. Journal of Agriculture of the 

University of Puerto Rico. 40(3): 203-204. 

Wolcott, G.N. 1960. Efficiency of ladybeetles (Coccinellidae: Coleoptera) in insect control. Journal of 

Agriculture of the University of Puerto Rico, 44(4): 

166-172. 

Wolcott, G.N. ; Martorell, L.F. 1942. The accidental 

introduction of a beneficial insect into Puerto Rico. 

Caribbean Forester. 3(2): 58-60. 

Wood, T.G. 1976. The role of termites (Isoptera) in 

decomposition processes. In: Anderson, J.M.; Macfadyen, A., eds. The role of terrestrial and aquatic 

organisms in decomposition processes. Oxford, UK: 

Blackwell Scientific Publications: 145-168. 

Wood, T.G. 1978. Food and feeding habits of termites. 

In: Brian, M.V., ed. Production ecology of termites 

and ants. Cambridge, UK: Cambridge University 

Press: 55-80. 
Puerto Rico. Washington, DC: U.S. Atomic Energy 

Commission. 1644 p. 

U.S. Department of Agriculture, Forest Service. 1972. 

Evaluation of tree insect and disease pests in Puerto 

Rico and St. Croix, V. I. Rep. 73-1-10. Atlanta, GA: 

U.S. Department of Agriculture, Forest Service, Forest Pest Management Group, State and Private Forestry, Southeastern Area. 19 p. 

U.S. Department of Agriculture, Forest Service. 1976. 

Insect and disease conditions - Puerto Rico and the 

United States Virgin Islands, 1976. Atlanta, GA: 

U.S. Department of Agriculture, Forest Service, Forest Pest Management Group, State and Private Forestry, Southeastern Area. 47 p. 

U.S. Department of Agriculture, Forest Service. 1981. 

Evaluation of insect and disease pests of trees in 

Puerto Rim. Rep. 81-1-29. Atlanta, GA: U.S. Department of Agriculture, Forest Service, Forest Pest 

Management Group, State and Private Forestry, 

Southeastern Area. 33 p. 

U.S. Department of Agriculture, Forest Service. 1982. 

Evaluation of tree disease and insect pests in Puertm 

Rico and the United States Virgin Islands. Rep. 

82-1-32. Atlanta, GA: U.S. Department of Agriculture, Forest Service, Forest Pest Management 

Group, State and Private Forestry, Southeastern 

Area. 41 p. 

U.S. Department of Agriculture, Forest Service. 1984. 

l'!ree insect and disease conditions: Puerto Rico and 

St. Croix, United States Virgin Islands. Rep. 84-1-7. 

Atlanta, GA: U.S. Department ofAgriculture, Forest 

Service, Forest Pest Management Group, State and 

Private Forestry, Southeastern Area. 47 p. 

Van Den Bussche, RA.; Willig, M.R.; Chesser, R.K.; 

Waide R.B. 1988. Genetic variation and systematics 

of four taxa of neotropical walking sticks (Phasmatodea: Phasmatidae). Proceedings of the 

Entomological Society of Washington. 90(4): 422- 

42 7. 

Villa, J.; Townsend, D.S. 1983. Viable frog eggs eaten by 

phorid fly larvae. Journal of Herpertology. 17: 27% 

281. 

Virkki, N. 1970. Karyotype rearrangements and malformation of gonads in the walking sticks (Phasmatoptera) of the El Verde radiation center. In: 

Odum, H.T.; Pigeon, R.F., eds. A tropical rain forest. 

Washington, DC: U.S. Atomic Energy Commission: 

G-51-62. 

Waller, D.A. 1988. Host selection in subterranean termites: factors affecting choice (Isoptera: Rhinotermitidae). Sociobiology. 14(1): 5-13. 

Weaver, P.L. 1988. Guarea guidonia (L.) Sleumer. SOITF-SM-17. New Orleans, LA: U.S. Department of 

Agriculture, Forest Service, Southern Forest 

Experiment Station. 7 p. 

Weaver, P.L.; Bauer, G.P. 1986. Growth, survival, and 

shoot borer damage in mahogany plantings in the 

Luquillo Forest in Puerto Rico. Turrialba. 36(4): 

509-522. 

Weigert, R.G. 19 70a. Energetics of the nest-building 

termite, Nasutitermes costalis (Holmgren), in a 

Puerto Rican forest. In: Odum, H.T.; Pigeon, R.F., 

eds. A tropical rain forest. Washington, DC: U.S. 

Atomic Energy Commission: 1-57-64. 

Weigert, R.G. 1970b. Effects of ionizing radiation on 

leaf fall, decomposition, and the litter microarthropods of a montane rain forest. In: Odum, 

H.T.; Pigeon, R.F., eds. A tropical rain forest. Washington, DC: U.S. Atomic Energy Commission: 

H-89-100. 

Weinbren, M.P.; Weinbren, B.M. 1970. Observations on 

the mosquito population in the irradiated forest at 

El Verde. In: Odum, H.T.; Pigeon, R.F., &. A tropical rain forest. Washington, DC: U.S. Atomic Energy 

Commission: E-159-168. 

