Group Title: Structure and composition of the aquatic invertebrate community inhabiting epiphytic bromeliads in south Florida and the discovery of an insectivorous bromeliad /
Title: Structure and composition of the aquatic invertebrate community inhabiting epiphytic bromeliads in south Florida and the discovery of an insectivorous bromeliad
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Title: Structure and composition of the aquatic invertebrate community inhabiting epiphytic bromeliads in south Florida and the discovery of an insectivorous bromeliad
Physical Description: ix, 78 leaves : ill. ; 28 cm.
Language: English
Creator: Fish, Durland, 1944-
Publication Date: 1976
Copyright Date: 1976
 Subjects
Subject: Aquatic invertebrates -- Florida   ( lcsh )
Bromeliaceae   ( lcsh )
Entomology and Nematology thesis Ph. D
Dissertations, Academic -- Entomology and Nematology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis--University of Florida.
Bibliography: Bibliography: leaves 65-72.
General Note: Typescript.
General Note: Vita.
Statement of Responsibility: by Durland Fish.
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Bibliographic ID: UF00098120
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000355727
oclc - 02686928
notis - ABZ3980

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STRUCTURE AND COMPOSITION OF THE AOUATIC INVERTEBRATE COMMUNITY
INHABITING EPIPHYTIC BROMELIADS IN SOUTH FLORIDA AND
THE DISCOVERY OF AN INSECTIVOROUS BROMELIAD











By

DURLAND FISH


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA
1976













ACKNOWLEDGMENTS

This study began as a course project in community ecology for

Dr. Archie Carr, and to him I express special thanks for his constant

enthusiasm and encouragement. Special thanks are also due to

Dr. Thomas J. Walker who served as my committee chairman until the last

quarter. His valuable suggestions, encouragement, and moral support

contributed much to the success of this study and to my graduate

experience at the University of Florida.

I sincerely thank the members of my Doctoral Committee,

Drs. D. H. Habeck, D. E. Weidhaas, J. J. Ewel, and J. J. Reiskind, for

their assistance and cooperation during the preparation of this manu-

script. I am particularly grateful to Dr. Weidhaas and the USDA-ARS

Insects Affecting Man Laboratory for providing financial assistance

throughout the course of this study through a research assistantship.

Transportation was provided by the Department of Entomology and lodging

was provided by the Archbold Biological Station, Lake Placid, the

University of Florida Agricultural Research and Education Center at

Homestead, and Dr. J. H. Frank of the Florida Medical Entomological

Laboratory in Vero Beach.

My wife Shirley faithfully assisted with all of the field work

under sometimes severe and uncomfortable conditions. The staff of the

Florida Medical Entomology Laboratory, the Everglades National Park, and

Dr. F. C. Craighead, Sr., and Mr. Richard Archbold provided information

on site locations. Mr. Ben Swendsen of Lykes Bros. Inc., provided access









to the Fisheating Creek Wildiife Management Area. Mr. Thomas Gibson of

the University of Arizona supplied valuable information and stimulating

discussion concerning carnivorous plants.

Dr. Lloyd Knutson of the Insect Identification and Beneficial Insect

Introduction Institute, USDA, and Dr. H. V. Weems of the Florida Depart-

ment of Agriculture and Consumer Services, Division of Plant Industry

assisted in coordinating insect identification efforts. The following

specialists provided identifications for the aquatic invertebrates:

Mr. W. M. Beck, Florida A&M University Chironomidae

Dr. H. L. Cromroy, University of Florida Anoetidae

Mr. Bill Hart, USNM Ostracoda

Dr. H. C. Huckett, Cornell University Muscidae

Dr. F. C. Thompson, USNM Syrphidae and Psychodidae

Dr. C. W. Sabrosky, USNM Aulacigastridae

Dr. W. A. Steffan, Bishop Museum (Honolulu) Sciaridae

Dr. W. W. Wirth, USNM Ceratopogonidae















TABLE OF CONTENTS


P

ACKNOWLEDGMENTS........... . . . . . .

LIST OF TABLES... . . . . . . . .

LIST OF FIGURES.......... . . . . . .

ABSTRACT . ... .. . ... . . . . . . .vi

INTRODUCTION...... . . . . . . .

CHAPTER I

INSECTIVORY IN CATOPSIS BERTERONIANA: A NEW
METHOD OF NUTRIENT PROCUREMENT IN THE
BROMELIACEAE . . . . . . ... . .

CHAPTER II

A SURVEY OF THE AQUATIC FAUNA INHABITING THE
LEAF AXILS OF EPIPHYTIC TANK BROMELIADS IN
SOUTH FLORIDA

Introduction . . . . . . . . . . .

Materials and Methods . . . . ... . .

Results and Discussion . . . . . . . .

Conclusion . . . . . . . . . . .

CHAPTER III

FACTORS INFLUENCING THE STRUCiURE OF AQUATIC
COMMUNITIES INHAEITING EPIPHYTIC BROMELIADS
IN SOUTH FLORIDA

Introduction . . . . . . .


age

ii

vi

'ii

ii
I


36














Collecting Sites .. . .. . . . .. .39

Sampling Program. . . . .. ...... ..41

Analytical Methods. ... . . .... ... .42

Results and Discussion. . . . . . .. .45

LITERATURE CITED . . ............... .65

APPENDIX ...... ..... .... .... .73

BIOGRAPHIC SKETCH ................. .. 78














LIST OF TABLES


Table Page

I Insect prey recovered from C. berteroniana leaf-axils
over 8 days at Gainesville, Florida . . . ... . 52

II Summary of site descriptions and collecting data of
bromeliad samples from south Florida . . . ... .53

III Distribution of aquatic invertebrates within 6 species
of Florida tank bromeliads. .. ... ..... . . 54

IV List of aquatic invertebrate species included in the
analysis. ... .. ... ........ ... ... . 55

V Two-way coincidence table showing abundances of aquatic
invertebrate species (with species numbers) within
bromeliad species at 3 sites and 4 seasons . ... .. 56














Li T OF FIGURES


Figure Pje

1 Visible and UV light photographs of bromeliads illuminated
by natural sunlight. .... . .. . . . t.

2 Location of bromeliad collection n sites in Florida .. 60

3 Sagittal (A) and cross-sections (B) of T. utricu-ata showing
inflated leaf axils that form water-holding chambers. . . c

4 Cluster analysis of euclidiar distances among structures of
the aquatic community inhabiting 2 bromeliad species, at
different times of the year and in 3 major south Florida
ecosystems . .... ...... .... .. . .. .









Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy


STRUCTURE AND COMPOSITION OF THE AQUATIC INVERTEBRATE COMMUNITY
INHABITING EPIPHYTIC BROMELIADS IN SOUTH FLORIDA AND
THE DISCOVERY OF AN INSECTIVOROUS BROMELIAD

By

Durland Fish

December 1976

Chairman: Dale H. Habeck
Major Subject: Entomology

A survey of the aquatic invertebrates inhabiting the water-

filled leaf axils of 6 species of epiphytic tank bromeliads occur-

ring in south Florida revealed 13 species of dipterous insects, 2

species of ostracods, a mite, a turbellarian, and an oligochaet worm.

The list represents 2 new families and 8 new species of bromeliad

inhabiting insects, including 7 undescribed new species. Most of those

species are widely distributed and abundant, and seem to be restricted

to the bromeliad habitat during their aquatic stages. A review of the

literature on aquatic fauna inhabiting bromeliads in the Neotronics

indicates that the Florida bromeliad fauna is of Neotropical origin but

is depauperate as compared to the faunal lists for Costa Rican and

Jamaican bromeliads.

A cluster analysis of community structure indicates that species

composition and abundance are characteristic of the ecosystem supporting

the bromeliad flora with the bromeliad species containing the aquatic

community having the least influence upon structure. Differences in

amounts and rates of throurhfall and leaf litter accumulation among

ecosystems supporting bromeliads are suggested to be important factors


viii









in determining community structure since most species of aquatic

invertebrates inhabiting bromeliads in south Florida seem to be

detritus feeders. Seasonal variation in community structure was

greatest in cypress and mangrove ecosystems and least in a tropical

hardwood hammock.

The epiphytic bromeliad Catopsis berteroniana (Shult) Mez was

found to be insectivorous. This species captures flying insects within

erect tube-like leaves which are lubricated with a fine white powder

that prevents escape. The white powder also reflects UV light which is

believed to attract the insects. Tie first report of insectivory in

the Bromeliaceae represents a new method of nutrient procurement foi

the family.














INTRODUCTION


Water-holding epiphytic bromeliads are characteristic features

of many ecosystems in the American tropics and subtropics, and their

abundance in certain areas of south Florida constitutes a major

feature of the total landscape. The existence of these plants poses

two important questions to the biologist: 1) How do they maintain

themselves in the total absence of soil? and 2) What is the nature of

the communities of aquatic organisms that inhabit them? Both of these

questions are addressed in this dissertation.

The first chapter describes an epiphytic bromeliad that was found

to be insectivorous. This discovery represents the first report of

insectivory in the Bromeliaceae and provides a new explanation for

nutrient procurement in this family in addition to those currently

proposed.

The results of a survey of aquatic invertebrates inhabiting the

bromeliads of south Florida are reported in the second chapter. This

survey indicates that the aquatic fauna is not evenly distributed

within all species of water-holding bromeliads or throughout the

total ranges of these species. An investigation into the causes of

differences in species composition and abundance of the aquatic

communities is reported in the third chapter.

These chapters are written in manuscript form and are intended

to be submitted with only minor changes in style and form for publica-

tion in major scientific journals.















CHAPTER I
INSECTIVORY IN CATOPSIS RERTERONIANA: A NEL METHOD
OF NUTRIENT PROCUREMENT INTHE ROMELIACEAE


The plant family Rromeliaceae contains over 2,000 described species

and constitutes a major portion of the epiphytic flora in the Neotropics.

In general, growth and maintenance of these plant communities is

not easily explained in the absence of a soil substrate, and nutrient

procurement by epiphytic bromeliads has been the subject of many

recent investigations (Benzing 1970a, 1970b, 1973, Benzing and Burt

1970, Benzing and Renfrow 1974). In adapting to a habitat of extreme

scarcity of nutrient salts, several strategies have evolved within

this family. The fundamental adaptations have been decreased dependence

upon root absorption of nutrients and an increased dependence upon

foliar procurement and foliar absorption of nutrients.

At least 3 methods of foliar procurement of nutrients have

been identified in the Bromeliaceae (Benzing 1973). 1) Tank bromeliads

have tightly overlapping leaf axils which are inflated to impound

varying amounts of rain water intercepted by channeled leaves in

a tight rosette configuration. They usually occur within or below

the forest canopy and intercept nutrients leached by rain from living

leaves in the canopy (Tukey 1970a, Benzing and Renfrow 1974). Falling

leaf litter also contributes to the nutrient pool which is ultimately

broken down by a variety of aquatic invertebrates and microorganisms








living in the leaf-axil water (Picado 1913, Laessle 1961, see also

Chapter 2). 2) Atmospheric bromeliads such as spanish moss (Tillandsia

usneoides), also absorb nutrients from canopy leachates but are

unable to impound water. These plants have a high surface/volume

ratio and possess large numbers of epidermal trichomes that act

as one-way valves for the absorption of dilute nutrient salts during

brief but heavy rains (Benzing and Dahle 1971). 3) Myrmecophytic

bromeliads such as T. butzii and T. caput-medusae have moderately

inflated leaf axils and an overall bulbous shape. The leaf axils

are unable to impound water but instead provide a habitat for colonies

of various species of ants. The foraging ants return nutrients

to the plant for brood rearing which, in addition to excretory products,

provide a nutrient pool for the plant (Benzing 1970a).

A fourth method of foliar procurement, that of insectivory,

is here described for the first time in the Bromeliaceae.

