COASTAL MYCOLOGY OF PUERTO RICO: A SURVEY AND BIOLOGICAL
ASPECTS OF MARINE, ESTUARINE, AND MANGROVE FUNGI
by
Ángel Manuel Nieves-Rivera
A dissertation submitted in partial
fulfillment of the requirements for the degree of
DOCTOR OF PHILOSOPHY
in
Marine Sciences
(Biological Oceanography)
UNIVERSITY OF PUERTO RICO
MAYAGÜEZ CAMPUS
May 2005
Approved by:
___________________________
Baqar R. Zaidi, Ph.D.
Member, Graduate Committee
Date
___________________________
Roy A. Armstrong, Ph.D.
Member, Graduate Committee
Date
___________________________
Juan G. González-Lagoa, Ph.D.
Member, Graduate Committee
Date
___________________________
Ernest H. Williams, Jr., Ph.D.
President, Graduate Committee
Date
___________________________
Lucy Bunkley-Williams, Ph.D.
Representative of Graduate Studies
Date
___________________________
Nilda E. Aponte, Ph.D.
Chairperson of the Department
Date
___________________________
José A. Mari Mutt, Ph.D.
Director of Graduate Studies
Date
ABSTRACT
This study was subdivided into the chapters listed below, which treated a number of
biological aspects of the coastal mycology of Puerto Rico (PR): (1) A review chapter on
mangroves and mangrove-associated plants was included for PR. (2) A checklist that
contains 604 taxa of fungi is provided. There were 13 new records for PR generated as a
result of the studies described herein. (3) Samples of sea foam and senescent leaves of
Avicennia germinans and Rhizophora mangle from two communities in an estuary known as
Rincón Lagoon in SW PR were assessed for the frequency of occurrence of 8 selected
filamentous fungi. Among 12 bags of sea foam and 1296 leaves screened (from February
2002 through January 2003), the samples consist of sporulating fungi and propagules,
respectively. A two-way ANOVA performed on 8 filamentous fungi (propagules and spore
counts, N = 432) in sea foam and ones isolated from mangrove leaves (colonies in A.
germinans, N = 576; in R. mangle, N = 576) showed that variability was significant (p >
0.05). Statistical analysis suggested that date/site and site were significant (p > 0.05),
although the date was insignificant (p < 0.05), rejecting the nule hypothesis.
(4)
Cladosporium oxysporum and C. sphaerospermum were isolated from seawater based on
their ability to use the polyaromatic hydrocarbons naphthalene (C10H8) and phenanthrene
(C14H10) as a sole carbon and energy source. (5) The planthopper Petrusa marginata
excretes a sugary honeydew upon which the fungus Asteridiella sepulta grows in the leaves
of A. germinans. The association of A. sepulta and P. marginata had not been noted. (6)
Field surveys in SW PR, and SW Florida (U.S.A.) from July 2001 throughout 2003, yielded
ii
14 spp. of manglicolous basidiomycetes of which 4 were new records. (7) Stemonitis
splendens is reported on R. mangle. This was the first report of this myxomycete on
mangroves in the Caribbean and the fifth report from R. mangle. (8) Aquatic fungi were
isolated from several subtrates from a river mouth estuary Manatí River, N of PR. Finally,
(9) a bibliography on coastal and marine biology primarily mycological in orientation is
presented.
iii
RESUMEN
Este estudio fue subdivido en los capítulos listados debajo y trató varios aspectos
biológicos de la micología costera de Puerto Rico (PR): (1) Se incluye un capítulo de
revisión en mangles y plantas asociadas a los mangles para PR. (2) Se provee una lista que
contiene 604 taxones de hongos. Se encontraron 13 nuevos registros para PR generados
como resultado de los estudios aquí descritos. (3) Muestras de espuma marina y hojas
seniles de Avicennia germinans y Rhizophora mangle de dos comunidades en un estuario
conocido como la Laguna Rincón en el SO de PR fueron evaluados en frecuencia de
ocurrencia de 8 hongos filamentosos selectos. De entre las 12 bolsas de espuma marina y
1296 hojas muestreadas (desde febrero 2002 hasta enero 2003), las muestras consisten de
hongos esporulantes y propágalos, respectivamente. Una ANOVA de dos vías que se hizo a
los 8 hongos filamentosos (conteo de propágulos y esporas, N = 432) en espuma marina y
para los aislados de hojas de mangle (colonias en A. germinans, N = 576; en R. mangle, N =
576) mostró que la variabilidad fue significativa (p > 0.05). El análisis estadístico sugirió
que la fecha/lugar y el lugar fueron significativos (p > 0.05), aunque la fecha fue
insignificante (p < 0.05), rechazando la hipótesis nula. (4) Cladosporium oxysporum y C.
sphaerospermum fueron aislados de agua de mar basándose en su habilidad de usar los
hidrocarburos poliaromáticos nafataleno (C10H8) y fenantreno (C14H10) como fuentes de
carbono y de energía. (5) El saltahojas Petrusa marginata excreta una secreción azucarada
sobre la cual crece el hongo Asteridiella sepulta en la superficie de las hojas de A.
germinans. La asociación de A. sepulta con P. marginata es poco conocida. (6) Los
iv
reconocimientos de campo en el SO de PR, y el SO de la Florida (U.S.A.) durante julio de
2001 y a través de 2003, produjeron 14 spp. de basidiomicetos manglícolas de los cuales 4
son nuevos registros. (7) Se informa a Stemonitis splendens en R. mangle. Este es el primer
informe de este mixomiceto en mangles en el Caribe y el quinto informe en R. mangle. (8)
Se aislaron hongos acuáticos de varios sustratos de la boca de río del estuario Río Manatí, al
N de PR. Finalmente, (9) se presenta una bibliografía sobre la biología costera y marina con
una orientación principalmente micológica.
v
© Ángel M. Nieves-Rivera, May 2005
ALL RIGHTS RESERVED
vi
“Chance makes a plaything of a man’s life.”
–
Seneca
(First Century A. D.)
“Recognosce notum, ignotum inspice.”
–
Motto of the British Mycological Society
“The most beautiful thing we can experience is the mysterious. It is the source of all true art
and science.”
–
Albert Einstein
(Physicist)
“The dream of man is to become God… The reasoning that taught man that he was clever
than the animals made him also aware of his own deficiencies.”
–
John M. Allegro
(Biblical Scholar)
“Skepticism is an attitude towards life, an ethical position. The heart of this position is not
to accept, but neither to reject anything a priori. No subject should be above discussion.
Every extraordinary claim requires extraordinary evidence.”
–
Motto of the Skeptical Society of Puerto Rico
vii
DEDICATION
I would like to dedicate this dissertation to my father, Víctor M. Nieves Seda, who
passed away on April 9, 2004, and to my mother, Adela Rivera Quiles, whose love and
friendship are truly an inspiration to continue graduate studies. Finally, to my stepfather,
Don Hiram Monteverde Rivera, who always taught me how to keep up with the good work
against all odds.
For the love of my life, my Isis, to my immortal beloved.
viii
ACKNOWLEDGMENTS
Thanks to God, my forefathers, and parents for their unconditional guidance and
support at all times. Sincere appreciation is extended to Ernest H. Williams, Jr. for serving
as major advisor during my graduate studies and for providing guidance, support, and
friendship during my stay at the University of Puerto Rico, Mayagüez.
I would like to thank Lucy Bunkley-Williams, Baqar R. Zaidi, Juan G. GonzálezLagoa, and Roy A. Armstrong, who served on my thesis committee and provided many
useful critiques and suggestions. Also thanks to all the faculty and staff of the Departments
of Marine Sciences and Biology, University of Puerto Rico, Mayagüez. In particular Nilda
E. Aponte, Carlos J. Santos-Flores, Carlos Betancourt, Julia S. Mignucci, Juan A. Rivero,
Ángel Berrios, José A. Mari Mutt, Nydia J. Rodríguez-Rodríguez, Jorge E. Corredor, Donato
Seguí Crespo, and Carolyn Rivera aided me greatly in my studies. I am grateful to Jan
Kohlmeyer and Briggite Kohlmeyer (Institute of Marine Studies, University of North
Carolina, Chapel Hill) for their assistance and encouragement. The statistical analysis was
performed after consultation with Miguel A. Rodríguez Montalvo, Ernesto O. Riquelme, and
Paul M. Yoshioka.
Thanks to Deborah Jean Lodge (USDA-Forest Service, Sabana Station, Luquillo) for
assistance and for granting me access to her library and fungal collections. Thanks to Steven
L. Stephenson (University of Arkansas at Fayetteville), Terry A. Tattar (University of
Massachusetts at Amherst), Leif Ryvarden (University of Oslo, Norway), Frank M. Dugan,
Jack D. Rogers (Washington State University, Pullman), Mary Catherine Aime (USDA-ARS,
Maryland), Ariel E. Lugo (USDA, Institute of Tropical Forestry, Río Piedras), Maren A.
Klich (USDA-ARS, Southern Regional Research Center, Louisiana), David W. Minter
(CABI Bioscience, Surrey), John Paul Schmit (University of Illinois, Urbana-Champaign),
Kevin D. Hyde, Stephen S. Pointing (Cryptogamic Herbarium, The University of Hong
Kong, China), and E. B. Gareth Jones (National Center for Genetic Engineering and
Biotechnology, Thailand) for the identification of species, loan of specimens or sending
ix
publications. I thank Carmen T. Acevedo, Conrado M. Calzada, and Sharon A. Cantrell for
their assistance and support.
I thank Arístides Armstrong, Ángel L. González, and Silverio Medina-Gaud
(Department of Crop Protection, University of Puerto Rico, Mayagüez) for the identification
of insects. Further insect taxonomy was kindly provided by Roy R. Snelling (Entomology,
Natural History Museum of Los Angeles County, California), and Dr. Stuart H. McKamey
(USDA, ARS Systematic Entomology Laboratory, Smithsonian Institution, Washington,
D.C.). The valuable help of Peter Rocafort (Department of Marine Sciences, University of
Puerto Rico, Mayagüez) in the endless task of digitalization of figures, remaking
illustrations, and good times is most appreciated (¡Gracias Peter!). Many thanks to
Monserrate “Taty” Casiano, Zulma Martínez, Nilda E. Ramírez, Jossie Moulier, Harry
Justiniano, and many other personel of Department of Marine Sciences, the main library, and
Department of Marine Sciences Collection for their help and patience. Thanks are extended
to Hernán Santos, Jorge Vélez-Juarbe, and María Ruiz-Yantín (Department of Geology,
University of Puerto Rico, Mayagüez) for their help in locating the trace fossils. Also, thanks
to Alan Graham (Missouri Botanical Garden, St. Louis, Missouri) for his assistance on Table
II. José L. Torres helped greatly with the elaboration of the PowerPoint presentation.
I thank the people of Sea Grant College Program of the University of Puerto Rico at
Mayagüez, the Hispanic Scholarship Fund, the Puerto Rico Experimental Program to
Stimulate Competitive Research (EPSCoR), and the University of Puerto Rico Alliance for
the Graduate Education and the Professorate Fellowship (Grant No. NSF/AGEP–HRD #
0302696) for partial funding my research and their unconditional support at all times. The
Puerto Rico Department of Natural and Environmental Resources provided logistical support
for this project. I would like to acknowledge my companions posted at Boquerón Wildlife
Refuge, Mona Island, Mayagüez Regional, and San Juan Offices for their help in pursuing
my research. Finally, for those who assisted, but that I forgot to mention here, thanks!
x
TABLE OF CONTENTS
List of Tables....................................................................................................................xiv
List of Figures....................................................................................................................xv
List of Symbols and Abbreviations..................................................................................xix
Chapter 1. Preface
Introduction ……......................................................................................................1
Objectives.................................................................................................................3
Literature Cited.........................................................................................................4
Chapter 2. New records of an exotic variety, fossil, and unusual structure in mangroves
of Puerto Rico
Abstract.......................................................................................................11
Resumen......................................................................................................11
Introduction.................................................................................................11
Conocarpus erectus var. sericeus: an introduced mangrove-associated
plant................................................................................................22
Aerial roots in Avicennia germinans.................................................24
Paleobotanical notes on mangrove-like plants...................................25
Literature Cited............................................................................................32
Chapter 3. History of marine mycology in Puerto Rico, including a checklist of coastal
and mangrove-associated fungi
Abstract.......................................................................................................41
Resumen......................................................................................................42
Introduction.................................................................................................43
Mangroves.......................................................................................43
Fungi...............................................................................................44
Previous Work.................................................................................46
Final Comments...........................................................................................50
Literature Cited..........................................................................................109
Chapter 4. Filamentous fungi in sea foam and mangrove senescent leaves in a tropical
estuary
Abstract.....................................................................................................140
Resumen....................................................................................................141
Introduction...............................................................................................142
Materials and Method
Study Sites.....................................................................................142
Field Studies
Sea Foam...........................................................................145
Mangrove Leaves...............................................................145
Laboratory Studies
Media.................................................................................146
xi
Observations and Herbarium Collections............................146
Statistical and Ecological Analyses.....................................146
Results and Discussion...............................................................................147
Environmental Features..................................................................147
Fungal Occurrence.........................................................................148
Literature Cited..........................................................................................151
Chapter 5. Characterization of Cladosporium oxysporum and C. sphaerospermum using
polyaromatic hydrocarbons (PAHs) as their sole carbon source in tropical
coastal seawater
Abstract.....................................................................................................179
Resumen....................................................................................................179
Introduction...............................................................................................179
Materials and Method................................................................................180
Results and Discussion...............................................................................181
Literature Cited..........................................................................................184
Chapter 6. Sooty mold-planthopper association on leaves of the black mangrove
Avicennia germinans in southwestern Puerto Rico
Abstract.....................................................................................................188
Resumen....................................................................................................188
Introduction...............................................................................................189
Materials and Method
Locality..........................................................................................191
Isolation of Sooty Mold.................................................................192
Insect Collection............................................................................192
Results.......................................................................................................193
Diagnostic......................................................................................193
Material Studied.............................................................................194
Distribution....................................................................................194
Insects............................................................................................194
Discussion.................................................................................................195
Literature Cited..........................................................................................197
Chapter 7. Manglicolous basidiomycetes of southwestern Puerto Rico and southwestern
Florida (U.S.A.)
Abstract.....................................................................................................206
Resumen....................................................................................................206
Introduction...............................................................................................207
Materials and Method................................................................................208
Results.......................................................................................................210
Discussion.................................................................................................215
Literature Cited..........................................................................................215
Chapter 8. The occurrence of Stemonitis splendens (Myxomycota: Stemonitales) on
Rhizophora mangle
xii
Abstract.....................................................................................................223
Resumen....................................................................................................223
Introduction...............................................................................................223
Materials and Method................................................................................224
Results and Discussion...............................................................................225
Literature Cited..........................................................................................226
Chapter 9. General Discussion........................................................................................229
Chapter 10. General Conclusions...................................................................................232
Chapter 11. Recommendations.......................................................................................233
Appendixes......................................................................................................................234
Appendix 1. Aquatic fungi from estuaries in Puerto Rico. I. Mouth of the Manatí
River
Abstract.....................................................................................................235
Resumen....................................................................................................235
Introduction...............................................................................................236
Materials and Method................................................................................236
Results and Discussion...............................................................................238
Literature Cited..........................................................................................240
Appendix 2. Bibliography on coastal and marine biology in Puerto Rico, with especial
emphasis on marine mycology
Introduction...............................................................................................253
Marine and Coastal Mycology....................................................................254
Marine Pollution, Biodegradation, and Biomedics…...................................273
General Mycology......................................................................................278
Marine Botany, General Botany, and Paleobotany......................................293
Biological Oceanography and General Biology...........................................312
Geological Oceanography..........................................................................338
Chemical Oceanography.............................................................................345
Physical Oceanography and Climatology....................................................349
Marine Archaeology and History...............................................................352
xiii
LIST OF TABLES
Table I. List of mangroves and mangrove-associated plants of Puerto Rico. Taxonomy and
nomenclature according to García-Molinari (1952), Schubert (1979), Little and
Woodbury (1980), Martorell et al. (1981), Tomlinson (1986), Más and GarcíaMolinari (1990), Francis and Lowe (2000), Little et al. (2001), Anonymous (2001),
Acevedo-Rodríguez (2003), and U. S. Department of Agriculture (2004)…...……..12
Table II Location of mangroves and mangrove-associated plants palynomorphs (PL) or
ichnofossils (IF) found in Puerto Rico. They are placed in order of geologic
time…………………………………………………………………………………..30
Table III. Checklist of previous and recent mycological collections (including marine—
obligate and facultative— and terrestrial fungi) in mangroves and mangroveassociated plants, coastal forests, estuaries, beaches, sand dunes, and marine habitats
in Puerto Rico………………………………………………………….…………….55
Table IV. Study sites in Boquerón Beach Inlet (BBI) and Boquerón Wildlife Refuge (BWR),
both part of the Rincón Lagoon, Boquerón Commonwealth Forest, Cabo Rojo,
southwestern Puerto Rico…………………………………...……………………..157
Table V. Monthly and annual environmental averages of study sites (Boquerón Beach Inlet:
Stations 1-3; Boquerón Wildlife Refuge: Stations 4-6) from Rincón Lagoon, Cabo
Rojo, southwestern Puerto Rico……………………..…………………………….158
Table VI. Monthly and annual averages of selected filamentous fungi isolated from sea foam
and mangrove leaves from collection sites in Rincón Lagoon, Boquerón
Commonwealth Forest, southwestern Puerto Rico during February 2002 through
January 2003…………………………………………………………….…………160
Table VII. Univariate analysis of variance (two-way ANOVA) of filamentous fungi collected
from sea foam and mangrove leaves (Avicennia germinans and Rhizophora mangle)
from Boquerón Beach Inlet (BBI) and Boquerón Wildlife Refuge (BWR) collection
sites at Rincón Lagoon, Boquerón Commonwealth Forest, southwestern Puerto Rico
during February 2002 through January 2003 (α = 0.05).………………....……..…164
Table VIII. Summary of the aquatic fungi (obligate and facultative marine fungi sensu
Kohlmeyer, 1974) recovered from samples of sea foam, leaf litter, and wood in the
mouth of the Manatí River, northern Puerto Rico………..………………..………245
Table IX. Summary of the aquatic fungi (obligate and facultative marine fungi sensu
Kohlmeyer, 1974) recovered from the Caribbean……...…………………..………247
xiv
LIST OF FIGURES
Figures 1A-C. A. Location of Puerto Rico in regards to the Greater Antilles. B. Bathymetry
of Puerto Rico (map courtesy of Aurelio Mercado and Harry Justiniano). C. Map of
southwestern Puerto Rico, showing the general collection area…..............................6
Figure 2. Studied sites in southwestern Puerto Rico, Cabo Rojo quadrangle …....................7
Figure 3. Studied sites in southwestern Puerto Rico, La Parguera quadrangle …..................8
Figure 4. Guayanilla Bay area, southwestern Puerto Rico, Guayanilla quadrangle….............8
Figures 5A-I. Examples of marine and estuarine microfungi isolated from Puerto Rico.
Ascospores: A. Arenariomyces triseptatus. B. Corollospora cf. colossa. C.
Corollospora filiformis. Conidia: D. Brachiosphaera tropicalis. E. Campylospora
sp. F-H. Clavatospora bulbosa. I. Camposporidium sp............................................9
Figures 6A-C. Example of a mushroom (Lepiota cf. erinana) recently collected from buried
wood in a saltpeter next to the Phosphorescent Bay, Lajas, Puerto Rico A. In situ. B.
Checking the lamellae. C. Basidiocarps in different stages of
development…...........10
Figures 7A–B. Rhizophora mangle (Rhizophoraceae). A. Leaves and seedling (La Parguera,
southwestern Puerto Rico). B. Aerial stilt roots, a common feature of this species
(Boquerón Commonwealth Forest, Cabo Rojo).......................................................37
Figures 8A–C. Avicennia germinans (Avicenniaceae) (La Parguera, southwestern Puerto
Rico). A. Flowers and leaves. B. Adult trees. C. Aerial roots................................37
Figures 9A–D. Conocarpus erectus var. sericeus (Combretaceae) from southwestern Puerto
Rico. A. Seedling from a commercial nursery from San Germán, Puerto Rico). B.
Used as ornamental in the Boquerón Beach-Cabin Complex, Cabo Rojo. C.
Pubescent leaves. D. Fruiting.……………...….…..………….................................38
Figures 10A–E. Santonian (Late Cretaceous) mangrove-like root casts reported by Santos
(1990, 1999) from the Cotuí Formation, Cabo Rojo-San Germán, southwestern
Puerto Rico. A–B. Magnetite bioturbated facies (lenses) where the roots are located.
C–E. Enlargement of the root casts. …………..………………..............................39
Figures 11A–F. Pleistocene (Late Quaternary) rhizoliths (LACMIP location 17768),
possibly root casts from the mangrove-associated plant Scaveola plumieri
(Goodeniaceae) in the aeolian fossilized dunes of Punta Jacinto, Playa Jobos, Isabela,
in northern Puerto Rico. A–B. Aeolian dune views. C–E. Rhizoliths in situ. F.
Profile
and
sectioned
views
of
weathered
rhizoliths.……………………………………….....……………40
Figure 12. Taxa of marine fungi (including estuarine) reported for Puerto Rico in published
and unpublished reports between 1900 and 2004. Years selected represent
publication date of the records. Numbers given are not cumulative. ..…………....132
Figure 13. Asteromassaria sp. (Ascomycota) on aerial roots of Rhizophora mangle in
Magueyes Island, southwestern Puerto Rico. ………..…………………................133
Figures 14A–B. Aspergillus niger (Mitosporic fungi) on Rhizophora mangle from La
Parguera Channels, southwestern Puerto Rico (Sc = Sporocarps)…......................134
xv
Figures 15A–B. Leaf spots and sporodochia (Sp) of Cercospora sp. (Mitosporic fungi) on a
leaf of Rhizophora mangle from La Parguera Channels, southwestern Puerto Rico.
A. Front. B. Back.…………..…………………………........................................135
Figures 16A–F. Cytospora rhizophorae (Ascomycota) deep orange cirri on prop roots or
trunks of Rhizophora mangle in southwestern Puerto Rico. A. Boquerón Wildlife
Refuge. B-D. Magueyes Island. E-F. Los Morrillos. …...…………………............136
Figure 17. Acervuli (Av) of Pestalotiopsis disseminata (Mitosporic fungi) on leaves of
Rhizophora mangle after growth in moist chambers from La Parguera Channels,
southwestern Puerto Rico…..……………….…………………………….….........137
Figures 18A–B. Phlebia sp. (Basidiomycota) on Rhizophora mangle wood from Boquerón
Wildlife Refuge, southwestern Puerto Rico……...……………………….…..........138
Figures 19A–D. Bolete (Fistulinella cf.) basidiocarp (Basidiomycota) found under the grove
of Myrica sp. (Myricaceae) next to Cabo Rojo’s lighthouse, Boquerón
Commonwealth Forest, southwestern Puerto Rico. A-B. Pileal view. C. Side view.
D. Pore view. …………………..………...............................................................139
Figure 20. Location of sampling sites at Rincón Lagoon, Boquerón Commonwealth Forest,
Cabo Rojo, southwestern Puerto Rico. Stations 1 to 3 are referred through the text as
‘Boquerón Beach Inlet’ (BBI) and the remaining stations (4 to 6) as ‘Boquerón
Wildlife Refuge’ (BWR)........................................................................................167
Figures 21A–D. Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico. A.
Aerial photo of the Boquerón Bay, including view of the Rincón Lagoon, Boquerón
Wildlife Refuge and Puerto Real. B–C. Rhizophora mangle (B) and Avicennia
germinans (C) forming channels in the Boquerón Wildlife Refuge. D. Boardwalk
prepared by the Puerto Rico Department of Natural Resources and brackish water
lagoon............................................................................................168
Figures 22A–C. Leaf baits and traps used for the isolation of estuarine fungi of Rincón
Lagoon, Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico. A.
Leaf baits (Rhizophora mangle) in 1.8 mm2 mesh plastic-screen packet contained in a
1.0 cm2 meshed aluminum envelope. B. Leaves of R. mangle culture in a Petri dish.
C. Potato dextrose agar (PDA) with fungal isolates................................................169
Figure 23. Annual rainfall (February 2002 to January 2003) in Magueyes Island Marine
Laboratories, La Parguera, southwestern Puerto Rico............................................170
Figure 24. Average air temperature (°C) (February 2002 to January 2003) in Magueyes
Island Marine Laboratories, La Parguera, southwestern Puerto Rico......................170
Figure 25. Average wind speed (m/s) and direction (February 2002 to January 2003) in
Magueyes Island Marine Laboratories, La Parguera, southwestern Puerto
Rico......................................................................................................................171
Figure 26. Average water level (m) (February 2002 to January 2003) in Magueyes Island
Marine Laboratories, La Parguera, southwestern Puerto Rico................................171
Figure 27. Average water temperature (°C) (February 2002 to January 2003) in Magueyes
Island Marine Laboratories, La Parguera, southwestern Puerto Rico......................172
xvi
Figure 28. Average water temperature (°C) of sampling sites at Rincón Lagoon, Boquerón
Commonwealth Forest, southwestern Puerto Rico.................................................172
Figure 29. Average salinity (g/L) of sampling sites at Rincón Lagoon, Boquerón
Commonwealth Forest, southwestern Puerto Rico.................................................173
Figure 30. Average pH of sampling sites at Rincón Lagoon, Boquerón Commonwealth
Forest, southwestern Puerto Rico..........................................................................173
Figure 31. Average dissolved oxygen (mg/L) of sampling sites at Rincón Lagoon, Boquerón
Commonwealth Forest, southwestern Puerto Rico.................................................174
Figure 32. Average alkalinity (mg/L) of sampling sites at Rincón Lagoon, Boquerón
Commonwealth Forest, southwestern Puerto Rico.................................................174
Figure 33. Number of fungal spores in sea foam per month during the year of sampling at
Boquerón Beach Inlet, Rincón Lagoon, southwestern Puerto Rico.........................175
Figure 34. Number of fungal spores in sea foam per month during the year of sampling at
Boquerón Wildlife Refuge, Rincón Lagoon, southwestern Puerto Rico..................175
Figure 35. Number of fungal colonies in Avicennia germinans leaves per month during the
year of sampling at Boquerón Beach Inlet, Rincón Lagoon, southwestern Puerto
Rico......................................................................................................................176
Figure 36. Number of fungal colonies in Avicennia germinans leaves per month during the
year of sampling at Boquerón Wildlife Refuge, Rincón Lagoon, southwestern Puerto
Rico......................................................................................................................176
Figure 37. Number of fungal colonies in Rhizophora mangle leaves per month during the
year of sampling at Boquerón Beach Inlet, Rincón Lagoon, southwestern Puerto
Rico......................................................................................................................177
Figure 38. Number of fungal colonies in Rhizophora mangle leaves per month during the
year of sampling at Boquerón Wildlife Refuge, Rincón Lagoon, southwestern Puerto
Rico......................................................................................................................177
Figure 39. Total percentage of fungal isolates from substrates (sea foam, leaves of Avicennia
germinans, and Rhizophora mangle) in the sampling sites at Rincón Lagoon,
Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico..............178
Figures 40A–G. Characteristic microscopic features of Cladosporium oxysporum (ATCC
MYA-3068) and C. sphaerospermum (ATCC MYA-3069) from coastal seawater.
A–D. Conidiophores of C. oxysporum on MEA. E. A group of oval to ellipticalshaped conidia and cylindrical ramoconidia of C. oxysporum at high magnification.
F. Ramoconidia and conidia of C. sphaerospermum on ½ V8 agar. G.
Conidiophores, ramoconidia, and conidia of C. sphaerospermum on MEA.
…..........................................................................................................................187
Figure 41. A. Map of the southwestern end of Puerto Rico, showing Cabo Rojo municipality
collection areas (*)…………….…………...……..…………………………..........202
Figures 42A–F. A. View of Los Morrillos, Boquerón Commonwealth Forest, Cabo Rojo,
Puerto Rico. B. Growth in situ of sooty mould (Asteridiella sepulta) on an upper
surface of leaf of the black mangrove Avicennia germinans. C-D. Leaves of black
mangrove Avicennia germinans, with a clean surface (left) and Asteridiella sepulta
xvii
infested surface (right). E-F. Growth of Asteridiella sepulta on the lower surface of
the front of the leaf of A. germinans, associated with infestation of the planthopper
Petrusa marginata. Photos taken at Los Morrillos, Boquerón Commonwealth Forest
……………………………………………………………….……...………..........203
Figures 43A–B. Asteridiella sepulta. A. Hyphae with perithecium and young ascostroma
bearing hyphal appendages, on leaves of the black mangrove Avicennia germinans
(Cabo Rojo, Puerto Rico). B. Spores……..……………………............................204
Figures 44A–B. Example of the sooty mold, Trimmatostroma sp. A. Perithecium or ascoma
with asci and ascospores inside. B. Dematiaceous hyphal subiculum (sooty mold) on
twig of the sapodilla (níspero) Malikara zapota (Mayagüez, Puerto
Rico)………………………………………………..……………...……… ….......205
Figures 45A–B. Sea grape Coccoloba uvifera, showing sooty mold (A) and with an
infestation of Petrusa marginata on lower surface (B). A. Dematiaceous hyphae
(Dh) and young ascostroma of Asteridiella sepulta and Trimmatostroma sp., on
leaves of the black mangrove Avicennia germinans surface. Photograph taken at
Playa Jobos, Isabela, Puerto Rico.……….……………………………...................205
Figures 46A–B. Forests of Avicennia germinans and Rhizophora mangle of the Aquatic
Preserves of Southwest Florida, located southwestern of Venice, at the Gulf of
Mexico, Florida (U.S.A.). A. Pneumatophores and trees of Avicennia germinans. B.
Aerial roots, buttresses, and trees of Rhizophora mangle.......................................220
Figures 47A–E. Manglicolous basidiomycete collected on southwestern Puerto Rico. A–B.
Crepidotus uber on dead upright Rhizophora mangle tree trunk (A–B, young
basidioma) and branch (C–E, mature basidioma) ……….………….……..............221
Figures 48A–H. Further manglicolous basidiomycetes collected on southwestern Puerto
Rico and Florida (U.S.A.). A-B. Psathyrella sp. (Pr = primordium; Bs = basidioma)
on Rhizophora mangle leaf litter. C. Lentinus crinitus. D. Coriolopsis floccosa. E.
Hexagonia hydnoides. F. Pycnoporus sanguineus. G. Tyromyces cf. chioneus on R.
mangle wood. H. Dacryopinax spathularia on R. mangle wood…..…………......222
Figures 49A–B. A. Sporangia of Stemonitis splendens (UARK #17308; a = side view, b =
bottom view) collected on Rhizophora mangle, Boquerón Commonwealth Forest,
Cabo Rojo, Puerto Rico, West Indies. B. Fruiting of Stemonitis splendens in New
Zealand…………………………...........................................................................228
Figure 50. Collection of sea foam samples from the mouth of Manatí River, north central
coast of Puerto Rico…..........................................................................................251
Figures 51A–H. Aquatic fungi isolated from samples collected of the mouth of Manatí
River, northern Puerto Rico. Ascospores: A. Arenariomyces triseptatus. B.
Astrosphaeriella aff. mangrovei. C. Corollospora cf. colossa. D. Corollospora
filiformis. E. Kirschsteiniothelia sp. F. Torpedospora radiata. Conidia: G.
Brachiosphaera tropicalis. H. Clavatospora bulbosa………...……………..........252
xviii
LIST OF SYMBOLS AND ABBREVIATIONS
alt.
=
altitude
ANOVA
=
Analysis of variance
AMNR
=
Ángel M. Nieves-Rivera
APSF
=
Aquatic Preserves of Southwest Florida, Florida
ATCC
=
American Type Culture Collection, Rockville, Maryland
BBI
=
Boquerón Beach Inlet
BCF
=
Boquerón Commonwealth Forest
BPI
=
U.S. National Fungus Collections, Maryland
BS
=
Bahía Sucia, Boquerón Commonwealth Forest
BWR
=
Boquerón Wildlife Refuge
°C
=
Celsius scale, Celsius degree or centigrade scale
CD-ROM
=
Compact disk-read only memory
cf.
=
(confer), compare with
CFMR
=
Center for Forest Mycology Research, U.S. Department of
Agriculture, Sabana Field Station, Puerto Rico
cm
=
centimeter(-s)
Co
=
coastal beaches (geologic nomenclature)
diam.
=
diameter
DNA
=
Deoxyribonucleic acid
DO
=
Dissolved oxygen
E
=
east
e.g.
=
(exempli gratia), for example
ENE
=
East-north-east
Fig(-s)., fig(-s).
=
Figure(-s) (in citations)
FL
=
Florida (U.S.A.)
g/L
=
grams per liter or g/1000 ml (= parts per thousand or ppt)
xix
GB
=
Guayanilla Bay, Guayanilla, Puerto Rico
G.P.S.
=
Global Positioning System
H0
=
Null hypothesis
HPLC
=
High Pressure Liquid Chromatograph
hr(-s)
=
hour(-s)
ibid.
=
(Ibidem), at the same place
MS
=
Institute of Marine Sciences, The University of North Carolina,
Chapel Hill
km
=
kilometer(-s) per hour
km2
=
square kilometer(-s)
KOH
=
5.0% aqueous solution of potassium hydroxide
LACMIP
=
Natural History Museum of Los Ángeles County, Department of
Invertebrate Paleontology, Los Ángeles, California
LM
=
Los Morrillos, Boquerón Commonwealth Forest
LPCF
=
La Parguera Commonwealth Forest
Lr
=
limestone rock lands (soil nomenclature)
LTER
=
Long Term Ecological Research
m
=
meter(-s)
Ma
=
million year(-s)
MAPR
=
Tropical Mycology Collection, University of Puerto Rico Herbarium,
Mayagüez
MEA
=
Malt extract agar
mg.ha–1.yr–1
=
milligrams per hectares per year
mg/L
=
milligrams per liter or 10–3 g/1000 ml (= parts per million or ppm)
MIML
=
Magueyes Island Marine Laboratories or Magueyes Island Field
Station, La Parguera
ml
=
milliliter(-s)
mm2
=
square milimeter(-s)
xx
MO
=
Herbarium, Missouri Botanical Garden, Saint Louis, Missouri
N
=
north
NJRR
=
Nydia J. Rodríguez-Rodríguez
NOAA
=
National Oceanographic and Atmospheric Administration (U.S.
Department of Commerce)
NY
=
Cryptogamic Herbarium, The New York Botanical Garden, Bronx
O
=
The Mycological Herbarium, The Botanical Garden, University of
Oslo, Norway
op. cit.
=
(opere citato), in the work mentioned
PAHs
=
Polyaromatic hydrocarbons or polycyclic aromatic hydrocarbons
PC
=
La Parguera Channels
PDA
=
potato dextrose agar
pl.
=
plate(-s) (in citations)
Pp., pp.
=
page(-s) (in citations)
PR
=
Puerto Rico
PRDNR
=
Puerto Rico Department of Natural Resources (old name of PRDNER)
PRDNER
=
Puerto Rico Department of Natural and Environmental Resources
Qb
=
beach deposits (geologic nomenclature)
QLE
=
Coefficient of light extinction
Qm
=
Holocene’s mangrove swamps (geologic nomenclature)
RL
=
Rincón Lagoon or “Laguna Rincón”, southwestern Puerto Rico
S
=
south
sp., spp.
=
species (singular and plural, respectively)
s. str.
=
(sensu stricto), in the strict sense; narrowly
SSE
=
South-south east
SSF
=
San Sebastián Formation
SWM
=
Sea water medium
Tf
=
tidal flats (soil nomenclature)
xxi
Ts
=
tidal swamps (Ts) (soil nomenclature)
UARK
=
Herbarium, University of Arkansas, Fayetteville, Arkansas
UPRMP
=
Geological Museum, Department of Geology, University of Puerto
Rico, Mayagüez Campus
UPRRP
=
Herbarium, University of Puerto Rico, Río Piedras Campus
USDA
=
United States Department of Agriculture
UTMC
=
University of Texas Myxomycete Collection, Texas
Vol(-s)., vol(-s).