Wetmore, A. 1916. Birds of Porto Rim. Bull. 326. Washington, DC: U.S. Department of Agriculture. 140 p. 

White, D.G. 1948. Bamboo culture and utilization in 

Puerto Rico [Circular 291. Mayagiiez, PR: U.S. 

Department Agriculture, Federal Experiment Station. 34 p. 

White, D.G.; Cobin, M.; Seguinot, P. 1946. The relation 

between curing and durability of Bambusa 

tuldoides. Caribbean Forester. 7t3): 253-267. 

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

Puerto Rico and St. Croix: establishment phase. 

Res. Note ITF-16. Rio Piedras, PR. U.S. Department 

of Agriculture, Forest Service, Southern Forest 

Experiment Station, Institute of %pica1 Forestry. 

11 p. 

Whitmore, J.L. 1978b. Myths mgarding H'sipyla and 

its host plants. In: Proceedings of the Ist symposium 

on integrated control of Hypsipyh; 1978 March 6- 

12; Turridba, Costa Rica. lnstituto Interamericano 

de Ciencias Agricolas-Centro Tropical de Ensefianza 

e Investigacibn: 1-4. 

Whitmore, J.L.; Hinojoea, G. 1977. Mahogany 

(Swietenia) hybrids. Res. Pap. ITF-23. Rfo Piedras, 

PR: U.S. Department of Agriculture, Forest Service, 

Southern Forest Experiment Station, Institute of 

Tropical Forestry. 8 p. 

Whitmore, J.L.; Medina, S. 1974. White peach scale 

attack on toon in Puerto Rim. Journal of Agriculture of the University of Puerto Rico. 58(2): 276- 

278. 

Williams, R.M. 1965. Infestation of Pinus caribaea by 

the termite Coptotermes niger Snyder. Proceedings 

of the 12th International Congress of Entomology; 

1964 [dates unknown]; London: 675-676. 

Willig, M.R.; Garrison, R.W.; Bauman, A.J. 1986. Population dynamics and natural history of a neotropical 

walking stick, Lamponius portoricerzsis &hn (Phasmatodea: Phasmatidae). The kas Journal of Science. 38(2): 121-137. 
Appendixes 
Appendix A. - Forest and Plantation %es and Their Associated or 

Potential Insect Pestse 

nee species Insect pest 

Andira inermis 

Anthcephalus chinensis 

Araucaria cunninghamii 

Bambusa spp. 

Bixa orellana 

Buchenauia capitata 

Byrsonima coriacea 

Casuarina equisetifolia 

Cecropia peltata 

Cedrela odorata 

Ceiba pentandra 

Cinchona spp. 

Citharexylum fruticosum 

Coccoloba pubescens 

Cordia alliodora 

Cupania americana 

Dacryodes occelsa 

Didymopanax morototoni 

Erythrina fusca 

Erythrina poeppigiana 

Erythrina spp. 

Eucalyptus robusta 

Diaprepes abbreuiatus 

Megalomge krugii 

Phostria originalis 

Platypus rugulosus 

Saissetia oleae 

Xyleborus ferrugineus 

Selenothrips rubrocinctus 

Octaspidiotus araucariae 

Asterolecanium bambusae 

Asterolecanium miliaris 

Dinoderus minutus 

Apate monacha 

Lepidoptera 

Nitidulidae 

Xyleborus volvulus 

Hypothenemus eruditus 

Megalome krugii 

Apate monacha 

Diphthera festiva 

Formicidae 

Icerya purchasii 

Nitidulidae 

Syllepte silicalis 

Ambmsiodmua lecontei 

Apate monacha 

Diaprepes abbreviatus 

Dikrellcl cedrelae 

Pulvinaria psiddii 

Diaprepes abbreviatus 

Qsdercus andreae 

Pseudaulacaspis pentagona 

Epicoris oedipodalis 

Schizocerina krugii 

Dictyla monotmpidia 

Omalegra cordiae 

Asterokcanium pustukzns 

Ambmsiodmus lecontei 

Monarthrum sp. 

Platypus compositus 

Platypus rugulosus 

&ate momha 

Hypothemus obscurus 

Me1 ipotis acantioides 

Megalome kruggii 

Semiothis difisata 

Phyllophaga portoricensis 

Terastia meticulosalis 

Xyleborus ferrugineus 

Agathodes designalis 

Terastia rneticubsalis 

Spodoptera frugiperdu 

Order+ 

Col. 

Lep. 

Lep. 

Col. 

Hom . 

Col. 

Thy. 

Hom. 

Horn. 

Hom. 

Col. 

Col. 

L~P. 

Col. 

Col. 

Col. 

GP. 

Col. 

L~PHym. 

Hom. 

Col. 

L~P. 

Col. 

Col. 

Col. 

Horn. 

Horn. 

Col. 

Hem. 

Horn. 

L~P. 

Hym. 

Hem. 

Horn. 

Horn. 

Col. 

Col. 

Col. 

Col. 

Col. 

Col. 

L~P. 

LeP. 

LeP. 

Col. 

Lep. 

Col. 

LeP. 

L~P. 

L~P. 