The epiphytic bromeliad Catopsis berteroniana ranging from

extreme south Florida to southern Brazil, hds all of the physical

attributes of a tank bromeliad. However, at least in south Florida,

it rarely occurs beneath the forest canopy, but instead it is found

attached to small branches in the very tops of canopy trees or on

dead trees and shrubs in open areas. The water-filled leaf axils

rarely contain leaf litter but are instead filled with the chitinous

remains of insects. This species is characterized by erect leaves

and a conspicuous white chalky powder covering the leaf bases on

both sides (Fig. 1A). Both of these features are important components

of its insectivorous nature.








The overlapping leaves form a series of tubes having steep sides,

and each contains about 10 ml of rain water. Insects are unable to

escape from these tubes because of the fine white chalk which effectively

lubricates the walls. (When the powder is removed with a brush the

insects easily escape.) This unique arrangement constitutes a passive

pitfall type trap for securing insect prey which ultimately drown in the

water. The pitfall type traps of C. berteroniana are functionally

similar to those of insectivorous pitcher plants (Nepentheaceae

and Sarraceniaceae), although in pitcher plants, the trap is formed

by a single leaf and escape is prevented by numerous downward pointing

trichomes on the internal leaf surface (Lloyd 1942, Fish and Carlysle

in preparation) instead of a lubricating powder.

The existence of proteolytic enzymes in the leaf axils of tank

bromeliads has been suspected for some time (Picado 1913) but has

not been investigated by modern biochemical techniques. However,

bacterial exoenzymes, introduced with insect prey, are an important

component of the digestive fluid of insectivorous pitcher plants

(Sarraceniaceae) (Plummer and Jackson 1963), and probably also occur

in the leaf axils of C.berteroniana.

Foliar absorption of nutrients in C. berteroniana has previously

been demonstrated by Benzing and Burt (1970) who also used P32 labelled

insects to demonstrate decomposition and subsequent foliar absorption

in the leaf axils of the epiphytic tank bromeliad Aechmea nudicaulis

(Benzing 1970b). Apparently most, if not all, tank bromeliads have

the capacity to break down and absorb both animal and plant matter

that accumulates within their leaf axils.

The white powder of the leaf bases might also be involved in

the recruitment of insect prey. Photographs of the same sunlit









plant taken with visible light (Fig. 1A) and with a Wratten 18-A UV

filter which absorbs visible light (400 nm.) allowing only UV light to

be recorded on the film, (Figs. B1, IC), shows that the white powder

strongly reflects UV light while the uncoated portion of the leaves absorb

UV light. In nature, UV light is an indication of open space since

natural objects absorb UV light and its only sources are the sun

and sky to which insects are believed to orient (Mazokhin-Porshnyakov

1969). With the basal portion of the entire plant acting as a mirror

in the UV end of the spectrum, flying insects sensitive to UV may

not be able to distinguish this part of the plant from the normal

UV radiation in the atmosphere, may collide with the plant, and fall

into the water-filled leaf axils. Since C. herteroniana normally

occurs only in exposed situations where they would be easily encountered

by flying insects, and since nectar secretions or other attractive sub-

stances were not evident, it is unlikely that it actively attracts insect

prey.

A passive method of capture is further suggested by the wide

diversity of insects captured by C. berteroniana in the field. Four

plants were transported from their natural habitat in Everglades

National Park to Gainesville, Florida, where they were affixed to

fence posts 1.5 m above the ground to facilitate observation. Freshly

captured insects were removed from the leaf-axils of the plants for 8

consecutive days. The catch totalled 136 specimens representing 8

insect orders in addition to spiders (Table 1). Hymenoptera, Diptera,

Coleoptera, and Lepidoptera constituted 87% of the prey. A high

incidence of parasites, predators, and phytophaqous insects indicates

that they were not attracted by a common food source.








Noninsectivorous tank bromeliads typically do not have erect leaves,

white powder, or UV reflecting surfaces (Fig. ID). Consequently they do

not recruit or have the capacity to retain insect prey. Also most tank

bromeliads usually harbor numerous living terrestrial arthropods such as

cockroaches, beetles, ants, scorpions, etc., within the older leaf axils

(Picado 1913, Laessle 1961, Fish,unpublished data) a feature never

observed in C. berteroniana.

The evolution of insectivory among epiphytic Bromeliaceae is most

likely the result of competition among the many species within this

family that have adapted to the nutrient poor epiphytic environment

from terrestrial ancestors (Pittendriqh 1948). Catopsis berteroniana

avoids direct competition with other tank bromeliads by being completely

independent of nutrients from the forest canopy in addition to being

independent from a soil root substrate. Such independence has been

achieved in at least two other families (Lcntibulariaceae and

Nepentheaceae) by giving rise to insectivorous epiphytic species

Utricularia montana (Taylor 1967), Nepenthes vetchei, and N.

reinwardtiana (Smythies 1965) directly from insectivorous ancestors

adapted to nutrient poor terrestrial environments.















CHAPTER II
A SURVEY OF THE AQUATIC FAUNA INHABITING THE LEAF AXILS OF
EPIPHYTIC TANK BROMELIADS IN SOUTH FLORIDA

Introduction

Many species in the plant family Bromeliaceae impound rain water

within tightly overlapping leaf axils and are referred to as tank brome-

liads. Most are true epiphytes which attach themselves to the trunks

and limbs of larger plants with strong fibrous roots. Their channeled

leaves and rosette configuration provide for efficient interception

of rain water which is funneled directly into the leaf-axil chambers

(Benzing et al. 1972). The impounded water can range from a few milli-

liters to several liters depending upon the size of the plant and degree

of leaf base inflation.

Water in the leaf axils contains varying amounts of nutrients

obtained from decomposition of allochthonous leaf litter and rain water

that has leached minerals and organic matter from living leaves in

the above forest canopy (Tukey 1970a, Benzing and Renfrow 1974). Epiphytic

tank bromeliads are totally dependent upon the contents of their leaf

axils for nutritional requirements in the absence of a root soil sub-

strate (Pittendrigh 1948, Benzing 1973, Benzing and Renfrow 1974) and

absorb nutrients directly through the leaf surface (Benzing 1970b,

Benzing and Burt 1970).

Foliar impoundment of nutrient laden water by epiphytic tank

bromeliads provides a unique arboreal habitat for a variety of aquatic










animal life ranging from protozoa to anuran tadpoles. The communities

of aquatic organisms associated with epiphytic tank bromeliads have

been studied by Picado (1913) in Costa Rica, Laessle (1961) in Jamaica,

and Maguire (1970) in Puerto Rico. These and other studies reviewed

by Maguire (1971) have shown that bromeliad communities are useful

in investigations of interspecific interactions, community structure,

colonization, dispersal, and other aspects of community ecology because

they are small, relatively simple, and conveniently sampled.

In addition to their ecological significance, bromeliad communities

are important as breeding sites for many species of blood-sucking insects

of potential public health importance. Larvae of at least 6 genera

of mosquitoes (Culicidae) are known to occur in bromeliads (Horsfall

1972) in addition to horseflies (Tabanidae) (Goodwin and Murdoch 1974)

and biting midges (Ceratopogonidae) (Wirth 1974). Very little is known

of the ecology of these important groups of bromeliad-breeding insects,

or how they are distributed among the hromeliad flora.

Of the more than 2,000 described species of bromeliads in the

Neotropics, 17 have become established in subtropical Florida (Craighead

1963). Several of these same species support communities of aquatic

invertebrates in the tropics but very little is known of the aquatic

bromeliad fauna in Florida. Only mosquitoes (3 spp.) (King et al.

1960), chironomids (3 spp.) (Beck and Beck 1966), and an ostracod (Tressler

1956) have been reported from Florida tank bromeliads and, except for

the mosquitoes, these reports are from single collections.

In view of the significance of bromeliad communities in ecological

studies and the public health significance of some of the fauna, it









is surprising that so little is known of the species composition of

these communities in general, and especially in south Florida where

the aquatic invertebrate fauna is relatively well known. The present

study represents the first attempt to compile a complete census of

aquatic metazoan invertebrates inhabiting the tank bromeliads of south

Florida, and to compare these findings with the results of similar

studies conducted in other areas of the Neotropics as reported in the

literature.

Materials and Methods

Only 7 of the 17 species of bromeliads occurring in south Florida

are considered true tank bromeliads; these are Tillandsia utriculata L,

T. fasciculata SWJ., T. valenzuelana A. Rich., Catopsis berteroniana

(Schult) Mez, C. floribunda (Brongh.) Smith, C. nutans (Sw) Griseb.,

and Guzmania monostachia (L.) Rushy. Several other species ot Florida

bromeliads are considered ephemeral tank bromeliads. These have very

small leaf axils that contain free water only briefly after heavy rain

and were not found to support aquatic organisms.

Over 360 bromeliads wer, sampled from 17 locations in south

Florida, (Fig. 2) between January 1974 and December 1975. A variety

of major plant communities supporting bromeliad populations were included

to insure a representative collection of the bromeliad fauna (Table 2).

Mere detailed floristic descr-ptions of the collecting sites are provided

by Davis (1943) and Craiqnead (1971).

The frequency that each bromeliad species was sampled is biased

toward the most abundant (Tab;e 3). Tillandsia utriculata and T.

fasciculata are by far the most abundant of the Florida tank bromeliads









10

and were consequently sampled heavily in comparison to the other 4

species. Tillandsia utriculata was sampled most frequently as it is

more widespread and occurred in most of the sampling sites. Catopsis

berteroniana and T. valen:uelana are not widespread but are locally

abundant in extreme south Florida, and were sampled on several occasions.

Catopsis floribunda and G. monostachia were sampled least frequently

since they were found at only one site. Catopsis nutans was not sampled

during this study because of its scarcity.

Truly random sampling was not feasible since it would require

the numbering of all plants in each location and subsequently collecting

them in random sequence. Although the sampling was admittedly biased

in favor of accessibility, special efforts were made to collect evenly

throughout the habitats, and trees were frequently climbed to obtain

specimens high in the canopy. Medium to large plants were purposely

selected because they were most likely to contain fauna and were easier

to identify.

Bromeliads were identified with a descriptive key by Craighead

(1963) and by comparison with herbarium specimens at the University

of Florida.

The maximum water holding capacity of each bromeliad species

was determined to assess its potential to support aquatic fauna.

Depending upon the species, 5 to 20 of the same plants selected for

faunal sampling were filed to capacity with tap water and then emptied

into a volumetric cylinder. The average water holding capacity of

the sample was calculated by dividing the total plan, volume by the









average number of leaves on the plants in the sample. The 3 measures --

total plant volume, number of leaves, and volume per leaf axil -- were

found to be useful in describing the physical characteristics of the

different bromeliad species. However, these measures are not intended

to be estimates of the actual water volumes present in the leaf axils

in nature since the sampled plants were rarely filled to capacity.

Individual plants were carefully removed from their attachment

sites to avoid spilling the leaf-axil water and immediately processed

in the field. Processing of the plants involved a modification of

the method devised by Frank et al. (1976) for removing the immature

stages of bromeliad mosquitoes. Each plant was inverted in a large

bucket containing sufficient water to cover the leaves. The leaf axils

were washed by a rapid up and down movement with the plant held by

its roots. After 30 sec of washing, the plant was removed and the

wash water strained through a fine mesh screen to remove the debris

and fauna. The collected material was rinsed from the screen into

an enamel pan and then rinsed into labelled plastic bags, each containing

the contents of one bromeliad. The bags were transported in an insulated

box containing ice to the laboratory in Gainesville where they were

stored at 80C until examination.

The efficiency of the sampling method was frequently checked by

carefully dismantling bromeliads leaf by leaf. The results indicated

that all faunal species were removed by the washing method except

Aschelminthes. Rotifers and nematodes were only occasionally retained

by the screen and consequently could not be included in the study.









Each sample was examined with a dissecting microscope at 7-30X

magnification after removing and rinsing the leaf litter and large

debris. Aquatic organisms were collected with a pipet ,nd either preserved

in 70% ethanol or set aside for rear-ng. Hearing proved to be a diffi-

cult task because of the number of species involved and the absence

of knowledge regarding their feeding habits or pupation requirements.