=
volume(-s) (in citations)
W
=
west
YPM
=
Yale Peabody Museum Herbarium, Yale University, Cincinnati
yy, yr, yrs
=
year(-s)
µm
=
micrometer(-s)
xxii
CHAPTER 1
PREFACE
“In the sea around the Pillars of Hercules where there is much water, fungi are produced
close to the sea, which people say have been turned to stone by the sun.”
–Theophratus (circa 372-287 B.C.)
INTRODUCTION
Aquatic fungi and fungal-like organisms, especially freshwater forms have been
studied for many years. The oomycetes, the most important group in early studies,
taxonomically speaking are a group derived phylogenetically from algae and with which they
were often confused (Ainsworth, 1976). The first representatives are zoosporic-like,
although zoospores were first reported in 1807 by Prévost in the terrestrial Albugo
(Peronosporales: Oomycota) (Ainsworth, 1976). Many important nineteenth-century works
were consolidated within the contribution “Kryptogamenflora der Mark Brandenburg” by M.
von Minden between 1911 to 1915, the latter being the forerunner of the later works of
Frederick K. Sparrow (1923) on Saprolegniaceae and Pythiaceae, also treated by William C.
Coker (1923) and Velma D. Matthews (1931) (Ainsworth, 1976). Since these early studies,
many details of their complicated life cycles have been revealed and been added to the
knowledge of the sexuality and cytology of the Oomycota. In the 1940’s, Cecil T. Ingold
described a series of mitosporic fungi today known as aquatic hyphomycetes (or Ingoldian
fungi) growing on decaying leaves in a British stream (Ingold, 1942; Ainsworth, 1976).
These fungi (mostly asexual forms of Ascomycota and Basidiomycota) are cosmopolitan in
distribution, and therefore can be isolated from temperate regions and tropics, including
estuaries (Kirk, 1969). However, our knowledge of these groups in general is increasing in
the coastal environment.
1
The study of marine fungi [obligate and facultative as defined by Kohlmeyer (1974)]
and marine mycology began early in the twentieth century with the report of some species by
Arthur D. Cotton in 1909 (Ainsworth, 1976). The first reports dealing with marine
pyrenomycetes were published by George K. Sutherland in 1915 and 1916 (Ainsworth,
1976). However, the report that caused major interest to the scientific community was the
paper on the marine fungi of New England and California by Elso S. Barghoorn and David
H. Linder in 1944 (Barghoorn and Linder, 1944). As a direct consequence, an expansion of
mycological studies has been carried out since the 1950’s, mostly due to popularity among
scientists from whom originated such monographs as “Fungi in oceans and estuaries”
(Johnson and Sparrow, 1961), “Recent advances in aquatic mycology” (Jones, 1976),
“Marine mycology: the higher fungi” (Kohlmeyer and Kohlmeyer, 1979), and “Marine
mycology: a practical approach” (Hyde and Pointing, 2000). Also, the “International marine
and freshwater mycology symposium” which occurs every 4 to 5 years and attracts an
increasing number of participants around the world is the source of new research on marine
fungi (Hyde and Pointing, 2000). Certainly, those new investigations, especially those on
biotechnological potentialities of marine fungal metabolites are of great interest to
pharmaceutical companies.
Traditionally, terrestrial and freshwater fungi have been the main subject for study by
Puerto Rican mycologists. Recently, coastal fungi (e.g., marine, estuarine, and manglicolous
fungi) have received some attention in Puerto Rico (Acevedo, 2001; Nieves-Rivera et al.,
2002). Human concern with oceanographic issues also account for the increased interest in
coastal organisms. Pollution in mangroves, estuaries, and local shores is common, thus the
roles of macro- and micromycetes in coastal environments are relevant. The topics reviewed
in this thesis included fungal nomenclature, taxonomy and morphology, and ecology. In
keeping with the growing concern over environmental issues, a chapter dealt with two locally
isolated microfungi (Cladosporium oxysporum and C. sphaerospermum) that have the
capability of degrading polyaromatic hydrocarbons (PAHs). It is therefore, my intention of
summarizing all available works, either published or unpublished, that dealt with coastal
2
fungi (obligate and facultative marine and terrestrial fungi) from Puerto Rico. This work
might be considered a supplement to the Caribbean mycological survey of Minter et al.
(2001). Most coast fungi collections are in some cases sporadic and interrupted by many
years. This study reported the incidence of fungi in coasts from Puerto Rico, a subtropical
island located between 18°00’–18°30’ N, 65°35’–67°15’ W, in the northeastern Caribbean
Sea (Figure 1A-C). The maps of southwestern Puerto Rico showing the general collection
sites (Cabo Rojo, La Parguera, and Guayanilla quadrangles) are included in Figures 2-4.
OBJECTIVES
The proposed study relies on the hypothesis that fungal communities from mangroves
and estuarine ecosystems in Puerto Rico are relevant as saprophytes and decomposers.
Obligate and facultative marine fungi (s. str. Kohlmeyer, 1974) are important to the
degradation of plant debris and have been found to be more diverse than previously
suspected. Although many ecological studies on manglicolous and estuarine fungi have been
accomplished in various parts of the globe, their ecology has been poorly documented by
previous researchers in Puerto Rico. The null hypothesis (H0) proposed is that sampling time
(mm-yy), site or location, and the combination of both sources (intersect) will not affect
significatively (p < 0.05) two filamentous fungal communities in a tropical estuary.
Specific objectives include:
1. To survey and report the taxonomic composition of marine, estuarine, and terrestrial costal
fungi (Ascomycota, Basidiomycota, Zygomycota, Chytridiomycota, and Mitosporic
fungi) and fungal-like organisms (Oomycota, Myxomycota, Plasmodiophoromycota)
associated with mangroves species (e.g., Rhizophora mangle, Avicennia germinans,
Laguncularia racemosa, and Conocarpus erectus) in marine, estuarine, and terrestrial
ecosystems of southwestern Puerto Rico (e.g., see Figures 5A-I and 6A-C);
2. To compare the ecological aspects (e.g., taxonomic composition y distribution patterns) of
the fungal communities that will be surveyed (see objective 1) with previous reports
3
from other subtropical and tropical mangal regions of the globe especially the
Caribbean.
3. Characterization by morphology of two species of Cladosporium (C. oxysporum and C.
sphaerospermum) isolated from coastal seawater and have the ability to use
polyaromatic hydrocarbons (PAHs) naphthalene and phenanthrene.
4. To contrast two filamentous fungal communities in a nutrient- and organic matter rich
brackwater lagoon bordered by mangroves versus the environmental parameters (e.g.,
water temperature, pH, salinity, dissolved oxygen, etc.).
5. Prepare a bibliography on coastal mycology of Puerto Rico with a particular emphasis on
tropical and subtropical marine, estuarine, and mangrove-associated fungi.
LITERATURE CITED
Acevedo, C. T. 2001. Marine fungi in Puerto Rico: endophytism and biodegradation. Ph.D.
Thesis, Puerto Rico, Río Piedras: University of Puerto Rico at Río Piedras and
Medical Sciences Campuses, Biology Inter Campus Doctoral Program. 85 pp.
Ainsworth, G. C. 1976. Introduction to the history of mycology. Cambridge University Press,
Cambridge. 359 pp.
Barghoorn, E. S. and D. H. Linder. 1944. Marine fungi: their taxonomy and biology.
Farlowia 1: 395-467.
Hyde, K. D. and S. B. Pointing (eds.). 2000. Marine mycology: a practical approach. The
University of Hong Kong Press, Hong Kong, China. 377 pp.
Ingold, C. T. 1942. Aquatic hyphomycetes of decaying alder leaves. Transactions of the
British Mycological Society 25: 339-417.
Johnson, T. W., Jr. and F. K. Sparrow, Jr. 1961. Fungi in oceans and estuaries. J. Cramer,
Weinheim, Germany. 668 pp., 17 pl.
Jones, E. B. G. (ed.) 1976. Recent advances in aquatic fungi. John Wiley & Sons, New York.
749 pp.
Kirk, P. W., Jr. 1969. Aquatic hyphomycetes on wood in an estuary. Mycologia 61: 177-181.
4
Kohlmeyer, J. 1974. On the definition and taxonomy of higher marine fungi.
Veröffentlichungen des Institus für Meerescforschung in Bremerhaven Supplement 5:
263-286.
Kohlmeyer, J. and E. Kohlmeyer. 1979. Marine mycology: the higher fungi. Academic Press,
New York. 690 pp.
Minter, D. W., M. Rodríguez Hernández and J. Mena Portales. 2001. Fungi of the
Caribbean: an annotated checklist. Published by the senior author and PDMS
Publishing, Middlesex, United Kingdom. 946 pp.
Nieves-Rivera, Á. M., T. A. Tattar and E. H. Williams, Jr. 2002. Sooty mould-planthopper
association on leaves of the black mangrove Avicennia germinans (L.) Stearn in
southwestern Puerto Rico. Arboricultural Journal 26:141-155.
5
Figures 1A-C. A. Location of Puerto Rico in regards to the Greater Antilles. B. Bathymetry
of Puerto Rico (map courtesy of Aurelio Mercado and Harry Justiniano). C. Map of
southwestern Puerto Rico, showing the general collection area.
6
Figure 2. Studied sites in southwestern Puerto Rico, Cabo Rojo quadrangle.
7
Figure 3. Studied sites in southwestern Puerto Rico, La Parguera quadrangle.
Figure 4. Guayanilla Bay area, southwestern Puerto Rico, Guayanilla quadrangle.
8
Figures 5A-I. Examples of marine and estuarine microfungi isolated from Puerto Rico.
Ascospores: A. Arenariomyces triseptatus. B. Corollospora cf. colossa. C.
Corollospora filiformis. Conidia: D. Brachiosphaera tropicalis. E. Campylospora
sp. F-H. Clavatospora bulbosa. I. Camposporidium sp.
9
Figures 6A-C. Example of a mushroom (Lepiota cf. erinana) recently collected from buried
wood in a saltpeter next to the Phosphorescent Bay, Lajas, Puerto Rico A. In situ. B.
Checking the lamellae. C. Basidiocarps in different stages of development.
10
CHAPTER 2
NEW RECORDS OF AN EXOTIC VARIETY, FOSSIL, AND UNUSUAL
STRUCTURE IN MANGROVES OF PUERTO RICO
ABSTRACT
A compilation of 125 plants routinely found in mangrove forests, the introduction of
a new variety of the mangrove-associated plant, Conocarpus erectus var. sericeus
(Combretaceae), unusual aerial roots in Avicennia germinans (Avicenniaceae), and several
new trace fossils (ichnofossils) of mangroves are reported for Puerto Rico. An actualized
checklist of scientific and common botanical names (English/Spanish), including 3 true
mangroves (A. germinans, Laguncularia racemosa, and Rhizophora mangle) and 122
mangrove-associates are herein reported for Puerto Rico.
RESUMEN
Un resumen de 125 plantas rutinariamente encontradas en los bosques de mangle, la
introducción de una nueva variedad de planta asociada al mangle, Conocarpus erectus var.
sericeus (Combretaceae), unas raíces aéreas inusuales en Avicennia germinans
(Avicenniaceae) y varios nuevos fósiles trazas (icnofósiles) de mangles son informados para
Puerto Rico. Se informa una lista actualizada de nombres científicos y comunes botánicos
(inglés/español), incluyendo 3 mangles verdaderos (A. germinans, Laguncularia racemosa y
Rhizophora mangle) y 122 asociados al mangle para Puerto Rico.
INTRODUCTION
A mangrove forest is considered a dynamic ecotone (or transition zone) between
terrestrial and marine habitats.
In its simplest sense, “mangrove” as used herein,
encompasses a group of woody, halophylic plants that occurs along sheltered tropical and
subtropical coastlines. Mangroves are derived from a variety of plant families and vary in
11
their dependence upon littoral habitats. Mangrove forests are also referred to as mangrove
swamps, tidal forests, tidal swamp forests or mangals. Caribbean mangroves range from 30°
N latitude (northern Florida) to 8° N latitude (northwest Colombia) and from 59° W
longitude (north of Guiana) to 89° W longitude (eastern Guatemala) (Tomlinson, 1986).
Until recently, four mangrove species were known to occur in Puerto Rico, the most widely
distributed of which are the red mangrove (Rhizophora mangle), black mangrove (Avicennia
germinans), white mangrove (Laguncularia racemosa), and buttonwood (Conocarpus
erectus) (Lugo and Snedaker, 1974; Cintrón et al., 1978) (Figures 7A-B, 8 A-C, and 9A-D).
Following Tomlinson (1986), only three species should be considered as “true mangroves”
(A. germinans, R. mangle, and L. racemosa) in Puerto Rico. Conocarpus erectus is
frequently considered a “true mangrove”, but Tomlinson (1986) suggested that is better
regarded as a mangrove associate because it lacks any of the biological features
(pneumatophores and vivipary) which characterize true mangroves; furthermore, it occurs in
inland communities. A number of other plants characteristically occur in association with
mangroves in Puerto Rico (Table I).
Table I. List of mangroves and mangrove-associated plants of Puerto Rico. Taxonomy and
nomenclature according to García-Molinari (1952), Schubert (1979), Little and Woodbury
(1980), Martorell et al. (1981), Tomlinson (1986), Más and García-Molinari (1990), Francis
and Lowe (2000), Little et al. (2001), Anonymous (2001), Acevedo-Rodríguez (2003), and
U. S. Department of Agriculture (2004).
TAXA
Family Aizoaceae
Sesuvium maritimum (Walt.) B.S.P.
COMMON NAMES
slender seapurslane;
verdolaga de mar
sea purslane, shoreline
seapurslane; yerba de vidrio,
verdolaga rosada
S. portulacastrum (L.) L.
Family Alismataceae
Echinodorus berteroi (Spreng.) Fassett
12
creeping burrhead; llantén de
agua
Sagitaria lancifolia L.
bulltongue arrowhead, arroz
leaf; flecha de agua, saeta de
agua
Family Annonaceae
Annona glabra L.
pond apple, alligator apple,
cork wood, mangrove
annona, bunta dog apple,
bunya, dog apple, monkey
apple; cayur, cayure, corazón
cimarrón, corcho, coyur
Family Apiaceae
Hydrocotyle bonariensis Comm.: Lam.
H. hirsuta Sw.
H. prolifera Kellogg
H. pusilla A. Rich.
H. umbellata L.
Family Apocynaceae
Rhabdadenia biflora (Jacq.) Muell.-Arg.
Family Aquifoliaceae
Ilex vomitoria Soland. in Ait.*
Family Avicenniaceae
Avicennia germinans (L.) Stearn
largeleaf pennyworth
yerba de clavo
whorled marshpennyworth
tropical marshpennyworth
marsh pennyworth,
manyflower
marshpennyworth; ombligo
de Venus, sombrerillo de
agua, yerba de cuarto
mangrove rubber vine,
mangrovevine; enredadera de
mangle
cassena, yaupon; yaupón
black mangrove, blackmangrove, olive mangrove,
salt pond, honey mangrove,
saltbush; mangle negro,
mangle prieto, mangle bobo,
chifle de vaca, prieto, salado,
siete cueros, mangle blanco
Family Batidaceae
Batis maritima L.
pickleweed, saltwort,
turtleweed; barilla, planta de
sal
Family Boraginaceae
Argusia gnaphalodes (L.) Heine
sea rosemary, sea lavender,
bay lavender; nigua de playa,
13
Heliotropium curassavicum L.
Heliotropium curassavicum L. var. curassavicum
Tournefortia filiflora (Ker-Gawl.) L. Benson
Family Cactaceae
Opuntia stricta (Haw.) Haw.
té de mar, temporana, yerba
cotorra
seaside heliotrope, salt
heliotrope; cotorrera de playa
seaside heliotrope, salt
heliotrope; cotorrera de playa
cold withe; nigua
erect pricklypear, prickly
pear, cactus; tuna, tuna brava,
higo de mar
Opuntia stricta (Haw.) Haw. var. dillenii (Ker-Gawl.) L. Benson
erect pricklypear, prickly
pear, cactus; tuna, tuna brava,
higo de mar
Family Casuarinaceae
Casuarina equisetifolia L.*
Australian pine, Australian
beefwood, beef wood, beach
sheoak; pino australiano,
casuarina, pino
Family Combretaceae
Conocarpus erectus L.
button mangrove,
buttonwood, buttontree,
button-tree; mangle botón,
mangle de botón, botoncillo
,
C. erectus var. sericeus Griseb.* ***
silver buttonwood, northern
buttonwood; mangle botón
plateado; botoncillo plateado
Laguncularia racemosa (L.) Gaertn. f.
white mangrove; mangle
blanco, mangle bobo
Terminalia catappa L.*
Indian almond, Indianalmond, tropical almond,
Malabar almond; almendra,
almendro
Family Compositae
Helianthus annuus L.*
common sunflower,
sunflower; girasol, mirasol
Family Convolvulaceae
Ipomoea imperati (Vahl) Griseb.
fiddle-leaf morningglory,
beach morning glory, beach
morning-glory; batatilla,
14
bejuco de costa, bejuco de
puerco de costa
I. pes-caprae (L.) R. Brown
beach morningglory, beach
morning glory, bayhops, bay
hops, goat foot; bejuco de
playa
I. pes-caprae (L.) R. Brown ssp. brasiliensis (L.) van Ooststr.*
Brazilian bayhops, Brazilian
bay hops; bejuco de playa
brasiliense
Family Cyperaceae
Cyperus imbricatus Retz.
shingle flatsedge; junco
C. unioloides R. Brown
uniola flatsedge
Eleocharis mutata (L.) Roem. & J.A. Schult.
scallion grass, scalion grass,
angled spikerush
Remirea maritima Aubl.
beachstar, beach sedge; junco
de playa
Scirpus tabernaemontani (K.C. Gmel.) Palla
softstem bulrush, bulrush;
junco
Torulinum filiforme (Sw.) C.B. Clarke
pajón de costa
Family Euphorbiaceae
Caperonia palustris (L.) St.-Hil.
sacatrapo
Croton astroites Ait.
wild marrow; maná, marán
C. betulinus Vahl
beechleaf croton
C. discolor Willd.
lechecillo
C. glandulosus L.
vente conmigo
C. humilis L.
pepper bush, pepperbush,
yerba bellaca
C. impressus Urb.
Puerto Rico croton
C. lobatus L.
lobed croton; croton lobulado
C. lucidus L.
firebush
C. microcarpus Ham.
money croton
C. poecilanthus Urb.**
saninon; sabilón
C. rigidus (Muell.-Arg.) Britton
yellow balsam; adormidera,
guayacanillo
C. stenophyllus Griseb.
West Indian croton
Hippomane mancinella L.
manchioneel, manchineel;
manzanillo
Family Goodeniaceae
Scaevola plumieri (L.) Vahl
inkberry, gullfeed; borbón,
borborón, coralillo
Family Gramineae (Poaceae)
15
Cenchrus brownii Roem. & J.A. Schult.
C. echinatus L.
C. gracillimus Nash
C. incerus M.A. Curtis
C. myosuroides Kunth
C. pauciflorum Benth.
Cynodon dactylon (L.) Pers.
Dactyloctenium aegypticum (L.) Willd.
Eustachys petraea (Sw.) Desv.
Paspalidium geminatum (Forsk.) Stapf
Paspalum vaginatum Sw.
Phragmites australis (Cav.) Trin.: Steud.
Spartina patens (Ait.) Muhl.
Sporobolus virginicus (L.) Kunth
Stenotaphrum secundatum (Walt.) Kuntze
16
slimbristle sandbur; abrojito
bur grass, southern sandbur;
abrojo, cadillo
slender sandbur; abrojo de
playa
sand burgrass; abrojo de
playa
big sandbur, spiked-burgrass; abrojo de espigas
sand burgrass, mat sandbur,
field sandbur; abrojo de
dunas, abrojo de playa
common Bermudagrass,
Bermuda grass; yerba
Bermuda, Bermuda común,
hala que te quedas, palo de
brujas, lao de brujas, Pepe
Ortiz
Egyptian grass, goose foot
grass, crowfoot grass, durban
crowfoot grass; yerba egipcia,
yerba de Egipto
pinewoods fingergrass;
stiffleaf eustachys; yerba de
dedo
water panicum, Egyptian
panicgrass; yerba de pantano
seashore paspalum, stiltgrass;
paspalum playero, cortadera
common reed, reed grass;
caña de indio, caña de
pantano
saltmeadow cordgrass, salt
grass; yerba de sal
seashore dropseed, sea-shore
rush grass, seashore rush
grass, sandcoach, beachgrass,
saltwater smutgrass; matojo
de burro, matojo de playa,
salaillo
Saint Augustine grass, St.
Augustine grass, St.
augustinegrass, running crabgrass; yerba San Agustín,
cinta, cinto, cintillo, grama,
grama blanca, grama dulce
Family Guttiferae
Calophyllum inophyllum L.*
Alexandrian laurel, ball nut,
kamani; María grande
Family Hydrocharitaceae
Thalassia testudinum Banks & Soland.: Koeing
Family Lecythidaceae
Barringtonia asiatica (L.) Kurz*
turtlegrass, thalassia;
thalassia, yerba tortuga,
palma de mar
fish poison tree; almendrota,
bonete de arzobispo, coco de
mar, mudilla
Family Leguminosaceae (Fabaceae)
Subfamily Caesalpinioideae
Caesalpinia bonduc (L.) Roxb.
gray nickers, yellow nicker;
mato de playa, mato azul,
haba de San Antonio
C. ciliata (Bergius: Wikstr.) Urb.
mato de playa, mato
C. culebrae (Britton & P. Wilson) Alain** smooth yellow nicker; mato
de playa culebrense
C. monensis Britton
black nicker; mato negro
C. portoricensis (Britton & P. Wilson) Alain**
brown nicker; mato marrón,
mato negro
Chamaecrista nictitans (L.) Moench
partridge pea; matojo de
playa
C. nictitans (L.) Moench ssp. nictitans
partridge pea; matojo de
playa
C. nictitans (L.) Moench ssp. nictitans var. aspera (Muhl.: Ell.) Irwin &
Barneby
partridge pea; matojo de
playa
C. nictitans (L.) Moench ssp. nictitans var. diffusa (DC.) Irwin & Barneby
partridge pea; matojo de
playa
C. nictitans (L.) Moench ssp. patellaria (DC.: Colladon) Irwin & Barneby
17
partridge pea; matojo de
playa
C. nictitans (L.) Moench ssp. patellaria (DC.: Colladon) Irwin & Barneby
var. glabrata (Vogel) Irwin &
Barneby
partridge pea; matojo de
playa
Pterocarpus officinalis Jacq.
swamp bloodwood,
dragonsblood tree; palo de
pollo, pterocarpus
Senna bicapsularis (L.) Roxb.
Christmas-bush,
Christmasbush, stiver bush,
styver-bush; hoja de sen, sen
del país
S. nitida (L.C. Rich.) H.S. Irwin & Barneby
hediondilla
Stahlia monosperma (Tul.) Urb.**
cóbana, cóbana negra,
polisandro
Subfamily Papilionoideae
Canavalia rosea (Sw.) DC.
baybean, beach bean, bay
bean, seaside bean; haba de
playa, canavalia, habichuela
playera, mato de playa
Dalbergia ecastophyllum (L.) Taub.
coin vine, maraymaray; palo
de pollo, maraimaray, siso
Machaerium lunatum (L. f.) Ducke
escambrón, palo de hoz
Family Lemnaceae
Lemna aequinoctialis Welw.
lesser duckweed, duck weed;
yerba de pato, lentejilla de
agua
Wolffiella lingulata (Hegelm.) Hegelm.
tongueshape bogmat;
wolffiella
Family Lentibulariaceae
Utricularia gibba L.
humped bladderwort; grasilla
Family Liliaceae
Yucca aloifolia L.*
aloe yucca, Spanish-bayonet;
aguja de Adán, bayoneta
española, mata de hueso
Family Malvaceae
Hibiscus tiliaceus L.
beach hibiscus; emajagua,
majagua
18
H. pernambucensis Arruda
Pavonia paludicola D.H. Nicols.: Fryxell
Thespesia populnea (L.) Soland.: Correa*
sea hibiscus; emajagua,
majagua
swampbush; cadillo de
ciénaga
seaside mahoe, portia tree,
Spanish cork, portiatree,
otaheita; emajagüilla
Family Mimosaceae
Prosopis pallida (Humb. & Bonpl.: Willd.) Kunth*
mesquite; bayahonda,
mesquite, algarroba del
Hawaii, kiawe
water dead and awake, water
neptunia; desmanto amarillo
Neptunia plena (L.) Benth
Family Myoporaceae
Bontia daphnoides L.
white alling; manzanilla,
mangle bobo
Family Myricaceae
Morella cerifera (L.) Small*
wax myrtle, southern
waxmyrtle, bay-berry, waxberry; arrayán, cerero
Family Najadaceae
Najas guadalupensis (Spreng.) Magnus
southern waternymph,
Guadalupe waternymph,
southern naiad, common
water nymph
Najas guadalupensis (Spreng.) Magnus ssp. guadalupensis
southern waternymph
N. marina L.
holly-leaf waternymph, hollyleaved waternymph, spiny
naiad, bushy pondweed
Family Orchidaceae
Psychilis krugii (Bello) Sauleda**
Krug’s butterfly orchid,
Krug’s peacock orchid
Remarks.—This endemic
orchid was common in the
mangroves and dry forests of
southwestern Puerto Rico,
but it has been almost
collected out of the
mangroves. Specimens
deposited in The New York
19
Botanical Garden herbarium
(NY) with accession numbers
59135 to 59149.
Family Palmaceae
Cocos nucifera L.*
Sabal causiarum (O.F. Cook) Becc.**
Family Polygonaceae
Coccoloba uvifera (L.) L.
coconut palm, coconut;
palma de coco(-s), cocotero,
coco
Puerto Rican palmetto,
Puerto Rico palmetto, hat
palm, hat palmetto, straw-hat
palm; palma de sombrero,
yarey
seagrape, sea grape; uvero,
uvera, uva de playa, uvas
Family Pteridaceae
Acrostichum aureum L.
golden leatherfern; helecho
de río, helecho de pantano,
palmita del río
giant leatherfern, giant fern,
swamp fern, inland
leatherfern; helecho de
pantano, helecho gigante de
pantano
water fern, Guayanan
waterclover; helecho de agua
A. danaeifolium Langsd. & Fisch.
Marsiela polycarpa Hook. & Grev.
Family Rhizophoraceae
Rhizophora mangle L.
red mangrove, American
mangrove, mangrove; mangle
rojo, mangle colorado,
mangle zapatero, mangle de
chifle
Family Rosaceae
Chrysobalanus icaco L.
coco-plum, cocoplum coco
plum; icaco, hicaco, jicaco
Family Rubiaceae
Ernodea littoralis Sw.
Family Ruppiaceae
Ruppia cirrhosa (Petag.) Grande
beach creeper, coughbush
spiral ditch-grass; yerba de
zanja espiral
widgeongrass, ditch-grass;
yerba de zanja
R. maritima L.
20
Family Scrophulariaceae
Bacopa monnieri (L.) Pennell
coastal water-hyssop, herb of
grace; yerba de culebra
Family Simaroubaceae
Suriana maritima L.
bay cedar, baycedar;
guitarrán, quitarán,
temporana
Family Typhaceae
Typha domingensis Pers.
southern cattail, cat tail;
eneas, enea, aeneas, yerba de
eneas
Family Umbelliferaceae
Centella erecta (L. f.) Fern
Hydrocotyle bonariensis Lam.
H. hirsuta Sw.
H. umbellata L.
erect centella; yerba de clavo
largeleaf pennywort
yerba de clavo
marsh pennywort; ombligo de
Venus, sombrerillo de agua,
yerba de cuarto
Family Verbenaceae
Lantana camara L.
L. camara var. aculeata (L.) Mold.
yellow sage; cariaquillo
pink sage, prickly sage;
cariaquillo espinoso
L. involucrata L.
button sage, wild sage;
cariaquillo Santa María,
Santa María
L. montevidensis (Spreng.) Briq.
trailing lantana, trailing
shrubverbena; cariaquillo
rastrero, cariaquillo de
canastas
Lippia nodiflora (L.) Michx.
common fog fruit, common
fiddlewood, northern fogfruit, cape-turkeytangle, capeweed, Turkey tangle fogfruit,
Turkey-tangle, fog fruit,
weed; cidrón, yerba de Sapo;
hierba de la Virgen María
________________________________________________________________________
* Exotic species; ** Endemic species; *** New record for Puerto Rico.
Mangroves are exceptionally adaptable and therefore can survive under a relatively
21
wide range of environmental conditions. Some species have been found in freshwater ponds
well above sea level. Environments suitable for mangrove growth occur where temperatures
are warm and frosts are infrequent, where the shoreline is protected from wave action and
pounding surf, or behind protective natural formations such as dunes (Martínez, 1988) and
on coastal plains where the influence of seawater occurs. They develop best where they also
receive terrestrial runoff and periodic flooding by river discharge (e.g., estuaries). Where the
tides are large and the coast is of low relief, salt water can intrude inland for long distances,
and mangrove coverage may be very extensive (Cintrón et al., 1978; Vázquez, 1983). One
major biological characteristic of mangrove forests is their homogeneity. Often, a single tree
species is monodominant over large areas.
The purpose of this study is to offer further information on the introduced species
Conocarpus erectus var. sericeus (Combretaceae), the unusual aerial roots formation in A.
germinans, and paleobotanical notes on mangrove-like plants in Puerto Rico.
CONOCARPUS ERECTUS VAR. SERICEUS: AN INTRODUCED MANGROVEASSOCIATED PLANT
The buttonwood mangrove (C. erectus) grows along the coast of west tropical Africa,
the tropic and subtropical coasts of the Atlantic, including the West Indies and the Pacific
coast of North and South America. A pubescent-leaf variety of C. erectus known as the
silver buttonwood, C. erectus var. sericeus, was reported originally from the Bahamas,
southern Florida, and northern Central America (Stemple, 1970; Miller and McRitchie, 1973;
Schubert, 1979). In recent times, C. erectus var. sericeus has been introduced into the
Greater and Lesser Antilles (Stearn, 1958; Stemple, 1970; Nieves-Rivera, unpubl. data).
This report is a new record for Puerto Rico and the mangrove habitat.
The study areas for the data herein reported were Boquerón Beach Inlet (BBI)
(18°00’95.7”N, 67°10’24.9”W), located west of the Boquerón Wildlife Refuge, Boquerón
Commonwealth Forest (BCF) and Magueyes Island Marine Laboratories (MIML)
(17°58’24.3”N, 67°02’71.8”W), located in La Parguera Commonwealth Forest), both in
22
southwestern Puerto Rico, at mean sea level. The general environment of BBI and MIML is
classified as a subtropical dry forest (Ewel and Whitmore, 1973). Further details on
climatology, ecology, geology, and edaphic formations of BBI and MIML are available in
Vázquez (1983), Volckmann (1984), Toro and Colón (1986), Torres-Figueroa (1993),
Vázquez and Kolterman (1998), Winter et al. (1998), and Nieves-Rivera et al. (2002).
Rhizophora mangle and A. germinans occasionally form thick coastal forests in BCF, as well
as on the rest of the southwestern coast of Puerto Rico (Cintrón et al., 1978; Lugo, 1989;
Vázquez and Kolterman, 1998) (Figures 7A-B and 8A-C).
Conocarpus erectus var. sericeus is frequently a shrub with alternate leaves which
generally lives on the landward side of the tidal mangrove swamps. The leaves are leathery
or pubescent, fleshy, lance-shaped or elliptical 3.0 to 7.6 cm long x 0.6 to 3.2 cm wide, longpointed at both ends, and yellow green on both sides. This species characteristically forms
small evergreen trees up to 6.0 m in height and 20 cm in trunk diameter, sometimes reaching
a larger size or occurring only as a low shrub, with a spreading crown. Stemple (1970)
reported three C. erectus morphotypes: one is pubescent or silky, sometimes with silvery
hairy foliage (type 1), glabrous throughout (type 2), combination of both or ‘cyclic’ (type 3).
The pattern formed by the hairs on the leaves in C. erectus var. sericeus is similar to that of
the coastal plants Spiraea ulmaria, Borrichia arborescens (Stemple, 1970), and to that of the
paramo-adapted plants Espeletia spp. and Senecio spp. (Rundel et al., 1994). In southern
Florida the variety with pubescent or silvery hairy foliage is usually grown as an ornamental.
This variety will grow on dry land well away from the seashores (Little et al., 2001).
Conocarpus erectus var. sericeus has been introduced in late 1970’s into Puerto Rico
as an ornamental and seedlings might be bought from commercial nurseries throughout the
island (Figures 9A–D). Schubert (1979) recommended 46 trees species useful for shade and
ornament in Puerto Rico and the Virgin Islands, including C. erectus var. sericeus. However,
he (Schubert, op. cit.) did not reported C. erectus var. sericeus as a new record for Puerto
Rico. Because of anthropogenic forces, such as plant commerce and human introductions,
the pubescent morphotype may intermingle with native coastal vegetation and eventually
23
become part of the landscape. In this way, C. erectus var. sericeus expands its distribution
throughout the Caribbean. So far, there is no attempt to restrict, control, or eradicate this
new variety in Puerto Rico.
Phytopathogens, such as fungi (e.g., Cylindrocladium
scoparium) might be introduced in Puerto Rico as a causal agent of a disease of silver
buttonwood as in Florida (see Miller and McRitchie, 1973).
AERIAL ROOTS IN AVICENNIA GERMINANS
Avicennia germinans is a small evergreen tree or shrub 3 to 12 m high, attaining a
trunk diameter of 30 cm in Puerto Rico, and characterized by a rounded crown of spreading
branches (Little et al., 2001) (Figures 8A–C). This species seems hardier than the other
mangroves, to which it is not related. It penetrates farther inland along rivers and in the
United States ranges farther northward, beyond the tropical zone. In Puerto Rico it appears
to withstand prolonged flooding better than white mangrove (Laguncularia racemosa) (Little
et al., 2001). The advancing thickets of mangroves with networks of roots collect and hold
silt, thus building up the shores.
In A. germinans, numerous pneumatophores often rise vertically from the long
horizontal roots in the mud under a tree, perhaps aiding in bringing air to the roots. Little et
al. (2001) reported masses of roots 7.6 to 46 cm long sometimes hanging in the air from the
upper part of large trunks, while discussing A. germinans general morphology. They (Little
et al., op. cit.) do not discussed location or quantity of these aerial root masses. A total of 15
pendulous aerial roots in two A. germinans trees were first noticed by oceanographer Jorge E.
Corredor (pers. comm.) during a walk to his office located on MIML the afternoon of May 8,
2003. MIML aerial roots of A. germinans were photographed by the senior author afterwards
(Figure 8C). These aerial roots were single or bifurcated, 1.4 to 16.0 cm long x 6.0 to 11.0
cm in diameter and protruded from large tree trunks. The morphology of aerial roots in A.
germinans was treated by Snedaker et al. (1981) and Tomlinson (1986). Avicennia
germinans aerial roots appear to occur only rarely. We could not determine what caused the
24
development of these roots in the present case. Snedaker et al. (1981) considered that this
development was a response to oil pollution.
PALEOBOTANICAL NOTES ON MANGROVE-LIKE PLANTS
The most common trace fossils (also known as ichnofossils or “Lebensspuren”,
‘living traces’ in German) left by plant activity are root traces or casts (rhizoliths), which
show the branching and irregularities of living root morphology. These casts also reveal
plant behavior, by showing growth. Their preservation taphonomy was probably the result of
early cementation around the original roots, followed by cementation of carbonate sand that
filled the mold (Martin, 1996). The fossilized mangrove-like plants of Puerto Rico have been
little studied. A total of 11 trace fossils (ichnofossils) of mangrove-associated plants are
herein reported for Puerto Rico (Table II).