Parts attacked 

Leaves 

Leaves 

Leaves 

Limbs (borer) 

Twigs, branches 

Bark 

Leaves 

?'wigs 

Culms 

Leaves 

Culms (borer) 

Wood (borer) 

Mesocarp 

Mesocarp 

Bark 

Dead pole 

Leaves 

Wood (borer) 

Trunk (borer) 

Seeds 

Leaves 

Seeds 

Leaves 

Dead branches 

Wood (borer) 

Leaves 

Leaves 

Branches, trunks 

Leaves 

Seeds 

. . . . . . .* 

Leaves 

Leaves 

Leaves 

Leaves 

Leaves 

Logs 

Logs 

Logs iborer) 

Logs (borer) 

Wood (bares) 

Dead branch 

Leaves 

hves 

Leaves 

Leaves 

Twigs, pods Dead wood 

Leaves, twigs 

mgs, pods 

Leaves 
ippendix A. - Forest and Plantation %es and Their Associated or 

Potential Insect Pests* - Continued 

Yee suecies Insect pest Order+ Parts attacked 

iyzygium jambos 

~raxinus uhdei 

7revillea robusta 

>uarea guidonia 

Yernandia sonora 

Meterotrichurn cymosum 

Yibiscus elatus 

Hymenaea courbaril 

lnga fagifol ia 

lnga quaternata 

lnga Vera 

Chionanthus domingensis 

ilammea americana 

Mangifera indica 

Ochroma lagopus 

Petitia domingensis 

Picramnia pentandra 

Prestoea montana 

Psidium guqjava 

Roystonea borinquena 

Spondias dulcis 

Swietenia macrophylla 

Swietenia mahogani 

Swietenia spp. 

Anastrepha suspensa 

Apate momha 

Myrrnelachista ramulorum 

Pseudaulacaspis pentagona 

Asterolecanium pustulans 

Hypothenemus trinitatis 

Ischnaspis longimstris 

Dictyla monotropidia 

Blepharornastix ebulealis 

Pseudaulacaspis pentagona 

Hypothenemus obscurus 

Megalopyge kruggii 

Myrmelachista ramulorum 

Platypus rdzeburgi 

Xyleborus afinis 

Myrmelachista ramulorum 

Agathodes tiesignalis 

Apate monacha 

Myrmehhista ramulorum 

Platypus excisus 

Platypus ratzeburgi 

'1E.tralopha scabridella 

Xyleborus afinia 

Apate monacha 

Poecilips sp. 

Anastrepha obliquu 

Xyleborus xylographus 

Oiketicus kirbyi 

Hyblaea puera 

Pilocmsis secernalis 

Apate monacha 

Cocotrypes carpophagus 

Hornaledm sabaldla 

Anastmpha sllspensa 

Diaprepes abbra,iatus 

Pseudaulacaspis pentagona 

Anastrepha suspensa 

Hypotknemlls crudiae 

Hypotknemus obscurus 

Hypothenenus seriatzls 

Xyleborus afinis 

Phyllophaga vandinei 

Anastrepha suspensa 

Diaprepres abbreviatus 

Hpsipyla grandella 

Ischnaspis longirostris 

Phyllophaga uandinei 

Xylosandrus compactus 

&ate monacha 

Polycesta porcada 

lhgoxylon aequak 

Dip. 

Col. 

Hym. 

Horn. 

Horn. 

Col. 

Horn. 

Hem. 

L~P. 

Horn. 

Col. 

L~P. 

Hym. 

Col. 

Cd. 

Hym. 

L~P. 

Col. 

Hym. 

Col. 

Cd. 

L~P. 

Col. 

Col. 

Col. 

Dip. 

Col. 

Lg3. 

L~P. 

LeP. 

Col. 

Col. 

L~PDip. 

Col. 

Horn. 

Dip. 

col. 

Col. 

Col. 

Col. 

Col. 

Dip. 

Col. 

LeP. 

Hom. 

Col. 

Col. 

Col. 

Col. 

Col. 

Fruits 

Wood (borer) 

Twigs 

'R-unk 

Leaves 

Dead branches 

Leaves 

Leaves 

Leaves 

. . . . . . . Wpods 

Leaves 

Twigs 

Bark 

Wood (borer) 

w 

'h@s (borer) 

wood (borer) 

Twigs 

Logs 

Logs, trunk 

Leaves 

Wood (borer) 

Wood (borer) 

Seeds 

Fruits 

Wood (borer) 

Leaves 

Leaves 

Leaves 

wood (borer) 

seeds 

Leaves 

F'ruits 

Seedling leaves . . . . . . . Fruits 

Dry fruits 

Dry fruits 

Dry fruits 

Fruits 

Leaves 

Fruits 

Leaves, roots 

Stems (borer) 

Leaves 

Leaves 

Wood (borer) 

wood (borer) 

Cambium, Heartwood 

Sapwood 
Appendix A. - Forest and Plantation &es and Their Associated or 

Potentidl ~nstX?t Pests" - Continued 

Tree species Insect pest Order+ Parts attacked 

lltLbebuia rigida 

%marindus indica 

Terminalia catappa 

Tetragastris balsamifera 

Thespesia grandiflora 

Thespesia populnea 

lbona ciliata 

'12.em.a lamarckianum 

Eulepte concordalis 

Pachymorphus subduciellus 

Rabela tabebuiae 

Eulepte concordalis 

Pachymorphus subductellus 

&ate monacha 

Hypothenemus obscurus 

Hypothenemus obscurus 

Oiketicus kirbyi 

Selenothrips rubrocinctus 

Hypothenemus eruditus 

Asterolecanium pustulans 

Dysdercus andreae 

Dysdercus sanguinarius 

Hypothenemus sp. 