Few specimens would pupate under laboratory conditions. To obtain

adequate numbers of adult specimens for identification, dark plastic

bags were placed over unemptied bromeliads in the field. Small plastic

vials fitted with filter paper cones were placed in an opening at the

tops of the bags. Emerging adults ttapned in the vials were used to

supplement the laboratory reared material.

Since it is difficult or impossible to view these organisms directly

in the leaf axils, observations on tie behavior and feeding habits

of some species were conducted in the laboratory under a variety of

experimental conditions. In general, predatory species were provided

with other bromeliad organisms for Drey, and particulate and filter-

feeding organisms were provided with naturally occurring leaf litter,

detritus, and bromeliad water.

Preserved specimens of both la'val and adult forms were submitted

to appropriate systematic specialists for identification or description.

Representative collections fromrt this study have been deposited in the

Florida State Collection or Arthropods and at the Archboid Biological

Station.








Results and Discussion

The measurements of total volume, number of leaves, and volume per

leaf axil varied considerably between the samples of different bromeliad

species (Table 2).

Tillandsia utriculata is the largest tank bromeliad in Florida

with an average total plant volume of 300 cc with exceptionally large

specimens holding over 700 cc. The leaf axils of this species are

moderately inflated with an average volume of 8.1 cc (Fig. 3). In

T. fasciculata the leaf axils are much less inflated and have an average

volume of only 1.5 cc, but its large number of leaves provides an

average total plant volume of 60 cc. Both T. utriculata and T.

fasciculata maintained free water in their leaf axils throughout the

year at all sites.

The smallest species sampled was T. valenzuelana having a total

plant volume of only 35 cc. It is unable to maintain free water within

its leaf axils during much of the dry season (December to April), but

frequent rains during the wet season (May to November) provide sufficient

water for small populations of a few species of aquatic organisms.

Some specimens of C. berteroniana also become dry but this species

seems to retain water much longer than T. valenzuelana during dry weather

possibly because of a relatively large leaf-axil volume of 10 cc.

Catopsis floribunda and G. monostachia were sampled only during the

wet season and it is not known if they maintain aquatic invertebrate

populations throughout the year.

The distribution of bromeliad fauna among the 6 bromeliad species

(Table 3) indicates that the number of faunal species found in each









species of bromeliad is related to the frequency that each bromeliad

species was sampled. However, the less frequently samoled species

are relatively rare and restricted to certain plant communities, whereas

T. utriculata and T. fasciculata are widespread throughout south Florica

and occur in a variety of plant communities. The numbers of invertebrate

species recorded for C. floribunda (7) and G. monostachia (6), which

were sampled from just one site, approximate the average 7.6 species

occurring per sample site for all sampling locations. The faunal lists

of these rarer bromeliad species probably would not be increased signifi-

cantly by increasing the sample size since the additional samples would

be from the same or very similar sites. Catopsis floribunda and G.

monostachia have average total plant volumes and leaf-axil volumes

larger than T. fasciculata in which 12 invertebrate species were found,

indicating that the relatively small numbers of faunal species recorded

for these species are a result of their restricted distribution and

not their size. Catopsis berteroniana supported more invertebrate

species (9) since it is more widely distributed and was sampled from

3 different plant communities (Table 2). However, plant size as well

as a restricted distribution may be important factors in limiting the

number of invertebrate species inhabiting T. valenzuelana. This smallest

bromeliad was abundant in only 2 locations. The most important point

learned from sampling the rarer bromeliad species is that they did

not support faunal species different from those found in the more common

tank bromeliads.

Over 39,000 specimens representing 18 species of aquatic inverte-

brates were collected from bromeliad leaf axils during the study.

The list includes the immature stages of 12 species of insects









representing 7 families of Diptera, an oligochaet worm, a turbellarian,

a mite, and 2 species of ostracods. Most species were either abundant

in one or more sampling sites or widely distributed among several sites.

The rarest species was represented by 29 specimens occurring in 6 samples.

Single specimens of 3 different unidentified dipterous larvae, each

found in only one sample appeared to be accidental, and were excluded

from the study.

The following list of Florida bromeliad inhabitants includes data

on collection sites, estimates of relative abundance, and observations

on the biology of each species. A brief review of the literature relevant

to the species or related forms is also presented in an attempt to

clarify its status as a regular bromeliad inhabitant and to provide some

indication of its possible origin.


CULICIDAE:

Wyeomyia mitchelli (Theobald) -- Sites 1-12 and 17

Wyeomyia mitchelli is the most abundant and most widely distributed

mosquito in Florida bromeliads. The average number of larvae from all

of the bromeliad samples was 12.2; however, one large specimen of T.

utriculata contained 339 larvae. Larval densities varied among sites

but few sites were completely free of larvae, and populations persisted

throughout the year reaching their maximum abundance in midsummer.

Wyeomyia mitchelli larvae are free-swimming filter feeders and

consume particulate matter and microorganisms suspended in the leaf-axil

water. They have a development time of 2-4 weeks (21C) and pupation

lasts 4-5 days.








Bruijning (1959) has proposed synonymy of mitchelli and other

similar species with medioalbipes Lutz from Brazil, which has been

retained by Stone (1969). However, Belkin et al. (1970) considers

mitchelli a distinct species and reports its distribution as Jamaica,

eastern Mexico, Cuba, Hispaniola, and south Florida. Obviously, mitchelli

is part of a large complex of Neotropical species morphologically similar

to medioalbipes.

In Jamaica, W. mitchelli larvae occur in bromeliads and also in

the leaf axils of terrestrial Araceae and in the flower bracts of Heliconia

spp. (Musaceae) (Belkin et al. 1970). However, in Florida, this species

has been collected only from bromeliads including exotic terrestrial

species such as Bilberqia sp., a common garden ornamental (Fish,unpublished

data).


Wyeomyia vanduzeei D.&K. -- Sites 1-3, 6-8, 12-14 and 17

Wyeomyia vanduzeei, a more distinct species, occurs in south Florida,

Cuba, Grand Cayman, and Jamaica (Belkin et al. 1970). The larvae have

been collected only from bromeliads throughout its range. The feeding

habits of W. vanduzeei are similar to those of W. mitchelli. Although

both species coinhabit the same bromeliads and are frequently found

together in the same leaf axils, there is no evidence of competitive

displacement between these 2 species. Wyeomyia mitchelli outnumbers

W. vanduzeei by 3 to 2 in the total from all samples, but in the mangrove

sites W. vanduzeei was more abundant and in other sites the more abundant

species varied seasonally.

Wyeomyia is a large genus of plant-axil-breeding mosquitoes com-

prised of at least 85 species (Lane 1953). Although 2 species, W.

smithii and W. hanei, have adapted to pitcher plants in temperate and








boreal North America, the genus is otherwise restricted to the American

tropics.

In Florida, Wyeomyia mosquitoes are not known to transmit disease,

but they are avid daytime biters of man and are a problem in some areas

when adult populations are high.


Toxorhynchites rutilus rutilus (Coq.) -- Sites 1, 2 and 17

The larvae of T. r. rutilus are predatory upon other mosquito

larvae but were rare inhabitants of bromeliads. The few specimens

found were restricted to the largest bromeliad T. utriculata possibly

because plants having smaller leaf-axil volumes do not contain enough

Wyeomyia larvae to support their development. They are not known to

have the ability to leave the water of one leaf axil in search of another

containing prey. Seabrook and Duffey (1946) reported, finding numerous

T. r. rutilus larvae in T. utriculata in several locations along the

east coast of Florida. This species is not restricted to bromeliads

and is also found in tree holes and artificial containers (Basham et

al. 1947). Toxorhynchites r. rutilus occurs throughout Florida and

ranges northward into South Carolina and Georgia where it is replaced

by T. r. septenioralis which is distributed throughout the Eastern

United States (Carpenter and LaCasse 1955).

At least 6 other species of Toxorhynchites have been collected

from bromeliads in Central and South America (Horsfall 1972). Picado

(1913) found T. superbus (D.MK) to be very abundant in Costa Rican

bromeliads, but Laessle (1961) did not find this genus in Jamaican

bromeliads. The specificity of Neotropical Toxorhynchites for the

bromeliad habitat remains to be determined.









Toxorhynchites r. rutilus is not considered to be an important

predator of Wyeomyia mosquitoes since it is not widely distributed

in Florida bromeliads.

Other mosquito genera occur in bromeliads with varying degrees

of specificity. Notable among those are Anopheles mosquitoes in the

subgenus Kertesia which are obligatory bromeliad inhabitants and are

known vectors of human malaria in many areas of South America (Forattini

1962). Picado (1913) reported 65 mosquito species in 5 genera occurring

in bromeliads in Costa Rica, and Laessle (1961) found 7 species of

3 genera in Jamaican bromeliads. Miller (1971) found mosquitoes to

be the most abundant insects in the bromeliads of St. John, Virgin

Island, but did not provide identifications. Mosquitoes are probably

the most important bromeliad inhabitants throughout the Neotropics

because of their diversity and abundance, as well as their public health

significance. They are the second most abundant insect family inhabiting

Florida tank bromeliads.


CHIRONOMIDAE:

Metriocnemus abdominoflavatus Picado -- Sites 1, 2, 4-10, 12-15 and 17

Metriocnemus is the most abundant bromeliad inhabitant and is

found in all species of Florida tank hromeliads. The overall density

was 18 larvae per plant for all plants sampled with a maximum of 183

in one large specimen of T. utriculata. The larvae of this species

do not build cases, as do many fresh-water chironomids, but instead

crawl freely about the settled detritus particles upon which they pre-

sumably feed. The larvae are usually most numerous in bromeliads inhab-

iting dry exposed sites and least abundant in those inhabiting shaded


hammocks.









Picado (1913) found M. abdominoflavatus to be the most abundant

chironomid in Tillandsia spp. in the central highlands of Costa Rica,

but rare in Bilbergia and Catopsis. Miller (1971) reported Metriocnemus

"possibly abdominoflavatus" to be more abundant in T. utriculata in

the drier areas of St. John than in Aechmea ligulata of the wet mountain-

ous areas. Picado (1913) also reported that these larvae can resist

desiccation for several days which suggests that this species may be

adapted to bromeliads inhabiting relatively dry environments.


Monopelopia tillandsia Beck & Beck -- Sites 2, 8 and 17

Monopelopia tillandsia was found only in east-central Florida

and only during late winter and scoring. The large orange colored larvae

are free swimming and predatory upon the larvae of an unidentified

Tanytarsini (Chironomidae). They move rapidly with a wild undulating

motion when disturbed. The pupae are motile but remain in the water.

Under laboratory conditions these larvae were unable to capture each

other or any organisms larger than themselves.

Monopelopia tillandsia was originally found by Beck and Beck (1966)

in T. utriculata in Florida. Both Laessle (1961) and Miller (1971)

reported unidentified predatory larvae in the closely related genus

Pentanura occurring in bromeliads. Laessle (1961) found them in all

major collecting areas in Jamaica, but Miller found them to be most

abundant in the wet mountains on St. John. Picado (1913) described

the orange colored larva of Isoplastus (=Ablabesmyia) costarricensis

Picado a Pentaneurini from Costa Rican bromeliads, but its relationship

to M. tillandsia cannot be ascertained because of the limited description

(Beck and Beck 1966).









Tanytarsini (unidentified) -- Sites 2, 8 and 17

This third chironomid species could not be precisely identified

because of the systematic disorder of this tribe. It occurs only in

T. utriculata and almost always in association with its predator, M.

tillandsia. It is much more numerous than M. tillandsia. More than

1,000 larvae were found in one sample with most samples containing

over 100.

The smaller bright red larvae of Tanytarsini strongly adhere to

leaf litter and other large particles in the leaf-axil water and appear

to graze upon growths of microorganisms. They construct cases out

of fecal material and detritus particles that provide protection from

their aggressive predator. Under laboratory conditions, Tanytarsini

removed from their cases were immediately consumed by M. tillandsia

without fail.