The earliest root casts in Puerto Rico were dated Late Cretaceous (probably
Santonian, 85.8 to 83.5 Ma; Santos, 1990, 1999). These plant ichnofossils (Figures 10A–E),
along with scattered large, near-vertical trace fossils (Skolithos s. str. Prothero and Schwab,
1996) are present on the top of bioturbated magnetite-rich units (magnetite lenses) of the
oldest facies of the Cotuí Limestone, Cabo Rojo-San Germán, in southwestern Puerto Rico
(Santos, 1990, 1999). The location of the Cotuí Limestone root casts is given at Table II and
was at 25.3 m elevation. Cretaceous root casts, have been tentatively interpreted to have
been caused by mangrove-like plants (e.g., Deltoidospora sp. [= Acrostichum] or the extinct
Brevitricolpites sp.), although recent palynological studies reported the origin of these two
candidates in the Eocene (Tomlinson, 1986; Rull, 1998). Another possible candidate is the
mangrove-associated palm Nypa which nowadays is widely distributed through southwestern
Asia and dates back to Late Cretaceous (Tomlinson, 1986; Rull, 1998). These casts (2 to 20
mm wide), which Santos suggested as were derived from some mangrove-like plants, possess
many anatomical traits also found in modern mangrove-associated plants (e.g., Acrostichum
spp.). The original wood material was decomposed and no longer exists; however, the
spaces were filled with soft sediment, easily removed by physical and biogeochemical
25
mechanisms (Figures 10C–E). These features are characteristic of intertidal to shallow
subtidal marine environments with high energy (active waves and currents) (Santos, 1990,
1999).
In the 1920s, Hollick (1928) began a paleobotanical survey in Puerto Rico, which was
summarized in his “Paleobotany of Porto Rico”, based on his field collections of seeds,
leaves, and wood macrofossils of various species (including mangrove species such as
Rhizophora) in the gray shale walls of the Collazo and Guatemala Rivers of the San
Sebastián Formation (SSF), of Oligocene in age (33.7 to 23.8 Ma). These plant species were
also collected in other localities around the island (Hollick, 1928). Hollick (op. cit.) reported
leaves of Rhizophora sp. (Rhizophora (?) doctrinalis) (Figure 5b, plate 82 of Hollick, 1928)
from station B (at the base of the first falls below the bridge) in the gray shales of the Collazo
River, SSF, northwestern Puerto Rico. A type (YPM 27218), paratype (YPM 27196), and
figure (YPM 27199) of R. (?) doctrinalis were deposited in the Yale Peabody Museum
(Table II).
Palynological studies in Puerto Rico were conducted by Graham and Jarzen (1969)
and Graham (1996, 2003), collecting at many of Hollick’s original Tertiary surveyed sites,
and adding new ones. Palynomorphs (pollen) of these plant species were collected in the
shales and organic-rich silty limestone layers of the San Sebastián and Lares Formations,
northwestern Puerto Rico.
In their study, Graham and Jarzen (1969) obtained 165
palynomorphs; 44 were identified and 15 are unknown. Graham (1996, 2003) summarized
Hollick’s works and demonstrated that there is further need for paleobotanical studies in the
region on the diversity and importance of the fossil plant record of Puerto Rico. Graham
(1995) carried out palynological studies in the Caribbean, with emphasis on the
diversification of the Gulf/Caribbean mangrove communities, especially before and after the
appearance of the Isthmus of Panama in the Pliocene (5.3 to 1.8 Ma) (s. str. Graham, 1992).
Although fossil pollen of Avicennia has not been collected in Puerto Rico, this
mangrove was the first to be found in the Late Miocene (11.2 to 5.3 Ma) of the Caribbean
(Graham, 1995), although Duke (1995) reported Avicennia in Early Miocene (23.8 to 16.4
26
Ma). Avicennia pollen is also common in the Quaternary (1.8 to 0.01 Ma) of Costa Rica,
Panama, and in northern South America (Müller et al., 1987; Graham, 1995). Microfossils
of Rhizophora first appeared in the Late Eocene (37.0 to 33.7 Ma), Avicennia in Late
Miocene, Laguncularia in the Pliocene (5.3 to 3.6 Ma), and Conocarpus in the Quaternary
(Graham, 1995).
More recent palynological surveys by Graham and Jarzen (1969) and Graham (1995,
1996, 2003), reported Puerto Rican Tertiary (Middle Oligocene) mangrove plant
microfossils. Mangrove pollen included Rhizophora sp. and Pelliciera sp. (Graham and
Jarzen, 1969; Graham, 1995, 1996, 2003). Palynomorphs of both mangrove species were
collected in the light gray and gray shales of SSF. Both mangrove species are typical of
coastal habitats with brackish or marine waters; however, Pelliciera rhizophoreae is now
limited to the Pacific coasts of Costa Rica, Panama, and both Pacific and Atlantic coasts of
Colombia (Tomlison, 1986).
Oligocene and Miocene lignitic rocks, traces of amber, and trace fossils (possibly
mangrove rhizoliths) also have been found in several geological formations of northern and
southern Puerto Rico (e.g., Juana Díaz Formation and Ponce Limestone s. str. Frost et al.,
1983) (Iturralde-Vinent, 2001; MacPhee and Wyss, 1990; MacPhee and Iturralde-Vinent,
1995; Nieves-Rivera, unpublished data, 2002) (Table II). The biostratigraphy of the (Holiday
Inn) outcrop, which is part of the Ponce Limestone, is Late Miocene in age is herein reported
(María Ruiz-Yantín, pers. comm., 2004). During Late Miocene times, this area was active
tectonically, whereas the northern part of Puerto Rico was passive. The outcrop has four
units. The first unit (Unit 1) shows a lagoon environment and it is 1.2 m in thickness. It is a
wackestone composed mainly of the foraminifer Miosorites cf. americanus, solitary corals,
Pecten with original shells, and internal gastropod molds. Unit 2 represents a reef front
environment and is 4.5 m thick. It is a massive unit (packstone) containing solitary corals in
growth position, colonial corals such as Diploria and Porites, unidentified burrows, Pecten,
crabs remains, M. cf. americanus foraminifers, rhizoliths of mangrove origin (LACMIP
locality 17772), and internal gastropods molds. It shows a coral framework. Unit 3
27
represents a reef crest environment and is 4.5 m in thickness. The corals are out of place and
are not as well cemented as in the underlying unit. There are solitary corals, Montastrea
annularis, Porites porites, brain corals that are probably Diploria. Also there are few
gastropods, bivalves, and crabs remains. There is an erosion surface between unit 3 and 4.
Unit 4 is 8.05 m in thickness and shows a lagoon environment similar to Unit 1 (María RuizYantín, personal communication, 2004).
Pleistocene eolianite terraces (s. str. Taggart, 1992) that occur in southwestern Mona
Island and northern Puerto Rico jut out from the coastline. The shoreline of Mona Island
terraces sometime shows “dead mangrove roots” that protrude from crannies and fissures,
and coral fossils (e.g., M. annularis, M. cavernosa, Diploria sp., Acropora palmata) are
found everywhere (Hernández-Ávila, 1970; Taggart, 1992). Punta Jacinto, located in Playa
Jobos in Isabela, northern Puerto Rico, is a typical example of aeolian fossilized dunes
having plant root casts (rhizoliths). The Isabela rhizoliths were deposited in the Natural
History Museum of Los Ángeles County, Department of Invertebrate Paleontology, Los
Ángeles, California (LACMIP). Pleistocene Isabela rhizoliths (see LACMIP location 17768)
are possibly from a mangrove-associated plant such as Scaveola sp. (Goodeniaceae) (Storrs
L. Olson, pers. comm., 2005) (Figures 11A–F) (Table I and II). Similar rhizoliths have been
found as reefs at Key Biscayne Bay in Florida (U.S.A.) (Hoffmeister and Multer, 1965), in
aeolian dunes of Hawaii (Olson and James, 1982), and San Salvador Island in the Bahamas
(Martin, 1996).
To date, the data on ichnofossils of mangrove-like plants are too poor to speculate on
such subjects as dispersal and island biogeography, especially in Puerto Rico. Most of the
taxonomic information presented herein was collected from fragmentary surveys from Puerto
Rico. However, the data herein suggest that the composition of fossilized mangrove-like
plants is more complex than previously suspected. Therefore, it would seem important to
continue studying such paleoenvironments, in order to contribute to the conservation and
knowledge of the paleoecology of mangrove-like plants of Puerto Rico.
28
In conclusion, a compilation of 125 plants routinely found in the mangrove forest, the
introduction of a new variety of mangrove-associated plant, Conocarpus erectus var. sericeus
(Combretaceae), unusual aerial roots in Avicennia germinans (Avicenniaceae), and several
new trace fossils (ichnofossils) of mangroves are reported for Puerto Rico. An actualized
checklist of scientific and common botanical names (English/Spanish), including 3 true
mangroves (A. germinans, L. racemosa, and R. mangle) and 122 mangrove-associates are
herein reported for Puerto Rico.
29
Table II. Location of mangroves and mangrove-associated plants palynomorphs (PL) or ichnofossils (IF) found in Puerto Rico. They
are placed in order of geologic time.
______________________________________________________________________________________________________
Taxa
Accession
Coordinates
Geological Geologic References
1
Numbers
Formation2
Time3
______________________________________________________________________________________________________
Mangrove-like root casts (IF)
18°04’72.0” N, 67°05’31.2” W
CM
LC
Santos (1990,
1999); NievesRivera (Unpubl.
data)
MO C-47, 15; ESF H-19
18°19’99.8” N, 66°56’91.8” W
SC
MO
Graham &
Jarzen (1969)
MO C-42, 4; ESF N-43
18°19’99.8” N, 66°56’91.8” W
SC
MO
Graham &
Jarzen (1969)
MO C-48, 1; ESF D-38, 2
18°19’99.8” N, 66°56’91.8” W
SC
MO
Graham &
Jarzen (1969)
Rhizophora (?) doctrinalis Hollick (IF)
YPM 27218 (type)
YPM 27196 (paratype)
YPM 27199 (figured)
18°19’99.8” N, 66°56’91.8” W
SC
MO
Hollick (1928);
Graham (1996)
Ilex sp. (PL)
Myrica sp. (PL)
Pelliciera sp. (PL)
Rhizophora sp. (PL)
-------
MO C-47, 1; ESF L-21, 3
30
18°19’99.8” N, 66°56’91.8” W
SC
MO
Graham &
Jarzen (1969)
MO A-12, 10; ESF U-34, 4
18°17’41.3” N, 66°53’41.1” W
SL
MO
17°58’41.2” N, 66°53’73.0” W
BL
MO
Graham &
Jarzen (1969)
Nieves-Rivera
(Unpubl. data)
-------, LACMIP 17772
17°58’94.7” N, 66°40’02.8” W
PC
LM
Olson and
James (1982);
Nieves-Rivera
(Unpubl. data)
Scaveola plumieri rhizoliths (IF)* LACMIP 17768
18°30’87.3” N, 67°04’50.1” W
ED
PL
Mangrove-like root casts (IF)
EF
QT
Nieves-Rivera
(Unpubl. data)
Hernández-Ávila
(1970); NievesRivera (Unpubl.
data)
Tournefortia sp. (PL)
Mangrove-like root casts (IF)*
Mangrove-like root casts (IF)*
-------
-------
18°04’85.4” N, 67°56’37.9” W
* = New records for Puerto Rico; ESF = England Slide Finder coordinates. 1Museum Abbreviations: LACMIP = Natural History
Museum of Los Ángeles County, Department of Invertebrate Paleontology, Los Ángeles, California; MO = Herbarium, Missouri
Botanical Garden, Saint Louis, Missouri; YPM = Yale Peabody Museum Herbarium, Yale University, Cincinnati. 2Geological
Formation: BL = Barrio Luna, Juana Díaz Formation, Guánica; CM = Cotuí Mountains, Cotuí Formation, Cabo Rojo-San Germán; ED
= Eolianite dunes, Punta Jacinto, Isabela; EF = Eolianite Formation, Carabinero, Mona Island; PC = Ponce Cement Quarry, Ponce
Limestone, Ponce; SC = Salto Collazo, San Sebastián Formation, San Sebastián; SL = Slope, junction of Roads PR-111 and PR-124,
San Sebastián Formation, Lares. 3Geologic Time: LC = Late Cretaceous (Santonian, 85.8-83.5 Ma); LM = Late Miocene (11.2-5.3
Ma); MO = Middle Oligocene (28.5 Ma); PL = Pleistocene (1.8-0.01 Ma); QT = Quaternary (1.8 Ma to 10,000 yr).
31
LITERATURE CITED
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Smithsonian Institution, Washington, D. C. 491 pp.
Anonymous. 2001. Guide to identify common wetland plants in the Caribbean area: Puerto
Rico and the U.S. Virgin Islands. A Joint Publication of the Commonwealth of
Puerto Rico Department of Natural and Environmental Resources (DNER), DOT
Federal Highway Administration, DTPW Puerto Rico Highway and Transportation
Authority, USDA Natural Resources Conservation Service, USDI Fish and Wildlife
Service and USVI Department of Planning and Natural Resources. Editorial de la
Universidad de Puerto Rico, Río Piedras, Puerto Rico. 268 pp.
Cintrón, G., C. Goenaga and J. González-Liboy. 1978. Ecología del manglar en una zona
árida: exposición al oleaje y estructura del manglar. In Departamento de Recursos
Naturales de Puerto Rico (ed.), Quinto simposio de los recursos naturales, pp. 57-86.
Estación Experimental Agrícola, Río Piedras, Puerto Rico.
Duke, N. C. 1995. Genetic diversity, distributional barriers and rafting continents—more
thoughts on the evolution of mangroves. Hydrobiologia 295: 167-181.
Ewel, J. J. and J. L. Whitmore. 1973. The ecological life zones of Puerto Rico and the U. S.
Virgin Islands. Forest Service Research Paper ITF-18. U.S. Forest Service, Institute
of Tropical Forestry, Río Piedras, Puerto Rico. 72 pp., 1 map.
Francis, J. K. and C. A. Lowe, eds. 2000. Bioecología de árboles nativos y exóticos de
Puerto Rico y las Indias Occidentales [Spanish translation of “Silvics of native and
exotic trees of Puerto Rico and the Caribbean islands”]. General Technical Report
IITF-15. U.S. Department of Agriculture, Forest Service, International Institute of
Tropical Forestry, Río Piedras, Puerto Rico. 582 pp.
Frost, S. H., J. L Harbour, D. K. Beach, M. J. Realini and P. M. Harris. 1983. Oligocene
reef-tract development, southwestern Puerto Rico (Sedimenta 9). The Comparative
Sedimentology Laboratory, Division of Marine Geology and Geophysics, Rosenstiel
School of Marine & Atmospheric Science, University of Miami, Florida. 144 pp.
32
Graham, A. 1992. Utilization of the isthmian land bridge during the Cenozoic—
paleobotanical evidence for timing, and the selective influence of altitudes and
climate. Review of Palaeobotany and Palynology 72: 119-128.
Graham, A. 1995. Diversification of Gulf/Caribbean mangrove communities through
Cenozoic time. Biotropica 27: 20-27.
Graham, A. 1996. Paleobotany of Puerto Rico: from Arthur Hollick’s (1928) scientific paper
to the present. Pp. 103-114 in J. C. Figueroa Colón, ed. The scientific survey of
Puerto Rico and the Virgin Islands: an eighty-year reassessment of the island’s
natural history. New York: Annals of the New York Academy of Science 776.
Graham, A. 2003. Historical phytogeography of the Greater Antilles. Brittonia 55: 357-383.
Graham, A. and D. M. Jarzen. 1969. Studies in Neotropical paleobotany. I. The Oligocene
communities of Puerto Rico. Annals of the Missouri Botanical Garden 56: 308-357.
Hernández-Ávila, M. L. 1970. Beach studies at Isla Mona. M.Sc. Thesis, University of
Puerto Rico, Mayagüez, Puerto Rico. 171 pp.
Hoffmeister, J. E. and H. G. Multer. 1965. Fossil mangrove reef of Key Biscayne, Florida.
Geological Society of America Bulletin 76: 845-852.
Hollick, C. A. 1928. Paleobotany of Porto Rico. Scientific Survey of Porto Rico and the
Virgin Islands. Vol. 7, part 2. New York Academy of Science, New York. Pp. 177393.
Iturralde-Vinent, M. A. 2001. Geology of the amber-bearing deposits of the Greater Antilles.
Caribbean Journal of Science 37: 141-167.
Little, E. L., Jr. and R. O. Woodbury. 1980. Rare and endemic trees of Puerto Rico and the
Virgin Islands. U.S. Department of Agriculture Conservation Research Report 27: 126.
Little, E. L., Jr., F. H. Wadsworth and J. Marrero. 2001. Árboles comunes de Puerto Rico y
las Islas Vírgenes, segunda edición revisada. Editorial de la Universidad de Puerto
Rico, Río Piedras, Puerto Rico. 764 pp.
Lugo, A. E. 1989. Los manglares de La Parguera. Acta Científica 3: 135-140.
33
Lugo, A. E. and S. C. Snedaker. 1974. The ecology of mangroves. Annual Review of
Ecological Studies 5: 39-64.
MacPhee, R. D. E. and A. R. Wyss. 1990. Oligo-Miocene vertebrates from Puerto Rico, with
a catalog of localities. American Museum Novitates 3141: 1-31.
MacPhee, R. D. E. and M. A. Iturralde-Vinent. 1995. Origin of the Greater Antillean land
mammal fauna, 1: New Tertiary fossils from Cuba and Puerto Rico. American
Museum Novitates 3141: 1-31.
Martin,
A.
J.
1996.
Plant
trace
fossils.
http:ww.emory.edu/COLLEGE/ENVS/research/ichnology/tf-plants.htm (Accessed
from World Wide Web in 15 November 2004).
Martínez, R. F. 1988. Las playas y dunas de Puerto Rico. Pp. i-viii, 1-114 in J. L. Vivaldi,
ed. Compendio enciclopédico de los recursos naturales de Puerto Rico, Tomo I, Vol.
IV,. Departamento de Recursos Naturales de Puerto Rico, Programa de Manejo de la
Zona Costanera de Puerto Rico. San Juan, Puerto Rico: Editorial Librotex, Inc.
Martorell, L. F., A. H. Liogier and R. O. Woodbury. 1981. Catálogo de los nombres vulgares
y científicos de las plantas de Puerto Rico. University of Puerto Rico Agriculture
Experiment Station Bulletin 263: 1-231.
Más, E. G. and O. García-Molinari. 1990. Guía ilustrada de yerbas comunes de Puerto Rico.
Colegio de Ciencias Agrícolas, Servicio de Extensión Agrícola, Universidad de
Puerto Rico, Mayagüez, Puerto Rico. 103 pp.
Miller, J. W. and J. J. McRitchie. 1973. Cylindrocladium scoparium, causal agent of a new
disease of silver buttonwood and its control. Plant Disease Reporter 57: 500-503.
Müller, J., E. di Giacomo and A. W. van Erve. 1987. A palynological zonation for the
Cretaceous, Tertiary and Quaternary of northern South America. American
Association of Stratigraphy and Palynology Contribution Series 19: 7-76.
Nieves-Rivera, Á. M., T. A. Tattar and E. H. Williams, Jr. 2002. Sooty mould-planthopper
association on leaves of the black mangrove Avicennia germinans (L.) Stearn in
southwestern Puerto Rico. Arboricultural Journal 26: 1-16.
34
Olson, S. L. and H. F. James. 1982. Prodromus of the fossil avifauna of the Hawaiian
Islands. Smithsonian Contribution to Zoology 365: 1-59.
Prothero, D. R. and F. Schwab. 1996. Sedimentary geology— an introduction to sedimentary
rocks and stratigraphy. W. H. Freeman and Company, New York. 575 pp.
Rull, V. 1998. Evolución de los manglares neotropicales: la crisis del Eoceno. Interciencia
23: 355-362.
Rundel, P. W., A. P. Smith and F. C. Meinzer, eds. 1994. Tropical alpine environments:
plant form and function. Cambridge University Press, Cambridge. 390 pp.
Santos, H. 1990. The stratigraphy, paleoenvironments, and biofacies of the Cotuí Limestone:
Cretaceous of southwestern Puerto Rico. M.Sc. Thesis, University of Colorado,
Boulder, Colorado. 153 pp.
Santos, H. 1999. Stratigraphy and depositional history of the Upper Cretaceous strata in the
Cabo Rojo-San Germán structural block, southwestern Puerto Rico. Ph.D.
Dissertation, University of Colorado, Boulder, Colorado. 185 pp.
Schubert, T. H. 1979. Trees for urban use in Puerto Rico and the Virgin Islands. U.S.
Department of Agriculture Forest Service General Technical Report SO-27:1-91.
Stemple, J. C. 1970. The distribution of pubescent leaved individuals of Conocarpus erectus
(Combretaceae). Rhodora 72: 544-547.
Snedaker, S. C., J. A. Jiménez and M. S. Brown. 1981. Anomalous aerial roots in Avicennia
germinans (L.) L. in Florida and Costa Rica. Bulletin of Marine Sciences 31: 467470.
Stearn, W. T. 1958. A key to West Indian mangroves. Kew Bulletin 1958: 33-37.
Taggart, B. E. 1992. Tectonic and eustatic correlations of radiometrically dated Late
Quaternary marine terraces on northwestern Puerto Rico and Isla de Mona, Puerto
Rico. Ph.D. Dissertation, University of Puerto Rico, Mayagüez, Puerto Rico. 252 pp.
Tomlinson, P. B. 1986. The botany of mangroves. Cambridge Tropical Biology Series.
Cambridge University Press, New York. 413 pp.
35
Toro, J. A. and J. A. Colón. 1986. Suplemento de información técnica para el plan de manejo
del área de planificación especial del suroeste- segmento de Boquerón. Oficina de
Zona Costanera, Area de Investigaciones Científicas, Departamento de Recursos
Naturales de Puerto Rico, San Juan. 83 pp., 1 map.
Torres-Figueroa, N. 1993. Modern sediments and Holocene history of Boquerón Bay. M.Sc.
Thesis, University of Puerto Rico, Mayagüez, Puerto Rico. 135 pp.
U. S. Department of Agriculture. 2004. Plants database. National Resources Conservation
Service, U. S. Department of Agriculture. http://plants.usda.gov/index.html
(Accessed from World Wide Web in 15 November 2004).
Vázquez, M. A. 1983. The effects of impounding on a mangrove forest. M.Sc. Thesis,
University of Florida, Gainesville. 117 pp.
Vázquez, O. J. and D. A. Kolterman. 1998. Floristic composition and vegetation types of the
Punta Guaniquilla Natural Reserve– Cabo Rojo, Puerto Rico. Caribbean Journal of
Science 34: 265-279.
Volckmann, R. P. 1984. Geologic map of the Cabo Rojo and Parguera quadrangles,
southwest Puerto Rico. U.S. Geological Survey, Miscellaneous Investigative Series
1984 G, Map I-1557.
Winter, A., R. S. Appeldoorn, A. Bruckner, E. H. Williams, Jr. and C. Goenaga. 1998. Sea
surface temperatures and coral reef bleaching off La Parguera, Puerto Rico
(northeastern Caribbean Sea). Coral Reefs 17: 377-383.
36
Figures 7A–B. Rhizophora mangle (Rhizophoraceae). A. Leaves and seedling (La Parguera, southwestern
Puerto Rico) (Boquerón Commonwealth Forest, Cabo Rojo).
Figures 8A–C. Avicennia germinans (Avicenniaceae) (La Parguera, southwestern Puerto Rico). A. Flowers
and leaves. B. Adult trees. C. Aerial roots). A. Leaves and seedling (La Parguera, southwestern
Puerto Rico) (Boquerón Commonwealth Forest, Cabo Rojo).
37
Figures 9A–D. Conocarpus erectus var. sericeus (Combretaceae) from southwestern Puerto
Rico. A. Seedling from a commercial nursery from San Germán, Puerto Rico). B.
Used as ornamental in the Boquerón Beach-Cabin Complex, Cabo Rojo. C.
Pubescent leaves. D. Fruiting.
38
Figures 10A–E. Santonian (Late Cretaceous) mangrove-like root casts reported by Santos
(1990, 1999) from the Cotuí Formation, Cabo Rojo-San Germán, southwestern
Puerto Rico. A–B. Magnetite bioturbated facies (lenses) where the roots are located.
C–E. Enlargement of the root casts.
39
Figures 11A–F. Pleistocene (Late Quaternary) rhizoliths (LACMIP location 17768),
possibly root casts from the mangrove-associated plant Scaveola plumieri
(Goodeniaceae) in the aeolian fossilized dunes of Punta Jacinto, Playa Jobos, Isabela,
in northern Puerto Rico. A–B. Aeolian dune views. C–E. Rhizoliths in situ. F.
Profile and sectioned views of weathered rhizoliths.
40
CHAPTER 3
HISTORY OF MARINE MYCOLOGY IN PUERTO RICO, INCLUDING A
CHECKLIST OF COASTAL AND MANGROVE-ASSOCIATED FUNGI
ABSTRACT
The history of marine mycology in Puerto Rico is described in general, but with
particular emphasis on coastal and mangrove-associated fungi. Until recently, there have
been few formal studies of the subject due to the lack of expertise, since there are few local
marine biologists interested in marine mycology. Published and unpublished studies in
marine mycology for Puerto Rico are summarized. A checklist that contains 604 taxa of
fungi, including 65 fungal-like organisms (oomycetes and myxomycetes) is provided. These
are divided among the principal divisions of fungi as follows: 23.7% Ascomycota, 40.7%
Mitosporic fungi, 21.7% Basidiomycota, 2.5% Zygomycota, 5.1% Oomycota, 0.5%
Chytridiomycota, 0.2% Plasmodiophoromycota, and 5.6% Myxomycota. The present study
found some potentially new species. Some of these fungi have not yet been identified to
species, thus there is a possibility that they might or not represent new species. There were
13 new records for Puerto Rico generated as a result of the studies described herein (Aigialus
cf. grandis, Asteromassaria sp., Calonectria morganii, Cochliobolus pallescens, Hysterium
sp., Leptosphaeria australiensis, Strigula sp. (Ascomycota), Curvularia robusta,
Exserohilum sp., Koorchaloma sp., Stemphylium cf. gracilariae, Trimmatostroma sp.
(Mitosporic fungi), and Halophytophthora sp. (Oomycota)). Ascomycetes found in the
intertidal and submerged parts of mangroves are undoubtedly the best known marine fungi
because of their wide geographic distribution. It was also found that there is a preponderance
of mitosporic fungi to be found in the sediments, and apparently this is not due to sampling
error. Most of the coastal and mangrove associated fungi in Puerto Rico are mitosporic
fungi. This might be caused by sampling bias. This bias may due that throughout the years,
41
most of the Puerto Rican mycologists are better trained in mitosporic fungi taxonomy than in
any other fungal or fungal-like organism taxa.
RESUMEN
Se describe la historia de la micología en Puerto Rico en general, con un particular
énfasis en los hongos costeros y asociados al manglar. Hasta recientemente, ha habido pocos
estudios formales en el tema debido a la falta de interés, ya que son pocos los biólogos
marinos locales que se interesan en la micología marina. Se resumen los estudios en
micología marina tanto publicados como inéditos para Puerto Rico. Se provee una lista que
contiene 604 especies de hongos, incluyendo 65 organismos parecidos a los hongos
(oomycetes y myxomycetes). Estos están divididos de acuerdo a las divisiones principales de
los hongos como sigue: 23.7% Ascomycota, 40.7% Hongos mitospóricos, 21.7%
Basidiomycota, 2.5% Zygomycota, 5.1% Oomycota, 0.5% Chytridiomycota, 0.2%
Plasmodiophoromycota y 5.6% Myxomycota, 25.7%. En el presente estudio se encontró
algunas nuevas especies potenciales. Algunos de estos hongos no han sido identificados
hasta especie, por lo tanto cabe la posibilidad de que pudieran o no ser nuevas especies. Se
encontraron 13 nuevos registros para Puerto Rico generados como resultado de los estudios
aquí descritos (Aigialus cf. grandis, Asteromassaria sp., Calonectria morganii, Cochliobolus
pallescens, Hysterium sp., Leptosphaeria australiensis, Strigula sp. (Ascomycota),
Curvularia robusta, Exserohilum sp., Koorchaloma sp., Stemphylium cf. gracilariae,
Trimmatostroma sp. (Hongos mitospóricos) y Halophytophthora sp. (Oomycota)).
Indudablemente, los ascomicetos encontrados en las partes intermareales y sumergidas de los
mangles son los hongos marinos mejor conocidos debido a su amplia distribución geográfica.
Se encontró que hay una preponderancia de encontrar hongos mitospóricos en los
sedimentos y aparentemente esto no se debe a errores de muestreo. La mayoría de los
hongos costeros y asociados al manglar en Puerto Rico son hongos mitospóricos. Esto puede
ser causado por parcialidad en los muestreos. Esta parcialidad puede deberse a que a través
de los años, la mayoría de los micólogos puertorriqueños han sido mejor entrenados en la
42
taxonomía de los hongos mitospóricos que en otros taxones fúngicos u organismos parecidos
a los hongos.
INTRODUCTION
Fungal distribution in coastal environments in the Caribbean is still poorly known,
although a considerable number of published reports exist. For example, summaries of
previous works in the Caribbean were included in Stevenson (1975), Nishida (1989), Lodge
(1996a, b), Minter et al. (2001), and Schmit and Shearer (2003). Puerto Rican fungal
collections from coastal habitats are in some cases sporadic and interrupted by many years.
The purpose of this study is to develop a better understanding of the history of marine
mycology, including saprophytic fungal communities, collected in sea foam, leaf litter, and
wood from mangroves and mangrove-associated plant species in marine, estuarine, and
terrestrial (coastal) ecosystems in Puerto Rico, a subtropical island located between the
coordinates 18°00’–18°30’ N and 65°35’–67°15’ W, in the northeastern Caribbean Sea.
Also, a partial checklist of marine, estuarine, and terrestrial fungi (with special interest for
coastal and mangrove-associated fungi) for Puerto Rico is provided.
MANGROVES
A mangrove forest is considered a dynamic ecotone (or transition zone) between
terrestrial and marine habitats. In its simplest sense, "mangrove" is used as a generic term
referring to a group of woody, halophylic plants that occur along sheltered tropical and
subtropical coastlines. Mangroves are derived from a variety of plant taxa and vary in their
dependence upon littoral habitats. Mangrove forests are also referred to as mangrove
swamps, tidal forests, tidal swamp forests or mangals. Caribbean mangroves range from 30°
N latitude (northern Florida) to 8° N latitude (northwest Colombia) and from 59° W
longitude (north of Guiana) to 89° W longitude (eastern Guatemala) (Tomlinson, 1986).
Only four mangrove species occur in the Caribbean; the most widely distributed are the red
mangrove (Rhizophora mangle), black mangrove (Avicennia germinans), white mangrove
43
(Laguncularia racemosa), and the buttonwood (Conocarpus erectus) (Lugo and Snedaker,
1974; Cintrón and Schaeffer-Novell, 1988).
Mangroves are exceptionally adaptable and therefore can survive under a relatively
wide range of environmental conditions. Some species have been found in freshwater ponds
well above sea level. Environments suitable for mangrove growth occur where temperatures
are warm and frosts are infrequent, where the shoreline is protected from wave action and
pounding surf, or behind protective natural formations like dunes (Martínez, 1988) and on
coastal plains where the influence of sea water is felt. They develop best where they also
receive terrestrial runoff and periodic flooding by river discharge (e.g., estuaries). Where the
tides are large and the coast is of low relief, salt water can intrude inland for long distances,
and mangrove coverage may be very extensive (Cintrón et al., 1978; Vázquez, 1983). One
major biological characteristic of mangrove forests is their homogeneity. Often, a single tree
species is monodominant over large areas.
Mangrove plants generate a large amount of litter in the form of branches,
inflorescences, leaves, twigs, and other debris. The estimated annual litter production for a
mangrove forest was somewhat similar for Puerto Rico (9.45 mg.ha–1.yr–1) and Florida (8.10
mg.ha–1yr–1) (Pool et al., 1975; Twilley et al., 1986). The contribution of mangroves debris is
the input of organic matter that which enriches the coastal ecosystem and in turn the fisheries
in what would be stagnant oligotrophic tropical waters.
FUNGI
Fungi are the most important microbes in the food web. They are defined as nonphotosynthetic microbes with nuclei, usually composed of threadlike hyphae, but sometimes
budding as in yeasts (Lodge, 1996a). According to Hyde et al. (2000), there are 444 marine
species described; however, Kirk et al. (2001) reported that fungi have a widespread
occurrence in the sea, with 800 to 1000 species known to be marine. The higher numbers in
Kirk et al. (ibid.) may include also the undescribed species (Ignoti). Species are parasites or
commensals of marine algae or animals; they are also saprophytes, species found on diverse
44
marine substrata. They have been found growing on seagrasses, protozoans, driftwood,
corals, and many other substrata. Although fungal spores accumulate in sea foam, the fungi
do not actually grow there. Fungal communities also occur in brackish water, salt marshes,
mangrove swamps, saltpeter beds, sand, dunes, and in coastal plains. Many marine fungal
spores have special appendages for attachment to substrata (Kohlmeyer and VolkmannKohlmeyer, 1991).
Marine fungi have been ecologically classified by Kohlmeyer (1974) into two major
groups, the obligate (type 1) and facultative marine fungi (type 2). He defined them as:
“obligate marine fungi are those that grow and sporulate exclusively in a marine or estuarine
(brackish water) habitat; facultative marine are fungi from freshwater or terrestrial areas able
to grow also in the natural marine environment...” (Kohlmeyer, 1974). In mangrove forests,
estuaries and salt marshes, fungi are considered to be extremely important in nutrient cycling
(Kohlmeyer et al., 1995; Hyde and Lee, 1995). Numerous studies have examined the
manglicolous fungal taxonomical composition, diversity and biogeography (Johnson and
Sparrow, 1961; Kohlmeyer and Kohlmeyer, 1979; Kohlmeyer and Volkmann-Kohlmeyer,
1991, 1998; Jones and Alias, 1997; Hyde and Pointing, 2000; Schmit and Shearer, 2003),
succession (Newell, 1976; Tan et al., 1989), vertical distribution (Hyde, 1988, 1989a, b;
Hyde et al., 1993), abundance or biomass (Newell, 1992; Newell and Fell, 1992), and a
summary to manglicolous fungi ecology in general was given by Hyde and Lee (1995).
Furthermore, many studies have examined the amount of litter production in mangrove
forests (Saenger and Snedaker, 1993; Bunt, 1995), the role of microorganisms in
decomposition of leaf litter (Fell and Master, 1973, 1980; Cundell et al., 1979; Robertson et
al., 1992) and their role in the mangrove trophic web (Heald, 1971; Odum and Heald, 1972,
1975; Fell and Master, 1980; Robertson et al., 1992). The involvement of fungi in the
breakdown of mangrove leaves and wood has also been documented (Hyde, 1990).
Recent attention has been directed towards mangrove tree mortalities, which are
caused by anthropogenic misuse and unfavorable environmental conditions (Lugo and
Snedaker, 1974; Saenger et al., 1983; Jiménez and Lugo, 1985; Anderson and Lee, 1995) as
45
well as by fungal diseases (Fomba and Singh, 1991; Tattar and Wier, 2002; Schmit and
Shearer, 2003), among other biotic (e.g., bacterial and viral diseases, etc.) and abiotic factors
(e.g., adverse climatic conditions, edaphic drastic changes, pollution, etc.). Although marine,
estuarine, and terrestrial manglicolous fungi have been extensively studied in various parts of
the world (Johnson and Sparrow, 1961; Kohlmeyer and Kohlmeyer, 1979; Rollet, 1981;
Hyde and Lee, 1995; Schmit and Shearer, 2003), in many Caribbean Islands these fungi are
poorly known.
PREVIOUS WORK
Our knowledge of marine fungi in the Caribbean in general and Puerto Rico in
particular is fragmentary.