Pectinophora gossypiella 

Hypothenemus eruditus 

Pectinophora gossypiella 

Pseudaulacaspis pentagona 

Pseudaulacaspis pentagona 

L~P. 

kP. 

Horn. 

L~P. 

Lep. 

Col. 

Col. 

Col. 

kP. 

Thy. 

Col. 

Horn. 

Hem. 

Hem. 

Col. 

Lep. 

Col. 

LeP. 

Hom. 

Horn. 

Leaves . 

Twigs (borer) 

Leaves 

Leaves 

Twigs (borer) 

Wood (borer) 

Seed pods 

Fruits 

Leaves 

Leaves 

Dry twigs 

Leaves 

Seeds 

seeds 

Seed pods 

234 pods 

-pods 

-pods 

Trunk 

TNnk, branches 

*See Martorell (1976) for a more complete list. 

tCol. = Coleoptera; Dip. = Diptera; Hem. = Hemiptera; Horn. = Homoptera; Lep. = Lepidoptera; Hym. = 

Hymenoptera; Thy. = Thysanoptera. 

*Unrecorded. 
Appendix B. - List of Important Entomological Studies in the 

Luquillo Mountains, Puerto Rico (complete 

citations are given in the literature cited 

section) 

Author 

Year of 

publication Topic 

Arendt, W. J. 

Benedict, F. F. 

Chudnoff, M; Goytia, E. 

Drewry, G. E. 

Garcia, J. 

Hlavac, T. F. 

Janzen,D. H. 

Lavigne, R. J. 

Lavigne, R. J; Drewry, G. 

Lister, B. C. 

Martod, L. F. 

Matthews, E. G. 

Nevling, L. I. 

Nichols, S. W. 

Plank, H. K. 

Plank, H. K. 

Weaver, P. L; Bauer, G. P. 

Wetmore, A. 

Wolcott, G. N. 

Wolcott, G. N. 

Bird parasitism 

Insect herbivory 

Service life of fencepoets 

List of insects from El Vde 

Fresh water ins& 

Carabidae morphoIogy and evolution 

General information on tropical insects 

Ants of Luquillo Forest 

Feeding habits of frogs and lizards 

Lizard insect food 

Food catalog of the ins& of Puerto Rico 

Scarab beetles of the Luquillo Mountains 

Insect pollination 

Carabidae biogeography 

Rhinocems beetles 

Insecticidal properties of plants 

Shoot borer in mahogany 

Insect bird predation 

Wood termite resistance 

Insects of Puerto Rico 
Appendix C. -Additional References 

Alexander, C.P. 1932. The crane-flies of Puerto Rico 

(Diptera). Journal of the Department of Agriculture 

of Puerto Rico. 16(4): 347-387. 

Describes several endemic crane-fly species of the 

Luquillo Mountains and suggests that there may 

be additional new species of Tipulidae confined to 

the Luquillo Mountains. Discussion of the 

possible origin of the Puerto Rican crane-flies and 

descriptions that aid in the species identification 

are also included. 

Barber, H.G. 1939. Insects of Porto Rico and the Virgin 

Islands - Hemiptera-Heteroptera (excepting the 

Miridae and Corixidae). New York Academy of Sciences Scientific Survey of Puerto Rico and the Virgin Islands. 14(3): 263--441. 

Information about families, subfamilies, and keys 

to the genera and species of Hemiptera. 

Bell, R.T. 1970. The Rhysodini of North America, Central America, and the West Indies (Coleoptera: Carabidae or Rhysodidae). Miscellaneous Publications of 

the Entomological Society of America. 6: 289-324. 

Descriptions of the two Rhysodini from Puerto 

Rim. Some notes on behavior and ecology are 

provided. 

Bell, R.T.; Bell, J.R. 1985. Rhysodini of the world, part 

4. Quaestiones Entomologicae. 21: 1-172. 

Descriptions of the two Rhysodini from Puerto 

Rim. 

Blake, D.H. 1941. New species of Chaetocnema and 

other chrysomelids (Coleoptera) from the West 

Indies. Proceedings of the Entomological Society of 

Washington. 43(8): 171-180. 

This and the following publications by Blake cover 

the taxonomy of the chrysomelids. 

Blake, D.H. 1943. New species of the genus Hadropodu 

Suffrian from the West Indies. Bulletin of the 

Museum Comparative Zoology. 92(8): 413-441. 

Blake, D.H. 1948. Six new species of West Indian 

Chrysomelidae. Proceedings of the Entomological 

Society of Washington. 506): 121-127. 

Blake, D.H. 1950. A new genus of flea-beetles from the 

West Indies. Psyche. 57(1): 10-25. 