Miller (1971) reported an unidentified Tanytarsus sp. (Tanytarsiri)

as being the most abundant chironomid on St. John, inhabiting both

T. utriculata and A. lingulata. Picado (1913) also found an unidentified

case-bearing chironomid in Aechmea spp. in Costa Rica.

When Tanytarsini and M. tillandsia are numerous in bromeliads

other fauna are much reduced or even absent. The red pigmentation

of these chironomids indicates a hemoglobin-oxygen transport system

which enables the larvae to survive at low dissolved oxygen levels

(Walshe 1950). They are usually found in plants that contain large

amounts of leaf litter and other organic debris in their leaf axils,

which may increase the associated microbial populations and subsequent

oxygen demand to the exclusion of much of the normal aquatic fauna

intolerant to low oxygen levels.









Chironomids seem to be major components of bromeliad communities.

Miller (1971) found them to be third in abundance in bromeliads on St.

John, and Smart (1938) lists chironomids as second in abundance in the

large terrestrial tank bromeliad, Brocchinia micrantha, in Guyana. Both

Laessle (1961) and Picado (1913) report chironomids to be abundant and

widespread in their studies of bromeliad fauna. In Florida, this

family comprises one-fourth of the total fauna and is the overall most

abundant group found in the bromeliad habitat.


PSYCHODIDAE:

Neurosystasis n. sp. -- Sites 1-12 and 17

This new species of psychodid fly is very common and usually quite

abundant in Florida tank bromeliads, averaging 16 larvae per plant from

all samples. The larvae are found adhering to the submerged leaf litter

on which they presumably feed. They are rather slow moving and frequently

rest with their posterior siphons exposed to the surface. The pupae are

motile and remain in the water.

This species is closely related to N. amplipenna (Knab) reared

from unspecified bromeliads in Cuba by Knab (1913a) who also reported

another psychodid Philosepedon fumata (Knab) from unspecified bromeliads

in Mexico. Other investigators have found unidentified and probably

undescribed psychodids from bromeliads. Laessle (1961) found them

in Vriesea sintenisci, Guzmania monostachia, and Hohenberoia sp. in

Jamaica. Miller (1971) reported them from T. utriculata and A. lingulata

on St. John where they represented only 1% of the total bromeliad fauna.

Psychodid larvae also occur in Aechmea nudicaulis in the Atlantic lowlands









in Costa Rica (Fish, unpublished data), as well as in unspecified

bromeliads in the central highlands (Picado 1913).


CERATOPOGONIDAE:

Forcipomyia seminole Wirth -- Sites 2, 3-8, 10, 12 and 17

The larvae of F. seminole are widely distributed among Florida tank

bromeliads, but never abundant. The average density from all plants

sampled was 0.2 larvae per plant, with the greatest density of 7 per

plant occurring in 2 samples of T. utriculata.

The spiny larvae resemble small caterpillars in appearance and are

usually found adhering to bromeliad leaves in a thin film just above the

leaf-axil water. When disturbed they quickly crawl into the water and

remain submerged indefinitely. Specimens reared to maturity in the

laboratory pupated on the sides of containers within 2 cm above the

water.

Forcipomyia seminole has only recently been described from light

trap collections near Vero Beach (site 17) and this report represents

the only records of its larval habitat. It is closely related to F.

pictoni, a widely distributed Neotropical species which breeds in rotting

cocoa pods in Costa Rica (Wirth 1976).


Forcipomyia (Warmkea) n. sp. -- Sites 1-7, 9-11, 15 and 17

This species of Forcipomyia is more common than F. seminole averaging

1.2 larvae per plant in all samples, with a maximum abundance of 16

larvae per plant. The small white larvae resemble chironomids in appear-

ance but are behaviorally different in that they do not freely swim in the

water but prefer to adhere to the sides of the containers as do F. seminole.








Forcipomyia is a very large genus of biting midges including both

temperate and tropical forms. The larvae are usually terrestrial or

semiaquatic and are found associated with decaying plant material such

as rotting logs, leaf litter, fruits, etc., but several species have

been reported from bromeliads and other water-holding plants (Wirth

and Stone 1971, Wirth 1974, 1975).

Other ceratopogonid qenera have also been reported from bromeliads.

Wirth and Blanton (1968, 1970) reported several species of Culicoides

from Guzmania sp. and unspecified bromeliads from Trinidad and Mexico,

and Laessle reported predatory Bezzia sp. from Hohenbergia so. in Jamaica.

Picado (1913) reported unidentified ceratopogonid larvae from unspecified

Costa Rican bromeliads and Smart (1938) reported them from Brocchinia

sp in Guyana. Miller (1971) found ceratopogonids to be abundant in

T. utriculata and A. lingulata on St. John, where they comprised over

32% of the total fauna.


SYRPHIDAE:

Meromacrus n. sp. -- Sites 2, 7 and 17

The larvae of Meromacrus n. sp. are relatively uncommon and found

only in large specimens of T. utriculata. Although they usually occur

singly in a plant, one sample yielded 19 first-instar larvae. Mature

larvae are the largest of the bromeliad inhabitants, measuring over 80

cm in length with their breathing tubes fully extended, which may explain

their absence from bromeliad species with small leaf-axil volumes. They

remain completely submerged in the leaf-axil water, feeding upon settled

organic matter with only their long breathing tubes reaching the surface

until just before pupation when they leave in search of a dry substrate.









This syrphid has previously been identified as M. ruficrus (Weidemann),

but is being described as new by Thompson (personal cuoii ,unication).

Although this is the first report of the larval habitat of Meromacrus

n. sp., its distribution records coincide perfectly with the range

of T. utriculata in Florida indicating that it may be restricted to

this habitat. With the exception of one temperate species, Meromacrus

is a Neotropical genus of primarily tree-hole breeding species.

From museum records, Thompson (personal communication) found that

6 syrphid genera have been reported from bromeliads, including Quichuana

and Leptomyia which appear to be restricted to this habitat, and another

species of Meromacrus from Brazil.

Picado (1913) reported Q. picadoi Knab as uncommon in Costa Rican

bromeliads. Various unidentified syrphid larvae represent a small

fraction of the total bromeliad fauna in the Virgin Islands (Miller

1971) and Guyana (Smart 1938). In the present study, 38 specimens

of Meromacrus n. sp. comprise less than 1% of the total bromeliad fauna

in south Florida.

AULACIGASTRIDAE:

Stenomicra n. sp. -- Sites 1-12, 15 and 17

This new species of Stenomicra represents a new family of bromeliad-

breeding insects. The small dorso-ventrally flattened larvae with

forked tails are predatory upon mosquitoes and possibly also chironomids.

They actively crawl upon the submerged leaf surfaces and among the

accumulated leaf litter in search of prey. Many specimens were reared

to maturity in the laboratory on Wyeomyia mosquito larvae. Development

is slow and the pupal stage lasts 19 days (range 18-20, N=5) at 210C.









Stenomicra n. sp. is never abundant, usually 1-5 per plant, but they

are widely distributed and occur in all species of tank bromeliads

in Florida.

Stenomicra is a pantropical genus with at least 12 species (Sabrosky

1975) and the larval habits are poorly known. In Hawaii, Swezey (1938)

reports S. orientalis (Malloch) larvae as predatory and occurring in

the water-filled leaf axils of Job's tear Coix lacrymajobi. This species

has subsequently been found in other water holding plants on the island

including screw pine (Pandanus sp.), sugar cane, and pineapple (Bromeliaceae)

(Williams 1939). Malloch (1927) also reported S. australis Mallock

from banana plants in Fiji and Sabrosky (1965) listed S. fascipennis

Mallock as being collected from screw pine in Guam. Unidentified Steromicra

larvae have been found in both bromeliad leaf-axils (Vriesea insignus

and G. monostachia) and those of elephant ear (Colocasia sp.) in Costa

Rica (Fish, unpublished data).

Although other published records indicate that Stenomicra is not

restricted to plant-held aquatic habitats, the larvae of these unusual

insects may not have been identified by previous workers because of

the difficulties involved in rearing, and may be widespread in bromeliads

and other similar habitats.


MUSCIDAE:

Neodexiopsis n. sp. -- Sites 1-3, 6-R, 11 and 17

The maggot-like larvae of Neodexiopsis n. sp. usually occur singly

within an entire plant sample. They are predatory with piercing mouthparts

and were reared to maturity on Wyeomyia mosquito larvae. The pupal

stage is long, averaging 18 days at 210C (range 16-19, N=5). Spiracles









on their blunt posterior ends enable these larvae to leave the water

of one leaf axil to search in others for prey. Such behavior was noted

in the laboratory when several larvae escaped from their containers

after consuming their prey.

Neodexiopsis is a large Neotropical qenus containing 65 species

(Huckett, personal communication), however, little is known of the

biology of the immature stages. Picado (1913) reported an unidentified

predatory muscoid larva from bromeliads (Aechmea sp.) in Costa Rica

which he placed in the genus Coenosia, a qenus from which Neodexiopsis

has been recently split (Snyder 1958). Synder (1958) observed that

collecting in habitats with moist soil often yields general adults

of Neodexiopsis spp. which indicates that the immature stages of other

species are also aquatic but that the genus is not restricted to bromeliads

of other plant-axil breeding sites.

Because they are relatively uncommon and rather difficult to rear

Neodexiopsis larvae may not have been identified by previous investi-

gators and may be more widely distributed in bromeliads than the litera-

ture indicates.


SCIARIDAE:

Corynoptera n. sp. -- Sites 2-5, 12 and 17

The occurrence of Corynoptera n. sp. in Florida tank bromeliads

represents a second new family of bromeliad-inhabiting insects. The

larvae of this fungus gnat are only moderately distributed and are

never very abundant. The average density in all plants sampled was

0.4 larvae per plant with a maximum of 13 found in a single sample.

They are easily mistaken for F. (Warmkea) n. sp. (Ceratopogonidae)

as the larval forms are very similar in size and color.









Corynoptera n. sp. presumably feeds upon fungi growing on the

decaying leaf litter accumulated in the bromeliad leaf axils as do the

larval forms of most other sciarid flies (Borror and DeLong 1971).


ACARI:

Anoetus n. sp. -- Sites 1, 2, 4-12 and 17

This new species of aquatic mite occurred sporadically among the

sites investigated, but when present, it appeared in most or all of the

plants sampled with an average density of 9 per plant. These small

white mites are found in various stages and are difficult to detect

when few in number. They attach to small particles of organic matter

and tend to hide in hollow twigs and folds of leaves.

Picado (1913) reports an unidentified aquatic mite in the genus

Tyroglyphus found on only one occasion in an unspecified bromeliad

species in Costa Rica. Many species formerly in this genus are now

placed in the family Anoetidae. It is quite possible that what Picado

actually found in Costa Rica was Anoetus. However, he provided no

description or illustration to support this assumption.

Anoetidae is a large family of mostly aquatic and semiaquatic

mites commonly found associated with dead organic matter (Krantz, 1970).

Two species of Anoetus occur in temperate pitcher plants; Anoetus

gibsoni (Nesbitt) in Sarracenia purpurea L. and A. hughsi Hunter and

Hunter in S. flava L. A terrestrial species of Anoetus has been shown

to feed on bacteria (Noble and Poe 1972) and Hunter and Hunter (1964)

suggest that A. qibsoni feeds on bacteria associated with decomposing

insects in the pitcher plant. Anoetus n. sp. probably has similar

feeding habits in bromeliads.









Anoetid mites have a resistant hypopal stage and are transported

between favorable habitats by attaching themselves to insects (Hughs and

Jackson 1958). Hunter and Hunter (1964) suggest that the pitcher plant

mosquito Wyeomyia smithii transports A. qibsoni among the leaves of

pitcher plants. Several insects could serve this function in bromeliads

in addition to Wyeomyia mosquitoes.