The fungi studied include marine fungi (Meyers, 1957;
Kohlmeyer, 1968, 1980; Kohlmeyer and Volkmann-Kohlmeyer, 1987; Acevedo, 1987, 1997,
2001; Calzada, 1988, 1991; Tattar et al., 1994; Wier et al., 1996, 2000; Minter et al., 2001;
Tattar and Wier, 2002; Schmit and Shearer, 2003; Nieves-Rivera and Santos-Flores, 2004),
terrestrial fungi in aquatic and coastal environments (Carvajal-Zamora, 1971a, b; HernándezVera, 1972, 1975, 1982; Stevenson, 1975; Hernández-Vera and Almodóvar, 1983, 1984;
Bunkley-Williams and Williams, 1994; Williams and Bunkley-Williams, 1996; Nagelkerken
et al., 1997a, b; Nieves-Rivera et al., 1998; Calzada, 1999; Nieves-Rivera, 1999, 2000a,
2002, 2003; Minter et al., 2001; Nieves-Rivera et al., 2002; Schmit and Shearer, 2003;
Nieves-Rivera and Stephenson, 2004; Ruiz-Suárez, 2004; Maldonado-Ramírez and TorresPratts, in press), yeasts (Valdéz-Collazo et al., 1987; Ricaurte, 1998; Ricaurte and Govind,
1999), and fungal-like organisms such as oomycetes (Rossy-Valderrama, 1955, 1956; GallerRimm, 1982).
During the Spanish governance of Puerto Rico no effort was made to study the
marine mycobiota. During the period from 1900 to mid-1950’s, some genera of halotolerant
fungi (e.g., Asteridiella spp., Cercospora spp., Gibberella spp.) having some parasitic
species and others that are commensals were reported by Heller (1900) throughout Weiss
(1950) (Figure 12). The first published records of species of true marine fungi from Puerto
46
Rico were those of Meyers (1957), Kohlmeyer (1968, 1980), Kohlmeyer and VolkmannKohlmeyer (1987), and Acevedo (1987) (Figure 12). The largest number of new records was
62 taxa reported by Hernández-Vera (1982, discussed below) (Figure 12). Other important
reports containing new records of marine fungi are quite recent (Nieves-Rivera 2004a; Ortiz
et al., 2004; Silva et al., 2004), adding 50 taxa to the 41 taxa reported by Hernández-Vera
(1972), the 32 taxa by Carvajal-Zamora (1971a, b), 30 taxa by Kohlmeyer and VolkmannKohlmeyer (1987) and Acevedo (1987), and 20 taxa by Minter et al. (2001) (Figure 12)
(Table III).
Stevenson (1975) reported 11 species of fungi from mangrove forests, mostly as
foliar diseases. He recorded Anthostomella rhizomorphae and Polyporus fulvocinereus (=
Datronia caperata) on R. mangle; Asteridiella sepulta on Avicennia nitida Jacq. (= A.
germinans); Asteridiella lagunculariae, Meliola nigra, Physalospora lagunculariae,
Schizothyrium lagunculariae, Spiropes capensis on L. racemosa; A. lagunculariae,
Cercospora conocarpi, S. capensis on C. erectus (Stevenson, 1975). He reported 6 new taxa
of halotolerant fungi (Figure 12) (Table III).
During 1979 to 1981, Hernández-Vera (1972, 1982) conducted a mycological survey
(mostly for Mitosporic fungi) covering the supralitoral, intertidal, and sublitoral zones in the
north and south coasts of Puerto Rico. He studied four beaches in the north (Guajataca,
Sardinera, Arecibo River, and Poza del Obispo) and two at the south (Caña Gorda Beach and
Enrique Reef at La Parguera). During the first year of study (1979-1980), a total of 1657
fungi were isolated and 1405 fungal isolated on the second year (1980-1981). The most
common taxon isolated at the three zones was Aspergillus spp. Of the 2974 fungal isolates
of both years, Hernández-Vera found 28 genera and 71 species, mostly mitosporic fungi,
zygomycetes, oomycetes (Stramenopilus fungi), a few yeasts (e.g., Saccharomyces and
Cryptococcus spp.), and an ascomycete (Neurospora crassa). Hernández-Vera (1982) also
developed a new culture medium (known as “Almodóvar Agar”) made of marine algae
(Acanthophora spicifera, Dictyota divaricata, Laurencia obtusa, Spyridia filamentosa, and
Ulva lactuca), agar, and sea water (35 g/L). He tested this seawater medium and the fungal
47
tolerance to salinity with the species he previously reported. Ninety-six percent (96%) of the
species tolerated salinities up to 45 g/L. It was found that the dinoflagellate Gonyaulax
tamarensis and the red alga Falkenbergia hillebrandii were found to be fungistatic to the
majority of the species previously reported (Hernández-Vera, 1972, 1982; Hernández-Vera
and Almodóvar, 1983, 1984).
Acevedo (1987) added a total of 30 genera and 41 species of obligate and facultative
marine fungi collected from driftwood, red mangrove and sandy beaches. She collected
driftwood from three reefs (Cayos Enrique, Laurel, and Turrumote), and sampled R. mangle
wood and beach sand of an offshore island (Isla Cueva), off the coast of La Parguera,
southwestern Puerto Rico. She also reported 18 species of manglicolous fungi (ascomycetes
and mitosporic fungi) from R. mangle (Table III).
In 1992, the USDA Forest Service, the Center for Forest Mycology Research, Forest
Products Laboratory in Madison Wisconsin, through a series of grants from the National
Science Foundation, began a fungal survey in the tropics and subtropics. The coordinator of
this project for Puerto Rico, the Forest Service botanist D. Jean Lodge, along with
mycologist Timothy J. Baroni of the University of New York at Cortland, and other
renowned mycologists, were surveying the fungi in the forests of Puerto Rico, especially
basidiomycetes. The project title was the “Basidiomycetes of the Greater Antilles” (Lodge et
al., 1998), and although it originally only considered the Caribbean National Forest El
Yunque, it later also included some coastal forests, mangroves, and beaches (e.g., Piñones
Commonwealth Forest). Some interesting new records and new basidiomycetous species
have been collected in beaches and sand dunes (Table III) (Lodge, 1996a; Lodge et al., 1998,
Cantrell and Lodge, 2000; Miller et al., 2000).
Nieves-Rivera et al. (1998) reported Schizophyllum commune and Hypoxylon Sect.
Hypoxylon in Avicennia nitida (A. germinans) and R. mangle, respectively, from the
Boquerón Wildlife Refuge, southwestern Puerto Rico. Other fungal checklists for coastal
environments of Puerto Rico and offshore islands have been documented in the coastal plains
48
(Nieves-Rivera et al., 1999), mangrove forests, and saltpeter bed margins (Stevenson, 1975;
Nieves-Rivera et al., 1998).
Calzada (1999) studied three phytopathogenic fungi, Pestalotiopsis disseminata,
Phoma eupyrena, and Pterosporidium rhizophorae, caused foliar signs (“signs” as defined by
Williams et al., 1993), such as spots and deterioration to the leaves in R. mangle of La
Parguera (Phosphorescent Bay and La Parguera channels). However, Aspergillus sp. and
Cladosporium sp. were found to be saprophytes (Calzada, 1999). Recently, Tattar et al.
(1994), Wier et al. (1996, 2000), and Tattar and Wier (2002) documented Cytospora
rhizophorae as a plant pathogen in R. mangle in the southwestern coast of Puerto Rico (Table
III).
Acevedo (2001) assayed marine endophytic, manglicolous, and lignicolous fungi
(e.g., Didymosphaeria rhizophorae (= Lineolata rhizophorae), Hydronectria tethys,
Hypoxylon oceanicum (= Halorosellinia oceanica), Lulworthia grandispora, Pestalotia sp.,
Xylaria spp., as well as other undetermined species of ascomycetes and mitosporic fungi) by
High Pressure Liquid Chromatograph (HPLC) for biotransformation of polycyclic aromatic
hydrocarbon (PAH) such as phenanthrene in algal and mangrove (R. mangle) substrata.
Phenanthrene is toxic and found in marine biota and petroleum. She found that 12 marine
fungi of the 30 assayed by HPLC to be able to biotransform phenanthrene (Table 4, page 49
of Acevedo, 2001); although she reported that those tests were inconclusive. Of particular
interest for microbiology are marine fungi and notably algal endophytes (e.g., Xylaria spp.)
which are potentially useful organisms for bioremediation in marine environments.
Tattar et al. (1994), Wier et al. (1996, 2000), and Tattar and Wier (2002) documented
the incidence of Cytospora rhizophorae as a plant pathogen in R. mangle in the southwestern
coast of Puerto Rico. The occurrence of C. rhizophorae cirri on Rhizophora spp. have been
reported in the Caribbean, including Puerto Rico (e.g., Magueyes Island, Los Morrillos, La
Parguera, and Boquerón Commonwealth Forest, southwestern Puerto Rico), Bahamas,
Florida, Guatemala, and Mexico (Kohlmeyer and Kohlmeyer, 1979; Shaw, 1989). Dieback
of seedlings and young plants of R. mangle caused by C. rhizophorae has been widely
49
studied in Puerto Rico (Tattar and Wier, 2002). The cirri of C. rhizophorae are at first
yellow, the orange, and those kept in herbaria dark red to almost black (Shaw, 1989) (Figures
5A–F). This species is halotolerant and considered an obligate and facultative marine fungus
on submerged and emerged host (Kohlmeyer and Kohlmeyer, 1979; Shaw, 1989).
Minter et al. (2001) gave a summary of manglicolous fungi of the Caribbean,
including Puerto Rico, updating the records of Stevenson (1975). More recently, the work of
Schmit and Shearer (2003) summarized worldwide manglicolous fungi, including early
collections in Puerto Rico and the Caribbean. Schmit and Shearer (op. cit.) reported 30
species of manglicolous fungi for Puerto Rico alone, showing the need for updated surveys.
FINAL COMMENTS
The checklist of coastal and mangrove-associated fungi is presented in Table III. This
checklist considers some works that were not reviewed by Minter et al. (2001) because of
they were not readily available. For example, some of these unpublished works (theses,
reports, and abstracts) are not available on the internet. However, many of the species on this
checklist have been reported since these earlier works were published. It seems likely that
many new coastal and mangrove-associated fungi remain to be found.
Ascomycetes found in the intertidal and submerged parts of mangroves are
undoubtedly the best known marine fungi because of their wide geographic distribution
(Hyde and Lee, 1995; Schmit and Shearer, 2003). It was also found that mitosporic fungi
dominates in the sediments, and apparently this is not due to sampling error (Schmit and
Shearer, op. cit.). Terrestrial fungi (mostly mitosporic fungi) are in general not the subjects
of most surveys according to Schmit and Shearer (2003). In Table III, most of the coastal
and mangrove associated fungi in Puerto Rico are mitosporic fungi. This might be caused by
sampling bias. This bias may due that throughout the years, most of the Puerto Rican
mycologists are better trained in mitosporic fungi taxonomy than in any other fungal or
fungal-like organism taxa.
50
The differences in substrate affinities among fungal and fungal-like organisms (e.g.,
oomycetes) are well documented. The ascomycetes, for instance, often possess appendages
and gelatinous sheaths which aid in their attachment to wood in the intertidal zone (Rees and
Jones, 1984; Kohlmeyer and Volkmann-Kohlmeyer, 1991). Mitosporic fungi are largely
lacking these appendages. This may explain why they are uncommon in the intertidal zone
and most common in sediments, where such appendages would not be needed for spores for
attachment (Schmit and Shearer, 2003). In general, basidiomycetes are rare in aquatic and
marine environments, which explain their restriction to wood above high tide. In the
terrestrial environment, basidiomycetes play an important role in the decay of plant material.
In coastal environments, however, almost all dead leaves and wood fall to the forest floor
and come into contact with seawater, which hampers the growth of most basidiomycetes.
Marine oomycetes, on the other hand, seem to be present on almost all dead intertidal
mangrove leaves, at least in the early stages of decay (Newell and Fell, 1995, 1997).
In this survey, we have found that several species potentially pathogenic to man (e.g.,
Aspergillus spp., Candida spp., Cryptococcus spp., Epidermophyton floccosum, Nectria
hematoccoca, Galactomyces geotrichum, Rhodotorula glutinis, Scopulariopsis brevicaulis,
Candida glabrata, and Trichosporon spp., Table III) that were reported in the literature,
similar to the study of González et al. (2000). However, species composition reported by
González et al. (2000) differs greatly due to the methodology and types of bait used.
González et al. (2000) isolated a total of 17 keratinophilic fungi, of which 13 were
hyphomycetes and 4 ascomycetes. González et al. (2001) found a larger variety of species in
Mexico; therefore, their checklist contains 47 ascomycetes, 14 mitosporic fungi, and 1
basidiomycete. In a Cuban survey, González et al. (2003) reported 29 marine fungi (25
ascomycetes and 4 mitosporic fungi), of which 19 were new records. In contrast, the present
checklist (Table III) contains 604 taxa of fungi, including 65 fungal-like organisms
(oomycetes and myxomycetes). These are divided among the principal divisions of fungi as
follows: 23.7 % Ascomycota, 40.7 % Mitosporic fungi, 21.7 % Basidiomycota, 2.5 %
Zygomycota, 5.1 % Oomycota, 0.5 % Chytridiomycota, 0.2 % Plasmodiophoromycota, and
51
5.6 % Myxomycota (Table III). The present study found some potentially new species.
Some of these fungi have not yet been identified to species, thus there is a possibility that
they might or not represent new species. There were 13 new records for Puerto Rico
generated as a result of the studies described herein (Aigialus cf. grandis, Asteromassaria
sp., Calonectria morganii, Cochliobolus pallescens, Hysterium sp., Leptosphaeria
australiensis, Strigula sp. (Ascomycota), Curvularia robusta, Exserohilum sp., Koorchaloma
sp., Stemphylium cf. gracilariae, Trimmatostroma sp. (Mitosporic fungi), and
Halophytophthora sp. (Oomycota)). The marine basidiomycetes Halocyphina villosa and
Nia vibrissa were expected to be found but were never encountered. This may be because
both marine basidiomycetes apparently require colder temperatures for growth than those
found in the waters of Puerto Rico.
Several phytopathogenic species have been commonly found in many of the
publications reviewed during the course of this survey (e.g., Cytospora spp.,
Helminthosporium spp., Pestaloriopsis disseminata, Phoma spp., Phytophthora palmivora,
Pythium sp., Uredo spp., Stemphylium spp., Table III). Also, 34 plasmodial slime molds
(myxomycetes) have been isolated from or collected in coastal plants, including mangroves.
This group, along with the cellular slime molds (Dictyosteliomycota and Protosteliomycota)
is the least studied group in coastal ecosystems, especially in mangrove forests. Although
freshwater myxomycetes have been reported (Shearer and Crane, 1986), there are as yet no
reports on this group from the marine environment.
Most of the marine fungal species collected are arenicolous (Arenariomyces cf.
majusculus, A. triseptatus, Corollosopora cf. colossa, C. filiformis, C. cf. pseudopulchella).
Arenariomyces triseptatus has been previously recorded from Cuba (González et al., 2003),
Mexico (Kohlmeyer, 1983), and Puerto Rico (Kohlmeyer and Volkmann-Kohlmeyer, 1987).
Halorosellinia oceanica, Lulworthia sp., and Torpedospora radiata are lignicolous species
(Kohlmeyer and Kohlmeyer, 1979). Other fungal species isolated in this survey are saprobes
living in parts of angiosperms, algae, drifting wood and other plant debris, including blades
of seagrasses.
52
Some fungal parasites have been found living on fishes (e.g., Paecilomyces spp.,
Saprolegnia spp.; Bunkley-Williams and Williams, 1994, 1995, 2004a-c; Williams and
Bunkley-Williams, 1996; Bunkley-Williams et al., 1998; Rand et al., 2000) or
commensalistic in the gut of living crustaceans (e.g., Asellaria ligiae and Enterobryus spp.;
Cafaro, 1999; White et al., 1999, 2000). Bunkley-Williams and Williams (1994, 1995)
reported saprolegniasis affecting sport fishes in artificial lakes (freshwater) and natural
lagoons (freshwater and brackish waters) of Puerto Rico. Examples of the latter are: San
José, Torrecillas, Piñones, Tortuguero, Mandri, Santa Teresa, Joyuda, and Rincón lagoons
(Bunkley-Williams and Williams (1994). Nieves-Rivera (2000a) isolated Aspergillus
fumigatus (= Fennellia flavipes), A. flavus, and Helminthosporium sp. (= Bipolaris sp.) from
the skin of a Caiman crocodilus (Crocodylia: Alligatoridae) from the Tortuguero Lagoon, a
freshwater and brackish natural lagoon located in Vega Baja, northern Puerto Rico. Cantrell
and Betancourt (1992, 1995) and Bunkley-Williams and Williams (pers. comm., 2004)
isolated Fusarium spp. from the freshwater prawn Macrobrachium rosenbergii, and
Bunkley-Williams and Williams (op. cit.) isolated Fusarium sp. rarely from freshwater fishes
in Puerto Rico.
The aquatic hyphomycete Tetraploa aristata is common in estuary Rincón Lagoon of
the Boquerón Wildlife Refuge, as reported by Kirk (1969) for Chesapeake Bay. This
microfungus is commonly found in river foam, sugarcane, and soil. Other geophilic (soilloving) mitosporic and chytrid fungi could be transported by Sahara dust and might
contribute to amphibian decline in Puerto Rico (Stallard, 2001; Burrowes et al., 2004).
Burrowes et al. (op. cit.) monitored the populations of Eleutherodactylus in Puerto Rico from
1989 through 2001; they found chytrid fungi in two species collected as early as 1976, being
the first report of chytrid fungus in the Caribbean. Analysis of weather data indicates
significant periods of drought and the decline of amphibians in Puerto Rico. There is a
possibility of a synergistic interaction between drought and the pathological effect of the
chytrid fungus on amphibian populations (Burrowes et al., 2004).
53
According to Dring (1980), the basidiomycete Clathrus crispus is the familiar West
Indian species, one of the longest known but strangely lacking an adequate description before
that by Wright (1949). Unfortunately, certain doubts arose whether Clathrus cancellatus (=
Clathrus ruber) collections were the same as C. crispus. However, we agree with Dennis
(1953), who considered that C. crispus is a tropical species with corrugated rims, different
from C. cancellatus which does not have corrugated rims, and according to Burk (1979), C.
cancellatus is not tropical (Nieves-Rivera et al., 1999). The polypore Datronia caperata was
originally described as Coriolopsis fulvocinera (and not Polyporus) from Cuba and the type
is a specimen of the common D. caperata, probably one of the most common species of
polypores in the tropical zone and rather variable as to pileus color and cover (pubescent
zones are changing with black glabrous ones according to growth conditions) (Leif
Ryvarden, personal communication, 2004).
In conclusion, the data on coastal and mangrove-associated fungi are too poor to
speculate on such subjects as dispersal and island biogeography, especially in Puerto Rico
and in the Caribbean. Most of the taxonomic information presented herein was collected
from fragmentary surveys in published and unpublished reports (e.g., technical reports,
theses, abstracts) from Puerto Rico. However, these data suggest that the taxonomic
composition of coastal and mangrove-associated fungi is more complex than previously
suspected. In other words, beaches, estuaries, sand dunes, and mangrove forests may support
a larger assemblage of fungal species than indicated by the previous published records.
Therefore, it would seem important to continue studying such habitats, in order to contribute
to the conservation and knowledge of the biodiversity of coastal and mangrove-associated
fungi of Puerto Rico.
54
Table III. Checklist of previous and recent mycological collections (including marine— obligate and facultative— and
terrestrial fungi) in mangroves and mangrove-associated plants, coastal forests, estuaries, beaches, sand dunes, and
marine habitats in Puerto Rico.1
______________________________________________________________________________________________________
Fungus
ASCOMYCOTA
Acanthostigma lantanae Theiss.
*Aigialus cf. grandis Kohlm. & S. Schatz
Substrata or
Host2
Lc
Bs, Ll, Sf
Antennospora caribbea Meyer
Df
A. quadricornuta (Cribb & J.W. Cribb) T.W. Johnson
Rh
Anthostosmella rhizomorphorae (Kunze) Berl. & Voglino, in Sacc.
Rh
Anthostosmella sp.
Pa, Un
Aphosphaeria sp. (s. str. Pérez Samot, 1986) = Apiosphaeria sp.
Apiosphaeria sp.
Am
Arenariomyces cf. majusculus Kohlm. & Volkm.-Kohlm.
Bs, Sf
A. trifurcatus (Höhn.) Kohlm.
Bs, Ll, Sf
A. triseptatus Kohlm.
Bs, Sf
55
References
Stevens (1917); Wellman (1961); Stevenson (1975);
Minter et al. (2001)
Nieves-Rivera (unpubl. data, Rincón Lagoon,
Boquerón Wildlife Refuge, southwestern Puerto
Rico)
Meyer (1957); Stevenson (1975); Minter et al.
(2001)
Kohlmeyer (1969); Stevenson (1975); Schmit and
Shearer (2003)
Stevens (1920); Seaver and Chardón (1926);
Kohlmeyer (1969); Stevenson (1975); Minter et al.
(2001); Schmit and Shearer (2003); Schmit (2004)
Lodge (1996a)
Pérez Samot (1986)
Nieves-Rivera and Santos-Flores (2004b)
Acevedo (1987)
Nieves-Rivera and Santos-Flores (2004a, b)
Asteridiella lagunculariae (Earle) Hansf.
Co, La
A. manca (Ellis & G. Martin) Hansf.
A. sepulta (Pat.) Hansf.
Mc
Av
Asterina coccolobae Ferd. & Winge
Cu
Heller (1900); Earle (1901); Stevens (1916); Toro
(1925); Seaver and Chardón (1926); Ryan (1926);
Stevens (1927, 1928); Seaver et al. (1932; Hansford
(1961, 1963); Kohlmeyer (1969); Stevenson (1975);
Minter et al. (2001); Schmit and Shearer (2003);
Schmit (2004)
Hansford (1961); Minter et al. (2001)
Stevens (1917); Chardón (1920); Toro (1925);
Seaver and Chardón (1926); Stevens (1927, 1928);
Hansford (1961, 1963); Kohlmeyer (1969);
Stevenson (1975); Minter et al. (2001); NievesRivera et al. (2002); Schmit and Shearer (2003)
Seaver and Chardón (1926); Stevenson (1975);
Francis and Lowe (2000); Minter et al. (2001)
Asterolibertia schroeteri (Rehm) Arx, in E. Müll. & Arx
*Asteromassaria sp.
Ch
Rh
Ryan (1924); Stevenson (1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, on bark of living aerial
Rhizophora mangle L. roots and trunk, south of
Magueyes Island, La Parguera, Lajas, southwestern
Puerto Rico; Figure 13).
Remarks.–Thallus deposited with the U.S. National
Fungus Collection (BPI) by ÁMNR, 29 November
2001, BPI 843791.
Astrosphaeriella aff. mangrovei (Kohlm. & Vittal) Aptroot & K.D. Hyde
Df, La, Ll, Sf
Acevedo (1987); Nieves-Rivera and Santos-Flores
(2004a, b)
Botryosphaeria quercuum (Schwein.) Sacc.
Ca, Cn, Ll
Phelps and Landgraf (1972); Stevenson (1975);
Minter et al. (2001)
Botryotinia allii (Sawada) W. Yamam., in Yamam., Oyasu & H. Iwasaki
56
Bs, Rh, Ss
Byssosphaeria schiedermayeriana (Fuckel) M.E. Barr
Cn, Un
*Calonectria morganii Crous, Alfenas & M.J. Wingf.
Ce
Minter et al. (2001); Nydia J. Rodríguez (unpubl.
data, from seawater in R. mangle roots, María Langa
Cay, Guayanilla Bay, Guayanilla, southwestern
Puerto Rico)
Remarks.–Strain deposited with the American Type
Culture Collection (ATCC) as Cladosporium
sphaerospermum Penzig. MYA-3069 (NJRR-1).
Stevenson (1975); Lodge (1996a); Minter et al.
(2001)
Nieves-Rivera (unpubl. data, Boquerón Beach and
Magueyes Island)
Remarks.– The anamorph of this species is
Cylindrocladium scoparium Morgan (s. str. Miller
and McRitchie (1973)).
Minter et al. (2001); Nieves-Rivera et al. (2002)
Capnodium sp.
Av, La, Tc
Ceratocystis paradoxa (Dade) C. Moreau = Endoconidiophora paradoxa
Chaetomastia cf. typhicola (P. Karst.) M.E. Barr
Bs, Sf
Stevens (1917); Chardón (1920); Seaver (1922);
Lodge (1996a); Nieves-Rivera and Santos-Flores
(2004b)
Remarks.–Minter et al. (2001) reported
Chaetomastia sp. in Puerto Rico (Río Grande) on
the branches of an undetermined plant.
Chaetomium globosum Kunze
Hp, Ss
Stevenson (1975); Minter et al. (2001); Silva et al.
(2004); Ortiz et al. (2004)
Chaetophoma sp.
Dv, Ll
Lodge (1996a); Acevedo (2001); Minter et al.
(2001)
Remarks.–Minter et al. (2001) reported
Chaetophoma sp. on Syngonium auritum (L.) Schott
57
*Cochliobolus pallescens (Tsuda & Ueyama) Sivan.
Ba, Bs, Ll, Sf, Ss
(= Xanthosoma undipes (C. Koch) C. Koch
(Araceae).
Nieves-Rivera (unpubl. data, Rincón Lagoon,
Boquerón Wildlife Refuge)
Schmit (2004)
Nieves-Rivera and Santos-Flores (2004a, b)
Nieves-Rivera and Santos-Flores (2004a, b)
Acevedo (1987)
Nieves-Rivera and Santos-Flores (2004b)
Corollospora cinnamomea Koch
Ss
C. cf. colossa Nakagiri & Toruka
Bs, Sf
C. filiformis Nakagiri, in Nakagiri & Toruka
Bs, Sf
C. maritima Werderm.
Bs, Sa, Sf
C. cf. pseudopulchella Nakagiri & Toruka
Bs, Sf
Coronopapilla aff. mangrovei (K.D. Hyde) Kohlm. & Volkm.-Kohlm.
Df, Sf
Nieves-Rivera and Santos-Flores (2004b)
C. cf. mangrovei (K.D. Hyde) Kohlm. & Volkm.-Kohlm. = Coronopapilla aff. mangrovei
Dactylospora haliotrepha (Kohlm. & E. Kohlm.) Hafellner
La
Acevedo (1987); Minter et al. (2001)
Dictyonella erysiphoides (Rehm) Höhn.
Cu
Stevenson (1975); Minter et al. (2001)
Didymosphaeria enalia Kohlm. = Verruculina enalia
D. rhizophorae Kohlm. & E. Kohlm. = Lineolata rhizophorae
Didymosphaeria sp.
Rh, Un
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Boquerón Wildlife Refuge and Mona
Island)
Diplotheca tunae (Spreng.) Starbäck
Od
Stevens (1917); Fitzpatrick (1927); Stevenson
(1975); Minter et al. (2001)
Dyrithium lividum (Fr.) M.E. Barr
Co
Barr (1994); Minter et al. (2001)
Emericella nidulans (Eidam) Vuill.
Bs
Hernández-Vera (1972, 1975); Minter et al. (2001)
E. unguis Malloch & Cain
Hp
Silva et al. (2004); Ortiz et al. (2004)
Endoconidiophora paradoxa (De Seynes) R.W. Davidson
Cn
Stevenson (1975); Francis and Lowe (2000); Minter
et al. (2001)
Endoxyla sp.
Un
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
58
Eutypella stellulata (Fr.) Sacc.
Fennellia flavipes B.J. Wiley & E.G. Simmons
Ca, Un
Stevenson (1975); Minter et al. (2001)
Bs, Hr, Ll, Rp, Ss Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Nieves-Rivera (2000a, b); Minter et
al. (2001)
Galactomyces geotrichum (E.E. Butler & L.J. Petersen) Readhead & Malloch
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Genicularia cytosporia (Dudd.) Rifai & R. C. Cooke
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Gibberella baccata (Wallr.) Sacc.
Bs, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Lodge (1996a)
G. fujikuroi (Sawada) S. Ito
Ma, Un
Weiss (1950); Stevenson (1975); Cantrell and
Betancourt (1995); Minter et al. (2001)
G. intricans Wollenw.
Ma, Ss, Un
Stevenson (1975); Cantrell and Betancourt (1995);
Minter et al. (2001)
G. subglutinans (E.T. Edwards) P.E. Nelson, Toussoun & Marasas
Bs, Ma, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Cantrell and Betancourt
(1995)
Halorosellinia oceanica (S. Schatz) Whalley, E.B.G. Jones, K.D. Hyde & T. Laessøe
Rh, Sf
Acevedo (2001); Nieves-Rivera and Santos-Flores
(2004b)
59
Halosphaeria cucullata (Kohlm.) Kohlm.
H. quadricornuta Cribbs & J.W. Cribbs
Bs, Rh, Sf
Nieves-Rivera and Santos-Flores (2004b)
Co, Rh, Un, Wd Kohlmeyer (1968, 1980); Stevenson (1975);
Kohlmeyer and Volkmann-Kohlmeyer (1987);
Acevedo (1987); Minter et al. (2001); Schmit
(2004)
H. salina (Meyers) Kohlm.
Ba
Acevedo (1987)
Halosphaeria sp.
Bs, Sf
Nieves-Rivera and Santos-Flores (2004a, b)
Hydronectria tethys Kohlm. & E. Kohlm.
Df, Rh
Acevedo (1987, 2001)
Hypocrea rufa (Pers.: Fr.) Fr.
Cn, Un
Stevenson (1975); Minter et al. (2001)
Hypoxylum culmorum Cooke = Kretzschmariella culmorum
H. quisquiliarum Mont.
Cn, Un
Miller (1961); Stevenson (1975); Minter et al.
(2001)
H. oceanicum S. Schatz = Halorosellinia oceanica
60
H. rubiginosum (Pers.: Fr.) Fr.
Hypoxylon section Hypoxylon
Hypoxylum sp.
*Hysterium sp.
Hysterographium sp.
Ba
Av, Rh
La, Ll, Un
Un
Un
Irene lagunculariae (Earle) Toro = Asteridiella lagunculariae
Irenina lagunculariae (Earle) F. Stevens = Asteridiella lagunculariae
Irenopsis molleriana (G. Winter) F. Stevens
Hi, Tr
I. molleriana var. major Hansf.
I. triumfettae (F. Stevens) Hansf. & Deighton
Kirschsteiniothelia aff. maritima (Linder) D. Hawksw.
Hi
Hi
Sf, Un
61
Stevenson (1975); Minter et al. (2001)
Nieves-Rivera et al. (1998)
Stevenson (1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, Magueyes Island)
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Magueyes Island)
Seaver et al. (1932); Poonyth et al. (2000); Minter et
al. (2001); Schmit and Shearer (2003); Schmit
(2004)
Minter et al. (2001)
Stevens (1916); Seaver and Chardón (1926);
Kohlmeyer (1969); Stevenson (1975); Poonyth et al.
(2000); Minter et al. (2001); Schmit and Shearer
(2003); Schmit (2004)
Minter et al. (2001); Nieves-Rivera and SantosFlores (2004a, b)
Krestzchmaria rugosa Earle = Penzigia cantareirensis
K. zonatum (Lév.) P.M.D. Martin
Cn, Un
Kretzschmariella culmorum (Cooke) Y.M. Ju & J.D. Rogers
Ba
Minter et al. (2001)
Stevenson (1975); Lodge (1996a); Minter et al.
(2001)
Kymadiscus haliotrephus (Kohlm. & E. Kohlm.) Kohlm. & E. Kohlm. = Dactylospora haliotrepha
Lecanidion atratum (Hedw.) Rabeh.
Cn, Un
Stevenson (1975); Minter et al. (2001)
Lecidea gymnocarpa Fink, in J. Hedrick
Cn
Hedrick (1930); Imshaug (1957); Minter et al.
(2001)
Lembosia coccolobae Earle
Cu
Chardón (1920); Seaver and Chardón (1926);
Stevenson (1975); Meurer-Grimes et al. (1992);
Minter et al. (2001)
L. tenella Lév.
Cu, Un
Earle (1904); Chardón (1921); Spegazzini (1923);
Ryan (1924); Stevenson (1975); Francis and Lowe
(2000); Minter et al. (2001)
Leptogium azureum (Sw.) Mont., in P.B. Webb & S. Berthelot
Cn, Un
Müller (1888); Imshaug (1957); Minter et al. (2001)
*Leptosphaeria australiensis (Cribb & J.W. Cribb) G.C. Hughes
Av, Rh
Nieves-Rivera (unpubl. data, on intertidal wood of
A. germinans and R. mangle roots, La Parguera
Channels and Magueyes Island)
Leptosphaeria sp.
Df, La, Rh
Acevedo (1987)
Lindra marinera Meyers
Bs
Acevedo (1987)
L. thalassiae Orpurt, Meyer, Boral & Simms
Th
Calzada (1988, 1991); Acevedo (1997)
Lindra sp.
Bs, Sf
Acevedo (1987); Nieves-Rivera and Santos-Flores
(2004b)
Lineolata rhizophorae (Kohlm. & E. Kohlm.) Kohlm. & Volkm-Kohlm.
Df, Rh
Acevedo (1987, 2001)
Lophionema bambusae Höhn.
Ba
Stevenson (1975); Minter et al. (2001)
62
Lophiostoma mangrovei Kohlm. & Vittal = Astrosphaeriella mangrovei
Lulworthia grandispora Meyers
Av, Rh, Th
Meyers (1957); Acevedo (1987, 2001); Minter et al.
(2001); Schmit and Shearer (2003)
L. medusa (Ellis & Everh.) Cribb & J.W. Cribb
Wd
Meyers (1957); Minter et al. (2001); Schmit and
Shearer (2003); Schmit (2004)
L. medusa var. biscaynia Meyers
Wd
Meyers (1957); Minter et al. (2001)
Lulworthia sp.
Av, Bs, Co, Cy, Hi, Rh, Sf, Th, Wd
Kohlmeyer (1980); Kohlmeyer and VolkmannKohlmeyer (1987); Minter et al. (2001); NievesRivera and Santos-Flores (2004b)
Maublancia uleana (Pazschke) Arx, in E. Müll. & Arx
Ch
Stevenson (1975); Minter et al. (2001)
Meliola ambigua Pat. & Gaillard
Lc, Le, Ls, Lv
Hansford (1961); Stevenson (1975); Minter et al.
(2001)
M. nigra F. Stevens
La
Stevens (1916, 1920); Chardón (1920); Seaver and
Chardón (1926); Hansford (1961, 1963); Kohlmeyer
(1969); Stevenson (1975); Poonyth et al. (2000);
Minter et al. (2001); Schmit and Shearer (2003);
Schmit (2004)
M. panici Earle
Se
Chardón (1920); Hansford (1961); Meurer-Grimes
et al. (1992); Stevenson (1975); Poonyth et al.
(2000); Minter et al. (2001); Schmit and Shearer
(2003)
M. triumfettae F. Stevens = Irenopsis triumfettae
Micropeltis ingae Bat. & Peres
Tc
Gómez-Acosta (1995); Minter et al. (2001)
Microsphaera sp.
Ad
Roure and Ramírez (1970); Betancourt et al. (1980)
Microthyrium lagunculariae G. Winter = Schizothyrium lagunculariae
Microthyrium sp.
Ch
Ryan (1924); Minter et al. (2001)
Moellerodiscus lentus (Berk. & Broome) Dumont
Cn, Ll, Un
Minter et al. (2001)
63
Monographella nivalis (Schaffnit) E. Müll.
Bs
Mycosphaerella chrysobalani Miles
Ch
Nectria calami (Henn. & E. Nyman) Rossman
N. episphaeria (Tode) Fr.
Cn, Un
Ss, Un
N. haematococca Berk. & Broome
Bs, Ma, Ss
N. inventa Pethybr.
N. suffula Berk & M.A. Curtis, in Berk.
Neurospora crassa Shear & B.O. Dodge
Ss
Cn, Ll, Un
Bs, Ss, Un
Neurospora sp.
Penzigia cantareirensis (Henn.) J.H. Mill.
Perisporina lantanae F. Stevens
Phylacia bomba (Mont.) Pat.
Phyllacora graminis (Pers.) Nitschke
Ad, Ma
Ba
Ls
Ca, Un
Se
64
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Miles (1917); Stevenson (1975); Minter et al.