Blake, D.H. 1951. A revision of the beetles of the genus 

Chalcosicya Blake (Chrysomelidae) from the West 

Indies. Bulletin of the Museum Comparative Zoology. 106(7): 287312. 

Blake, D.H. 1952. Six new species of Megistops with 

keys to the known species (Coleoptera). Psyche. 

59(1): 1-l2. 

Blake, D.H. 1953. The chrysomelid beetles of the genus 

Strabala Chevrolat. Proceedings of the United 

States Natural Museum. 103(3319): 121-134. 

Blake, D.H. 1964. Notes on new and old species of 

AZticinae (Coleoptera) from the West Indies. Proceedings of the United States Natural Museum. 

115(3477): 8-29. 

Blake, D.H. 1970. A review of the beetles of the genus 

Metachroma Chevrolat (Coleoptera-Chrysomelidae). 

Smithsonian Contributions to Zoology. 57: 1-109. 

Bohart, R.M.; Stange, L.A. 1965. A revision of the 

genus Zethus Fabricius in the Western Hemisphere. 

University of California Publications in Entomology. 

Berkeley, CA: University of California Press: 12-147. 

Subspecies of Zethus rufinodis found in the 

Caribbean are discussed. 

Bright, D.E. 1985. Studies on West Indian Scolytidae 

(Coleoptera). Entomologische Arboretum Museum 

George Frey. 34: 169-187. 

A checklist of the West Indian scolytids. 

Caldwell, J.S.; Martorell, L.F. 1950a. Review of the 

Auchenorhynchous Homoptera of Puerto Ria. 1: 

Cicadellidae. Journal of Agriculture of the University of Puerto Rico. 34(1): 1-132. 

A review of the Auchenorhynchous Homoptera of 

Puerto Rim. 

Caldwell, J.S.; Martorell, L.F. 1950b. Review of the 

Auchenorhynchous Homoptera of Puerto Rico. 2: 

The Fulgoridae except the Kinnaridae. Journal of 

Agriculture of the University of Puerto Rico. 34(2): 

133-269. 

A review of the Auchenorhynchous Homoptera of 

. herto Rico. 

Caldwell, J.S.; Martorell, L.F. 1951a. A brief review of 

the Psyllidae of Puerto Rim (Homoptera). Annals 

Entomological Society of America. 44(4): 603-613. 

A review of the Psyllidae of Puerto Rico. 

Caldwell, J.S.; Martorell, L.F. 1951b. New leafhoppers 

from Puerto Rico (Cicadellidae: Homoptera). Journal of Agriculture of the University of Puerto Riw. 

35(2): 88-89. 

A review of the new leafhoppers from Puerto 

Rim. 

Colbn, M. 1981. Contribution to the systematic8 of the 

diaspidids (Homoptera: Diaspididae). Mayagiiez, PR: 

University of Puerto Rim. 199 p. M.S. thesis. 

The taxonomy, host plants, predators, and parasites of 77 homopteran species are discussed. 

Cornstock, W.P. 1944. Insects of Porto Rico and the 

Virgin Islands - Lepidoptera (suborder) Rhopalocera 

(superfamily) Papilionoidea (true butterflies) 

(superfamily) Hesperioidea (skippers). New York 

Academy of Sciences Scientific Survey of Puerto 

Rico and the Virgin Islands. l2(4): 421-622. 

Descriptions and notes on the natural history of 

the Puerto Rican Lepidoptera (suborder Rhopalocera). 

Curran, C.H. 1928. Insects of Porto Rico and the Ti 

gin Islands: Diptera or two-winged flies. New York 
Academy of Sciences Scientific Survey of Puerto 

Rico and the Virgin Islands. 11(1): 3-118. 

Taxonomic keys and description of species of the 

order Diptera including species found in the 

Luquillo Mountains. 

Curran, C.H. 1931. First supplement to the Diptera of 

Puerto Rico and the Virgin Islands. American 

Museum Novitates. 456: 1-23. 

Taxonomic keys and description of species of the 

order Diptera including species found in the 

Luquillo Mountains. 

Davis, W.T. 1928. The cicadas of Puerto Rico with a 

description of a new genus and species. Journal of 

the New York Entomological Society. 36: 29-34. 

A review of the cicadas of Puerto Rico. 

Drake, C.J.; Maldonado, J. 1954. Puerto Rim waterstriders (Hemiptera). Proceedings of the Biological 

Society of Washington. 67: 219-222. 

Checklist of the Puerto Rican water striders 

(Hebridae, Hydrometridae, Gerridae, and 

Veliidae). It also includes information on collecting sites. A new species of water strider on the 

Island is described, and it is suggested that many 

species remain undiscovered. 

Drewry, G.E. 1969. Animal diversity. In: Jordan, C.F.; 

Drewry, G.E., eds. The rain forest project. Annual 

Rep. 129. Rio Piedras, PR: Puerto Rico Nuclear Center: 107-121. 

Key to the Muscidae family at El Verde (Anthomyiidae and Muscidae). 

Equihua, A.; Atkinson, T.H. 1987. Catdogo de Platypodidae (Coleoptera) de Norte y Centroamerica. 

Folia Entomol6gica Mkicana. 72: 5-31. 

A checklist of the Central American pin-hole 

borers (Platypodidae). 