OSTROCODA:

Metacypris maracaoensis (Tressler) -- Sites 3, 5 and 13

This ostracod is abundant in T. fasiculata in the tropical hardwood

hammocks of Everglades National Park, averaging over 100 per plant. It

is less common in C. floribunda and G. monostachia and rarely found in

T. utriculata.

This species was originally described from an unidentified bromeliad

in the Big Cypress Swamp of south Florida and had since been reported

from unspecified bromeliads in Puerto Rico (Tressler 1956, Maguire

1970). Laessle (1961) found 3 species of ostracods to be abundant in

Jamaican bromeliads; M. laesslei (Tressler), M. bromeliarum (Muller),

and Candonopsis anisitsi (Daday). Other ostracods reported from bromeliads

are C. kingsleii in Puerto Rico (Tressler 1941) and 2 undescribed species

from Costa Rica (Picado 1913).

Although distribution data for all of these species are fragmentary,

M. bromelarium, M. maracaoensis, and M. laesslei are presently known

only from bromeliads. Their method of transport among these relatively

isolated aquatic habitats is unknown.








Podocopa (unidentified sp.) -- Sites 3, 9 and 13

An unidentified ostracod frequently occurs with M. maracaoensis in

the bromeliads of the tropical hardwood hammocks. Since only juvenile forms

were found, identification was impossible.


OLIGOCHAETA:

Naididae (unidentified) -- Sites 1-5, 6-10, 12 and 17

Unidentified oligochaet worms are widely distributed among Florida

tank bromeliads with an average density of 10 per plant for all samoles.

They are most numerous in the oak hammocks and cypress swamps where

their densities average nearly 50 per plant in T. utriculata, but are

rare or absent in tropical hardwood hammocks.

These worms do not form tubes, as do many fresh water forms, and

are found buried in the water-saturated detritus of the older leaf axils.

Like the chironomids M. tillandsia and Tanytarsini, these olipochaet

worms are red, suggesting a hemoglobin-oxygen transport system which

would enable them to survive under near-anaerobic conditions.

Picado (1913) reports Aulphorus superterrenus Michlsn. (Naididae)

as being very abundant in Vriesea sp. in Costa Rica, and Laessle (1961)

reports similar forms in unspecified Jamaican bromeliads.

Very little is known of the ecology or systematics of bromeliad-

inhabiting oligochaet worms, and it cannot be determined from the

literature if certain groups or species are specific for this habitat.


TURBELLARIA:

(Unidentified)

An unidentified rhabdocoel flatworm is the rarest inhabitant of

tank bromeliads in south Florida. One to 5 specimens were consistently









found in samples of 10 T. utriculata from the mangrove swamp in Everglades

National Park throughout the year, but were only occasionally found at

other sites.

Picado (1913) found Geoplana picadoi Beauchamp, Rhynchodemus

bromelicola Beauchamp, R. costarricensis Beauchamp and Geocentrophora

metameroides (Beauchamp) in Costa Rican bromeliads and Laessle (1961) found

G. metameroides (Beauchamp) and G. applanata (Kennel) in Jamaican bro-

meliads. Although flatworms seem to be common bromeliad inhabitants in

the Neotropics, very little is known of their systematics or ecology.

Many turbellarians feed on small aquatic organisms which they trap

in mucus secretions (Pennak 1953) and some have been observed to be

predatory upon the eggs and larvae of mosquitoes (Jenkins 1964, Medved

and Legner 1974). Because of the small size of the species occurring in

Florida tank bromeliads, it would probably prey only upon the egg stages

of other bromeliad fauna.

It is difficult to compare the species composition of the aquatic

fauna inhabiting tank bromeliads in south Florida with what has been

reported from bromeliads in the Neotropics. Only the studies of Picado

(1913) in Costa Rica and Laessle (1961) in Jamaica attempt to list all

of the organisms that were found inhabiting bromeliads; but Picado

(1913) also lists terrestrial species which frequently leads to confusion

in comparing only the aquatic fauna. Interpretation of his results is

further complicated by the name changes that have occurred at various

taxonomic levels within the past 60 years which cause serious problems

in updating his list with valid species ndmes. Also, many important

species are not identified in Picado's study and several insect families

are mentioned only by name.


~









Laessle's (1961) study provides more recent data, but unfortunately,

only half of the organisms are identified to species. Maquire (1970)

identified only the ostracods in his study of Puerto Rican bromeliads.

Miller (1971) identified only the chironomid genera that inhabit the

bromeliads of St. Johns. Many other scattered reports on aquatic broneliad

fauna include identification of no more than a few species of special

interest to the investigators with only passing mention of the other

fauna present.

Precise identifications to the species level are sometimes difficult

to obtain since most of the organisms are in immature stages and must be

reared to adult usually without any prior knowledge of their feeding

habits or pupation requirements. Also the systematics of many major

groups, including mosquitoes, are badly in need of revision and willing

capable specialists for some of the minor groups are difficult to

locate.

However, precise species determination of all inhabiting organisms

is essential in studying the origin, evolution, and biogeography of

bromeliad-inhabiting invertebrate communities, and in assessing the

specificity of the fauna for the bromeliad habitat.


Conclusion

The aquatic invertebrate fauna inhabiting the leaf axils of Florida

tank bromeliads is composed of 18 species, most of which are abundant

and widely distributed in several bromeliad species. Fourteen inverte-

brate species have been positively identified, including 7 new species,

and 4 species remain unidentified pending the actions of systematic

specialists and in some cases the acquisition of additional specimens.









Large invertebrate species such as the predatory mosquito T. r.

rutilus and the syrphid fly Meromacrus n. sp. seem to be restricted to

the largest bromeliad species T. utriculata, but the size and shape of

the bromeliad species seems to be less important than its distribution

in determining the species composition of the aquatic inhabitants.

Tillandsia utriculata and T. fasiculata are widely distributed throughout

south Florida and, although quite different in structure and total plant

volume, support the largest numbers of invertebrate species. Other

bromeliad species are more restricted to certain plant communities and

support fewer inhabitant species.

While direct comparison between the Florida bromeliad fauna and the

bromeliad fauna reported from other areas cannot be made to any great

extent at the species level, certain conclusions can be drawn from the

systematic information that is presently available in the literature.

It is apparent from comparing the results of the present study with

those of Picado (1913) and Laessle (1961) that the invertebrate fauna

inhabiting tank bromeliads in south Florida is relatively depauperate.

Aquatic insects are represented by the single order Diptera in Florida

bromeliads while Picado (1913) lists the insect orders Odonata, Hemiptera,

Coleoptera, and Plecoptera as well as Diptera from Costa Rican bromeliads.

In the Diptera he also lists the families Stratiomyidae, Tabanidae,

Tipulidae, Anisopidae, and Borboridae which are not present in Florida

bromeliads. Picado's total list of aquatic insects include well over

100 species as compared to 13 in Florida bromeliads.

Laessle (1961), in studying the island bromeliad fauna of Jamaica,

reports 3 insect orders and 2 families of Diptera not found in Florida








bromeliads with a total of 33 species of aquatic insects. Both Picado

(1913) and Laessle (1961) also report additional species of oligochaet

worms, ostracods, turbellarians, and water mites not present in Florida

bromeliads.

Many theories could be proposed to account for the depauperate

nature of the Florida bromeliad fauna including those reviewed by Pianka

(1966) and Baker (1970) relating to increased species diversity in the

tropics. Also, in view of the insular nature of subtropical south

Florida, theories on island biogeography outlined by MacArthur and

Wilson (1967) might also be considered. However, at present too little

is known of the ecological roles of the various bromeliad inhabitants or

of the evolutionary relationship between the bromeliad fauna of different

areas to advance or discredit any of these theories.

There is considerable evidence suggesting that epiphytic tank

bromeliads in south Florida support an aquatic fauna derived for the

most part from Neotropical ancestors specifically adapted to the bromeliad

habitat. Much of the aquatic fauna inhabiting Florida tank bromeliads

is systematically related to Neotropical bromeliad fauna reported in the

literature. The taxonomic levels of relatedness include 4 species at

the species level, 6 species at the generic level, and 13 species at the

family level. It is very probable that a closer systematic relationship

would become apparent if more species determinations were available for

the Neotropical fauna.

The aquatic stages of the Neotropical bromeliad fauna are reported

to be exclusively restricted to the bromeliad habitat by Calvert (1911),

Knab and Malloch (1912), Champion (1913), Knab (1913b), and Picado









(1913). Specialized morphological adaptations are recognized in bromeliad-

inhabiting dragonflies (Calvert and Calvert 1917), crane flies (Alexander

1912), and syrphid flies (Knab 1913c). Although gross morphological

adaptations were not observed in any of the Florida bromeliad fauna, it

is evident from the existing distribution records that most species

exhibit a degree of habitat specificity that is characteristic of Neo-

tropical bromeliad fauna.

Only one identified species, T. r. rutilus, is known to occuoy

other habitats because of its high incidence in tree holes as reported

by Basham et al. (1947) and its low incidence in bromeliads as reported

in the present study, this Dredatory mosquito may be considered an

opportunistic colonizer of bromeliads in south Florida. Aquatic stages

of the remaining 13 identified species have been found only in bromeliads,

including 8 species which have been found only during the course of this

study. Since most of these species are widely distributed and frequently

abundant in bromeliads, it seems unlikely that they would also occupy

alternative aquatic habitats to any great extent and totally escape

notice. With the exception of T. r. rutilus, there is no evidence from

this study that subtropical or temperate species from other aquatic

habitats have adapted to bromeliads in south Florida.

The presence of a diverse aquatic fauna restricted to the widespread

but discrete habitat of epiphytic bromeliads provides a unique opportunity

to study basic concepts of community ecology such as interspecific

interactions, energy flow, and species of diversity as well as the

evolution of communities. On a large scale these aquatic communities

might be useful in experimental studies of island biogeography in areas


I





35


such as the Caribbean Islands and in investigations into the causes of

latitudinal gradients in species diversity in continental areas.

Hopefully this study will generate renewed interest in the aquatic

communities inhabiting epiphytic bromeliads and foster increased coopera-

tion between field workers and systematicists in establishing a sound

systematic basis for future investigations concerning the many evolu-

tionary and ecological aspects of these unusual aquatic communities.















CHAPTER III
FACTORS INFLUENCING THE STRUCTURE OF AQUATIC COMMUNITIES
INHABITING EPIPHYTIC BROMELIADS IN SOUTH FLORIDA


Introduction

Communities of aquatic organisms that inhabit the leaf axils of

water-holding tank bromeliads are important components of Neotropical

ecosystems. The many epiphytic bromeliad species provide a unique

aquatic habitat, elevated and discontinuous from surface waters, and

increase the overall spacial heterogeneity of available aquatic habitats

for a given land area.

The total water volume contained in these arboreal habitats can

be considerable. Hazen (1966) reported densities of 4 bromeliads per

meter length of tree branch for water-holding Guzmania sp. at a site

in Costa Rica. Mature specimens of G. monostachia hold an average of

180 ml of free water (Fish, unpublished data), and considering a con-

servative 15% of the total bromeliad population as being mature, the

total epiphytic free water for this site is estimated to be over 100

ml per meter length of tree branch. Many larger epiphytic bromeliad

species hold vast amounts of water with capacities ranging from 2 liters

(Laessle 1961) to over 20 liters (Picado 1913). It would not take many

of these large bromeliads to approximate the water volume of a small

terrestrial pond.

Picado (1913) first recognized the magnitude of the total

aquatic habitat provided by epiphytic bromeliads by equating








them as a whole to a large fractionated swamp extending throughout

tropical America.

The aquatic fauna inhabiting epiphytic tank bromeliads is exten-

sive. Nearly all of the major groups of fresh water invertebrates

have been reported from this habitat including 6 orders of aquatic

insects, snails (Gastropoda), crabs (Decapoda) as well as tadpoles

(Anuara) (Calvert 1911, Picado 1913, Laessle 1961). Complete faunal

lists are rare but Picado (1913) reported over 130 species of aquatic

invertebrates from Costa Rican bromeliads, and Laessle (1961) reported

over 60 species from Jamaican bromeliads.