(2001)
Rossman (1983); Minter et al. (2001)
Carvajal-Zamora (1971a, b); Stevenson (1975);
Minter et al. (2001)
Remarks.–This species was also known as
Fusarium aquaeductuum Lagerh. var. medium
Wallenw. (s. str. Booth (1971) and Minter et al.
(2001)).
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982; Hernández-Vera and Almodóvar
(1983, 1984); Cantrell and Betancourt (1995);
Lodge (1996a); Minter et al. (2001)
Carvajal-Zamora (1971a, b)
Stevenson (1975); Minter et al. (2001)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
Roure and Ramírez (1970); Betancourt et al. (1980);
Cantrell (1991); Cantrell and Betancourt (1992)
Stevenson (1975); Lodge (1996a); Minter et al.
(2001)
Hansford (1946); Minter et al. (2001)
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
Garman (1915); Chardón (1920); Minter et al.
(2001)
P. minuta Henn.
Hi
P. paspalicola Henn.
Pv
P. sphaerosperma G. Winter
Physalospora lagunculariae Rehm
Pleospora sp.
Poronia oedipus (Mont.) Mont.
Cc, Cs
La
Co, Rh
Un
Pterosporidium rhizomorphae (Kunze: Curr.) W.H. Ho & K.D. Hyde
Rh
Chardón (1920); Seaver and Chardón (1926);
Stevenson (1975); Minter et al. (2001)
Stevens (1917); Chardón (1920); Fitzpatrick (1927);
Orton (1944); Stevenson (1975); Minter et al.
(2001)
Chardón (1920); Orton (1944); Stevenson (1975);
Minter et al. (2001)
Stevens (1917); Seaver and Chardón (1926);
Stevenson (1975); Minter et al. (2001)
Acevedo (1987); Lodge (1996a); Minter et al.
(2001); Nieves-Rivera (unpubl. data, on R. mangle
submerged wood and mangrove mud, Magueyes
Island)
Toro (1924); Stevenson (1975); Guzmán (1986);
Nieves-Rivera et al. (1998); Minter et al. (2001)
Ho and Hyde (1996); Poonyth et al. (2000); Schmit
and Shearer (2003); Schmit (2004)
P. rhizophorae (Vizioli) W.H. Ho & K.D. Hyde
Rh
Puiggarina lagunculariae (Rehm) Speg. = Physalospora lagunculariae
Pyrenographa sp.
Un
Pyrenula sp.
Rhytidhysterium rufulum (Spreng.: Fr.) Speg.
Rosellinia corticium (Schwein.) Sacc.
Un
Ca, Tc, Un
Ca
65
Kohlmeyer (1969); Fell and Master (1973); Ho and
Hyde (1996); Calzada (1999); Poonyth et al. (2000);
Schmit and Shearer (2003); Schmit (2004)
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
Stevenson (1975); Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Saccardoella rhizophorae K.D. Hyde
Saccardoella sp.
Saccharomyces cerevisiae Meyen: E. C. Hansen
Rh, Un
Co, Df, Un
Bs
Schizothyrium lagunculariae (G. Winter) Arx, in E. Müll. & Arx
La
Scutellinia sp.
Sphaeria zonata Lév. = Kretzschmaria zonatum
Spiropes capensis (Thüm.) M.B. Ellis
*Strigula sp.
Telimena ecastophylli (Lév.) Cif.
Thyridaria sp.
Thyronectria pseudotrichia (Schwein.) Seeler
Torpedospora radiata Meyers
Trichocladium sp. = Microsphaera sp.
Trichomerium coccolobae Bat. & Cif.
T. ornatum Bat. & Cif.
Varicosporina ramulosa Meyers & Kohlm.
Verruculina enalia Kohlm. & Volkm.-Kohlm.
Xylaria adscendens Fr.
Cn, Ll, Ss, Un
Co, La, Me
Un
De
Un
Cn, Un
Co, Df, Hi, Sf, Th
Cu
Mc
Bs, Sp
Df, Rh
Lp, Un
66
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Mona Island)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Earle (1901); Stevens (1917); Seaver and Chardón
(1926); Stevenson (1975); Minter et al. (2001)
Pfister (1979); Minter et al. (2001)
Stevens (1917, 1919); Spegazzini (1923); Stevenson
(1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, Mona Island)
Stevenson (1975); Minter et al. (2001)
Minter et al. (2001); Nieves-Rivera (unpubl. data,
Mona Island)
Chardón (1920); Stevenson (1975); Minter et al.
(2001)
Acevedo (1987); Nieves-Rivera and Santos-Flores
(2004a, b); Nieves-Rivera (unpubl. data, Rincón
Lagoon, Boquerón Wildlife Refuge)
Stevenson (1975); Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Acevedo (1987, 2001)
Acevedo (1987); J. Kohlmeyer (pers. comm., 2004)
Lodge (1996a); Acevedo (2001); Minter et al.
(2001)
X. arbuscula Sacc.
X. feejeensis (Berk.) Fr.
X. multiplex (Kunze) Fr.
X. scruposa (Fr) Fr.
Xylaria sp.
Unknown sp. 1
Unknown sp. 2
Unknown sp. 3
Unknown sp. 4
Ba
Stevenson (1975); Lodge (1996a); Minter et al.
(2001)
Un
Lodge (1996a); Nieves-Rivera et al. (1999)
Remarks.–Britton (1915) reported as Xylaria sp.
Misidentified as Pterula capilaris (Lév.) Sacc.
(sensu Saccardo (1888)) by Nieves-Rivera (1996)
Lp, Ss
Lodge (1996a); Acevedo (2001); Minter et al.
(2001)
Cn, Un
Stevenson (1975); Minter et al. (2001)
Af, As, Ch, Cu, Db, Dd, Go, Gs, Ip, Lp, Pg, Rh, Sp
Acevedo (2001); Minter et al. (2001); Rodríguez et
al. (2004); Nieves-Rivera (unpubl. data, on
intertidal R. mangle wood, Magueyes Island)
Bs, Sf
Nieves-Rivera and Santos-Flores (2004b)
Sf
Nieves-Rivera and Santos-Flores (2004b)
Rh
Nieves-Rivera (unpubl. data, on R. mangle dead
wood, Magueyes Island)
Rh
Nieves-Rivera (unpubl. data, on R. mangle dead
wood, Magueyes Island)
MITOSPORIC FUNGI (ANAMORPHIC FUNGI)
67
Acremonium murorum (Corda) W. Gams
Acremonium sp.
Bs
Ad, Am, Tm
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Bunkley-Williams and
Williams (per. comm., 2004)
Alternaria humicola Oudem. = Alternaria tenuissima
A. tenuissima (Kunze) Wiltshire
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Alternaria sp.
Ad, Bs, Ec, Ll, Rh, Sf, Ss
Roure and Ramírez (1970); Betancourt et al. (1980);
Acevedo (1987); Lodge (1996a); Minter et al.
(2001); Ortiz Rivera and Semidey Laracuente
(2004); Nieves-Rivera (unpubl. data, on intertidal R.
mangle wood, Magueyes Island)
Anguillospora cf. longissima (Sacc. & P. Syd.) Ingold Bs, Ll, Rf, Sf, Un Lodge (1996a); Santos-Flores and BetancourtLópez (1997); Minter et al. (2001); Nieves-Rivera
and Santos-Flores (2004b)
Antennariella persea Bat., Nascim. & Cif.
Cn
Stevenson (1975); Minter et al. (2001)
Apiocarpella sp.
Am
Pérez Samot (1986)
Arthrinium sp.
Un
Acevedo (2001)
Articulospora tetracladia Ingold
Rf, Sf
Santos-Flores and Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004b)
Aschersonia cubensis Berk. & M.A. Curtis
Cn
Stevenson (1975); Minter et al. (2001)
A. turbinata Berk.
Cn
Stevenson (1975); Minter et al. (2001)
Aschersonia sp.
Am
Pérez Samot (1986)
Aspergillus caespitosus Raper & Thom
Hp
Silva et al. (2004); Ortiz et al. (2004)
A. candidus Link
Bs, Hp, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001); Silva et al. (2004); Ortiz et al. (2004)
68
A. clavatus Desm.
Bs, Ss
A. flavipes (Bainier & R. Sartory) Thom & Church
Hp, Ss
A. flavus Link
Bs, Hp, Rp, Ss
A. fumigatus Fresen. = Fennellia flavipes
A. funiculosus G. Sm.
Bs, Ss
A. humicola Chaudhuri & Sachar
Bs, Ss
A. luchuensis Inui = Aspergillus niger var. awamori
A. nidulans (Eidam) Winter = Emericella nidulans
A. niger Tiegh.
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Minter et al. (2001); Silva et al. (2004); Ortiz et al.
(2004)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Lodge (1996a); NievesRivera (2000a); Minter et al. (2001); Silva et al.
(2004); Ortiz et al. (2004); Ruiz-Suárez, (2004)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Bs, Rh, Ll, Ss, Tm Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001); Ruiz-Suárez, (2004); BunkleyWilliams and Williams (pers. comm., 2004);
Nieves-Rivera (unpubl. data, on intertidal R. mangle
wood and leaves, Magueyes Island and mouth of the
Manatí River; Figures 14A–B)
A. niger Tiegh. var. awamori (Nakaz.) Al-Musallam
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
A. oryzae (Ahlb.) Cohn
Bs
Hernández-Vera (1972, 1975); Minter et al. (2001)
A. okazakii Okazaki
Bs
Hernández-Vera (1982) ; Hernández-Vera and
Almodóvar (1983, 1984)
69
A. ostianus Wehmer
A. ruber (J. König, Spieck. & Bremer) Thom & Church
Hp
Bs
A. sydowii (Bainier & R. Sartory) Thom & Church
Bs, Gf, Gv, Ss
A. terreus Thom, in Thom and Church
Bs, Ll, Rh, Ss
Silva et al. (2004); Ortiz et al. (2004)
Hernández-Vera (1982) ; Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1982; Hernández-Vera and
Almodóvar (1983, 1984); Ernesto Weil (pers. obs.,
2000, Enrique Cay, La Parguera); Minter et al.
(2001); Nieves-Rivera (2002); Greetchen Díaz and
Rafael Montalvo-Rodríguez (pers. comm., 2004,
hypersaline lagoons of Salinas Bay, refer to “Fig. 1”
of Montalvo-Rodríguez et al., 1997: pp. 99)
Hernández-Vera (1972, 1975, 1982) ; HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001); Ruiz-Suárez, (2004); Nieves-Rivera
(unpubl. data, on submerged R. mangle wood and
leaves, Magueyes Island)
A. unguis (Émile-Weil & L. Gaudin) Thom & Raper = Emericella unguis
A. ustus (Bainier) Thom & Church
Rh, Ll
Minter et al. (2001); Nieves-Rivera (unpubl. data,
on submerged R. mangle wood and leaves,
Magueyes Island)
A. versicolor (Vuill.) Tirab.
Bs, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001)
A. violaceus-fuscus Gasperini
Bs, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
Aspergillus sp.
Ad, Am, Bs, Ch, Cn, Cu, Ga, Hp, Ip, Lh, Ll, Ma, Ms, Rh, Ss, Te, Th, Tm, Tu
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Acevedo (1987, 2001);
Cantrell (1991); Cantrell and Betancourt (1992);
70
Vargas et al. (1998); Calzada (1999); Minter et al.
(2001); Silva et al. (2004); Ortiz et al. (2004);
Rodríguez et al. (2004); Ruiz-Suárez, (2004)
Atractilina parasitica (G. Winter) Deighton & Piroz.
Ls
Stevens (1917); Stevenson (1975); Minter et al.
(2001)
Aureobasidium sp.
Ad, Go, Gx, Ss, Th Betancourt et al. (1980); Lodge (1996a); Acevedo
(2001); Minter et al. (2001)
Beauveria bassiana (Bals.-Criv.) Vuill.
Cu
Wolcott (1948, 1955a, b); Stevenson (1975); Minter
et al. (2001)
B. brogniartii (Sacc.) Petch
Ss
Carvajal-Zamora (1971a, b)
Beltrania rhombica Penz.
Bs, Cu, Ll, Sf
Lodge (1996a); Santos-Flores & Betancourt-López
(1997); Minter et al. (2001); Nieves-Rivera (unpubl.
data, Rincón Lagoon, Boquerón Wildlife Refuge)
Bipolaris cynodontis (Marignoni) Shoemaker
Cd
Theis (1953); Stevenson (1975); Minter et al.
(2001)
Bipolaris sp.
Rp, Un
Nieves-Rivera (2000a); Acevedo (2001)
Blodgettia bornetii E.P. Wright
Wd
Kohlmeyer (1980); Minter et al. (2001); Schmit
(2004, isolated from algae?)
Botryoderma lateritium Papendorf & H.P. Upadhyay
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Brachiosphaera tropicalis Nawawi, in Descals
Rf, Sf
Santos-Flores & Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004a, b); Nieves-Rivera (unpubl. data, Rincón
Lagoon, Boquerón Wildlife Refuge)
Camarosporium roumeguerii Sacc.
Df
Acevedo (1987)
Camposporidium sp.
Sf
Lodge (1996a); Santos-Flores & Betancourt-López
(1997); Minter et al. (2001); Nieves-Rivera and
Santos-Flores (2004b)
Campylospora sp. (s. str. Santos-Flores & Betancourt-López, 1997)
71
Df, Hi, Ll, Rf, Sf
Santos-Flores & Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004a, b)
Candida glabrata (H.W. Anderson) S.A. Mey. & Yarrow, in Yarrow & S.A. Mey.
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
C. guillermondi (Castell.) Langeron & Guerra
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1982); Hernández-Vera and Almodóvar (1983,
1984); Minter et al. (2001)
C. krusei (Castell.) Berkhout
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1982); Hernández-Vera and Almodóvar (1983,
1984); Stevenson (1975); Minter et al. (2001)
C. parapsilosis (Ashford) Langeron & Talice
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
C. pseudotropicalis (Castell.) Basgal
Bs, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
C. tropicalis (Castell.) Berkhout
Bs, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Candida sp.
Ad
Roure and Ramírez (1970)
Catenophora sp.
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Cephalosporium acremonium Corda
Bs, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Lodge (1996a); Minter et
al. (2001)
C. asperum Harz
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
C. coremioides Raillo = Verticillium lecanii
72
C. curtipes Sacc.
Bs, Ss
C. lecanii Zimm. = Verticillium lecanii
Cephalosporium sp.
Cercospora chrysobalani Ellis & Everh.
Bs
Ch
C. conocarpi Chupp & A.S. Mull.
Co
C. hibisci Tracy & Earle = Pseudocercospora abelmoschi
C. hibiscina Ellis & Everh.
C. ipomoeae G. Winter
Hi
Ip
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Acevedo (1987)
Weiss (1950); Chupp (1953); Stevenson (1975);
Minter et al. (2001)
Chupp (1953); Stevenson (1975); Minter et al.
(2001)
Stevenson (1975); Minter et al. (2001)
Toro (1931); Chupp (1953); Stevenson (1975);
Minter et al. (2001)
Cercospora sp.
Av, Rh, Se, Tc, Tp Hansford (1946); Minter et al. (2001); NievesRivera (unpubl. data, Mona Island; Figures 15A–B)
Chaetomella sp.
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Cladosporium cladosporoides (Fresen.) G.A. de Vries Bs, Hr, Ss, Un
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Nieves-Rivera (2000b); Minter et al. (2001); RuizSuárez, (2004)
C. herbarum (Pers.) Link
Bs, Ss, Un
Stevens (1917); Weiss (1950); Wellman (1961);
Stevenson (1975); Minter et al. (2001); RuizSuárez, (2004)
C. oxysporum Berk. & M.A. Curtis, in Berk.
Ba, Bs, Cn, Rh, Ss
Minter et al. (2001); Nieves-Rivera (unpubl. data,
from seawater in R. mangle roots, Los Morrillos and
Bahía Sucia, Cabo Rojo, southwestern Puerto Rico)
Remarks.–Strain deposited with the ATCC as
Cladosporium oxysporum Berk. & M.A. Curtis
MYA-3068 (ÁMNR-7).
73
C. sphaerospermum Penz. = Botryotinia allii
C. werneckii Horta = Phaeoannellomyces werneckii
Cladosporium sp.
Ad, Am, Av, Bs, Hp, Lh, Ma, Ms, Rh, Sp, Ss, Td, Te, Tm, Tu
Roure and Ramírez (1970); Carvajal-Zamora
(1971a, b); Betancourt et al. (1980); Pérez Samot
(1986); Acevedo (1987, 2001); Cantrell (1991);
Cantrell and Betancourt (1992); Vargas et al.
(1998); Calzada (1999); Minter et al. (2001);
Sánchez and Santos (2004); Silva et al. (2004);
Ortiz et al. (2004); Ruiz-Suárez, (2004)
Clavatospora bulbosa (Anastasiou) Nakagiri & Tubaki Bs, Rf, Sf
Nieves-Rivera and Santos-Flores (2004a, b)
Coniosporium shiraianum (Syd.) Bub.
Ba
Stevenson (1975); Minter et al. (2001)
Coniothyrium sp.
Am
Pérez Samot (1986)
Cryptococcus laurentii (Kuff.) Skinner
Bs, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
C. neoformans (San Felice) Vuill.
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Curvularia pallescens Boedijn = Cochliobolus pallescens
*C. robusta Kilp. & Luttr.
Bs, Ll, Sf
Nieves-Rivera (unpubl. data, Rincón Lagoon,
Boquerón Wildlife Refuge)
C. subulata (Nees) Boedijn
Ss
Carvajal-Zamora (1971a, b)
Curvularia sp.
Ad, Bs, Df, Ec, Lh, Ll, Ms, Sf, Ss, Td, Te, Tm, Tu
Roure and Ramírez (1970); Betancourt et al. (1980);
Acevedo (1987); Vargas et al. (1998); Minter et al.
(2001); Ortiz Rivera and Semidey Laracuente
(2004); Nieves-Rivera and Santos-Flores (2004b);
Sánchez and Santos (2004); Ruiz-Suárez, (2004);
Nieves-Rivera (unpubl. data, La Parguera Channels
and Phosphorescent Bay in Lajas, southwestern
Puerto Rico)
74
Cylindrocarpon sp.
Bs, Cu, Ll, Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Minter et al. (2001)
Cylindrocladium scoparium Morgan (s. str. Miller and McRitchie, 1973) = Calonectria morganii
Cytospora palmicola Berk. & M.A. Curtis
Cn
Stevenson (1975); Minter et al. (2001)
C. rhizophorae Kohlm. & E. Kohlm.
Ba, Rh
Acevedo (1987); Tattar et al. (1994); Wier et al.
(1996, 2000); Tattar and Wier (2002); J. Kohlmeyer
(pers. comm., 2004); Nieves-Rivera (unpubl. data,
on R. mangle prop roots, Magueyes Island, Los
Morrillos, and Boquerón Commonwealth Forest,
southwestern Puerto Rico; Figures 16A–F).
Remarks.–The orange to deep orange cirri of
Cytospora sp. (= Cytospora rhizophorae Kohlm. &
E. Kohlm.) on R. mangle prop roots (south of
Magueyes Island, La Parguera, Lajas, southwestern
Puerto Rico) was deposited with BPI by ÁMNR, 15
March 2001, BPI 843790.
Cytospora sp.
Ll
Lodge (1996a)
Dactylaria sp.
Bs, Cu, Ll, Un
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Dendryphiella arenaria Nicot
Bs, Dd, Sa, Sp, Th Hernández-Vera (1982); Acevedo (1987, 2001);
Hernández-Vera and Almodóvar (1983, 1984)
Dictyochaeta sp.
Cn, Cu, Ll, Un
Minter et al. (2001)
Dictyonella erysiphoides (Rehm) Höhn.
Cu
Stevenson (1975); Minter et al. (2001)
Dictyosporium sp.
Cu, Ll, Un
Minter et al. (2001)
Diheterospora sp.
Am
Pérez Samot (1986)
Diplocladiella scalaroides G. Arnaud
Cu, Ll, Rf, Sf, Un Santos-Flores & Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004b); Nieves-Rivera (unpubl. data, Rincón
Lagoon, Boquerón Wildlife Refuge)
75
Diplocladiella sp.
Cu, Sf
Diplococcium spicatum Grove
Bs, Ss
Diplodia epicocos Cooke
Cn
D. natalensis Pole-Evans = Lasiodiplodia theobromae
D. oraemaris Linder
Df
Diplodia sp.
Bs
Diploidia sp. = Septoidium sp.
Dreschlera gigantea (Heald & F.A. Wolf) S. Ito
Cd
Dreschlera sp.
Bs, Ch, Cu, Ip, Ss
Endocalyx melanoxanthus (Berk. & Broome) Petch
Cn, Ll, Un
Epidermophyton floccosum (Harz) Langeron & Miloch.
Bs
Exophiala werneckii (Horta) Arx = Phaeoannellomyces werneckii
*Exserohilum sp.
Bs, Ll, Rh, Sf
Lodge (1996a); Santos-Flores & Betancourt-López
(1997); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Francis and Lowe (2000)
Acevedo (1987)
Acevedo (1987)
Stevenson (1975); Minter et al. (2001)
Rodríguez et al. (2004); Ruiz-Suárez, (2004)
Stevenson (1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, Rincón Lagoon,
Boquerón Wildlife Refuge)
Fusarium aquaeductum (Radl. & Raben.) Sacc. = Nectria episphaeria
F. aquaeductuum Lagerh. var. medium Wallenw. = Nectria episphaeria
F. chlamydosporium Wollenw. & Reinking
Ma
Cantrell and Betancourt (1995)
F. episphaerica (Tode) W.C. Snyder & H.N. Hansen = Gibberella baccata
F. epishaericum (Cooke & Ellis) Sacc. = Gibberella baccata
F. equiseti (Corda) Sacc. = Gibberella intricans
F. lateritium Nees = Gibberella baccata
F. moniliforme J. Sheld. = Gibberella fujikuroi
F. moniliforme J. Sheld. var. subglutinans Wollenw. & Reinking = Gibberella subglutinans
F. neoceras Wollenw. & Reinking = Gibberella subglutinans
F. nivale (Fr.) Sorauer = Monographella nivalis
F. oxysporum Schltdl.
Bs, Hr, Un
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Nieves-Rivera (2000b);
Minter et al. (2001)
76
F. roseum Link.: Fr.
Bs, Ss
F. semitectum Berk. & Ravenel, in Berk.
Hp, Ma
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Cantrell and Betancourt (1995); Silva et al. (2004);
Ortiz et al. (2004)
F. solani (Mart.) Sacc. = Nectria haematococca
Fusarium sp.
Ac, Ad, Am, Bs, Df, Ec, Go, Hi, Lh, Ll, Hp, Ma, Ms, Sf, Ss, Td, Te, Tm, Tu, Un
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Acevedo (1987, 2001);
Cantrell (1991); Cantrell and Betancourt (1992);
Lodge (1996a); Vargas et al. (1998); Minter et al.
(2001); Ortiz Rivera and Semidey Laracuente
(2004); Sánchez and Santos (2004); Silva et al.
(2004); Ortiz et al. (2004); Nieves-Rivera and
Santos-Flores (2004b); Ruiz-Suárez, (2004);
Bunkley-Williams and Williams (pers. comm.,
2004)
Fusicoccum microspermum Har. & P. Karst.
Tc
Stevenson (1975); Minter et al. (2001)
Fusoma sp. (nom. dub.)
Ma
Cantrell (1991); Cantrell and Betancourt (1992)
Gelatinosporium sp.
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Geotrichum candidum Link = Galactomyces geotrichum
Geotrichum sp.
Bs, Ch, Cu, Ip, Ss, Tm
Rodríguez et al. (2004); Ruiz-Suárez, (2004);
Bunkley-Williams and Williams (per. comm., 2004)
Gilmaniella sp.
Ad
Betancourt et al. (1980)
Gliocladium roseum Bainier
Bs, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001)
G. virens J.H. Mill., Giddens & A.A. Foster
Ss
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
77
Almodóvar (1983, 1984); Lodge (1996a); Minter et
al. (2001)
Gliomastix murorum (Corda) S. Hughes = Acremonium murorum
Graphium squarrosum Ellis & Langl.
Ba
Harposporidium sp. = Harposporium sp.
Harposporium sp.
Bs
Helicomyces roseus Link = Tubeufia cylindrothecia
H. torquatus L.C. Lane & Shearer
Bs, Ll, Sf
Helicorhoida sp. = Helicorhoidion sp.
Helicorhoidion sp.
Helicosporium griseum Berk. & M.A. Curtis, in Sacc.
Ad
Stevenson (1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Santos-Flores & Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera (unpubl. data, Rincón
Lagoon, Boquerón Wildlife Refuge)
Betancourt et al. (1980)
Ba, Bs, Cn, Hi, Ll, Un
Minter et al. (2001)
Helicosporium sp.
Ba, Cn, Hi, Ll, Un
Minter et al. (2001)
Helminthosporium anomalum J.C. Gilman & E.V. Abbott Bs, Ss
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
H. velutinum Link
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Helminthosporium sp.
Ad, Bs, Cd, Cn, Ss, Un
Theis (1953); Roure and Ramírez (1970);
Betancourt et al. (1980); Acevedo (1987); Minter et
al. (2001)
78
Heterosporium sp. = Cladosporium sp.
Hirsutella saussurei (Cooke) Speare
Cn, Un
Wolcott (1948); Stevenson (1975); Minter et al.
(2001)
Roure and Ramírez (1970)
Histoplasma sp.
Ad
Hormodendron sp. = Hormodendrum sp.
Hormodendrum sp. = Cladosporium sp.
Hortaea werneckii (Horta) Nishim. & Miyaji. = Phaeoannellomyces werneckii
Humicola brevis (J.C. Gilman & E.V. Abbott) J.C. Gilman
Ss
Carvajal-Zamora (1971a, b)
Hyaloflorea sp. = Cylindrocarpon sp.
Hyalopicnic sp. = Hyalopycnis sp.
Hyalopycnis sp.
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Khuskia oryzae H.J. Huds.
Bs, Cu, Hr, Ip, Sf, Un
Stevenson (1975); Lodge (1996a); Nieves-Rivera
(2000b); Minter et al. (2001); Rodríguez et al.
(2004); Nieves-Rivera (unpubl. data, Rincón
Lagoon, Boquerón Wildlife Refuge)
*Koorchaloma sp.
Sr
Nieves-Rivera (unpubl. data, Magueyes Island)
Lasiodiplodia theobromae (Pat.) Griffon & Maubl.
Ba, Ca, Cn, Hi, Tc Francis and Lowe (2000); Minter et al. (2001)
Lemonniera pseudofloscula Dyko, in Descals, J. Webster & Dyko
Ll, Rf, Sf
Santos-Flores and Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004b)
Leprieurina radiata Toro
Ch
Stevenson (1975); Minter et al. (2001)
Macrophoma sp.
Av, Df, Rh
Acevedo (1987) Ortiz Rivera and Semidey
Laracuente (2004);
Macrophomina sp.
Ec
Ortiz Rivera and Semidey Laracuente (2004);
Melanconium saccharinum Penz. & Sacc. = Papularia vinosa
Microdochium nivale (Fr.) Samuels & I.C. Hallett = Monographella nivalis
79
Microxyphium aciculiforme Cif., Bat. & Nascim.
M. atrocarpi Bat., Nascim. & Cif., in Bat. & Cif.
Monodyctis sp.
Mycovellosiella lantanae (Chupp) Deighton
Myrothecium sp.
Nigrospora sphaerica (Sacc.) E.W. Mason
Cn, Ll
Cn, Ll
Ba, Cn, Cu, Ll, Un
Lc
Stevenson (1975); Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Minter et al. (2001)
Toro (1931); Chupp (1953); Wellman (1961);
Stevenson (1975); Minter et al. (2001)
Ec
Ortiz Rivera and Semidey Laracuente (2004);
Cc, Ch, Cu, Ip, Un Stevenson (1975); Minter et al. (2001); Rodríguez
et al. (2004)
N. oryzae (Berk. & Broome) Petch = Khuskia oryzae
Nigrospora sp.
Ad, Am, Ss
Nodulisporium sp.
Ad, Lp, Ss
Ochroconis humicola (G.L. Barron & L.V. Busch) de Hoog & Arx
Tm
Odiodendrum griseum Robak = Oidiodendron griseum
Oidiodendron griseum Robak
Bs
Paecilomyces javanicus (K. Friedericks & Bally) A.H.S. Br. & G. Sm.
Ca
P. lilacinus (Thom) Samson
Ss, Tm
Paecilomyces sp.
Ss
80
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Minter et al et al. (2001)
Betancourt et al. (1980); Lodge (1996a); Acevedo
(2001); Minter et al et al. (2001)
Bunkley-Williams and Williams (per. comm., 2004)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Stevenson (1975); Minter et al. (2001)
Bunkley-Williams et al. (1998); Rand et al. (2000);
Minter et al. (2001); Bunkley-Williams and
Williams (per. comm., 2004)
Remarks.–Rand et al. (2000) reported it as the
causative agent of tilapia-wasting disease in
brackish and freshwater environments.
Carvajal-Zamora (1971a, b); Williams and BunkleyWilliams (1996)
Papularia vinosa (Berk. & M.A. Curtis) E.W. Mason
Ba
Papularia sp.
Penicillium aculeatum Raper & Fennell
P. albicans Bainier
Am
Ss
Bs
P. brevicompactum Dierckx
P. canadense G. Sm.
Bs, Ss
Bs
P. canescens Sopp
Bs
P. chermesium Biourge
Bs
P. citrinum Thom
Bs, Ss
P. coeruleoviridae Biourge
P. corylophilum Dierckx
Ss
Bs, Ss
P. digitatum (Pers.) Sacc.
Bs, Ss
P. duclauxii Delacr.
P. fellutanum Biourge
Bs
Bs, Ss
81
Seaver and Chardón (1926); Stevenson (1975);
Minter et al. (2001)
Pérez Samot (1986)
Carvajal-Zamora (1971a, b)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975, 1982); Hernández-Vera and
Almodóvar (1983, 1984); Lodge (1996a); Minter et
al. (2001)
Carvajal-Zamora (1971a, b)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001)
P. frequentans Westling
Bs, Ss
P. funiculosum Thom
Bs, Ss
P. godlewiskii K.M. Zalessky = Penicillium jensenii
P. janczewskii K.M. Zalessky
Ss
P. jensenii K.M. Zalessky
P. nigricans Bainier: Thom = Penicillium janczewskii
P. ochro-chloron Biourge
Bs, Ss
P. oxalicum Currie & Thom
P. paxilli Bainier
Bs, Ss
Bs, Ss
P. pulvillorum Turfitt = Penicillium simplicissimum
P. roqueforti Thom
P. roqueforti Thom var. carneum Frisvad
Ll
Av, Rh
P. sclerotiorum T.H. Beyma
Bs, Ss
P. simplicissimum (Oudem.) Thom
P. spinulosum Thom
Bs, Ss
Ll, Rh
P. steckii K.M. Zalessky = Penicillium citrinum
P. waksmani K.M. Zalessky
Penicillium sp.
Ss
Bs
Carvajal-Zamora (1971a, b); Hernández-Vera
(1982); Hernández-Vera and Almodóvar (1983,
1984)
Hernández-Vera (1972, 1975); Lodge (1996a);
Minter et al. (2001)
Carvajal-Zamora (1971a, b); Stevenson (1975);
Minter et al. (2001)
Carvajal-Zamora (1971a, b)
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Minter et al. (2001)
Lodge (1996a)
Nieves-Rivera (unpubl. data, on intertidal A.
germinans and R. mangle wood and leaves,
Magueyes Island and Boquerón Wildlife Refuge)
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Lodge (1996a); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, on intertidal R. mangle
wood and leaves, Magueyes Island)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Ad, Am, Bs, Ch, Cn, Cu, Ga, Hp, Hr, Ip, Lh, Ll, Ma, Ms, Ss, Td, Te, Tm, Tu, Un
82
Periconia byssoides Pers.
Cn, Hi, Ll, Un
Periconia prolifica Anastasiou = Halosphaeria cucullata
Periconia sp.
Hi, Hp
Pestalotia adusta Ellis & Everh.
P. coccolobae Ellis & Everh. = Pestalotiopsis versicolor
P. disseminata Thüm. = Pestalotiopsis disseminata
P. gibberosa Sacc.
P. palmarum Cooke = Pestalotiopsis palmarum
P. pezizoides de Not.
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Acevedo (1987, 2001);
Cantrell (1991); Cantrell and Betancourt (1992);
Vargas et al. (1998); Nieves-Rivera (2000b); Minter
et al. (2001); Sánchez and Santos (2004); Silva et
al. (2004); Ortiz et al. (2004); Rodríguez et al.
(2004); Ruiz-Suárez, (2004)
Stevenson (1975); Minter et al. (2001)
Ch
Minter et al. (2001); Silva et al. (2004); Ortiz et al.
(2004)
Guba (1961); Stevenson (1975); Minter et al. (2001)
Cn
Guba (1961); Stevenson (1975); Minter et al. (2001)
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Pestalotia sp.
Ad, Am, Bs, Bt, Cn, Cu, Ps, Rh, Ss, Td
Carvajal-Zamora (1971a, b); Hernández-Vera
(1972, 1975); Betancourt et al. (1980); Pérez Samot
(1986); Lodge (1996a); Acevedo (2001); Minter et
al. (2001); Sánchez and Santos (2004); NievesRivera (unpubl. data, on Cocos nucifera L.,
Pseudophoenix sargentii H. Wendland, and Thrinax
morrisii H. Wendland, Mona Island)
Pestalotiopsis disseminata (Thüm.) Steyaert
Av, Bs, Ll, Rh, Sf, Tc
Stevenson (1975); Calzada (1999); Minter et al.
(2001); Nieves-Rivera (unpubl. data, on R. mangle
83
P. palmarum (Cooke) Steyaert
Cn, Ps
P. versicolor (Speg.) Steyaert
Cn, Cu
leaves, Magueyes Island, La Parguera Channels, Los
Morrillos, and Boquerón Wildlife Refuge; Figures
17A-B)
Remarks.–This species is common on R. mangle
leaves after wet chamber.
Stevens (1917); Weiss (1950); Guba (1961);
Stevenson (1975); Francis and Lowe (2000); Minter
et al. (2001)
Stevens (1917); Guba (1961); Stevenson (1975);
Lodge (1996a); Francis and Lowe (2000); Minter et
al. (2001)
Rodríguez et al. (2004)
Pestalotiopsis sp.
Ch, Cu, Ip
Peyronellaea sp. = Phoma sp.
Phaeoannellomyces werneckii (Horta) McGinnis & Schell
Bs, Hp, Ss, Tm Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Lodge (1996a); Silva et
al. (2004); Ortiz et al. (2004); Bunkley-Williams
and Williams (per. comm., 2004)
Phialophorophoma litoralis Linder
Df, La, Rh
Acevedo (1987)
Phoma eupyrena Sacc.
Rh
Calzada (1999)
P. humicola J.C. Gilman & E.V. Abbott
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Phoma sp.
Ad, Am, Av, Df, Ll, Ls, Ma, Rh, Ss
Roure and Ramírez (1970); Kohlmeyer (1980);
Betancourt et al. (1980); Pérez Samot (1986);
Acevedo (1987); Cantrell (1991); Cantrell and
Betancourt (1992); Lodge (1996a); Minter et al.
(2001)
Phomopsis cocöes Petch
Cn
Stevenson (1975); Minter et al. (2001)
84
Phyllosticta bonduc F. Stevens
Cb
P. coccolobae Ellis & Everh., in A.S. Hitchcok
Cu
P. lantanicola Sacc.
Pithomyces sp.
Podoxyphium ampullaceum Bat. & H. Maia, in Bat. & Cif.
Lc, Lv
Ad
Stevenson (1975); Minter et al. (2001)
Young (1916); Seaver and Chardón (1926);
Kohlmeyer (1969); Stevenson (1975); Minter et al.
(2001)
Pyricularia grisea (Cooke) Sacc.
Se
Theis (1953); Stevenson (1975); Minter et al.
(2001)
Rhodotorula glutinis (Fresen.) F.C. Harrison
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Rhodotorula sp.
Bs, Ss
Hernández-Vera (1972, 1975); Lodge (1996a);
Minter et al. (2001)
Rhynchosporium sp.