Fisher, W.S. 1935. New cerambycid beetles from Puerto 

Rico. Journal of Agriculture of the University of 

Puerto Rico. 19(2): 51-63. 

ltYelve new species of Cerambycidae from Puerto 

Rico are described. Additional references to 

Puerto Rican Cerambycidae and Buprestidae 

families can be found in "An annotated bibliography of Puerto Rican entomology" (Leonard 

1933) and in Wolcott (1948b). 

Flint, O.S. 1964. The caddisflies (Trichoptera) of 

Puerto Rico. Tech. Pap. 40. Rio Piedras, PR: University of Puerto Rico, Agricultural Experiment Station: 5-80. 

A taxonomical account of the caddisflies of Puerto 

Rico and information on the natural history of 

some species. 

Forbes, W.T.M. 1930. Inseds of Porto Rico and the 

Virgin Islands- Heteroeera or moths (excepting the 

Noctuidae, Geometridae, and Pyralidae). New York 

Academy of Sciences Scientific Survey of Puerto 

Rico and the Virgin Islands. 12(1): 4-171. 

A review of the taxonomy and natural history of 

moths (suborder Heterocera). 

Forbes, W.T.M. 1931. Supplementary report on the Heterocera or moths of Porto Rico. Journal of the 

Department of Agriculture of Puerto Rico. 15(4): 

339-394. 

A review of the taxonomy and natural history of 

moths (suborder Heterocera). 

Fox, I. 1946. A review of the species of biting midges or 

culicoides from the Caribbean region. Annals of the 

Entomological Society of America. 39(2): 248-258. 

~onomic keys to the females and geographical 

distributions of species of biting midges or 

culicoides. 

Fox, 1. 1957. The insect family Japygidae (Order Thysanura) in Puerto Rico. Journal of Agriculture of 

the University of Puerto Rico. 41(1): 35-37. 

Discussion on the Japygidae family in Puerto 

Rico. 

Garcia, J. 1938. An ecological survey of the fresh water 

insects of Puerto Rico. Journal of Agriculture of the 

University of Puerto Rim. 22(1): 43-97. 

Mayflies (order Ephemeroptera) are reported as 

uncommon in the lowlands, but numerous along 

the creeks in the higher areas. 

Garrison, R.W. 1986. Diceratobasis melanogaster Spec. 

Nov., a new darnsel-fly from the Dominican Republic 

(Zygoptera: Coenagrionidae) with taxonomic and 

distributional notes on the Odonata of Hispaniola 

and Puerto Rico. Odonatologica. 15(1): 61-67. 

Swen new records from Puerto Rico are reported, 

increasing the number of known Odonata species 

to 49. A key for most neotropical coenagrionidslacking postocular spots is included. 

Grimaldi, D.A. 1988. Relicts in the Drosophilidae (Diptera). In: Liebherr, J.K., ed. Zoogeography of Caribbean insects. Ithaca, NY: Cornell University Press: 

183-213. 

A list of the endemic species of Antillean Drosophilidae. Many records from the Luqdlo Mountains are included. Drosophila carcinophila 

larvae feed on the exudate in nephric gmoves of 

the black land crab, Gecarcinus ruricula, at El 

Yunque. 

Mots, E.B. 1932. Insects of Puerto Rico and the Viigin 

Islands: Odonata or dragon flies. New York Academy 

of Sciences Scientific Survey of Puerto Rico and the 

Virgin Islands. 14(1): 3-107. 

Descriptions of and keys to the species of dragonflies of Puerto Rico. 

Leonard, M.D. 1933. An annotated bibliography of 

Puerto Rican entomology. Journal of the Department of Agriculture of Puerto Rico. 17(1): 5-96. 

References to former taxonomic works for Puerto 

Rican entomology. 

Liebherr, J.K. 1986. Barylaus, new genus (Coleoptera: 

Carabidae) endemic to the West Indies with Old 

World afinities. Journal New York Entomological 

Society. 940): 83-97. 

The new genus was erected based on two species: 
Colpodes estriatus from Puerto Rico and C. puncticeps from the Dominican Republic. 

Liebherr, J.K., ed. 1988. Zoogeography of Caribbean 

insects. Ithaca, NY: Cornell University Press. 285 p. 

Publications on the zoogeography of the following 

groups: Lygaeidae (Hemiptera), Auchenorrhynchous Homoptera, Carabidae (Coleoptera), Polycentropodidae (Trichoptera), Drosophilidae 

(Diptera), Formicidae (Hymenoptera), and Halictidae (Hymenoptera). 

Maldonado, J. 1969. The Miridae of Puerto Rico 

(Insecta, Hemiptera). %h. Pap. 45. Rio Piedras, 

PR: University of Puerto Rico, Agricultural Experiment Station. 133 p. 

The furst systematic study of the Puerto Rican 

Miridae, which includes discussion on the species 

from the Luquillo Mountains. Description of each 

genus is given. Keys to the neotropical species (if 

the number of species is not large) are provided. 

Otherwise, a key to the Puerto Rican species is 

included. 

Maldonado, J. 1980. The genus Jorbetus Distant, 1884 

(Hemiptera: Miridae: Orthotylinae). Journal of 

Agriculture of the University of Puerto Rico. 64(3): 

304-309. 