Early investigators of the bromeliad fauna noted that nearly all

aquatic species are restricted to this habitat and are distinct from

allied forms found in other fresh water environments (Calvert 1911,

Champion 1913, Picado 1913). These observations have been subse-

quently supported by more recent studies, (Laessle 1961, see also

Chapter II) although the exact degree of habitat specificity for all

bromeliad inhabiting organisms remains to be determined.

The arboreal aquatic habitat provided by epiphytic tank bromelaids

and its occupation by a diverse and unique aquatic fauna results in

overall increased animal species diversity per unit of land area for many

areas in the Neotropics. Previous investigations of the aquatic community

inhabiting bromeliads have been primarily descriptive and little is known

of its structure and dynamics or of factors that influence species com-

position. However, Laessle (1961) investigated some of the physical

and chemical properties of bromeliad water in Jamaica and found

that bromeliads growing in full sunlight supported an alqae-based









food chain whereas these growing in shaded situations supported a

detritus-based food chain.

The results of a survey of the aquatic fauna associated with

epiphyLic tank bro:niliads in south Florida are presented in Chapter II.

This study reports 18 species of aquatic invertebrates inhabiting 6

species of tank bromeliads sampled from 17 site locations. However,

all 18 species were rievr round in a single site, and the average number

of species found per site was only 7.6. Samoles were taken from different

species of bromeliads, at different tires of the year, and from a variety

of majcr ecosystems occurring in south Florida. Any or all of these

factors Imry influence the structure and composition of the aquatic

community inhabiting the bromeliads. Therefore a special sampling

program was initiated to determine the effects upon the community strLucture

(in terms of species composition and abundance) of 1) the bromeliad species

providing the aquatic habiLat, 2) seasonal changes, and 3) the ecosyst-em

supporting the brT~aied flora.

The subtropical climate of south Florida represents the northern

limits for 6 species of epiphytic tank bromeliads which are also widely

distributed throughout the Caribbean Islands and Central and South Americp.

Most of the aquatic invertebrate fauna inhabiting these bromeliads are

also of tropical origin, including several species that are endemic to

south florida ei:e Chapter II'. However, Pcosystems occurring in souT.h

Florida we of both temperate (cypress swamps) and tropical origin

(mangr-ove swamps and tropical hardwood harrmmocks). Despite this

mixture of temperate and tropical ecosystems, most forested areas in

s:c-th F'1oriid supci'ort poi.l',tiorns of epiphytic tank bromeliads and the










factors in Fluencing the structure of aiquatic communities inl.hbitinqj

brcielicads in subtropical snuth Florida m[ay be operative throughout

the Neotrcpics as well.

Collecting Sites

Epiphytic tank bromeliads occur in a variety of Florida ecosyst.em-

and the species composition of epiphyte communities varies among them.

Iwo specie, of tank broimeliads, T Tllarnsi utriculata and F. msciculeta,

art partii-uariy abundant and widespread in south Fio oria and exhibit

the least site preference of any of the tank hromeliads. Both species

have bec!! shown to support a lerce and diverse aquatic fauna inh.bItirg

their leaf axils (see Chapter i). The presence of one or both of

these srec;es in separate and distinct ecosystems were the major criti;-ia

for site selection.

Three study sites were established that met these criteria: 1) a

cypress swamp supporting both T. utriculata and T. fasciculata, 2)

a mangrove swanp supporting T. 'triculta, aind 3) a tropical iard wood

hammock supporting T. fasciculata. In this manner the effects of the

bro,,eliiad species upon the aquatic community structure could be determined

at the cypress site, and the effects of the ecosystem could be determined

by co.~:aring the community structure of each bromeliad species at the

cypress site with either the mangrove site or the tropical hardwoods

site.




This site is located in the Fisheating Creek W'ildlife Management

Area, Glades County, which contains a large stand of bald cypress

L-xuJil Jis't-ch'11 extending over 40 km along the Fisheatiig Creek
f3iood p Iin. SPr!pl ing w;as restricted to a 1/2-ha area adjacent to










the strLear bed about 2 km west of the ismin campground. This area was

chosen because of its large population of both T. utriculata and T.

fas"icula '.J. tstimatcs of their ide-issities ranged from 2 to ?1 specimens

per tree (average 6.,) with T. fasciculeta outnumbering T. utriculata by

apprcxiirately I to 1. Other broaeliads ate also abundant at this site,

especially T. ;0 libsiana and T'. !sneoid'Ls which do not hold water.


Minoruove _Sw!rr_ Site

Ihis site is located adjiacnt to the Flaningo road near Sn~aKe

Bight Trail in Evcirgladi~e Nlational Park. White rmngrove Laquncularia

raccmose ;l.d buttonwood Conocarous ere Ict occur in a nearly continuous

strand alo ng the enban!kment of the road in this area as well as in small

clu ips distributed throughout the surrounding batis marsh. Tillanusia

utri culatl is aburn.dt along several km on both sides of the road and

in the many t'ee clunps in the batis marsh. However, sampling was

restricted to an approximate 100 ,m length of the roadside strand where

epiphyte ; ensities are the highest, and T. utriculata occurs at an

average density. of 1.5 piar .s nai tre.:e. Other epiphytes are co;minio'

and include T. fasciculata, T. flexuosa, T. pruinosa, and orchids

(E _i _r,.r; ssp ).


Trc.nica! liar,'dood Hammock Si4e

Cl pop's !Ha-:o.k is located 1.5 km north of the Missile Base

roA'd in r Everglades Natione l P.r.. It is tynical of the 125 or more

tropical hardw:oid hi3irclk~ n l1 !i: pinielands area of the Park that de-

veop on sliu'htly higher elevations of limestone rock (Craighead 1971).

Ci-p:' ii H::,.K is relati ely asmnll (1/2 ha) but supports a late









poipultiol of T. fasjccuaia at d ensities of approximately 3 plants

per tree and is less than 200 m from a much larger hammock supporting

an equivalen'1 t epiphytic flore.

Tropical ha;-d..ood hammocks of this region support a mixture of

tropical and temperate trees such a.; live oak Quercus virlnijuina, Cgubo

limbo Bursea siimaruba, poiso',/o'jd I'er:topium toxiferuw,, mahogany S' itenia

rmahoqani, an'd dove plum Coccoluob diversifolia. Other epiphytes are

abundant at:d include T. valenzulana, T. setacae, and _pidendrum spp.

Sampipling Prgram

Epiphytic tank bromeliads proviJe natural sampling units and a

series of plants provide replicate samples of the aquatic community

inhabiting the brci.naiads in a particular site. Each collection con-

sisted of 10 specimens eacn of 1. utriculata and 10 of T. f.scicuelaa

from tli-: cypress ritc and iu of T. utriculata from the mangrove site.

Only 5 T. fasciculate were sampled or each occasion from the tropical

hardwoods site because of the possible adverse effects that plant ;e:,oval

might have upon; the tutal brom;eliad fauna occurring in this relatively

small area.

All 3 siLes vierc visited within 2 weeks at 3--month intervals froi;

ept:br, ~7 i13 to Jiune 1975, so seasonal charges in cormmunity structure

could hi d-etrmined thr: ghout one coiiplete year.

Samping jr; ceaiueres and '-he processing of invertebrate fauna are

descr;rib in d:tai3 in Chapter II and will only be summarized here.

Bromnelid: coKlr; :rt te sampled in a truly random fashion but were

coec..tl ee n c' thr'ouhiut ji A .ch sit usi ig rree-c imbing spurs when

necessary t(; Cibl ii si .ciiu cs above reach from the ground. The plants









were r' t''., 'td f'ro, their atte .;hn,! t sites and the aquatic fauna contained

in e-ch was washed from Lhe Nlef axils. Each sample was kept in a

separate labelled plastic hag during storage and transportation, in

the laboratory ali aquatic organisms found in each bromerlia sample were

iderntifitd and the num:hers of individuals ir, each soeLcis counted to

determine the community structure represented in each sample. Protozoa

and A;Ciel;iithes (rotifei s and nnematodes) were not efficiently samnpli'

by the niethods employed and were consequerlntly eliminated frcm the study.

The replicate sani-ples from each bromelaid species and each site were

averaged to represent a total community structure for each collecting

occasion.

AnayIti cal Met hods
Satisfactory and readily e 'ailable methods of comparing the structure

of biological coiTrunities have recen*Cy been developed as a result of

applying ecological oata to nrumierical analytical techniques introduced

by Sokal and S:;sl.h (1963). Nurerical methods are frequently employed

in con arative studies of plant c),nnuriities (Whittaker 1962, 1973;

ticlntoish 1967), but their application in the analysis of animal commurni-

ties s a been ho ls coiimon (Cliffoid and Stephenson 1975)

Numorical methods are particularly useful in defining csseimblaes

of organisms and rl-ating these asseamblage~ to environmental or biotic

factors, Stephenson et al. (1970, 1974) successfully used nunm rical

methods 4in rela-i0ng recu;'ring groups of marine benthic fauna to

diffe,-reces in physical characteristics of bottom sfrdiments in Mtlreto1

bay, Australia, and FoNar and MicCo am0 (1963) were able to associate

groups of zOcpln kton with differences in itei r mass types in thie Nor'th

Pacific. iUsinh similar methods, Kikkawa (l1 o) was able to demonstrate









that bird coimiunities of similar species composition in Eastern

Pustralia viere associated with structurally similar plant formations.

Because of their proven usefulness in determining species groups which

can be related to other factors, numerical methods were employed to

analyze the collection data from the bromeliad samples.

Numerical analysis of ecological communities involves 2 pro-

cedures. The first procedure is to measure similarities (or dis-

similarities) among collections of org;niisl;'s, and the secoEd is to

organize or classify the collections on the basis of their simila'rties

(or dissimilarities).

Methods of comparing the structures of biological communities

have only been recently adopted in ecological studies. Previously

species cormosition in terms of presence and absence data was the

only criteria used in measuring similarity between communities, and

measures such as Jaccard's Co-efficient (Jaccard 1908) and Sorensen's

Index (Sorensen 1948) were used to quantify comparisons (Odum 1950,

Kikkaa.a 1958, Roback et al. 1959). while e these measures are still

successfully used in plant e'.ui.jgy (Williams et al. 1970, Goodall 1973),

these ;Qeasur,:s iare freaueitly unsatisfactory in studies of animals

because they ignore e he els.tiv't ebjnda-ne of each species present anong

the communi" lies to be compai',d.

Some animals, and especially insects, are extremely ontile and the

presence of one individual ir, .a c(rm-;unity is not always ecologically

meaningful, but wol'. d carry the equivalenr- weight of hundreds or thou-

sands of individuals in tei is u' presence or absence data. Furthermore,

measures based on presence rnd c'ser.ce ditc from communities of vsry









similar species composition tut drastically different ir. abundanc.e 'oul:

provide little information as to their true likeness.

To lain the most information from the sampling efforts, both species

composition and abundance were used in the analysis of bromeliad communities

Euclidean distance was adopted as a dissimilarity measure using species

abundances as attributes (Clifford and Stephenson 1975). Distances

were calculated by the equation:

n )2 1/2
D- r ^ x x
jk =1 ^ k
where Xi -; the number of individuals in species i of co-munities j a"d k.

The community structure data wis :stndardized as a percent of

the total for each collection and reduced to exclude species that com-

prised less than 0.5% of the total fauna for all collections. Justi-

fication and details on dita reduction and standardization can be found

in Clifford and Stephenson (i975). The collection data ware arranged on

a 9 x 16 metrix (Table V) and euclidean distances were calculated among

the 16 collections in terms of the abundances of 9 species.

In the second procedure the community collections were arranged

in a hierarchical classification based on euclidean distances. Tnis

procedure is available in the preproogrammed Statistical Analysis

System (SAS) (Barr et a!. i976) which was employed in the analysis.