Ad
Betancourt et al. (1980)
Sclerotium portoricense F. Stevens
Cd
Stevens (1917); Stevenson (1975); Minter et al.
(2001)
Scolecobasidium humicola G.L. Barron & L.V. Busch = Ochroconis humicola
Scopulariopsis brevicaulis (Sacc.) Bainier
Bs
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
Scopulariopsis sp.
Bs, Ss
Carvajal-Zamora (1971a, b); Acevedo (1987)
Seimatosporium sp.
Ad
Betancourt et al. (1980)
Pseudocercospora abelmoschi (Ellis & Everh.) Deighton
Cn, Hi, Ll
Hi
Stevens (1917); Seaver and Chardón (1926);
Stevenson (1975); Poonyth et al. (2000); Minter et
al. (2001); Schmit and Shearer (2003); Schmit
(2004)
Young (1915); Seaver and Chardón (1926);
Stevenson (1975); Francis and Lowe (2000); Minter
et al. (2001)
Stevens (1917); Wellman (1961); Stevenson (1975);
Minter et al. (2001)
Betancourt et al. (1980)
85
Septoidium sp.
Septoria fici-indicae Voglino
Ad
Od
S. lantanae Garman
Lc
Spegazzinia tessarthra (Berk. & M.A. Curtis) Sacc.
Sphaeropsis subglobosa Cooke, in Sacc.
Sphaeropsis sp.
Ba, Bs, Ss
Ba
Am, Ba, Bs, Ss
Spicaria javanica K. Friedericks & Bally = Paecilomyces javanicus
Sporotrichum schenckii Matr. = Beauveria brogniartii
S. schenckii Matr. var. fioccoi C.W. Dodge = Beauveria brogniartii
Sporotrichum sp.
Ad
Stachybotrys sp.
Ad
Stemphylium botryosum Wallr.
Bs, Rh, Sf, Ss
*S. cf. gracilariae E.G. Simmons
Rh, Sf
Roure and Ramírez (1970)
Stevens (1917); Stevenson (1975); Minter et al.
(2001)
Garman (1915); Wellman (1961); Stevenson
(1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Stevenson (1975);
Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Stevenson (1975);
Pérez Samot (1986); Minter et al. (2001)
Roure and Ramírez (1970)
Roure and Ramírez (1970)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Nieves-Rivera (unpubl. data, Rincón Lagoon,
Boquerón Wildlife Refuge)
Stemphylium sp.
Ad
Roure and Ramírez (1970)
Tetraploa aristata Berk. & Broome
Bs, Cn, Ll, Sf, Un Stevenson (1975); Santos-Flores and BetancourtLópez (1997); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Rincón Lagoon, Boquerón Wildlife
Refuge)
Tetraploa sp.
Td
Sánchez and Santos (2004)
Thermomyces sp.
Bs, Ss
Hernández-Vera (1972, 1975); Minter et al. (2001)
Thielaviopsis paradoxa (De Seynes) Höhn. = Ceratocystis paradoxa
Torula sp.
Ad, Cn, Un
Roure and Ramírez (1970); Betancourt et al. (1980);
Minter et al. (2001)
Torulopsis glabrata (H.W. Anderson) Lodder & N.F. deVries = Candida glabrata
86
Trichoderma album Preuss
Bs
T. glaucum E.V. Abbott
Bs
T. harzianum Rifai
T. koningii Oudem., in Oudem. & C.J. Koning
Ss
Bs, Ss
T. lignorum (Tode) Harz = Trichoderma viride
T. longibranchiatum Rifai
T. viride Pers.
Ss
Bs, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Carvajal-Zamora (1971a, b)
Carvajal-Zamora (1971a, b); Hernández-Vera
(1982); Hernández-Vera and Almodóvar (1983,
1984); Lodge (1996a); Minter et al. (2001)
Carvajal-Zamora (1971a, b)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Lodge (1996a);
Minter et al. (2001)
Trichoderma sp.
Ad, Am, Bs, Hr, Ma, Ss, Un
Roure and Ramírez (1970); Betancourt et al. (1980);
Pérez Samot (1986); Acevedo (1987); Cantrell
(1991); Cantrell and Betancourt (1992); NievesRivera (2000b); Minter et al. (2001)
Trichophyton rubrum (Castell.) Sabour.
Bs, Ss, Un
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
T. cf. mentagrophytes (C.P. Robin) Sabour. in Saccardo Hr, Ss, Un
Carrión (1930, 1935); Kesten et al. (1932); Carrión
and Silva (1944a, b); Stevenson (1975); NievesRivera (2000b); Minter et al. (2001)
Trichosporium cutaneum (Küchenm. & Rabenh.) Vuill. = Trichosporon cutaneum
T. pullulans (Lindner) Diddens & Lodder = Trichosporon pullulans
Trichosporon cutaneum (Beurm., Gougerot & Vaucher) N. Ota
Bs, Ss
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984); Minter et al. (2001)
87
T. pullulans (Lindner) Diddens & Lodder
Trichosporon sp.
*Trimmatostroma sp.
Tripospermum roupalae (Syd.) S. Hughes
Triscelophorus acuminatus Nawawi
Triscelophorus sp.
Tubercularia pulvurulenta Speg.
Tubeufia cylindrothecia (Seaver) Höhn.
Valsa chlorina Pat.
Verticicladium effusum Earle
Verticillium lateritium Berk. = Nectria inventa
V. lateritium (Ehrenb.) Rabenh. = Nectria inventa
Bs
Ad
Av, Ba, La, Un
Cn, Un
Bs, Rf, Sf
Sf
Rh
Bs, Rf, Sf
Cn
Cu
88
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Roure and Ramírez (1970)
Nieves-Rivera (unpubl. data, on living leaves of A.
germinans and L. racemosa, Piñones
Commonwealth Forest at northern Puerto Rico, and
from Boquerón Commonwealth Forest, Los
Morrillos, Magueyes Island, and La Parguera
Channels at southwestern Puerto Rico)
Remarks.–Dematiaceous mycelium on living leaves
of A. germinans, deposited with the U.S. National
Fungus Collection by ÁMNR, 7 March 2002, BPI
863548.
Stevenson (1975); Minter et al. (2001)
Santos-Flores & Betancourt-López (1997); Minter
et al. (2001); Nieves-Rivera and Santos-Flores
(2004b)
Nieves-Rivera and Santos-Flores (2004b)
Acevedo (1987)
Toro (1924); Santos-Flores & Betancourt-López
(1997, reported as Helicomyces cf. roseus Link);
Minter et al. (2001); Nieves-Rivera and SantosFlores (2004b)
Stevenson (1975); Minter et al. (2001)
Francis and Lowe (2000)
V. lecanii (Zimm.) Viégas
Bs, Cn, Ss
Stevenson (1975); Hernández-Vera (1982);
Hernández-Vera and Almodóvar (1983, 1984);
Minter et al. (2001)
Verticillium sp.
Cu, Dd, Go, Ss, Th, Un
Acevedo (2001); Minter et al. (2001)
Zalerion maritimum (Linder) Anastasiou
Bs, Rh, Th, Wd Schmit and Shearer (2003); Nieves-Rivera (unpubl.
data, Rincón Lagoon, Boquerón Wildlife Refuge)
Mycelia Sterilia (Agonomycetales)
Ad, Bs, Ch, Cu, Hp, Ip, Lm, Ma, Ms, Te, Tm, Tu
Betancourt et al. (1980); Acevedo (1987); Cantrell
(1991); Cantrell and Betancourt (1992); Vargas et
al. (1998); Silva et al. (2004); Ortiz et al. (2004);
Rodríguez et al. (2004); Nieves-Rivera (unpubl.
data, Rincón Lagoon, Boquerón Wildlife Refuge,
Boquerón Commonwealth Forest, Los Morrillos,
Magueyes Island, and La Parguera Channels at
southwestern Puerto Rico)
Unknown sp. 1
Df, Sf
Nieves-Rivera and Santos-Flores (2004b)
Unknown sp. 2
Ll, Sf
Nieves-Rivera and Santos-Flores (2004b)
Unknown sp. 3
Sf
Nieves-Rivera and Santos-Flores (2004b)
Unknown sp. 4
Df, Sf
Nieves-Rivera and Santos-Flores (2004b)
BASIDIOMYCOTA
Acanthosphysium mirabile (Berk. & M.A. Curtis) Parmasto Un
Lodge et al. (2000); Minter et al. (2001)
Agaricus johnstonii Murrill
Ba, Cn, Ll, Ss, Un Stevenson (1975); Nieves-Rivera (1996); NievesRivera et al. (1999); Minter et al. (2001)
Alboleptonia sp.
Ss, Un
Lodge et al. (2000); Minter et al. (2001)
Aleurodiscus mirabilis (Berk. & M.A. Curtis) Höhn. = Acanthosphysium mirabile
Amanita cf./aff. ingrata Pegler
Ss
Guzmán (1986); Nieves-Rivera (unpubl. data, Mona
Island)
89
Amanita sp.
Athelia rolfsii (Curzi) C.C. Tu & Kimbr.
Auricularia cornea Ehrenb., in Nees
A. delicata (Fr.) Henn.
A. polytricha (Mont.) Sacc. = Auricularia cornea
Battarrea stevenii (Libosch.) Fr.
B. phalloides (Dicks.) Pers. = Battarrea stevenii
Boletus earlei (Murrill) Overh., in Seaver & Chardón
Bs, Cu, Ll, Ss
Ba
Tp, Un
Un
Ss
Bs, Ss
Boletus section Luridi
Boletus sp.
Calvatia cyathiformis (Bosc.) Morgan
Cantharellus cinnabarinus Schwein.
Ceriporia xylostromatoides (Berk.) Ryvarden
Ss
Bs, Cu, Ss
Ss
Cu, Ss
Ca, Un
Chlorophyllum molybdites (G. Mey.: Fr.) Massee
Bs, Ss
Clathrus baumii (Henn.) E. Fisch.
Ss, Un
90
Remarks.–Misidentified as Tricholomopsis aff.
humboldtii Singer, Ovrebo & Halling by NievesRivera et al. (1999)
Nieves-Rivera et al. (1999); Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Lowy (1952, 1971); Lodge (1996a); Lodge et al.
(2000); Minter et al. (2001); Nieves-Rivera (unpubl.
data, Magueyes Island and La Parguera Channels)
Lowy (1952, 1971); Nieves-Rivera et al. (1998);
Minter et al. (2001)
Nieves-Rivera et al. (1998); Minter et al. (2001)
Murrill (1921); Seaver (1925); Seaver & Chardón
(1926); Fitzpatrick (1927); Stevenson (1975);
Minter et al. (2001)
Nieves-Rivera et al. (1999)
Minter et al. (2001)
Nieves-Rivera et al. (1998); Minter et al. (2001)
Murrill (1910); Pegler (1983); Minter et al. (2001)
Lowe (1966); Stevenson (1975); Ryvarden (1985);
Nieves-Rivera (1996); Nieves-Rivera et al. (1998,
1999); Minter et al. (2001)
Guzmán (1986); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Caño Corazones, Mayagüez)
Lodge (1996a); Minter et al. (2001)
Remarks.–This species have a lilac volva (Lodge,
1996a).
C. crispus Turpin
Coleosporium ipomoeae (Schwein.) Burrill
Cn, Cu, Rh, Ss
Is
Collybia sp.
Cn, Ss, Tp
Copelandia cyanescens (Berk. & Broome) Singer
Coprinus cf./aff. ephemerus (Bull.: Fr.) Fr.
Dg, Ss, Un
Ca, Ss, Un
C. cf. jamaicensis Murrill (s. str. Dennis (1970))
C. cf. plicatilis (M.A. Curtis: Fr.) Fr.
Un
Ss
91
Dennis (1953); Dring (1980); Nieves-Rivera et al.
(1999); Minter et al. (2001) Lodge (1996a);
Maldonado-Ramírez and Torres-Pratts (in press)
Arthur (1915(b), 1917, 1924); Stevens (1917);
Roure (1962, 1963); Stevenson (1975); Minter et al.
(2001)
Nieves-Rivera et al. (1999); Minter et al. (2001);
Nieves-Rivera (unpubl. data, Magueyes Island,
southwestern Puerto Rico)
Remarks.–White mycelium on decaying leaves of
Thespesia populnea, deposited with the U.S.
National Fungus Collection by ÁMNR, 17 March
2004, BPI 863547.
Guzmán (1986); Navarro and Betancourt (1992)
Nieves-Rivera (1996); Nieves-Rivera et al. (1999);
Minter et al. (2001)
Nieves-Rivera et al. (1999); Minter et al. (2001)
Bor de Garrido (1969); Stevenson (1975); NievesRivera (1996); Nieves-Rivera et al. (1998, 1999);
Minter et al. (2001)
Coprinus sp.
Coriolopsis flocossa (Jungh.) Ryvarden
Ba, Cn, Ll, Sb, Un Pegler (1983); Dennis (1970); Minter et al. (2001)
Ca, Hm, Rh
Ryvarden (1984); Minter et al. (2001); NievesRivera et al. (2004)
C. fulvocinera Murrill = Datronia caperata
Coriolopsis sp.
Corticium debile Berk. & M.A. Curtis: Massee
Crepidotus uber (Berk. & M.A. Curtis) Sacc.
Crinipellis septotricha Singer
La
Un
Rh
Ca, Cn, Cu, Un
Crinipellis sp.
Ca, Ll, Un
Cyathus striatus (Huds.) Pers.
Cn, Ss, Un
Cyathus sp.
Dacryopinax spathularia (Schwein.) G.W. Martin
Datronia caperata (Berk.) Ryvarden
Earliella scabrosa (Pers.) Gilb. & Ryvarden
Tc, Un
Ca, Cn, Rh, Un
Ca, Rh, Un
Cn, Pt, Tc, Un
Flammula earlei Murrill
Fomes lividus (Kalchbr.) Sacc. = Perenniporia tephroporia
Fomitopsis nivosa (Berk.) Gilb. & Ryvarden
Ganoderma australe (Fr.) Pat.
Cn
Cn
Ba, Cn, Un
92
Minter et al. (2001)
Burt (1925); Stevenson (1975); Nieves-Rivera
(1996); Nieves-Rivera et al. (1999); Minter et al.
(2001)
Nieves-Rivera et al. (2004)
Pegler (1983); Nieves-Rivera (1996); Nieves-Rivera
et al. (1999); Minter et al. (2001)
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Magueyes Island)
Stevenson (1975); Nieves-Rivera et al. (1998);
Minter et al. (2001)
Minter et al. (2001)
Bresadola et al. (1893); Heller (1900); Britton
(1915); Stevens (1917); Lowy (1971); Stevenson
(1975); Nieves-Rivera (1996); Nieves-Rivera et al.
(1998, 1999, 2004); Minter et al. (2001)
Murrill (1915); Seaver (1925); Fitzpatrick (1927);
Stevenson (1975); Minter et al. (2001)
Ryvarden (1985); Stevenson (1975); Minter et al.
(2001)
Stevenson (1975); Minter et al. (2001)
Seaver & Chardón (1926); Lowe (1975); Stevenson
(1975); Lodge (1996a); Minter et al. (2001)
Minter et al. (2001)
G. lucidum (Curtis) P. Karst.
Un
Stevenson (1975); Ryvarden (1985); Nieves-Rivera
et al. (1998); Minter et al. (2001)
G. resinaceum Pat.
Cn, Hm, Un
Phelps and Landgraf (1972); Stevenson (1975);
Ryvarden (1985); Minter et al. (2001)
G. zonatum Murrill
Ci
Stevenson (1975); Minter et al. (2001)
Gloeophyllum striatum (Sw.: Fr.) Murrill
Ca, Rh
Britton (1915); Murrill (1915); Stevenson (1975);
Nieves-Rivera et al. (1998, 1999, 2004); Minter et
al. (2001)
Gymnopilus sp.
Ba, Ca, Cn, Un Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1999); Minter et al. (2001)
Heterochaete sp.
Un
Minter et al. (2001); Nieves-Rivera et al. (1999)
Hexagonia hydnoides (Sw.: Fr.) M. Fidalgo
Ca, Cm, Cu, Hm, Rh, Un
Britton (1915); Stevenson (1975); Ryvarden (1984);
Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1998, 1999, 2004); Minter et al.
(2001)
Hygrocybe acutoconica (Clem.) Singer = Hygrocybe persistens
H. persistens (Britzelm.) Britzelm.
Bs, Cu, Ss
Cantrell and Lodge (2000); Minter et al. (2001)
Hymenochaete sp.
Cu
Minter et al. (2001)
Hymenogaster sp. (?)
Cn, Cu, Ss
Nieves-Rivera (1996); Nieves-Rivera et al. (1999)
Inonotus porrectus Murrill
La
Stevenson (1975); Minter et al. (2001)
Kuehneola malvicola Arthur
Pa
Seaver and Chardón (1926); Roure (1962, 1963);
Minter et al. (2001)
Lachnella candida (Pers.) G. Cunn.
Ba
Stevenson (1975); Minter et al. (2001)
Lactarius coccolobea O.K. Miller & Lodge
Bs, Ss
Lodge et al. (2000); Miller et al. (2000)
Lactocollybia sp. (L. cf. angiospermarum Singer)
Cn, Ss, Tp, Un Lodge et al. (2000); Minter et al. (2001); NievesRivera et al. (1999); Nieves-Rivera (unpubl. data,
Magueyes Island and La Parguera Channels)
93
Laetiporus persicinus (Berk & M.A. Curtis) Gilb.
Lentinus bertieri (Fr.) Fr.
Lentinus crinitus (L.: Fr.) Fr. = Panus crinitus
Lenzites elegans (Spreng.: Fr.) Pat.
Un
Cn, Un
Ca, Cn, Rh, Un
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Magueyes Island)
Pegler (1983); Minter et al. (2001)
Bresadola et al. (1893); Britton (1915); Stevenson
(1975); Ryvarden (1984, 1985); Lodge (1996a);
Nieves-Rivera (1996); Nieves-Rivera et al. (1998,
1999)
Lepiota sp.
Ba, Cu, Hi, Ll, Ss Lodge (1996a); Nieves-Rivera et al. (1999); Minter
et al. (2001)
Leucoagaricus hortensis (Murrill) Pegler
Ss, Un
Nieves-Rivera et al. (1998); Minter et al. (2001)
Leucocoprinus birnbaumii (Corda) Singer
Cn, Ll, Un
Bor de Garrido (1969); Stevenson (1975); Guzmán
(1986); Lodge (1996a); Nieves-Rivera (1996);
Nieves-Rivera et al. (1999); Minter et al. (2001)
Marasmiellus coilobasis (Berk.) Singer
Cn, Un
Pegler (1983); Minter et al. (2001)
M. semiustus Berk. & M.A. Curtis = Marasmiellus semiustus
M. cf. semiustus (Berk. & M.A. Curtis) Singer
Ca, Cn, Un
Lodge (1996a); Nieves-Rivera et al. (1999); Minter
et al. (2001)
Remarks.—Lodge (1996a) reported Marasmiellus
semiustus (Berk. & M.A. Curtis) Singer var. sabali
Singer on palm.
Marasmiellus sp.
Ca, Cn, Ll, Pt, Ss Minter et al. (2001)
Marasmius haematocephalus Mont.
Cn, Cu, Ll, Un Dennis (1970); Pegler (1983); Minter et al. (2001)
M. pallescens Murrill
Cn, Un
Murrill (1915); Stevenson (1975); Nieves-Rivera et
al. 1998; Minter et al. (2001)
M. sacchari Wakker
Cn, Un
Stevenson (1975); Minter et al. (2001)
Morganella fuliginea (Berk. & M.A. Curtis) Kriesel & Dring
Cn, Ll, Un
Guzmán (1986); Stevenson (1975); Minter et al.
(2001)
94
Panaeolus antillarum (Fr.) Dennis
Dg, Ss, Un
P. cyanescens Berk. & Broome = Copelandia cyanescens
P. sphinctrinus (Fr.) Quél.
Dg, Ss, Un
Stevenson (1975); Guzmán (1986); Navarro and
Betancourt (1992); Nieves-Rivera et al. (1998);
Minter et al. (2001)
Bor de Garrido (1969); Navarro and Betancourt
(1992)
Panus crinitus (L.) Singer
Ca, Rh
Bresadola et al. (1893); Britton (1915); Bor de
Garrido (1969); Stevenson (1975); Guzmán (1986);
Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1999, 2004); Minter et al. (2001)
Remarks.—Lodge (1996a) reported this species as
Lentinus swartzii Berk. (= Lentinus crinitis (L.: Fr.)
Fr. sensu Berk.). Basidiocarps deposited with the
U.S. National Fungus Collection by ÁMNR, 8 May
2002, BPI 863546.
Perenniporia tenuis var. tenuis (Schwein.) Ryvarden
Ca
Stevenson (1975); Minter et al. (2001)
Remarks.– The name Polyporus tenuis Sacc. is
nomen illegitimum and so P. tenuis Klotzsch. The
taxonomic concept behind Saccardo’s name is
unclear (L. Ryvarden, pers. comm., 2004).
P. tephroporia (Mont.) Ryvarden
Ca, Cn
Stevenson (1975); Minter et al. (2001)
Phanaerochaete crassa (Lév.) Burdsall
Un
Burt (1920); Welden (1975); Stevenson (1975);
Pegler (1983); Nieves-Rivera (1996); Nieves-Rivera
et al. (1999)
Phellinus dependens (Murrill) Ryvarden
Ca, Un
Britton (1915); Lowe (1957); Stevenson (1975);
Ryvarden (1985); Nieves-Rivera (1996); NievesRivera et al. (1998, 1999); Minter et al. (2001)
P. gilvus (Schwein.) Pat.
Ba, Ca, Cn, Cu, Tc, Un
95
Stevenson (1975); Guzmán (1986); Ryvarden
(1990); Minter et al. (2001)
P. merrillii (Murrill) Ryvarden
Rh
Nieves-Rivera et al. (2004)
P. rimosus (Berk.) Pilát
Hm
Ryvarden (1985); Minter et al. (2001)
Phellinus sp.
Ba, Co, Cu, La, Ll, Rh, Un
Minter et al. (2001)
Phlebia spp.
Rh, Ss
Minter et al. (2001); Nieves-Rivera et al. (2004;
Figures 18A–B);
Phlebopus beniensis (Singer & Digilio) Heinem. & Rammeloo
Ss, Un
Lodge et al. (1998)
Phylloporia chrysites (Berk.) Ryvarden
Ca, Un
Britton (1915); Murrill (1915); Stevenson (1975);
Ryvarden (1985); Lodge et al. (2000); NievesRivera (1996); Nieves-Rivera et al. (1998, 1999);
Minter et al. (2001)
P. frutica (Berk. & M.A. Curtis) Ryvarden
Ca, Un
Britton (1915); Murrill (1915); Stevenson (1975);
Ryvarden (1984); Nieves-Rivera (1996); NievesRivera et al. (1999); Minter et al. (2001)
Pleurotus djamor (Fr.) Boedijn
Ba, Ca, Cn, Cu, Rh, Sb
Guzmán et al. (1993); Lodge (1996a); Ortiz Santana
(1997); Nieves-Rivera et al. (1998, 1999); Minter et
al. (2001)
P. eugrammus (Mont.) Dennis
Cn, Un
Stevenson (1975); Pegler (1983); Guzmán (1986);
Minter et al. (2001)
P. flabellatus Berk. & Broome
Cn, Un
Pegler (1983); Minter et al. (2001)
Pleurotus sp.
Rh
Minter et al. (2001)
Pluteus sp.
Cn, Hi, Ss
Nieves-Rivera et al. (1999); Minter et al. (2001)
Polyporus fulvocinereus (Murrill) Overh., in Seaver & Chardón = Datronia caperata
Polyporus gilvus (Schwein.) Fr. = Phellinus. gilvus
Polyporus lignosus Klotzsch in Fr. = Rigidoporus microporus
96
P. nivosellus (Murrill) Sacc. & Trotter = Fomitopsis nivosa
P. porrectus (Murrill) Sacc. & Trotter = Inonotus porrectus
P. tricholoma Mont.
Un
P. zonalis Berk. = Rigidoporus lineatus
Polyporus sp.
Prospodium tuberculatum (Speg.) Arthur
Cn, Ll, Ss, Un
Lc, Lv
Psathyrella spp.
Ss, Ll, Rh, Un
Psilocybe cubensis (Earle) Singer
Dg, Ss, Un
P. subcubensis Guzmán
Dg, Ss, Un
Puccinia cenchri Dietel & Holw., in Holw.
Cc, Ct
P. cynodontis Lacroix, in Desm.
Cd
P. heterospora Berk. & M.A. Curtis, in Berk.
Pa
P. lantanae Farl.
P. stenotaphricola J. Walker
Lc, Le, Ls, Lv
Se
97
Heller (1900); Stevenson (1975); Ryvarden (1984,
1985); Guzmán (1986); Lodge (1996a) ; NievesRivera (1996); Nieves-Rivera et al. (1999); Minter
et al. (2001)
Ryvarden (1984); Minter et al. (2001)
Arthur (1925); Kern and Whetzel (1926); Stevenson
(1975); Minter et al. (2001)
Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1999, 2004); Minter et al. (2001)
Seaver and Chardón (1926); Stevenson (1975);
Guzmán (1986); Navarro and Betancourt (1992);
Minter et al. (2001); Guzmán et al. (2003)
Navarro and Betancourt (1992); Guzmán et al.
(2003)
Arthur (1915b, 1916, 1917); Theis (1953); Roure
(1962, 1963); Stevenson (1975); Minter et al.
(2001)
Arthur (1916, 1917); Theis (1953); Stevenson
(1975); Minter et al. (2001)
Seaver and Chardón (1926); Roure (1962, 1963);
Minter et al. (2001, reported on a leaf of the swamp
mallow Pavonia paludicola D.H. Nicols.: Fryxell (=
Pavonia scabra K. Presl. (Malvaceae)
Arthur (1915c, 1917); Kern (1932); Roure (1962,
1963); Stevenson (1975); Minter et al. (2001)
Stevenson (1975); Minter et al. (2001)
Pycnoporus sanguineus (L.: Fr.) Murrill
Ca, Cn, Rh, Un
Bresadola et al. (1893); Britton (1915); Dennis
(1970); Stevenson (1975); Guzmán (1986); Lodge
(1996a); Nieves-Rivera (1996); Nieves-Rivera et al.
(1998, 1999, 2004); Minter et al. (2001)
Rigidoporus lineatus (Pers.) Ryvarden
Ba, Ca, Cn, Un Stevenson (1975); Ryvarden (1984, 1988); Lodge
(1996a); Minter et al. (2001)
R. microporus (Sw.) Overeem
Ca, Cn, Un
Stevenson (1975); Ryvarden (1984); Lodge (1996a);
Minter et al. (2001)
Rigidoporus sp.
Cn, Ss, Un
Minter et al. (2001)
Russula cremeolilacina var. coccolobicola Pegler, in Pegler & Singer
Cu, Ss
Lodge et al. (2000, at Piñones Commonwealth
Forest near Loíza); Minter et al. (2001)
R. littoralis Pegler = Russula littoralis
R. littoralis Romagn., Chevassut & Privat, in Romagn. Bs, Cu, Ss
Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1999); Miller et al. (2000); Minter et
al. (2001)
Russula sp.
Bs, Cu, Ss
Lodge (1996a); Minter et al. (2001)
Schizophyllum commune Fr.: Fr.
Ac, Av, Ba, Cn, La, Rh, Un
Bresadola et al. (1893); Britton (1915); Cooke
(1961); Bor de Garrido (1969); Pegler (1983);
Guzmán (1986); Dennis (1970); Stevenson (1975);
Lodge (1996a); Nieves-Rivera (1996); NievesRivera et al. (1998, 1999, 2004); Minter et al.
(2001)
Sclerangium bermudense (Coker) D.A. Reid
Bs, Cu, Ss
Guzmán (1970, 1986); Reid (1977); Nieves-Rivera
(1996); Nieves-Rivera et al. (1999); Minter et al.
(2001)
S. bermudense (Coker) D.A. Reid var. trinitense Reid
Bs, Ss
Reid (1977)
98
Scleroderma stellatum Berk.
Bs, Cu, Ss
Guzmán (1970, 1986); Reid (1977); Nieves-Rivera
(1996); Nieves-Rivera et al. (1999); Minter et al.
(2001)
Sclerotium rolfsii Sacc. = Athelia rolfsii
Solenia candida Pers.: Fr. = Lachnella candida
Sphaerobolus stellatus Tode: Pers.
Ba, Un
Stevenson (1975); Minter et al. (2001)
Sporisorium cenchri (Lagerh.) Vánky
Cc
Stevenson (1975); Minter et al. (2001)
Thellephora sp.
Bs, Cu, Ll, Ss, Un Minter et al. (2001)
Trametes corrugata (Pers.) Bres. = Earliella scabrosa
T. cubensis (Mont.) Sacc.
Cn, Un
Stevenson (1975); Minter et al. (2001)
T. elegans (Spreng.: Fr.) Fr.). = Lenzites elegans
T. maxima (Mont.) A. David & Racjchenb.
Ch
Fitzpatrick (1927); Stevenson (1975); Minter et al.
(2001)
T. nivosella Murrill = Fomitopsis nivosa
T. villosa (Fr.) Kriesel
Co
Stevenson (1975); Minter et al. (2001)
Tremellostereum dichroum (Lloyd) Ryvarden
Cu, Ss, Un
Minter et al. (2001)
Trogia cantarelloides (Mont.) Singer
Un
Pegler (1983); Nieves-Rivera (1996); Nieves-Rivera
et al. (1999); Minter et al. (2001)
Tulostoma berkeleyi Lloyd
Ss
Stevenson (1975); Minter et al. (2001)
T. exasperatum Mont.
Ss
Stevenson (1975); Minter et al. (2001)
T. cf. meridionale J.E. Wright
Ss
Nieves-Rivera et al. (1998)
T. portorricense J.E. Wright
Bs
Wright (1987); Minter et al. (2001)
T. volvulatum I.G. Borshch., in Sorokĭn
Ss
Stevenson (1975); Minter et al. (2001)
Tyromyces cf. chioneus (Fr.) P. Karst.
Rh
Nieves-Rivera et al. (2004)
Tyromyces sp.
Rh
Ryvarden (1984); Minter et al. (2001)
Uredo coccolobae Henn.
Cu
Arthur (1917); Seaver and Chardón (1926);
Thurston and Kern (1933); Roure (1962, 1963);
Stevenson (1975); Francis and Lowe (2000); Minter
et al. (2001)
99
U. ignava Arthur
Ba
Seaver and Chardón (1926); Stevenson (1975);
Minter et al. (2001)
U. uvifera P. Syd. & Syd. = Uredo coccolobae
Uromyces tenuicutis McAlpine
Sv
Ustilago affinis Ellis & Everh., in Cockerell
Se
Arthur (1915a), 1916, 1917); Theis (1953);
Stevenson (1975); Minter et al. (2001)
Whetzel and Kern (1926); Theis (1953); Stevenson
(1975); Minter et al. (2001)
Lodge et al. (2000); Minter et al. (2001)
Nieves-Rivera (unpubl. data, next to Cabo Rojo’s
lighthouse, Boquerón Commonwealth Forest,
southwestern Puerto Rico)
Remarks.–This unknown yellowish bolete
(Basidiomycota: Boletaceae) was found under a
grove of Myrica sp. (Myricaceae), Figures 19A–D.
Volvariella sp.
Unknown sp. 1
Ll, Ss, Un
Ll
ZYGOMYCOTA
100
Absidia sp.
Asellaria ligiae Tuzet & Manier
Basidiobolus sp.
Enterobryus spp.
Gongronella sp.
Haplosporangium sp. = Mortierella sp.
Mortierella sp.
Mucor bacilliformis Hasselt.
M. christianiensis Hagem
M. globosus A. Fisch.
M. jansseni Lendn.
Mucor sp.
Am
Li
Am
Uc
Am
Am, Bs, Ss
Ss
Bs, Ss
Bs, Ss
Bs
Am, Bs, Ma, Ss
Phycomyces sp.
Rhizopus oryzae Went & Prins. Geerl.
R. stolonifer Ehrenb. & Fr.
Bs, Ss
Bs, Ss
Ss
Syncephalastrum racemosum Cohn: J. Schröt, in Cohn
Bs, Ss
OOMYCOTA (CHROMISTA)
Achlya americana Humprey
Bs, Ss
A. bisexualis Coker & Couch
Ss
101
Pérez Samot (1986)
White et al. (2000)
Pérez Samot (1986)
White et al. (2000)
Pérez Samot (1986)
Hernández-Vera (1972, 1975); Pérez Samot (1986);
Minter et al. (2001)
Carvajal-Zamora (1971a, b); Pérez Samot (1986)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Hernández-Vera (1972, 1975, 1982); HernándezVera and Almodóvar (1983, 1984); Minter et al.
(2001)
Hernández-Vera (1982); Hernández-Vera and
Almodóvar (1983, 1984)
Pérez Samot (1986); Cantrell (1991); Cantrell and
Betancourt (1992); Ruiz-Suárez, (2004)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Carvajal-Zamora (1971a, b); Stevenson (1975);
Minter et al. (2001)
Hernández-Vera (1972, 1975); Minter et al. (2001)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Minter et
al. (2001)
De Santiago Serrano (1999)
A. cambrica (Trow) T.W. Johnson
A. caroliniana Coker
A. conspicua Coker
Bs, Ss
Bs, Ss
Bs, Ss
A. debaryana Humprey
A. diffusa J.V. Harv.: T.W. Johnson, in T.W. Johnson
Bs, Ss
Bs, Ss
A. dubia Coker
Bs, Ss
A. flagellata Coker
Bs, Ss
A. klebsiana Pieters
Bs, Ss
A. orion Coker & Couch
A. primoachlya Beneke
A. prolifera Nees = Achlya prolifera
A. prolifera Pringsh.
Bs, Ss
Bs, Ss
Bs, Ss
A. rodrigueziana F.T. Wolf
A. treleaseana (Humphrey) Kauffman
Achlya sp.
Albugo ipomoeae-panduratae (Schwein.) Swingle
Bs, Ss
Ss
Tm
Ip
102
Galler-Rimm (1982)
Galler-Rimm (1982)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Minter et
al. (2001)
Galler-Rimm (1982)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Minter et
al. (2001)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Minter et
al. (2001)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Longcore
(1996); Minter et al. (2001)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); Minter et
al. (2001)
Galler-Rimm (1982)
Galler-Rimm (1982)
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Galler-Rimm (1982); De
Santiago Serrano (1999); Minter et al. (2001)
Galler-Rimm (1982)
De Santiago Serrano (1999)
Bunkley-Williams and Williams (per. comm., 2004)
Stevens (1917); Tucker (1927); Weiss (1950);
Stevenson (1975); Minter et al. (2001)
A. ipomoeae-pes-caprae Cif.
Ip
Stevens (1917); Stevenson (1975); Minter et al.
(2001)
Aphanomyces laevis de Bary
Ss
De Santiago Serrano (1999)
Brevilegnia crassa Rossy-Vald.
Bs, Ss
Rossy-Valderrama (1955, 1956); Johnson (1956);
Stevenson (1975); Minter et al. (2001)
B. unisperma Coker & Braxton
Bs, Ss
Galler-Rimm (1982)
Dictyuchus monosporus Leitg.
Bs, Ss
Galler-Rimm (1982); De Santiago Serrano (1999)
D. pseudodictyon Pringsh.
Bs, Ss
Rossy-Valderrama (1955, 1956); Stevenson (1975);
Galler-Rimm (1982); De Santiago Serrano (1999);
Minter et al. (2001)
*Halophytophthora sp.
Rh
Nieves-Rivera (unpubl. data, Magueyes Island)
Phytophthora palmivora (E.J. Butler) E.J. Butler
Cn, Un
Weiss (1950); Phelps and Landgraf (1972);
Stevenson (1975); Francis and Lowe (2000); Minter
et al. (2001)
P. nicotianae var. parasitica (Dastur) G.M. Waterh.
Cn, Hi, Op
Weiss (1950); Stevenson (1975); Minter et al.
(2001)
Pythium spp.