Revision of the genus Jobertus, including J. gracilentus, collected in the Luquillo Mountains. 

Maldonado, J.; Navarro, C.A. 1967. Additions and corrections to Wolcott's "Insects of Puerto Rico." 

Caribbean Journal of Science. 7: 45-64. 

The approximate number of insects found in 

Puerto Rico is indicated and the presence of the 

order Zoraptera on the island is reported. The 

zorapterans were collected in a rotting log in the 

Luquillo Mountains. The order Zoraptera is 

mostly found in tropical regions and contains 

approximatdy 25 described species. 

Mari, JA. 1976. The genera of Collembola (Insecta) in 

Puerto Rico; keys, diagnosis, and general comments. 

Journal of Agriculture of the University of Puerto 

Rim. 60(1): 113-128. 

Keys for the Collembola families and genera of 

Puerto Rico. The keys and diagnostic criteria are 

elaborated for the nonspecialist. 

Mari, JA. 1987. Puerto Rican species of Paronellidae 

(Insecta: Collembola). Caribbean Journal of Science. 

23: 400-416. 

A discussion of the members of the Paronellidae. 

Additional species of the genera Dicranocentruga 

may be discovered by additional collecting in the 

Luquillo Mountains. 

Medina, S. 1961. The Thysanoptera of Puerto Rico. 

Tech. Pap. 32. Rio Piedras, PR: University of Puerto 

Rico, Agricultural Experiment Station. 159 p. 

Keys to families, genera, and species for the Thysanoptera of Puerto Rico. 

Medina, S. 1978. The Muscidae (Diptera) of Puerto 

Rico. Iowa State University. 308 p. Ph.D. thesis. 

Keys to subfamilies, genera, and species of Muscidae (sensu lato) of Puerto Rico, describing several species from the Luquillo Mountains. 

Medina, S.; Martorell, L.F. 1973. New insect records for 

Puerto Rico. Journal of Agriculture of the University of Puerto Rico. 57 (3): 247-254. 

This paper complements Maldonado and Navarro 

(1967). 

Miller, L.D.; Miller, J.Y. 1989. The biogeography of 

West Indian butterflies (Lepidoptera: Papilionoidea, 

Hesperioidea): a vicariance model. In: Woods, CA., 

ed. Biogeography of the West Indies: past, present 

and future. Gainesville, FL: Sandhill Crane Prurss: 

229-262. 

A combination of vicariance/dispersal model for 

the evolution of the Antillean butterfly fauna is 

presented. 

Moxey, C.F. 1972. The stick-insects (Phasmatodea) of 

the West Indies- their systematics and biology. 

Cambridge, MA: Harvard University. 211 p. Ph.D. 

thesis. 

The most recent publication dealing with Puerto 

Rican walking stick taxonomy. 

Needham, J.G. 1941. Life history notes on some West 

Indian coenagrionine dragonflies (Odonata). Journal 

of Agriculture of the University of Puerto Rico. 25 

(3): 1-19. 

Information on the life history of four genera of 

damselflies in the West Indies. Also contains keys 

to the adults and nymphs, but without refmce 

to collection localities from the Luquillo Mountains. 

Nosek, J. 1978. Protura: a new insect order record for 

Puerto Rico with description of new species. Journal 

of Agriculture of the University of Puerto Rico. 

62(2): 133-138. 

A new insect order for Puerto Rico. 

Osborn, H. 1935. Insects of Porto Rico and the Virgin 

Islands: Homoptera (excepting the Sternorhynchi). 

New York Academy of Sciences Scientific Survey 

of Puerto Rico and the Virgin Islands. 14(2): 

111-260. 

Information on host plants, economic importance, 

and the effects of tropical storms on Homoptera 

abundance. Also contains taxonomic keys and 

information on natural history for some species. 

Ramos, J.A. 1957. A review of the Auchenorhynchous 

Homoptera of Puerto Rico. Journal of Agriculture of 

the University of Puerto Rico. 41(1): 3847. 

Review of the families Kinnaridae, Cercopidae, 

Membracidae, and Cicadidae. %onomic keys for 

the identification of species, notes on the distribution, abundance, and host plants are provided. 

This work complements the works on the 

Cicadellidae, Psyllidae, and Fulgoroidea conducted by Caldwell and Martorell (1950a, 1950b, 

1951a, 1951b). 

Ramos, S.J. 1982. Checklist of the butterflies of Puerto 
Rico (Lepidoptera, Rhopalocera, West Indies). Caribbean Journal of Science. 17: 59-68. 

A checklist of the butterflies of Puerto Rico. 

Riley, N.D. 1975. A field guide to the butterflies of the 

West Indies. New York: Demeter Press. 224 p. 

Information on the life history and distribution of 

Puerto Rican butterflies. 

Ruiz, M.J. 1989. Revisicin del g6nero Penichrophorus 

Richter (Homoptera: Membracidae, Cerescini). Mayagiiez, PR: University of Puerto Rico. 104 p. M.S. 

thesis. 

A review of the genus Penichmphorus (Homoptera: Membracidae). 