The cluster procedure performs a hierarchial cluster analysis based on

an algorithm outlined by Jchnson (1967). Clusters are formed using

the futheLst-ncighbor technique (Sneath and Sokal 1973). This SAS

progiamr is particularly convenient because it also computes euclidean

distance for the cluster analysis. The resulting dendogram (Figure 4)









sho'" the relationship bet,,een entities (cotirunities) based upon

attributes (species End abuiidance) in teris of dissimilarity.


Results and Discussion

Five of the 18 species of aquatic invertebrates known to occur in

Florida tank bironmeliads did not occur in any of the 3 sampling sites.

An additional 4 species were rare, each representing less than 0.5, of

the total fauna collected and were excluded from the analysis in the

data reduction process. A total of 9 species of aquatic organisms

were used in the final analysis (Table IV). The data matrix (Table V)

cf the mean abundance for each species in each of the 16 collections

represents e total cf over 13,000 organisms.

The cluster analysis of euclide.an distance dissinilarity ireas-

ures iaong the 16 collections (Figure 4) shows that the greatest

differences in community structure occurred among the 3 sites. The

sites did not cluster until a fusion level of 1.27 was reached, almost

twice the distance of the highest seasu.el clustering at 0.70. Seasonal

differences were greater in mangro"e arid cypress sites at fusion levels

of 0.69 and 0.71, respectively, than in the tropical hardwood site at

fusion level 0.22. The coiamuncity structures contained'in T. Utricul;ts

and T. f;scicualeiad show the least- difference among the 3 factors con-

sidered. iil!andsia utriculata fused with T. fasciculata in the spring,

su!:'i;ier, and fall at the 0.20 level before fusing with T. fasciculata

in thle :inter at 0.38.

It is evident from this analysis that the ecosystem supporting

the ,ra';e!iad flora has a greater effect upon the structure of the









aquatic community inhabiting the bromeliads than dees either season

or brromliad species. Although nicroclimate may have some effect upon

the aquatic community, the major Factor to be considered is the nutrient

inout to water contained in the bromeliads.

Epiphytic bromreliads inLtercept dissolved nutrients leached by rain

from the above forest csnopy as twell as allochthonous leaf litter,

which are iioi',unded within the inflated: leaf axils (TIukey 1970b, Benzi;,g

and RRei-fro,, 19?1). T;;s pojl of dilute nutrients and decomposing leaf

litter is ultimately util i. b .' the plant through foliar absorption

(Benzing 197L':,, :..: and Burt 1970), but these substances also

serve as nutric.:t sou:'-c for the inhabiting aquatic fauna. Nutrients

leached from forest canopies ary both quantitatively and qualitatively

among both tepcrate aid tropical' ecci.sY.sleis (Tukey 1970a, b, Bnrnhard-

Reverst '1975) and impeiundments of these nutrients within bromeliad

lea axils will certainly reflect these differences. Likewise, both

the quanrtity ard quality of leaf litter entering tank bromeliads will

affect the chemical and nutrient composition of the leaf-axil water.

The amount and rate of leaf litter accumulation will vary among dif-

ferent ecusy'tems depending upon tree species composition and

especially between deciduous and everpraen forests. Also, the material

leached from the decomposing leaf litter 'ill very among oacsystems

depending upon tree species composition (Nykvist 1952).

Although canopy leachates (throughfal ) and leaf litter wer:e

nct measured during this study, it can be ass;u' ed that significant

diffe!r'.e!ces in bcth the quantity and quality of these mn'teria's reist

ailing thei 3 different ecosystemssttudnr d.









Differences in structure of the ac6Citic community inhabitirc; bro-

medliads are likely the result of difFerences in nutrient input. All of

the aquatic fauna inhabiting tank bromeliads in south Florida seem to be

detritus feeders (see Chapter 11). Dissolved nutrients from both

throughfall and leaf litter decomposition provide a substrate for the

growth of bacteria aid protozoa (Parsons and Seki 1970, and Slater 1954),

which are in turn food sources for mosquito larvae and ostracods (Yonge

1928 and Cleients 1969). Particulate leaf litter also serves as a

substrate for microbi~ growths which are in turn grazed by other

aquatic organimns (Kaushik ac. Hynes 1968), although many aquatic

insects feed directly upon particulate leaf litter (Cummnins 1973).

The chemical and nutrient composition of streams in both temperate

and tropical ecosystems are dlso influenced by throughfali and leaf

litter from surrounding forests (McCall 1970, Reichle 1975, Soil 1975)

which directly affect the structures of inhabiting invertebrate

comnuniities (Nelson and Scott 1962, Minshall 1957). In this respect

aquatic communities of brcTiolids and those of streams are quite

similar in trophic structure and energy flow because both communities

are treophically dependent uporn 0on outside source of detritus. However,

the aquatic cor:munities inhabiting bromeiiads are likely to be much

more sensitive to changes in ecosystems than stream communities because

of their closer proximity to the nutrient source.

The cluster analysis also shows differences in the structure of

the aqumctic community inhabiting bromeliads at different seasons in

both th mangrove aid cypress sites. These seasonal changes may he

due to seasonal changes in nutrient input. Cypress trees are deciduous









and provide a massive input -i leaf litter in the fall months (Carter

et a!. 173) and thereafter throughfall nutrients would. be unavaileble

to the brorleliads un+il tre spi ing growth provides new leaves. 'lan-

grove ecosystems have a period of maximum leaf litter production

but also maintain leaves throughout the year. Tropical hardwood hammocks

are predominat.ely evergreen and pr3djce relatively constant inputs of

both throughfall nd leaf litter wiich may explain the greater seasonal

similarities amonGg the. conmuitiy structure of brcmelirds inhabiting the

ecosystem.

It is difficult to separate the climatic factors from the bio-

logical characteristics of each ecosystem in explaining the seasonal

variation in comi:nunity structure. Rainfall and temperature obviously

have some effect upon the aquatic organisms because of the pronounced

cool dry season (November to April) and the warm wet season (May to

October) in south Florida. Seasonal variation in rainfall would

influence the rates of throuphfall input as well as the total water

volumes available to the aquatic organismis. I!Wter volumes were not.

measured at tlt time of collection, but water was observed to be perm:a-

nontly ,-eintcined in the bromeliad leaf axils at all sites throughout

thi year.

Seasonal variation in temperature may also affect the aquatic

fauna directly. Low; temperatures will prolong the development times

of the aquatic stages, and light frosts may kill a certain proportion

of the Idult iisect populations resulting in a temporary population

decline. Marked seasonal variation i! the number of adult bromeliad-

bred c i'1,^ mosquitoes was observed at the cypress site with maximum










ad.ilt popal.'tioat appoaririn in iidsumimer. PAults were rare in the

winter months a!tnough larval populations were still high, and the area

had beet siuih ;e-ted to a series of frosts during this time. However,

mosquito larval populations completely disappeared from bromeliads

at the rangrove site during the winter with no frost having occurred

in the area during ithe entire win.ler.

Tropical hardwood hamrrr'cks tend to buffer temperature extremes

ard are warier in the winter nrid cooler in the sumner than the sur-

rounding area (Craighead 1971) which may also have contributed to the

increased stability observed in the aquatic community inhabiting the

bromeliads irn this site.

The species if bromeliads supporting the aquatic community seems

to hav'e less influe,]ce upon community structure than either seasons or

ecosystemis. This is indicated by fusion of the T. utriculata corn-

munity with that of T. fasciculata at a lower level than that at

which ;he seasonal fusions occurred. The 2 bromeliad species differ

markedly in both size and structure. Tillandsia utriculata holds an

average maximum water volume of 300 mnil with each leaf axil holding

an average of 3.1 ml whereas T. fasciculata holds a maximum, average

of 60 mni with an average leaf axil volume of only 1.5 mi (see

Chapter Il). But despite thEse size differences both brireliad species

contain aquatic communities which are structural ly very similar.

Leaf-txil size may actually be more important in determining the

c!mmuniiiy stucture than this analysis indicates. The two largest

irsect species inhabiting Florida tank broreliads seem to be restricted










to T. u iculot; in other areas (see Chaptzr II) but these are rare

ard did not occur in any of the study sites. Leaf-axil size is a

significant factor in the distribLtion of Odonata and large Coleoptera

anong fCsta Rican brenli ads (Fisa, unpub listed data) and may have a

greater iiflueoce '~pon cor'm'nity structure in the Neotropics where

there is morc diversity in the size and shape of bromeliads T1hn in

south Florida.

Conclus ion

The numerical analysis of the composition and abundance of the

aquatic invertebrate species inhabiting the epiphytic tank bormeliads

in south Florid indicates that community structure is more character-

istic of the ecosystem supporting the bromeliad flora than the species

of :romwiiad that supports the community. This implies that the rela-

tionship between the aquatic fauna and the bromeliad host plant is not

as specific as phytophagous arthropod-plant relationships. Although

the fauna inhabiting bromeliad leaf axils seem to be specific for bro-

neliads there is little evidence that this specificity occurs at the

level of b:-oneliad species. It appears that the aquatic fauna has

adapted to a unique epirhytic habitat provided by the Brocieliaceae and

occupy all sunitable water-holding for's regardless of the bronimelied

species.

The sioq:ificai)ce of throuqhFall and leaf litter accumulation in

det'r-.mining thEf structure of the arquaitic comnrunities inhabiting hr

me!Qids noay have further impliicaicns in understanding the structu.

of other hcterotrophic caqurtic communities such as those that inhabit

rot hoies in t '-es, artificial containers, and small ground pools.






51


Coiiceivably the'.e coniunit-is wuld be siili cr-ly affected by differences

in quantity and quality of nut'.iernt input. It is apparent that a more

thcr:ogh understanding of ec-syster nutrient cycles is essential in

understanding the ecology of the many small but important aquatic

communities that populate both temperat: and trceical forests.









Table 1. Insect prey recover ed from C. br-tpronriiaa leaf-axiis over 8
dys (29 i!;re to 2 July'') at rOipisvil', io-r-ida.


Plant Number

1 2 3 4 Total


6 5 4 13 28

6 12 4 4 26

4 8 4 3 i9

9 2 2 5 18

2 3 1 2 8

0 0 2 0 2


Total 29 30 19 27 105


Insect Orderi


HymenrCpo tera

Diptera

Lepidcptera

Coleoptera

Homopterra

Neuiroptera

ArFchnida

Hef-iptera

Orthoptera
















Table II. Surrnary of site descriptions and collecting data of bronalidd samples from south Florida


Site No.


Location


1 Fishheating Creek Wildlife Management

Area, 1.5 ki west of campgrounds





2 Schewy's Har ioc, 10 km west of

Vero Beach'



3 Everglades liational Park 1.5 kmi

northeast of Myrazek Pond

4 Everglades National Park,

Clapp's Hiartack

5 Homestead, Fuch's Hai..lock

Sanctuary



6 8 km west of North Port

Charlotte east of 1S 41

7 Highlands Cou.ty, 5 km north ut

S 70; cn south side SR 29

8 Christmas, south side US 50

9 9 km, S.W. lIur osted, south side

SR 27

10 Colller-Senir.ole State Parkl south

side US 41

11 Ponce Inlet, 13 kn, south of Daytona

Beach

12 US 411, 16 kn, south-cst of

West Palii Bech

13 Everglades trtional Park

).5 Liu north of sile base PJ.

14 Evergladus hItiorial Park

13 1ni nor-th of F'ohogany IJl.oecs

15 Colliir County, Jet. US 41

and SR 94i

16 Collier Celrty, south bend

on SR 94 (l oo Rd. )

17 Vero BeaLi. Fla. Meidlcal

Entololojy Lab.