Bs, Ss
Galler-Rimm (1982)
Saprolegnia ferax (Gruith.) Thuret
Ss
Nieves-Rivera and Betancourt-López (1994); De
Santiago Serrano (1999)
S. parasitica Coker
Ms
Bunkley-Williams and Williams (1994)
S. torulosa de Bary
Bs, Ss
Rossy-Valderrama (1955, 1956); Stevenson (1975);
Galler-Rimm (1982); Minter et al. (2001)
Saprolegnia spp.
Ao, Ht, Lh, Lm, Mf, Ms, Rm, Te, Tm, Tu
Bunkley-Williams and Williams (1994, 1995);
Williams and Bunkley-Williams (1996); Minter et
al. (2001); Nieves-Rivera (unpubl. data, Boquerón
Wildlife Refuge)
103
CHYTRIDIOMYCOTA
Allomyces spp.
Bs, Ss
Rhizophydium carpophilum Zopf
Unknown sp. 1
Bs, Ss
Am
Rossy-Valderrama (1955, 1956); Stevenson (1975);
Galler-Rimm (1982); Minter et al. (2001)
Galler-Rimm (1982)
Burrowes et al. (2004)
PLASMODIOPHOROMYCOTA
Woronina polycystis Cornu
Bs, Ss
Galler-Rimm (1982)
MYXOMYCOTA
Arcyria cinerea (Bull.) Pers.
Av, Un
Seaver and Chardón (1926); Hagelstein (1932);
Lodge (1996a); Minter et al. (2001); Novozhilov et
al. (2001); Stephenson and Nieves-Rivera (unpubl.
data, Magueyes Island)
Seaver and Chardón (1926); Hagelstein (1932);
Minter et al. (2001); Novozhilov et al. (2001)
A. incarnata (Pers.) Pers.
Un
Badhamia gracilis (T. Macbr.) T. Macbr., in T. Macbr. & G.W. Martin
Mi, Pr
Ceratiomyxa fruticulosa (O.F. Müll.) T. Macbr.
Comatricha irregularis Rex
C. longa Peck
Ca, Cn, Ll, Un
Cn
Pt, Un
C. typhoides (Bull.) Rostaf. = Stemonitopsis typhoides
104
Lodge (1996a); Minter et al. (2001); Nieves-Rivera
(unpubl. data, Magueyes Island)
Seaver and Chardón (1926); Hagelstein (1932); Farr
(1976); Nieves-Rivera and Santos-Flores (1998);
Minter et al. (2001)
Hagelstein (1927, 1932); Stevenson (1975); Farr
(1976); Minter et al. (2001)
Seaver and Chardón (1926); Hagelstein (1927,
1932); Stevenson (1975); Minter et al. (2001)
Craterium leucocephalum (Pers.) Ditmar
Cn, Ll, Un
Seaver and Chardón (1926); Hagelstein (1932);
Stevenson (1975); Farr (1976); Minter et al. (2001)
Cribaria intricata Schrad.
Ca, Un
Hagelstein (1932); Nieves-Rivera and Santos-Flores
(1998); Minter et al. (2001); Novozhilov et al.
(2001)
C. violacea Rex
Tc, Un
Hagelstein (1927); Stevenson (1975); Farr (1976);
Minter et al. (2001)
Cribaria sp.
Av
Stephenson and Nieves-Rivera (unpubl. data,
Magueyes Island)
Dictydium cancellatum (Batsch) T. Macbr.
Ca, Un
Hagelstein (1932); Nieves-Rivera and Santos-Flores
(1998); Minter et al. (2001)
Diderma effusum (Schwein.) Morgan
Av, Cn, Ll, Un
Stevenson (1975); Farr (1976); Minter et al. (2001);
Stephenson and Nieves-Rivera (unpubl. data,
Magueyes Island)
Didymium squamulosum (Alb. & Schwein.) Fr.
Cn, Ll, Un
Seaver and Chardón (1926); Hagelstein (1927,
1932); Stevenson (1975); Farr (1976); Minter et al.
(2001)
Echinostelium minutum de Bary, in Rostaf.
Av
Stephenson and Nieves-Rivera (unpubl. data,
Magueyes Island)
Fuligo septica (L.) F.H. Wigg.
Ba, Ca, Cn, Ss, Un Seaver and Chardón (1926); Hagelstein (1927,
1932); Stevenson (1975); Farr (1976); NievesRivera and Santos-Flores (1998); Minter et al.
(2001)
Hemitrichia serpula (Scop.) Rostaf.: Lister
Cn, Ll, Un
Seaver and Chardón (1926); Hagelstein (1932);
Stevenson (1975); Farr (1976); Minter et al. (2001)
H. stipitata (Massee) T. Macbr.
Cn, Un
Stevenson (1975); Minter et al. (2001)
Herpotrichia albidostoma (Peck) Sacc.
Cn, Un
Chardón (1921); Seaver (1922); Stevenson (1975);
Farr (1976); Minter et al. (2001)
H. schiedermayeriana Fuckel = Byssosphaeria schiedermayeriana (Ascomycota)
105
Licea operculata (Wingate) G.W. Martin
Lycogala sp.
Pt, Un
Ca, Cn, Un
Metatrichia vesparium (Batsch) Nann.-Bremek.
Perichaena chrysosperma (Curr.) Lister
Cn, Cu, Un
Av, Un
P. depressa Lib.
Physarella oblonga (Berk. & M.A. Curtis) Morgan
Physarum gyrosum Rostaf.
P. nutans Pers.
P. polycephalum Schwein.
P. pusillum (Berk. & M.A. Curtis) G. Lister
Ac, Av, Cn, Un
Cn, Ss, Un
Pr
Un
Un
Mi, Pr, Un
Physarum sp.
Av, Un
Stemonitis axifera (Bull.) T. Macbr.
Ca, Un
S. splendens Rostaf.
Cn, Rh, Un
Stemonitis sp.
Stemonitopsis typhoides (Bull.) T. N. Lakh. & Mukerji
Tc
Ag, Pt, Un
106
Stevenson (1975); Farr (1976); Minter et al. (2001)
Nieves-Rivera and Santos-Flores (1998); Minter et
al. (2001)
Stevenson (1975); Farr (1976); Minter et al. (2001)
Lodge (1996a); Minter et al. (2001); Stephenson
and Nieves-Rivera (unpubl. data, Magueyes Island)
Hagelstein (1932); Stevenson (1975); Farr (1976);
Minter et al. (2001); Stephenson and Nieves-Rivera
(unpubl. data, Magueyes Island)
Hagelstein (1932); Stevenson (1975); Farr (1976);
Minter et al. (2001)
Clark et al. (2004)
Hagelstein (1932); Novozhilov et al. (2001)
Hagelstein (1932); Novozhilov et al. (2001); Minter
et al. (2001)
Hagelstein (1932); Minter et al. (2001); NievesRivera (unpubl. data, Magueyes Island)
Minter et al. (2001); Stephenson and Nieves-Rivera
(unpubl. data, Magueyes Island)
Nieves-Rivera and Santos-Flores (1998); Minter et
al. (2001)
Seaver and Chardón (1926); Hagelstein (1932);
Stevenson (1975); Farr (1976); Minter et al. (2001);
Nieves-Rivera and Stephenson (2004); NievesRivera (present work)
Minter et al. (2001)
Seaver and Chardón (1926); Hagelstein (1932);
Minter et al. (2001); Novozhilov et al. (2001)
Tubifera ferruginosa (Batsch) J.F. Gmel.
Cn, Tc, Un
Seaver and Chardón (1926); Hagelstein (1932);
Stevenson (1975); Farr (1976); Nieves-Rivera and
Santos-Flores (1998); Minter et al. (2001)
______________________________________________________________________________________________________
* New records for Puerto Rico.
1
Notes: Some of the plants mentioned in this partial checklist, such as Bambusa vulgaris, Casuarina equisetifolia, Coccoloba
uvifera, Cocos nucifera or cacti, are not considered as mangroves or plants associated with mangroves. However, much of their
plant debris has been found as natural rafts (e.g., driftwood, floating leaves, propagules, etc.) in beaches, estuaries, and river mouths
due to storms, aeolian and sea currents (Hedgpeth, 1994). It is possible that plant’s mycobiota might be transported into littoral or
pelagic environments, many of the fungi mentioned in this checklist are common on these and other substrata.
2
Substrata or hosts: Plants found in mangroves (including mangrove-associated plants), sand dunes and coastal plains are named
according to Martorell et al. (1981) and Craig (1991). Marine algal taxonomy was based on Ballantine and Aponte (1997).
Abbreviations: Ac = Acrostichum spp. (Polypoidiacea); Ad = Air dust; Af = Amphiroa fragilissima (Rhodophyta); Ag = Agave
americana L. (Agavaceae); Am = Amphibians (Eleutherodactylus spp.); Ao = Astronotus ocellatus (Pisces: Chiclidae); As =
Acanthophora spicifera (Rhodophyta); Av = Avicennia germinans (Verbenaceae); Ba = Bambusa vulgaris (Gramineae); Bs =
Beach sand; Bt = Bryothamnion triquetrum (Rhodophyta); Ca = Casuarina equisetifolia (Casuarinaceae); Cb = Caesalpinia bonduc
(Caesalpiniaceae); Cc = Cenchrus echinatus (Gramineae); Cd = Cynodon dactylon (Gramineae); Ce = Conocarpus erectus var.
sericeus (Combretaceae); Ch = Chrysobalanus icaco (Rosaceae); Ci = Calophyllum inophyllum (Guttiferae); Cm = Coccoloba
microstachya (Polygonaceae); Cn = Cocos nucifera (Palmaceae); Co = Conocarpus erectus (Combretaceae); Cs = Cenchrus
myosuroides (Gramineae); Ct = Cenchrus tribuloides (Gramineae); Cu = Coccoloba uvifera (Polygonaceae); Cy = Cymodocea
manatorum (Potamogetonaceae); Db = Dasya balillouviana (Rhodophyta); Dd = Dictyota dichotoma (Phaeophyta); De = Dalbergia
ecastophyllum (Fabaceae); Df = Driftwood; Dg = animal dung (mostly bovine or equine); Dv = Dictyota divaricata (Phaeophyta);
Ec = Eichhornia crassipes (Pontederiaceae); Ga = Gelidiella acerosa (Rhodophyta); Gf = Gorgonia flabellum (Cnidaria:
Gorgoniidae); Go = Galaxaura oblongata (Rhodophyta); Gr = Gracilaria sp. (Rhodophyta); Gs = Galaxaura subverticillata
(Rhodophyta); Gv = Gorgonia ventalina (Cnidaria: Gorgoniidae); Gx = Galaxaura sp. (Rhodophyta); Ha = Harrisia portoricensis
(Cactaceae); Hi = Hibiscus tiliaceus (Malvaceae); Hm = Hippomane mancinella (Euphorbiaceae); Hp = Hypersaline pool; Hr = hair
(either animal and human); Ht = Helostoma temmincki (Pisces: Anabantidae); Ip = Ipomoea pes-caprae (Convolvulaceae); Is =
107
Ipomoea stolonifera (Convolvulaceae); La = Laguncularia racemosa (Combretaceae); Lc = Lantana camara (Verbenaceae); Le =
Lantana camara var. aculeata (Verbenaceae); Lh = Lepomis macrochirus (Pisces: Centrarchidae); Li = Ligia sp. (Isopoda: Ligidae);
Ll = Leaf litter; Lm = Lepomis auritus (Pisces: Centrarchidae); Lp = Liagora pinnata (Rhodophyta); Ls = Lantana sp.
(Verbenaceae); Lv = Lantana involucrata (Verbenaceae); Ma = Macrobrachium rosenbergii (Decapoda: Palaemonidae); Mc =
Myrica cerifera (Myricaceae); Me = Meliola spp. (Ascomycota: Meliolaceae); Mf = Micropterus salmoides floridanus (Pisces:
Centrarchidae); Mi = Melocactus intortus (Cactaceae); Ms = Micropterus salmoides salmoides (Pisces: Centrarchidae); Od =
Opuntia dillenii (Cactaceae); Op = Opuntia spp. (Cactaceae); Pa = Pavonia spicata (Malvaceae); Pg = Pandina gymnospora
(Phaeophyta); Ps = Pseudophoenix sargentii (Palmaceae); Pt = Pterocarpus officinalis (Fabaceae); Pr = Pilosocereus cf. royeni
(Cactaceae); Pv = Paspalum vaginatum (Gramineae); Rf = River foam; Rh = Rhizophora mangle (Rhizophoraceae); Rm = Rivulus
marmoratus (Pisces: Aplocheilidae); Rp = Reptile (Caiman crocodylus); Sa = Sargassum sp. (Phaeophyta); Sb = Sabal palmetto
(Palmae); Se = Stenotaphrum secundatum (Gramineae); Sf = Sea foam; Sp = Sargassum polyceratium (Phaeophyta); Sr = Spartina
sp. (Gramineae); Ss = Sandy soil; Sv = Sporobolus virginicus (Gramineae); Tc = Terminalia catappa (Combretaceae); Td = Typha
domingensis (Typhaceae); Te = Tilapia rendalli (Pisces: Chiclidae); Th = Thalassia testudinum (Hydrocharitaceae); Tm = Tilapia
mossambica (Pisces: Chiclidae); Tp = Thespesia populnea (Malvaceae); Tr = Triumfetta spp. (Tiliaceae); Tu = Tilapia urolepis
hornorum (Pisces: Chiclidae); Uc = Uca spp. (Decapoda: Ocypodidae); Un = Undetermined dead wood; Wd = Wood panel (Pinus)
as bait.
108
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131
Year
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Number of Species Reported
Figure 12. Taxa of marine fungi (including estuarine) reported for Puerto Rico in published
and unpublished reports between 1900 and 2004. Years selected represent
publication date of the records. Numbers given are not cumulative.
132
Figure 13. Asteromassaria sp. (Ascomycota) on aerial roots of Rhizophora mangle in
Magueyes Island, southwestern Puerto Rico.
133
Figures 14A–B. Aspergillus niger (Mitosporic fungi) on Rhizophora mangle from La
Parguera Channels, southwestern Puerto Rico (Sc = Sporocarps).
134
Figures 15A–B. Leaf spots and sporodochia (Sp) of Cercospora sp. (Mitosporic fungi) on a
leaf of Rhizophora mangle from La Parguera Channels, southwestern Puerto Rico.
A. Front. B. Back.
135
Figures 16A–F. Cytospora rhizophorae (Ascomycota) deep orange cirri on prop roots or
trunks of Rhizophora mangle in southwestern Puerto Rico. A. Boquerón Wildlife
Refuge. B-D. Magueyes Island. E-F. Los Morrillos.
136
Figure 17. Acervuli (Av) of Pestalotiopsis disseminata (Mitosporic fungi) on leaves of
Rhizophora mangle after growth in moist chambers from La Parguera Channels,
southwestern Puerto Rico.
137
Figures 18A–B. Phlebia sp. (Basidiomycota) on Rhizophora mangle wood from Boquerón
Wildlife Refuge, southwestern Puerto Rico.
138
Figures 19A–D. Bolete (Fistulinella cf.) basidiocarp (Basidiomycota) found under the grove
of Myrica sp. (Myricaceae) next to Cabo Rojo’s lighthouse, Boquerón
Commonwealth Forest, southwestern Puerto Rico. A–B. Pileal view. C. Side view.
D. Pore view.
139
CHAPTER 4
FILAMENTOUS FUNGI IN SEA FOAM AND MANGROVE SENESCENT
LEAVES IN A TROPICAL ESTUARY
ABSTRACT
Samples of sea foam and senescent leaves of Avicennia germinans and Rhizophora
mangle from an estuary known as Rincón Lagoon (Boquerón Beach Inlet = BBI; Boquerón
Wildlife Refuge = BWR) in southwestern Puerto Rico were assessed for the frequency of
occurrence of eight selected filamentous fungi. Among 12 bags of sea foam and 1296 leaves
screened (108 leaves per month from February 2002 through January 2003), the samples
consist of sporulating fungi and propagules, respectively. The species Aigialus sp. (A. cf.
grandis) (Ascomycota) and Penicillium roqueforti var. carneum (Mitosporic fungi) are new
records for Puerto Rico. Cochliobolus pallescens (Boquerón Beach Inlet [BBI] = 50.51%;
Boquerón Wildlife Refuge [BWR] = 50.98%), Exserohilum sp. (BBI = 16.95%; BWR =
15.37%), Tetraploa aristata (BBI = 11.74%; BWR = 14.26%), and Alternaria sp. (BBI =
10.88%; BWR = 9.53%), were frequently found in sea foam, while P. roqueforti var.
carneum and Aspergillus niger were frequently found in mangrove leaves (A. germinans and
R. mangle) in both BBI and BWR, followed closely by Pestalotiopsis disseminata, and
Alternaria sp. The two-way ANOVA performed to filamentous fungi (propagules and spore
counts, N = 432) in sea foam and the ones isolated from mangrove leaves (colonies in A.
germinans, N = 576; in R. mangle, N = 576) showed that variability was significant (p >
0.05) between the two communities studied (BBI and BWR). Statistical analysis suggested
that date/site was significant (p > 0.05), although date was insignificant (p < 0.05) in regards
to H0. Thus, this analysis rejects H0, in other words, it is unlikely that chance is operating to
produce observed differences and the evidence is strong enough to support the significant
effect, one that is probably not due to chance.
140
RESUMEN
Muestras de espuma marina y hojas seniles de Avicennia germinans y Rhizophora
mangle de un estuario conocido como la Laguna Rincón (Isleta Playera de Boquerón = BBI;
Refugio de Vida Silvestre de Boquerón = BWR) en el suroeste de Puerto Rico fueron
evaluados en frecuencia de ocurrencia de ocho hongos filamentosos selectos. De entre las 12
bolsas de espuma marina y 1296 hojas muestreadas (108 hojas por mes desde febrero 2002
hasta enero 2003), las muestras consisten de hongos esporulantes y propágalos,
respectivamente. Las observaciones de las carnadas (espuma marina y hojas de mangles y
escombros incubado mostraron que estos ocho hongos filamentosos pertenecen a los
ascomicetos (2 especies) y hongos mitospóricos (6 especies). Las especies Aigialus sp. (A.
cf. grandis) (Ascomycota) y Penicillium roqueforti var. carneum (Hongos mitospóricos) son
registros nuevos para Puerto Rico. Cochliobolus pallescens (Isleta de Playa de Boquerón
[BBI] = 50.51%; Refugio de Vida Silvestre de Boquerón [BWR] = 50.98%), Exserohilum sp.
(BBI = 16.95%; BWR = 15.37%), Tetraploa aristata (BBI = 11.74%; BWR = 14.26%) y
Alternaria sp. (BBI = 10.88%; BWR = 9.53%), son encontrados frecuentemente en espuma
marina, mientras que P. roqueforti var. carneum y Aspergillus niger han sido aislados
frecuentemente en las hojas de mangle (A. germinans y R. mangle) en ambos BBI and BWR,
seguidos de cerca por Pestalotiopsis disseminata y Alternaria sp. El ANOVA de dos vías
que se hizo a los hongos filamentosos (conteo de propágulos y esporas, N = 432) en espuma
marina y para los aislados de hojas de mangle (colonias en A. germinans, N = 576; en R.
mangle, N = 576) mostró que la variabilidada fue significativa (p > 0.05) entre las dos
comunidades estudiadas (BBI y BWR). El análisis estadístico sugirió que la fecha/lugar fue
significativa (p > 0.05), aunque la fecha fue insignificante (p < 0.05) con respecto a H0. Por
lo tanto, este análisis rechaza la H0, en otras palabras, es improbable que el azar esté
operando para producer diferencias observadas y la evidencia es lo suficientemente fuerte
para sostener el efecto es significativo, uno que es probablemente no debido al azar.
141
INTRODUCTION
Mangrove plants generate a large amount of litter in the form of branches,
inflorescence, leaves, twigs, and other debris. In general, the estimated annual litter
production in mangrove forests was slightly higher for Puerto Rico (9.45 mg.ha-1.yr-1) than
Florida (8.10 mg.ha-1yr-1) (Pool et al., 1975; Twilley et al., 1986). The contribution of
mangroves debris is the input of organic matter that enriched the coastal ecosystem and in
turn the fisheries (Cintrón and Schaeffer-Novelli, 1988).
Fungal distribution in marine and estuarine habitats are still poorly known, although a
number of published reports and summaries of previous studies in the Caribbean were
included in Stevenson (1975), Nishida (1989), Minter et al. (2001), and Schmit and Shearer
(2003). Filamentous fungi (Carvajal-Zamora, 1971; Hernández-Vera, 1972, 1975, 1982;
Stevenson, 1975; Hernández-Vera and Almodóvar, 1983, 1984; Minter et al., 2001; NievesRivera and Santos-Flores, in press), yeasts (Valdéz-Collazo et al., 1987), and fungi-like
organisms such as oomycetes (Rossy-Valderrama, 1956; Galler-Rimm, 1982) are known to
occur in beaches close to river mouths and estuaries of Puerto Rico. Most estuarine and
marine collections are in some cases sporadic. The purpose of this study is to understand the
diversity and frequency of occurrence of eight selected filamentous fungi in marine foam and
mangrove leaf litter from a river mouth of Puerto Rico, a subtropical island located between
the coordinates 18°00’-18°30’ N and 65°35’-67°15’ W, in the northeastern Caribbean Sea.
MATERIALS AND METHOD
Study Sites
The Rincón Lagoon (RL) (also known as Boquerón Lagoon s. str. Anonymous, 1905)
is located about 97 km southwestern of San Juan, Puerto Rico (Figure 20). This irregular
brackish water lagoon (7.5 km2 and 0.75 m of average depth) is open to the sea by a deep
canal (0.2 km long by 0.1 km wide, being deeper at the mouth, where it reach 2.0 m deep)
(Vázquez, 1983; Negrón-González, 1988; Aliaume et al., 1997). Candelas et al. (1973) and
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Negrón-González (1988) surveyed RL and BWR, and found that RL has a depth of 1.3 to 2.0
m, a visibility obtained with the Secchi disk of 0.6 m and the coefficient of light extinction
(QLE) of 2.51. RL has a shoreline development index of 2.10 (Negrón-González, 1988). It
receives, through its drainage channel, a continuous freshwater input from two streams (Los
Llanos and Boquerón Streams) and adjacent fields or, occasionally, from an impoundment
when gates overflow. The impoundment area (17 km2), nowadays known as the Boquerón
Wildlife Refuge (BWR) is part of the Boquerón Commonwealth Forest (BCF) (Figures 21AD), is filled with freshwater and was constructed in 1963-1964 (PRDNR, 1976). The BWR
impoundment was constructed to replace the loss of wetland habitat caused by the
eutrophication of Cartagena Lagoon and the loss of the natural lagoons, Guánica and
Anegado which were being drained for agriculture purpose at the time. To control the flow
of freshwater from the east, and seawater from the west, into and out of the impoundment a
total of six floodgates were constructed along the dikes (PRDNR, 1976; Vázquez, 1983).
Today the impoundment serves as both a waterfowl and hunting refuge. The geology of the
region was treated by Volckmann (1984), USDA (1993), Torres-Figueroa (1993), and
Barreto-Orta (1997).
The salinity of RL fluctuated between 30 to 39 g/L, especially at the bottom of the
mouth of the lagoon; however, salinity changes had been as low as 8.0 g/L or as high as 40.2
g/L caused by terrestrial runoff or evaporation (Carvajal-Zamora, 1976; Negrón-González,
1988). The dissolved oxygen of RL was 3.7 to 58 mg/L and has a water temperature of 29.5
to 32.7°C (Candelas et al., 1973; Negrón-González, 1988). Nutrient concentrations at a
depth of 0.5 to 1.0 m for RL was reported as total phosphorous (0.01 to 0.09 mg/L),
phosphate (< 0.01 to 0.023 mg/L), total nitrogen (0.17 to 0.24 mg/L), nitrate (< 0.001 to 0.01
mg/L), nitrite (0.004 to 0.016 mg/L), and ammonium (< 0.05 mg/L) (Negrón-González,
1988). In general, the water column is considered to be mixed and the oxygen concentrations
are low because of anthropogenic pollution (Negrón-González, 1988).
Mangroves are represented by R. mangle, A. germinans, Laguncularia racemosa,
Conocarpus erectus, and Thespesia populnea which form thick coastal woodland
143
surrounding the lagoon (Table IV) (Figures 21A-D). The terrestrial vegetation of BCF was
treated by Toro and Colón (1986), Vázquez and Kolterman (1998), and Nieves-Rivera et al.,
(2002). Negrón González (1988) reported 35 genera of phytoplankton in RL, including
Clostrium, Synechococcus, and Phormidium. The bioluminescent dinoflagellate Pyrodinium
bahamense is also present in RL. Fish reported in RL included 15 species, from these,
Oreochromis mossambicus was the most abundant species, followed by Centropomus
undecimalis, Mugil curema, and Caranx latus (Negrón-González, 1988; Aliaume et al.,
1997).
Field Studies
Six stations (subdivided in 3 replicates per station or two communities) were selected
in the shores of RL and delimited by Global Position System (G.P.S.) (Table IV). Stations 1
to 3 are referred through the text as ‘Boquerón Beach Inlet’ (BBI) and the remaining stations
(4 to 6) as ‘Boquerón Wildlife Refuge’ (BWR) (Figures 20 and 21A-D). During February
2002 through January 2003, intertidal sea foam and senescent leaves of A. germinans and R.
mangle were collected from studied sites. Following each harvest, chemical parameters were
taken from study sites. LaMotte® Saltwater Aquarium Kit (Model AG-104), Hach®
Saltwater Aquaculture Test Kit (Model FF-3), and LaMotte® Limnology Kit (Model AM-02)
were used to determined water chemical parameters.
Weather conditions at the time of sampling (February 2002 through January 2003)
were obtained from the nearest weather station, which was located at Magueyes Island, PR
(Station Identification Number 9759110, established on 1 December 1954), located at 17°
58.3’N, 67° 2.8’W, La Parguera, Lajas, southwestern Puerto Rico (NOAA, 2004) and a local
weather station (WEATHER.CNN.COM, 2004). Further meteorological studies have been
carried out in southwestern Puerto Rico which are pertinent to our study (Ewel and
Whitmore, 1973; Glynn, 1973; Ravalo et al., 1986; Winter et al., 1998).
144
Sea Foam—Intertidal sea foam was collected with a sterile spoon and stored in sterile plastic
bags (NASCO, Inc.) monthly (from February 2002 through January 2003). The foam was
allowed to condense into liquid and a solution of 1.0% cotton-blue in lactophenol (1 ml of
solution per 25 ml of foam) was added to stain and preserve the spores (Iqbal and Webster,
1973; Santos-Flores and Betancourt-López, 1997). Sea foam samples were stored in a
refrigerator (4°C) to keep them fresh until examination. These fresh samples were baited in
the laboratory with pieces of sterilized balsa wood.
Mangrove Leaves—A total of 1296 unspoiled, senescent leaves of A. germinans and R.
mangle were collected from the lower part of the crown and used as bait. The leaves were
dipped in 70% ethanol for 5 seconds, immersed in 10% hypochlorite for 90 seconds and
finally rinsed in sterile water for 10 seconds. The leaves were dried in a NESCO food
dehydrator (FD-1010 Gardenmaster®), and UV irradiated for 24 hrs. A set of three leaves of
A. germinans and 3 leaves of R. mangle were separately enclosed in two 1.8 mm2 mesh
plastic-screen packet (as in Padgett, 1976) (Figure 22A-C). Preliminary experiments with
various types of traps revealed that the flat design used here, where the leaves were tightly
sandwiched between two layers of screening, allowed maximum exposure to the water and
resulted in lesser build-up of sediment in the mesh. The packets were next arranged into
groups, each of which contained enough packets of each species to serve as one of triplicates
of each parameter to be measured when the group was harvested. All packets of a group
were tied together with a nylon line and anchored to the underwater prop roots of R. mangle.
Leaf baits and traps were prepared according to Padgett (1976).
Mangrove leaves were transported to the laboratory in sterile polythene bags, washed
thoroughly in running water and processed within 24 hrs of collection. Leaf baits were
placed in wet chambers at room temperature (25°C) following Agrios (1997: pp. 255-258)
method to induce fungal sporulation. A sterilized square (~ 1 cm3) of potato dextrose agar
(PDA) was placed on each mangrove leaves while in wet chambers (wet chambers as
described in Agrios, 1997). Leaves were incubated at room temperature (25±2°C) in sterile
145
Petri dishes with sterile freshwater. Each agar square was screened for the presence of fungal
structures within a month of sampling. The Petri dishes were re-hydrated with sterile water
once every two weeks.
Laboratory Studies
Media—According to Johnson and Sparrow (1961), filtered aged seawater is recommended
for primary media preparation, although we used local filtered aged brackish water and on
pure cultures filtered fresh water. The medium used in this study was potato dextrose agar
(PDA) (Difco Laboratories, 1998), prepared with 39 g PDA in 1.0 L of water, 10%
kasugamycin (hydrochloride), and 10% streptomycin, pH 5.6 ± 0.2 at 25°C.
Observations and Herbarium Collections —Microscopical observations were achieved by
using semipermanent slides mounted in lactophenol with cotton blue. The illustrated keys
are described elsewhere (Ellis, 1971, 1976; Ellis and Ellis, 1985; Barnett and Hunter, 1987;
Kohlmeyer and Volkmann-Kohlmeyer, 1991; Santos-Flores and Betancourt-López, 1997;
Hyde and Sarma, 2000). Permanent slides were prepared according to Volkmann-Kohlmeyer
and Kohlmeyer (1996). Voucher specimens or slides of all species collected were deposited
in the Center for Forest Mycology Research at Sabana Field Station (CFMR) and the
National Fungus Herbarium (BPI).
Statistical and Ecological Analyses—A two-way analysis of variance (ANOVA) (Steel and
Torrie, 1980; Sokal and Rohlf, 1995) was employed in a design completely randomized with
2 x 12 factorial treatments. The first factor to study was the two collection sites (BBI, BWR)
and the second was sampling time (12 months). The two-way ANOVA test was used to
determine any relationship between the chemo-physical parameters versus the occurrence of
eight selected filamentous fungi in collection sites of the estuary. Computer statistical
analysis was performed by using SPSS Graduate Pack 10.0 for Windows. The null
hypothesis (H0) proposed for this study is that sampling time (mm-yy), site or location, and
146
the combination of both sources (intersect) will not affect significatively (p < 0.05) two
filamentous fungal communities in a tropical estuary. I used a Type I error of 5% (α = 0.05)
for all hypotheses tested (Portney and Walkins, 1993).
RESULTS AND DISCUSSION
Environmental Features— Atmospheric and chemical environmental parameters
were taken from study sites (Table V). Barometric pressure minimum (1009.4 mb) and
maximum (1015.6 mb) values were obtained from BBI on July and May respectively (Table
V). The air temperature minimum (24.8°C) and maximum (36.0°C) were recorded from
BWR in October and January (Table V). The relative humidity obtained from BWR had a
lower value (71.0%), while the highest (88.0%) on March was obtained from BBI on January
(Table V). The annual rainfall in Magueyes Island Marine Laboratories (MIML) (La
Parguera, southwestern Puerto Rico) showed a minimum (0 mm in July 2002) and a
maximum (99.12 mm in April 2002) values, respectively (Figure 23). Therefore MIML was
selected because it has the same xeric conditions as BWR and its proximity to the study site.
MIML climatic data for February 2002 to January 2003 was summarized in Figures 23-27.
In general, seasonal variation in mean seawater temperature follows closely that of
the atmosphere, ranging between 25 to 30°C, and reaching a maximum in August-October
and a minimum in February (NOAA and PRDNR, 1984) (Figure 27). BWR showed a
minimum and maximum values for surface water temperature (0 through 60 cm depth) of
24.0°C in May and 31.0°C in April (Table V; Figure 28). In general, annual variations in
surface water temperature in southwestern Puerto Rico are generally less than 15.3°C (Coker
and González, 1960; NOAA and PRDNR, 1984). Increased winds in the spring appear to be
related to a slight decline in the surface thermal structure (Glynn, 1973) and for the sampling
year of this study, the wind direction was mostly from the southwest (Figure 25).
Seawater salinity values normally vary less than 2.3 g/L in southwestern Puerto Rico
(NOAA and PRDNR, 1984). The lesser value (12 g/L) for salinity was obtained from BWR
on April and May, while the higher value was 38.0 g/L from BBI on February (Figure 29).
147
Salinity data obtained in BWR indicated a fluctuation of 12.0 g/L (minimum) through 34.0
g/L (maximum) between February and May respectively (Table V, Figure 29). These salinity
values are similar to offshore values indicating that circulation of the inner shelf region is
adequate to maintain open water salinity conditions (NOAA and PRDNR, 1984). Tropical
storms in the region usually lower salinities considerably, thereby temporarily stressing
marine communities (NOAA and PRDNR, 1984). Runoff from heavy rainfall, resuspension
of bottom sediments by wave action and plankton blooms occasionally create turbid
conditions. Water clarity within southwestern Puerto Rico varies only from about 1.0 m near
the shore to about 30 m at the edge of the shelf (NOAA and PRDNR, 1984).
Minimum (6.2) and maximum (8.6) values of pH were obtained from BBI for
October and December, respectively (Figure 30). Two secondary peaks (6.39 and 8.59) were
obtained for BWR and BBI in October, respectively (Figure 30). In December, the minimum
value of dissolved oxygen for BWR was 4.4 mg/L, while in October the maximum value was
8.5 mg/L for BBI (Figure 31). In December, the minimum value of alkalinity for BBI was
127.44 mg/L, while in February the maximum value was 575.81 mg/L for BWR (Figure 32).
Fungal Occurrence—According to the two-way ANOVA performed to the communities of
filamentous fungi (propagules and spore counts, N = 432) in sea foam and the ones isolated
from mangrove leaves (colonies in A. germinans, N = 576; in R. mangle N = 576), fungal
variation was significant (p > 0.05) between the two communities studied (BBI and BWR)
(Table VII). Statistical analysis suggested that intersept (date/site) was to be significant (p >
0.05), although date was insignificant (p < 0.05) in regards to H0 (Table VII). Thus, this
analysis reject H0, in other words, it is unlikely that chance is operating to produce observed
differences and the evidence is strong enough to support the significant effect, one that is
probably not due to chance.
In sea foam samples from BBI and BWR, the microfungus Cochliobolus pallescens
had the higher spore count (115.67 spores) in November, followed by Exserohilum sp. with
13.67 spores at the same month (Table VI, Figure 33). Similarly, at the same dates in BWR,
148
C. pallescens had the higher spore count (144.00 spores), followed by Exserohilum sp.
having 35.00 spores (Figure 34). Smaller peaks are noticeable in July (C. pallescens with
31.33 spores) and April (C. pallescens with 20.00 spores) (Figure 34).
In the A. germinans leaf baits from BBI, contained 72.33 colonies of Penicillium
roqueforti var. carneum in May and a second peak (52.33 colonies) in November (Figure
35). The second most abundant fungus was Aspergillus niger with 54.67 colonies and a third
smaller peak for Pestalotiopsis disseminata with 32.00 colonies, both in May (Figure 35). In
BWR, 98.33 colonies of P. roqueforti var. carneum were isolated in November and 70.67
colonies in May (Figure 36). Both peaks were followed by A. niger with 65.33 colonies in
May and 59.00 colonies in November (Figure 36).
From leaf baits of Rhizophora mangle in BBI 83.33 colonies were isolated of P.
roqueforti var. carneum in May and 56.33 colonies in November (Figure 37). The second
most abundant fungus was A. niger with 66.00 colonies in May and 38.67 colonies in
November (Figure 37). Two more peaks are noticeable, one by C. pallescens (21.00
colonies) in November and the other by P. disseminata (20.67 colonies) in May (Figure 37).
In BWR, P. roqueforti var. carneum had 100.00 colonies isolated in November and 78.67
colonies in May, and a third peak (47.33) appeared in February, and a fourth (25.33 colonies)
in August (Figure 38). Pestalotiopsis disseminata also showed a small peak (15.67 colonies)
in August (Figure 38). All these fungi were abundant in May and November (Figure 38)
possibly because of the run-off input to RL by usually high rainfall (compare with Figure 23)
in April to June and occurred again in August to October (Figure 38).