Schauss, W. 1940. Insects of Porto Rico and the Virgin 

Islands. Moths of the family Noctuidae. New York 

Academy of Sciences Scientific Survey of Puerto 

Rico and the Virgin Islands. 12(2-3): 177-417. 

Description of the moths of the family Noctuidae. 

Selander, R.B.; Bouseman, J.K. 1960. Meloid beetles 

(Coleoptera) of the West Indies. Proceedings of the 

United States Natural Museum. lll(3428): 197- 

226. 

Comments on the scarcity of this family in the 

West Indies and the origin of the Caribbean meloids. Meloidae larvae are parasitic on grasshopper 

egg pods and the contents of nesting cells of wild 

bees. They are generally phytophagous as adults. 

Because of the complexity of their ecology, the 

Meloidae face problems of dispersal, and it is 

expeded that they will be poorly represented in 

areas having a prolonged history of isolation. 

Selander and Bouseman postulated that phoresy 

(especially on bees) was a prerequisite to successful colonization of meloid beetles on the Caribbean islands. 

Selander, R.B.; Bouseman, J.K. 1961. Supplementary 

records of meloid beetles (Coleoptera) of the West 

Indies. Entomological News. 72(7): 190-191. 

Records of meloid beetles in the West Indies. 

Smith, C.F. 1970. Notes on the genus Picturaphis and 

related genera with a new species of Pictumphis 

from Puerto Rico (Aphididae: Homoptera). Journal 

of Agriculture of the University of Puerto Rico. 

54(4): 683-688. 

Description of a new species of Picturaphis from 

the Luquillo Mountains. 

Smith, D.R.; Lavigne, R.J. 1973. nYo new species of 

ants of the genera 'Zhpinorna Forester and Paratrechina Motschoulsby from Puerto Rico (Hymenoptera: Formicidae). Proceedings of the Entomological 

Society of Washington. 75(2): 181-187. 

Description of Paratrechina cisipa as a new 

species based on material collected near El Verde. 

Smith, M.R. 1936. The ants of Puerto Rico. Journal of 

Agriculture of the University of Puerto Rico. 20(4): 

819-875. 

Keys to the Puerto Rican ants and notes on the 

natural history of the species. 

Snelling, Roy R. 1986. New synonymy in Caribbean 

ants of the genus Leptothorax (Hymenoptera: Formicidae). Proceedings of the Entomological Society 

of Washington. 88(1): 154-156. 

Review of the genus Lepthotonuc found .in the 

Luquillo Mountains. 

Snyder, F.M. 1957. Puerto Rican NeodQiopsis (Diptera: Muscidae: Coenosiinae). Journal of Agriculture 

of the University of Puerto Rico. 41(3): 207-229. 

Description of flies of the genus Neoderiopsis, 

which are limited to the higher elevations of the 

island of Puerto Rico. The majority of the holotypes for the descriptions of 10 new species came 

from the Luquillo Mountains. 

Soto, F.N. 1988. Nuevos dicirtumidos de Puerto Rico 

(Insects: Collembola: Dicyrtomidae): New Dicyrtomidae species from Puerto Rico (Ins&. Collembola: Dicyrtomidae). Caribbean Journal of Science. 

24: 60-70. 

Four additional Collembola species for Puerto 

Rico are reported. 

Steyskal, G.C. 1973. The Clusiidae of Puerto Rico and 

the Virgin Islands (Diptera). Studia Entomologica. 

16(1-4): 439--444. 

Description of and general information on the 

family Clusiidae in Puerto Rim. 

Telford, H.S. 1973. The Syrphidae of Puerto Rico. 

Journal of Agriculture of the University of Puerto 

Rim. 57(3): 217-246. 

A taxonomic review of the Syrphidae. Also includes keys, illustrations of some species, and 

notes on the natural history of the group. 

%avers, J.R. 1938. Mayflies of Puerto Rico. Journal of 

Agriculture of the University of Puerto Rico. 22(1): 

5-42. 

Keys to the mayflies of Puerto Rico (for both adults 

and nymphs) and descriptions of new Puerto Rican 

species. Many of the specimens collected by 'havers were from the Luquillo Mountains. 

Tulloch, G.S. 1937. The mosquitoes of Puerto Rico. 

Journal of Agriculture of the University of Puerto 

Rico. 21(4): 137-167. 

Information on the natural history and economic 

importance of Puerto Rican mosquitoes 

(Culicidae). Health problems caused by mosquitoes are discussed. 'Ihonomic keys are presented. 

Wheeler, M.R.; Takada, H. 1963. A revision of the 

American species of Mycodrosophila (Diptera: Drosophilidae). Annals of the Entomological Society of 

America. 56(3): 392-399. 

- HRvision of the genus iviycodmsophila describii-ig 

new species. Includes records from the Luquillo 

Mountains. 
Torres, Juan A. 1994. Insects of the Luquillo Mountains, Puerto Rico. 

Gen. Tech. Rep. SO-105. New Orleans, IA U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 53 p. 

The literature on forest entomology in the Luquillo Mountains is 

reviewed, and information on basic insect ecology, insect pests, and 

systematics is provided. 

Keywords: Bark beetles, herbivorous, termites, tropical forests, 

wood. 

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