* Total samples indicratrl y n.,c,'rs in ( ).

t Destroyed by fire iLbrualy 1/'1


Description


Cypress Swamp






Palni-Haple Hanilock





Buttonwood-Mangrove

Swamp





T roicl ock




Oak Hailock



Bty Snwip



Rdy SwariI

IroSpica l Ha rdwud



Oirk acock



Oak P uauock



Lypress Si, p



Scrub lruttonwuod



Scrub I arlanruve



Custard Apple S'iallp



Custard Apple Swarp



Oak hliu luck


Species


Oate(s)


T. utriculata (40)* 7 Sep 74, 14 Dec 74,

22 Mar 75. 21 Jun 75

T. fasclculat (40) 7 Sep 74. 14 Dec 74,

22 Mar 75, 21 Jun 75

1. utriculata (30) 6 Feb 74, 31 Pug 74,

30 Nov 74

T. fasclculata (10) 30 Nov 75

1. utriculata (40) 14 Sep 74, 2 Jan I/

22 Apr 75, 21 Jl 75

T. fascicilata (!p) 15 Sep 74, 2 Jar. 7-,

22 Apr 75, 22 J.1 75

C. berteronlana (5) 10 Aug 74

C. floribunda (5) 10 Aug 74

G. mionostachia (5) 10 Aug 74

1. utriculata (10) 21 Jul 74



I. utriculata (10) 1 Sep 74



T. utriculata (10) 25 Jun 14

T. utriculata (10) 29 Jun 74

T. fasciculaia (101 29 J.,i 74

l. utriculata (10) 9 Aug 74

T. fasciculata (10) 9 Aug 74

i. utriculata (10) 17 Oct 74



i. fasciculata (10) 22 Sep 75



C. ti terorianr (10) 2 Jan 75, 2P Oct 1P



C. tbrteronlana (5) 23 Apr 75



T. valenzuelana (10) 23 Mar 75



T. valenzuelana (10) 4 Jan 75



T. utriculata (504) R Fell 74, 24 r r !5













Table III. Distribution of aquatic invertebrates within 6 species of
Florida tank bromeliads


Bromeliad species


Ave. total volume (cc)

Ave. no. leaves

Ave. vol. per leaf axil (cc)

No. specimens sampled

Wyeomyia mitchelli

W. vanduzeei

Toxorhynchites r. rutilus

Metriocnemus abdolninoflavatus

Monopelopia tillandsia

Tanytarsini (unidentified)

Neurosystasis n. sp.

Forcipomyia seminole

F. (Warmkea) n. sp.

Meromacrus n. sp.

Stenomicra n. sp.

Neodexiopsis n. sp.

Corynoptera n. sp.

Anoetus n. sp.

Metacypris maracaoensis

Podocopa (unidentified)

Naididae (unidentified)

Turbellaria (unidentified)

Total species


I*-

300

37

8.1

220+

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X







18


60 129

40 13

1.5 10

90 20

X

X X


Eo




43

4.2

5


X X


X
X X X X
x




X X X X X
x x x x x

x

x x
x x


x x x
X X





x x
X X X

X X

X X X



12 9 7 6 3





























0


U

0.
1 .
ci
VI I i 4 -


q OVI
a) > ), V-




I C D 41 r'


C 0ra


SO cr I l

-C U
Ct ) i rI







4- 4 &
o U4 Ci


a41 >, 0 3



:r C, cU Ci CD C

, *- Q ID 1 C- I 0) C S-o
-C-' L >J >) 1- 0i '- a

C) C) C .. CC (r
















4.1-



C-^


4-'



-J -) I





I A'I C'- C) n Lo iC r-U In o


C I i
re a
h-
























1 cj C


-cl

*t7
'A c 2 c' n- r--C- cCO C



S- O l i CO -I C
4i r- r, -
I i I .- _







C. -
Sc- C jc
: I 41
















0 it--
C 1 CO. C,) *0
'2-

r- I- C-


C,! 1 '3 -'






C-) cc cc)




'n-


















I./ C- 0 .0 v ., )
C4 oI -1 r' O C
Ii, rI i3 l ,

0 r C- F c

C) -0 II> (5 I .Jr r- C- L3 .1-' '"


C '- C I i f_ -.

C1 0 C` (A ^ CO 0'

O'C : I i I) C) Q
IA 1 I I CO 3I -
>, 'F" I COl IC I (i C" r> C1 /CI C /I



C '1 *O ( -


- .CC i- CO I) M I C ) C





i- -Q





- C, II C- 0.
i- 71 'F (7' )









Figure 1. Visible arnd iV light photo? ihs of bromeliads illuminated
by natural sunl eight.

A. Visible light photograph ?F insectivorlos C. berteroniara.

B. UV light photograph of same specimen showing strong reflectance
by white powder on the leaf bases.

C. UV reflecting powder also provides a lubricated leaf surface
which prevent: insects from escaping the leaf axils.

D. UV light photograph of T. utriculata, a typical noninsectivorous
tank bromeiliad showing r.o UV reflecting leaf surface.













fI




















Figure 2 Locations of broeiiliad collection sites in Florida










0 9


0


100km




















rn
-s-
C,1-




n











o



*0
-S



O La


; cI

1-























0
C>


















-C





ou
3


























na
I)'























C-A-
<-i








^--

0l












0--











--1i
C/l




















OP









Figure 4. fluster analysis of cuclidian distances among structures of
the aquatic community inhabiting 2 bromeliad species (U =. T. utriculata,
F = T. fasciculata) at different time of the year and in 3 major south
Florida ecosystems.






















TROPICAL HARDWOODS


F, FALL

F, I N T ER

F, SPR II G

F. SUM MER

U. SPR I N G

U. FALL

U. WI ENTER

U, SUMMER

U, S U M ME R

U, I INTER

U. F A L L

F. I N T E R

F, FALL

F, SPRING

F, S U Ml E R

U. S PR I N G


0,1 0.2 0.3 0.4 0.5 0,6 0.7 0,3 0.9 1.0 1.1 1.2 1.3 I.N
EUCLIDIArI DISTANCE









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APPENDIX
COLLECTION DATA USED IN THE CLUSTER ANALYSIS IN CHAPTER III


4-'


nj











Fisheating Creek
T, tricul ata

Plant no. 141 4 2 64 4 1
142 1 27 19 3






143 25 70 3 1 68
(3l 3













144 148

145 2 9 47 1
746 7 16 40 !55
Plat o 4




147 7 13 40 4 1,2
148 2 47 36 2
149 1 16 48
150 34 89 60 132
14 DEC 74
222 1i 9 46 6 12 42
223 7 5 34 12 4
224 21 30 58 6 4
225 4 28 112


227 6 2 19 2 7
147 7 13 40 4 692
227 21 47 36 7 2









228 33 104 161 21 322
229 22 10 15 7 8 1
230 4 12 64 37 7











APPENDIX CONKTINUED
4-1
'ZI
ll >
,r2 r


ri
z- (U.
4-I' 0' ,,
3 CM L C-


>1 r-L S.- 0
i- -- 3 SV as! -
e -t- : *r

>iI I c
r- a c L
,I Q'i u" *
i- 0 l aj >^ (U +-i CJ V-i!{ <
>>r c c ol nJ n ; 3
El a s.-L l *r r o : o cu 4'
c; >r o a .. c vrj Lm n
E- a:1 I-- h: in- ,! ^ (s L O s:


22 Nar 75
Plant no. 311
312
313
314
315
316
317
318
319
320
21 JUN 75
Plant no. 371
372
373
374
375
376
377
378
379
380
Y. fasciculata
7 SEP 74
Plant no. 151
152
153
15~
15r
156
157
158
159
160


(U

(U r
(U
rL
(U
2: I-


1 2
1 18
1
2
3
1 5
3 8
3 6
4 5
1

6 1
29 17
16
7 23
12 1,5
52 22
17 22
16 18
19
16 3


3 11
1
1
9 40
1

7
12
11
5


71
102
1 39
38
39
1 222
230
79
3 36
46

1
2 35
1 42
16
2 18
1 14
84
1 45
11
63


9
1
71
68
10
35
34
91
49
68


__~ ______I_ __













APPENDIX CONTINiJED
















14 DEC 74
Plant no. 231
232
233
234
235
236
237
238
239
240
22 Mar 75
Plant no. 301
302
303
304
305
306
307
303
309
310
20 JUN 75
Plant no. 181
182
183
184
185
18F
187
1S8
189
190


Z3
4L'

4)
c WIJ .. i
Co *n .
LO CL
,'; .( o a ,0- >1 "
l r L S-
0) ~ ~ ~ ~ C Coa\ G c~ ,
L5 Cc La o c
't 5.- Co PI L'SL
4-1 4 I D ,*
( .i -' vc s 0 Co .1 1 ) "

0*0 S I -- C 0
Qj > C U I -'
C(0 0 .41lCO C C
S.1 ( 0l C "
( K Ll- U- ^ 0, l/ 2 <:


4 2
11


7 1


26
2 63
3 66
6
1


1
9
3 5


c11 '
5-~
H~


_~L__I~I II__II___________I_-111-1-










APPENDIX CONTINUED


CJ


rt3
s- -
u 4-' C '
*r' .C' U



>, n -r o c
rl rc rc^ -
C)!~ C)i L



(U a, X +-? !-
c2 ct





51 CI i i


i




S 'I n


41 ci S
tQ


ENP Mangrove Site
T. utriculata
14 SEP 74
Plant no. 165
166
!67
168
169
170
171
172
173
174
2 JAN 75
Plant no. 251
252
253
254
255
256
257
258
259
260
22 MAR 75
Plant no. 331
332
333
334
335
336
337
338
339
340


2


5 125
41
4 28
24
6 72
17 141
3 49
7 52
3 26
16


4 1
3 168
35
13
3
3 74
38
1


12




1 96



12
4
1 36
28


112
50
3 28
23
4 1 37
3 70
36
2 6 44
3 32
3 22

18
92
1 149
5 109
77
38
3 84
4 101
1 29
4

37
25
12?
1 15
38
7 6
1 163
8 6
25
52











APPENDIX CONTINUED


j71j
li
rri
4 4-0"
o U)
M
LI ~v ClC
^ !--J 0 ~ *r C Cul
s- Ut o ru n: *
!, -Cl .- 0

EI C, s l - ,~ L n
a; i -Q ^ r- r-'
il a' ir m LI-Q- r
tjj
Ei _: 1 .i-i v


LC1Ln
.,- I CL) *- ^5 A C ^ .
mlr c= ct c E= ^ ''
*r- ~ ~ ~ ~ ~ o -0 >*^ -> r c -


21 JUL 75
Plant no. 411
412
413
414
415
416
417
418
419
420
Tropical Har-du/oo:l.
15 SEP 74
Plant no. 175
176
177
178
179
2 JAN 74
Plant no. ?26
262
263
264
265
22 fl-VR 75
Plant no. 341
342
343
344
345
21 JUL 75
Plant no. 421
422
423
424
425


2 44
10
1 13
1 19
3 12
27
1 14
19
1 6
12 53


62
1 36
24
62
1 80
70
8
25
15
88


11 1
1

1 1


4 1


1 4
12






78







BIOGRAPHIC SKETCH

The author was born on November 23, 1944, in Berwick, Pennsylvania

and graduated with a B.S. Degree in Biology with a Chemistry minor from

Albright College, Reading, Pennsylvania in 1966. From 1966 to 1970 he

was employed by the Pennsylvania Department of Environmental Resources

where he served for one year as County Health Officer and three years

as Regional Vector Control Coordinator. He received an M.S. Degree in

Entomology from the University of Massachusetts, Amherst in 1973.














I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality as a dissertation for the degree of
Doctor of Philosophy.



Dale H. Habeck, Chairman
Professor Entomology




I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation :nd is fully
adequate in scope and quality as a dissertation for the degree of
Doctor of Philosophy.



Donald E. Weidhaas, Cochairman
Adjunct Professor of Entomology





I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality as a dissertation for the degree of
Doctor of Philosophy.

c X

.-John J. Ewel
Assistant Professor of Botany













I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality as a dissertation for the degree of
Doctor of Philosophy.



Jonathan J. Reiskind
Associate Professor of Zoology








This dissertation was submitted to the Graduate Faculty of the
College of Agriculture and to the Graduate Council, and was
accepted as partial fulfillment of the requirements for the
degree of Doctor of Philosophy.

December 1976


Dean College of Agri-culture
*?


Uean, Graduate School




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