From the samples collected, I selected eight filamentous fungi, of which two belong
to the ascomycetes and six to the mitosporic fungi (Table VI). The species Aigialus sp. (A.
cf. grandis) (Ascomycota) and Penicillium roqueforti var. carneum (Mitosporic fungi) were
new records for Puerto Rico. Cochliobolus pallescens (BBI = 50.51%; BWR = 50.98%),
Exserohilum sp. (BBI = 16.95%; BWR = 15.37%), Tetraploa aristata (BBI = 11.74%; BWR
= 14.26%), and Alternaria sp. (BBI = 10.88%; BWR = 9.53%), were frequently found in sea
foam, while P. roqueforti var. carneum and Aspergillus niger were frequently found in
149
mangrove leaves (A. germinans and R. mangle) in both BBI and BWR, followed closely by
Pestalotiopsis disseminata, and Alternaria sp. in abundance (Figure 39, Table VII).
Other fungal species were also isolated from BBI and BWR sea foam and leaf litter:
Pleospora sp., Torpedorpora radiata (Ascomycota), Beltrania rhombica, Brachiosphaera
tropicalis, Camposporidium sp., Curvularia robusta, Curvularia sp., Diplocladiella
scalaroides, Helicomyces torquatus, Stemphylum aff. gracilariae, and Zalerion cf.
maritimum (Mitosporic fungi). Camposporidium sp. conidia resembled Camposporidium
spp. reported from Río Sonadora and Quebrada Jiménez next to El Verde LTER Field
Station, Puerto Rico (Hamilton, 1973: Figs. 36 and 39) and the one isolated from Río Manatí
mouth by Nieves-Rivera and Santos-Flores (2004). Some specimens of B. tropicalis showed
a halo around the central cell, while other B. tropicalis were typical, as reported by NievesRivera and Santos-Flores (in press). Aquatic hyphomycetes are frequently found in estuarine
habitats (Johnson and Sparrow, 1961; Kohlmeyer and Kohlmeyer, 1979). For example, Kirk
(1969) reported two lignicolous aquatic hyphomycetes (Clavatospora stellatacula and
Tetraploa aristata) adapted to seawater conditions on Chesapeake Bay. Tetraploa aristata
have been previously isolated from dead stalks and leaves of sugar cane Saccharum
officinarum in Puerto Rico (Minter et al., 2001). Stemphylium botryotium was isolated from
Boquerón Public Beach by Hernández-Vera (1972, 1975). The BBI and BWR specimens are
similar in spore shape to Stemphylium gracilariae although its size is somewhat smaller, thus
I named it Stemphylium aff. gracilariae.
In conclusion, few marine ascomycetes were isolated probably due to the selectivity
of the isolation method employed, substrates, environmental conditions, and geomorphology
of the lagoon. Ascomycetes and mitosporic fungi (mostly aquatic hyphomycetes) are found
mixed in foam, leaf litter, and beach sand of RL. Most of the species studied are terrestrial
hyphomycetes that might have been introduced by the river flow or by land run-off into
riverine or brackish stagnant waters. All species isolated in this study are saprobes living in
parts of angiosperms, mangroves, debris, or in the blades of seagrasses.
150
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156
Table IV. Study sites in Boquerón Beach Inlet (BBI) and Boquerón Wildlife Refuge (BWR), both part of the Rincón Lagoon,
Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico.
________________________________________________________________________________________
Locality
Collection Sites
Coordinates
Mangroves and Associates
________________________________________________________________________________________
BBI
Station 1
18°00’68.4” N, 67°10’32.2” W
BBI
Station 2
18°00’67.8” N, 67°10’31.4” W
BBI
Station 3
18°00’47.1” N, 67°10’65.2” W
BWR
Station 4
18°01’07.2” N, 67°09’94.6” W
BWR
Station 5
18°00’95.4” N, 67°09’99.4” W
BWR
Station 6
18°00’65.3” N, 67°10’12.2” W
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Thespesia populnea
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Acrostichum spp.
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Thespesia populnea
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Thespesia populnea
Avicennia germinans
Laguncularia racemosa
Rhizophora mangle
Thespesia populnea
________________________________________________________________________________________
157
Table V. Monthly and annual environmental averages of study sites (Boquerón Beach Inlet: Stations 1-3; Boquerón Wildlife Refuge:
Stations 4-6) from Rincón Lagoon, Cabo Rojo, southwestern Puerto Rico.
Boquerón Beach Inlet (BBI)
Date
(mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Annual
Average
Barometric
Pressure
(mb)
Air
Temperature
(°C)
Relative
Humidity
(%)
Salinity
(g/L)
Water
Temperature
(°C)
pH
Turbidity
(JTU)
Dissolved
Oxygen
(mg/L)
Alkalinity
(mg/L)
Ammonium
(mg/L)
Nitrate
(mg/L)
Nitrite
(mg/L)
1012.33
1013
1014
1009.41
1015
1015.57
1010.89
1010.67
1013
1013
1014
1012
25
25.07
25.56
32
32.83
34.33
32.78
32.5
33
27.83
27
25.17
76
84.07
78.33
83.33
80.22
74.33
78.22
80.89
73.33
75.78
83.33
71.33
37.78
37.72
38
38
37
37
35.11
34.33
34
34
33
36.39
26.22
27.85
24
31.22
30
30.67
29.89
28.96
29.89
29.54
27.66
26.67
7.79
7.72
8
7.88
8.23
8.17
7.92
7.88
8.59
7.77
6.2
8.53
37.22
28.33
10
60
31.11
6.67
43.33
20
26.67
0
53.33
8.33
6.92
8.06
7.13
7.09
7
6.01
5.54
6.47
5.27
6.94
8.37
6.49
161.63
142.04
146.93
149.3
143.96
144.56
131.96
138.3
140.22
139.22
127.44
131
0
0
0
0
0
0.03
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1012.74
29.42
78.26
36.03
28.55
7.89
27.08
6.77
141.38
0.00
0.00
0.00
Boquerón Wildlife Refuge (BWR)
Date
(mm-yy)
Barometric
Pressure
(mb)
Air
Temperature
(°C)
Relative
Humidity
(%)
Salinity
(g/L)
Water
Temperature
(°C)
pH
Turbidity
(JTU)
Dissolved
Oxygen
(mg/L)
Alkalinity
(mg/L)
Ammonium
(mg/L)
Nitrate
(mg/L)
Nitrite
(mg/L)
Feb-02
Mar-02
Apr-02
May-02
1014
1014.33
1013
1009.89
26
30.83
29
29.56
0
87
79.89
80.78
12
22.67
20.33
17.63
31.07
30.33
30
28.94
8.31
7.94
8.17
7.54
100
100
100
100
5.22
6.56
7.7
6.51
575.81
167.22
406.04
223.96
0
0
0
0
0
0
0
1
0
0
0
0.02
158
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Annual
Average
1014.89
1013.33
1010.22
1010.67
1011
1012.33
1014
1012.67
28.28
34
33
32.56
36.17
27.24
31.56
24.5
77.89
75.67
76.33
77.11
75.33
78.76
79
77.93
31.44
27.99
24.22
22.33
29.56
30.28
32.33
33.67
28.33
30.89
29.85
29.78
30.2
30.11
28.67
27
7.93
8.17
8.01
8.08
6.39
7.86
8.07
7.48
100
100
100
95
86.67
77.78
86.67
100
5.87
5.35
5.62
5.86
4.33
6.86
7.4
7.28
299.33
274.44
412.41
141.56
285.81
300.96
133
133.89
0
0
0
0
0
0
0
0
0.98
1.1
0
0
0
0
0
0
0.02
0
0
0
0
0
0
0
1012.53
30.23
72.14
25.37
29.60
7.83
95.51
6.21
279.54
0.00
0.26
0.00
159
Table VI. Monthly and annual averages of selected filamentous fungi isolated from sea foam and mangrove leaves from collection sites in
Rincón Lagoon, Boquerón Commonwealth Forest, southwestern Puerto Rico during February 2002 through January 2003.
Sea Foam/Boquerón Beach Inlet1
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Total = 29.14
Aigialus sp.
Alternaria sp.
0.33
4.67
2.33
2.00
0.67
1.00
2.00
1.00
1.00
1.00
0.67
1.33
1.50 = 5.15 %
4.33
2.33
0.67
0.67
2.00
9.00
1.33
1.67
1.67
8.33
5.67
0.33
3.17 = 10.88 %
Cochliobolus
pallescens
3.33
2.67
3.00
5.00
13.33
23.67
2.33
0.33
4.33
115.67
0.33
2.67
14.72 = 50.51 %
Exserohilum sp.
10.00
6.33
4.33
2.00
1.33
8.00
3.33
2.67
2.33
13.67
1.33
4.00
4.94 = 16.95 %
Pestalotiopsis
disseminata
1.00
0.67
0.33
1.33
0.33
1.00
0.33
1.33
1.00
6.00
2.67
0.67
1.39 = 4.77 %
Tetraploa
aristata
0.33
2.00
1.00
1.33
3.33
11.00
1.33
6.67
9.00
4.00
0.33
0.67
3.42 = 11.74 %
Pestalotiopsis
disseminata
0.67
3.67
0.67
0.67
3.00
Tetraploa
aristata
5.00
17.33
3.33
3.33
4.67
Sea Foam/Boquerón Wildlife Refuge1
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Aigialus sp.
Alternaria sp.
1.00
12.67
2.67
0.00
1.67
3.00
6.33
8.00
3.00
1.33
Cochliobolus
pallescens
8.33
11.67
20.00
8.67
11.67
160
Exserohilum sp.
4.00
6.00
12.67
3.33
2.67
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Total = 42.29
5.67
0.00
0.00
0.67
1.33
3.00
0.00
2.39 = 5.65 %
13.00
1.00
0.00
2.33
9.00
1.00
0.33
4.03 = 9.53 %
31.33
1.67
0.33
5.33
144.00
13.33
2.33
21.56 = 50.98 %
7.67
0.33
0.33
0.00
35.00
6.00
0.00
6.50 = 15.37 %
0.33
0.00
0.00
1.00
9.00
2.33
0.00
1.78 = 4.21 %
10.00
0.00
6.67
13.33
7.00
1.67
0.00
6.03 = 14.26 %
Avicennia germinans Leaves/Boquerón Beach Inlet
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Total = 80.90
Aigialus sp.
Alternaria sp.
Aspergillus niger
Cochliobolus
pallescens
Exserohilum
sp.
Pestalotiopsis
disseminata
Penicillium
roqueforti2
Tetraploa
aristata
1.67
0.67
1.00
5.67
5.67
3.00
3.33
2.33
3.67
0.00
4.00
0.00
2.58 = 3.19 %
3.00
8.67
3.67
9.00
5.33
1.33
5.00
13.33
8.67
0.33
1.67
1.33
5.11 = 6.32 %
45.00
29.33
15.67
54.67
29.67
26.67
23.33
12.67
22.67
46.67
31.67
13.67
29.31 = 36.23 %
3.67
1.33
3.00
5.00
15.00
2.67
0.33
1.00
4.33
14.33
3.33
2.00
4.67 = 5.77 %
0.67
0.33
3.33
0.33
0.00
0.67
2.00
0.00
0.67
0.33
0.33
0.00
0.72 = 0.89 %
5.67
15.33
2.00
32.00
2.33
6.33
0.33
9.67
10.00
17.33
3.67
5.33
9.17 = 11.33 %
28.67
30.33
33.33
72.33
20.67
18.33
23.67
29.00
3.33
52.33
25.00
11.67
29.06 = 35.92 %
0.33
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.28 = 0.35 %
Avicennia germinans Leaves/Boquerón Wildlife Refuge
161
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Total = 73.87
Aigialus sp.
Alternaria sp.
Aspergillus niger
Cochliobolus
pallescens
Exserohilum
sp.
Pestalotiopsis
disseminata
Penicillium
roqueforti2
Tetraploa
aristata
1.33
0.00
0.67
3.00
0.33
0.00
0.33
0.00
0.00
0.00
0.00
0.00
0.47 = 0.64 %
3.00
1.67
1.00
1.67
0.67
4.33
0.67
2.33
2.33
0.00
0.00
1.00
1.55 = 2.10 %
45.00
18.67
18.67
65.33
33.33
24.00
23.33
7.67
24.67
59.00
47.67
13.33
31.72 = 42.94 %
0.67
0.33
4.67
1.00
0.33
2.33
0.33
0.67
0.33
2.67
2.33
0.33
1.33 = 1.80 %
0.00
0.00
3.00
1.67
1.00
1.00
2.00
1.33
0.00
3.67
4.00
0.00
1.47 = 1.99 %
2.33
1.00
0.33
4.67
1.67
2.33
3.00
1.33
1.67
0.33
1.33
0.00
1.67 = 2.26 %
51.67
41.33
48.00
70.67
14.67
12.00
14.00
4.33
4.00
98.33
19.33
29.67
34.00 = 46.03 %
0.00
0.00
0.00
0.33
0.33
0.00
0.00
0.00
0.33
0.00
1.00
0.00
1.66 = 2.25 %
Rhizophora mangle Leaves/Boquerón Beach Inlet
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Aigialus sp.
Alternaria sp.
Aspergillus niger
Cochliobolus
pallescens
Exserohilum
sp.
Pestalotiopsis
disseminata
Penicillium
roqueforti2
Tetraploa
aristata
3.67
0.67
8.00
8.00
5.00
3.00
4.33
3.33
7.00
0.00
3.33
0.33
6.67
9.00
6.00
9.67
8.33
0.33
3.33
9.67
13.67
2.00
4.67
0.33
33.67
37.67
14.33
66.00
25.33
27.33
22.67
7.67
29.00
38.67
24.67
17.67
4.33
1.33
2.33
3.67
11.00
6.33
0.33
3.33
2.67
21.00
4.00
6.33
0.33
1.00
4.33
2.33
0.00
2.00
4.00
0.33
1.67
0.67
1.00
0.00
6.67
13.00
2.00
20.67
5.67
7.00
2.00
8.67
8.33
10.67
11.67
4.67
29.33
23.00
25.00
83.33
23.67
24.33
25.33
26.33
6.00
56.33
12.00
12.33
0.33
0.00
0.67
0.33
0.33
0.00
0.00
0.00
0.00
0.00
0.00
0.00
162
Total = 83.25
3.89 = 4.68 %
6.14 = 7.38 %
28.72 = 34.50 %
5.55 = 6.67 %
1.47 = 1.77 %
8.42 = 10.11 %
28.92 = 34.74 %
0.14 = 0.17 %
Rhizophora mangle Leaves/Boquerón Wildlife Refuge
Date (mm-yy)
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Total = 80.70
Aigialus sp.
Alternaria sp.
Aspergillus niger
Cochliobolus
pallescens
Exserohilum
sp.
Pestalotiopsis
disseminata
Penicillium
roqueforti2
Tetraploa
aristata
2.33
0.00
2.33
8.00
0.00
0.67
1.67
1.67
3.00
0.00
1.00
0.00
1.72 = 2.13 %
6.67
2.67
5.67
3.00
1.00
3.00
3.67
3.33
6.33
0.00
0.00
1.67
3.08 = 3.82 %
33.67
39.33
32.33
48.33
22.00
25.33
22.67
5.33
30.33
76.67
30.33
18.67
32.08 = 39.75 %
1.67
1.00
6.00
1.33
0.67
2.00
0.33
1.67
2.33
7.33
5.33
1.00
2.56 = 3.17 %
0.00
0.00
7.67
4.00
1.00
1.67
4.00
5.67
0.00
1.67
4.67
0.00
2.53 = 3.14 %
4.00
1.67
1.00
9.00
5.67
2.33
15.67
1.67
3.67
2.33
6.00
0.00
4.42 = 5.48 %
47.33
31.67
46.67
78.67
15.33
12.00
25.33
18.33
6.67
100.00
16.00
8.67
33.89 = 42.00 %
0.00
0.00
1.00
0.67
0.67
0.00
0.00
0.00
2.00
0.67
0.00
0.00
0.42 = 0.52 %
1
Aspergillus niger and Penicillium roqueforti var. carneum conidia were not counted from sea foam due to difficulties in isolation and
identification of conidia by morphology alone.
2
Although named Penicillium roqueforti in this table, it should be noticed that this species was identified as Penicillium roqueforti var.
carneum (M. Klich, pers. comm., 2004).
163
Table VII. Univariate analysis of variance (Two-way ANOVA) of filamentous fungi
collected from sea foam and mangrove leaves (Avicennia germinans and Rhizophora
mangle) from Boquerón Beach Inlet (BBI) and Boquerón Wildlife Refuge (BWR)
collection sites at Rincón Lagoon, Boquerón Commonwealth Forest, southwestern
Puerto Rico during February 2002 through January 2003 (α = 0.05).
Univariate Analysis of Variance (SEA FOAM)
Between-Subjects Factors
Value Label
Date
Site
N
FEB 02
36
MAR 02
36
APR 02
36
MAY 02
36
JUN 02
36
JUL 02
36
AUG 02
36
SEP 02
36
OCT 02
36
NOV 02
36
DEC 02
36
JAN 03
36
1
BWR
2
BBI
216
216
Tests of Between-Subjects Effects
Dependent Variable: Seafoam spore count
Source
Intercept
Hypothesis
Error
DATE
Hypothesis
Error
SITE
Hypothesis
Error
Type III Sum
of Squares
15301.021
df
517.891
1
Mean Square
15301.021
1
517.891a
24260.507
11
2205.501
101463.581
419
242.157b
517.891
1
101463.581
419
a. MS(SITE)
b. MS(Error)
164
517.891
242.157b
F
29.545
Sig.
.116
9.108
.000
2.139
.144
Univariate Analysis of Variance (Avicennia germinans)
Between-Subjects Factors
Value Label
Date
N
FEB 02
48
MAR 02
48
APR 02
48
MAY 02
48
JUN 02
48
JUL 02
48
AUG 02
48
SEP 02
48
OCT 02
48
NOV 02
48
DEC 02
48
JAN 03
Site
1
2
48
BWR
288
BBI
288
Tests of Between-Subjects Effects
Dependent Variable: Fungal spore count in Avicennia
Source
Intercept
Hypothesis
Error
DATE
Hypothesis
Error
SITE
Hypothesis
Error
Type III Sum
of Squares
52689.377
1
Mean Square
52689.377
153.141
1
153.141a
13583.686
11
1234.881
563
275.056b
154856.797
df
153.141
1
154856.797
563
a. MS(SITE)
b. MS(Error)
165
153.141
275.056b
F
344.059
Sig.
.034
4.490
.000
.557
.456
Univariate Analysis of Variance (Rhizophora mangle)
Between-Subjects Factors
Value Label
Date
N
FEB 02
48
MAR 02
48
APR 02
48
MAY 02
48
JUN 02
48
JUL 02
48
AUG 02
48
SEP 02
48
OCT 02
48
NOV 02
48
DEC 02
48
JAN 03
Site
48
1
BWR
2
BBI
288
288
Tests of Between-Subjects Effects
Dependent Variable: Fungal spore count in Rhizophora
Source
Intercept
Hypothesis
Error
DATE
Hypothesis
Error
SITE
Hypothesis
Error
Type III Sum
of Squares
60475.007
1
Mean Square
60475.007
14.694
1
14.694a
14667.660
11
1333.424
563
260.122b
146448.639
df
14.694
146448.639
1
563
a. MS(SITE)
b. MS(Error)
166
14.694
260.122b
F
4115.501
Sig.
.010
5.126
.000
.056
.812
Figure 20. Location of sampling sites at Rincón Lagoon, Boquerón Commonwealth Forest,
Cabo Rojo, southwestern Puerto Rico. Stations 1 to 3 are referred through the text as
‘Boquerón Beach Inlet’ (BBI) and the remaining stations (4 to 6) as ‘Boquerón
Wildlife Refuge’ (BWR).
167
Figure 21A–D. Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico. A.
Aerial photo of the Boquerón Bay, including view of the Rincón Lagoon, Boquerón
Wildlife Refuge and Puerto Real. B–C. Rhizophora mangle (B) and Avicennia
germinans (C) forming channels in the Boquerón Wildlife Refuge. D. Boardwalk
prepared by the Puerto Rico Department of Natural Resources and brackish water
lagoon.
168
Figure 22A–C. Leaf baits and traps used for the isolation of estuarine fungi of Rincón
Lagoon, Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico. A.
Leaf baits (Rhizophora mangle) in 1.8 mm2 mesh plastic-screen packet contained in a
1.0 cm2 meshed aluminum envelope. B. Leaves of R. mangle culture in a Petri dish.
C. Potato dextrose agar (PDA) with fungal isolates.
169
120
99.12
100
82.8
Precipitation (mm)
80
60
54.48
40.32
40
36.48
34.32
27.36
17.04
20
11.52
7.44
3.36
0
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm/yy)
Figure 23. Annual rainfall (February 2002 to January 2003) in Magueyes Island Marine Laboratories, La
Parguera, southwestern Puerto Rico.
Figure 24. Average air temperature (°C) during February 2002 to January 2003 in Magueyes Island Marine
Laboratories, La Parguera, southwestern Puerto Rico (graph courtesy of NOAA/NOS/CO-OPS).
170
Figure 25. Average wind speed (m/s) and direction (true) during February 2002 to January 2003 in Magueyes
Island Marine Laboratories, La Parguera, southwestern Puerto Rico (graph courtesy of
NOAA/NOS/CO-OPS).
Figure 26. Water level (m) during February 2002 to January 2003 in Magueyes Island Marine Laboratories, La
Parguera, southwestern Puerto Rico (graph courtesy of NOAA/NOS/CO-OPS).
171
Figure 27. Average water temperature (°C) during February 2002 to January 2003 in Magueyes Island Marine
Laboratories, La Parguera, southwestern Puerto Rico (graph courtesy of NOAA/NOS/CO-OPS).
35
30
Water Temperature (°C)
25
20
15
10
Boquerón Beach Inlet
5
Boquerón Wildlife Refuge
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 28. Average water temperature (°C) of sampling sites at Rincón Lagoon, Boquerón Commonwealth
Forest, southwestern Puerto Rico.
172
40
35
30
Salinity (g/L)
25
20
15
10
Boquerón Beach Inlet
5
Boquerón Wildlife Refuge
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 29. Average salinity (g/L) of sampling sites at Rincón Lagoon, Boquerón Commonwealth Forest,
southwestern Puerto Rico.
10
9
8
7
pH
6
5
4
3
2
Boquerón Beach Inlet
1
Boquerón Wildlife Refuge
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 30. Average pH of sampling sites at Rincón Lagoon, Boquerón Commonwealth Forest, southwestern
Puerto Rico.
173
9
8
Dissolved Oxygen (mg/L)
7
6
5
4
3
2
Boquerón Beach Inlet
1
Boquerón Wildlife Refuge
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 31. Average dissolved oxygen (mg/L) of sampling sites at Rincón Lagoon, Boquerón Commonwealth
Forest, southwestern Puerto Rico.
700
Boquerón Beach Inlet
600
Boquerón Wildlife Refuge
Alkalinity (mg/L)
500
400
300
200
100
0
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 32. Average alkalinity (mg/L) of sampling sites at Rincón Lagoon, Boquerón Commonwealth Forest,
southwestern Puerto Rico.
174
140.00
Aigialus sp.
Alternaria sp.
120.00
Cochliobolus pallescens
Exserohilum sp.
Number of Spores
100.00
Pestalotiopsis disseminata
Tetraploa aristata
80.00
60.00
40.00
20.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 33. Number of fungal spores in sea foam per month during the year of sampling at Boquerón Beach
Inlet, Rincón Lagoon, southwestern Puerto Rico.
160.00
Aigialus sp.
Alternaria sp.
140.00
Cochliobolus pallescens
Exserohilum sp.
120.00
Number of Spores
Pestalotiopsis disseminata
Tetraploa aristata
100.00
80.00
60.00
40.00
20.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 34. Number of fungal spores in sea foam per month during the year of sampling at Boquerón Wildlife
Refuge, Rincón Lagoon, southwestern Puerto Rico.
175
80.00
Aigialus sp.
Alternaria sp.
70.00
Aspergillus niger
Cochliobolus pallescens
Exserohilum sp.
60.00
Pestalotiopsis disseminata
Number of Colonies
Penicillium roqueforti
Tetraploa aristata
50.00
40.00
30.00
20.00
10.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 35. Number of fungal colonies in Avicennia germinans leaves per month during the year of sampling at
Boquerón Beach Inlet, Rincón Lagoon, southwestern Puerto Rico.
120.00
Aigialus sp.
Alternaria sp.
Aspergillus niger
100.00
Cochliobolus pallescens
Exserohilum sp.
Pestalotiopsis disseminata
Penicillium roqueforti
Number of Colonies
80.00
Tetraploa aristata
60.00
40.00
20.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 36. Number of fungal colonies in Avicennia germinans leaves per month during the year of
sampling at Boquerón Wildlife Refuge, Rincón Lagoon, southwestern Puerto Rico.
176
90.00
Aigialus sp.
Alternaria sp.
80.00
Aspergillus niger
Cochliobolus pallescens
70.00
Exserohilum sp.
Pestalotiopsis disseminata
Penicillium roqueforti
Number of Colonies
60.00
Tetraploa aristata
50.00
40.00
30.00
20.00
10.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 37. Number of fungal colonies in Rhizophora mangle leaves per month during the year of sampling at
Boquerón Beach Inlet, Rincón Lagoon, southwestern Puerto Rico.
120.00
Aigialus sp.
Alternaria sp.
Aspergillus niger
100.00
Cochliobolus pallescens
Exserohilum sp.
Pestalotiopsis disseminata
Penicillium roqueforti
Number of Colonies
80.00
Tetraploa aristata
60.00
40.00
20.00
0.00
Feb-02
Mar-02
Apr-02
May-02
Jun-02
Jul-02
Aug-02
Sep-02
Oct-02
Nov-02
Dec-02
Jan-03
Date (mm-yy)
Figure 38. Number of fungal colonies in Rhizophora mangle leaves per month during the year of sampling at
Boquerón Wildlife Refuge, Rincón Lagoon, southwestern Puerto Rico.
177
70
Boquerón Beach Inlet
59.2
Percent of Fungal Isolates (%)
60
Boquerón Wildlife Refuge
52.27
50.78
47.73
50
49.22
40.8
40
30
20
10
0
Sea Foam
Avicennia germinans
Rhizophora mangle
Figure 39. Total percentage of fungal isolates from substrates (sea foam, leaves of Avicennia
germinans, and Rhizophora mangle) in the sampling sites at Rincón Lagoon,
Boquerón Commonwealth Forest, Cabo Rojo, southwestern Puerto Rico.
178
CHAPTER 5
CHARACTERIZATION OF CLADOSPORIUM OXYSPORUM AND C.
SPHAEROSPORUM USING POLYAROMATIC HYDROCARBONS (PAHs) AS
THEIR SOLE CARBON SOURCE IN TROPICAL COASTAL SEAWATER
ABSTRACT
Two species of Cladosporium (C. oxysporum and C. sphaerospermum) were isolated
from surface coastal seawater based on their ability to use the polyaromatic hydrocarbons
(PAHs) naphthalene (C10H8) and phenanthrene (C14H10) as a sole carbon and energy source.
Although both Cladosporium spp. are cosmopolitan species, both species are new records to
mangrove forests of Puerto Rico. These two species may be of value in the bioremediation
of natural oil spills or other contaminants in tropical environments.
RESUMEN
Se aislaron dos especies de Cladosporium (C. oxysporum y C. sphaerospermum) de
agua de mar superficial costera basándose en su habilidad de usar los hidrocarburos
poliaromáticos (PAHs) nafataleno (C10H8) y fenantreno (C14H10) como fuentes de carbono y
de energía. Aunque ambos Cladosporium spp. son especies cosmopolitas, ambas especies
son nuevos registros para los bosques de mangles de Puerto Rico. Estas dos especies pueden
ser de valor en la bioremediación de derrames naturales de aceite u otros contaminantes en
ambientes tropicales.
INTRODUCTION
Polyaromatic hydrocarbons or polycyclic aromatic hydrocarbons (PAHs) are formed
primarily as products from the combustion of fossil fuels. The United States Environmental
Protection Agency lists 16 PAHs as priority pollutants (Zaidi and Imam, 1999). In marine
environments, due to their hydrophobic nature and low water solubility, PAHs are readily
179
adsorbed by particulate matter and tend to accumulate in sediments (Means et al., 1980).
Because of their toxic, mutagenic, or carcinogenic properties, high concentrations of PAHs
are harmful to the marine biota and human health (Stegeman, 1977). In Puerto Rico, for
example, PAHs pollution has induced nuclear mutations for chlorophyll deficiency in trees of
Rhizophora mangle (Corredor et al., 1995).
The fate of most petroleum substances in the marine environment is ultimately
defined by their transformation and degradation due to microbial activity (Bragg et al., 1994).
Microorganisms are the primary means of degrading PAHs naturally. About a hundred
known species of microorganisms (e.g., bacteria and fungi) are able to use oil components to
sustain their growth and metabolism. In pristine areas, their proportions usually do not
exceed 0.1 to 1.0% of the total abundance of heterotrophic bacterial communities; in polluted
areas, however, this proportion increases to 1.0 to 10.0% (Atlas and Bartha, 1997). Most of
the information on degradation of PAHs has being derived from pure cultures that were
isolated from temperate environments (Kirk and Gordon, 1988; Cerniglia and Sutherland,
2001; Qi et al., 2002). Very little information is available on isolation of fungal strains
capable of degrading PAHs in subtropical marine environments (Acevedo, 2001).
Kohlmeyer & Kohlmeyer (1979) and González et al. (1998) have classified members
of the genus Cladosporium as facultative marine fungi. These fungi from freshwater or
terrestrial areas are capable of growing in the marine environment.
Cladosporium
oxysporum and C. sphaerospermum are typically geophilic (soil loving) and cosmopolitan
(deVries, 1952). In this paper, we report the identification and characterization of C.
oxysporum and C. spherospermum (from Cabo Rojo and Guayanilla, respectively, coastal
waters of southwestern Puerto Rico) two species that are capable of degrading PAHs
naphthalene (C10H8) and phenanthrene (C14H10).
MATERIALS AND METHOD
Guayanilla Bay (GB) is located on the south coast of Puerto Rico about 35 km east of
the island’s southwestern corner (Figure 4). This bay was the site of one of the largest
180
petrochemical complexes in the world until these were shut down in 1982 (Zaidi and Imam,
1999). Many studies have determined the fate of PAHs at this bay after closure of industrial
complex (Zaidi et al., 2003). Bahía Sucia (BS) and Los Morrillos (LM) are part of the
Boquerón Commonwealth Forest, Cabo Rojo, and were described by Tosterson et al. (1977)
and Nieves-Rivera et al. (2001). On March 18, 1973, the tanker Zoe Colocotronis ran
aground on a reef 4.8 km off La Parguera (southern Puerto Rico) releasing 1.01 million
gallons of Venezuelan (Tijuana) crude oil on BS shores (Nadeau and Bergquist, 1977;
Corredor et al., 1990).
Naphthalene (99% purity), phenanthrene (> 96% purity), and agarose type VII (low
gelling temperature) were obtained from Sigma Chemical Co., St. Louis, and Noble agar
from Difco Laboratories, Detroit. Seawater and sediment samples were collected from BS,
GB, and LM bays, stored in a refrigerator and used within 2-3 weeks after collection of the
isolates. Isolates were grown on sea water medium (SWM) (Bogart and Hemmingsen, 1992)
and on malt extract agar (MEA) (Ho et al., 1999). Wet mounts were observed at 25-60 x and
100-1000 x with stereo and compound microscopes. Fungal isolates were deposited in the
American Type Culture Collection, ATCC.
Identification of C. oxysporum and C.
sphaerospermum was based on deVries (1952), Ellis (1971), and Ho et al. (1999). In
addition to the authorities cited, other descriptions in a number of other published works
(e.g., Stevens (1981), de Hoog et al. (2000), Samson et al. (2000), Flannigan et al. (2001),
Wang and Zabel (1990)) were useful.
RESULTS AND DISCUSSION
Characters of colonies and morphological structures of ATCC MYA-3068 and MYA3069 on MEA (Figures 40A-G) were generally consistent with descriptions of C.
sphaerospermum and C. oxysporum respectively, in both Ellis (1971) and Ho et al. (1999).
MYA-3068 keys to C. oxysporum in Ho et al. (1999) and in Ellis (1971) C. oxysporum
resembles C. tenuissimum, but the latter lacks the intercalary nodes. The defining character
of this species, rather long conidiophores routinely approaching 500 µm in length and
181
possessing intercalary conidiogenous nodes, was observed in culture. Conidia of MYA-3069
(NJRR-1) were intermediate in size between those described for C. sphaerospermum in Ellis
(1971) and those in Ho et al. (1999). Cladosporium sphaerospermum has been isolated on
extreme halophilic environments (Kis-Papo et al., 2001), in organic chemicals such as
toluene (Weber et al., 1995), and after the fallout of Chernobyl (Zhdanova et al., 2000).
Isolates were grown on SWM and on MEA (Ho et. al. 1999). Characters of colonies
and morphological structures of MYA-3068 and MYA-3069 on MEA (Figures 40A-G) were
generally consistent with descriptions of C. sphaerospermum and C. oxysporum respectively,
in both Ho et al. (1999) and Ellis (1971). Conidia of NJRR-1 were intermediate in size
between those described for C. sphaerospermum in Ellis (1971) and those in Ho et al.
(1999). Characters on SWM were indistinguishable from those on MEA with the exception
of growth rate, which were faster in MEA (2 mm/week) versus the slower SWM (1
mm/week).
Cladosporium oxysporum Berk. & M.A. Curtis, 1869.
Colonies on MEA (Ho et al., 1999) 43-44 mm diameter in 10 days at ca. 25 °C, deep
olive green, lighter and concentrically banded toward margin, surface almost velutinuous to
flocculose. Reverse dark greenish black, lighter at margin. Conidiophores macronematous,
olive, smooth, mostly ca. 300-900 x 3.5-4.5 µm (up to 7.5 µm at nodes), straight, mostly
unbranched, with (0-) 1-5 conspicuously swollen internodes and a swollen apical node
(Figures 40A-D). Conidia with 1-3 scars smooth, olive, mostly oval to elliptical,
occasionally limoniform, mostly (3.5-) 4-6 x 2.5-3.5 µm, the larger conidia grading into
ramoconidia (Figure 40E). Ramoconidia smooth, olive, elliptical to cylindrical with 3-5
scars; up to ca. 20 x 4 µm (Figure 40E). Scars protuberant, dark, on conidia, ramoconidia
and conidiophores. Strain deposited as ATCC MYA-3068 (ÁMNR-7); seawater in R.
mangle roots, Los Morrillos and Bahía Sucia, Cabo Rojo, Puerto Rico, 4 November 2002, Á.
M. Nieves-Rivera.
182
Cladosporium sphaerospermum Penz., 1882.
Colonies on MEA (Ho et al., 1999) 25-26 mm and 36-37 mm on half strength V8
agar (½ V8, Stevens, 1981) at 10 days at circa 25°C; dark olive green, velvety, powdery, and
reverse blackish green. Conidiophores olivaceous, macronematous, straight to flexous, 0-1 (3) branched, intercalary or terminal, smooth, septate, up to ca. 160 µm long, usually circa 30
to 125 µm, up to 3 µm in diameter (slightly expanded at apices), not geniculate. Conidia in
simple or branched chains, globose or subglobose to limoniform, mostly aseptate, moderately
verrucose, olive, mostly 3.5-6.5 x 3.0-4.5 µm; abscission scars darkened, protuberant
(Figures 40F-G). Ramoconidia subglobose to cylindrical, olive, typically aseptate, 7.5-17.5
(-25.0) x 3.5-4.5 µm wide (Figure 40G). Hyphae septate, olivaceous, smooth, up to 3 µm
wide. Strain deposited as ATCC MYA-3069 (NJRR-1); seawater in R. mangle roots, María
Langa Cay, Guayanilla Bay, Guayanilla, Puerto Rico, 31 January 2002, N. J. RodríguezRodríguez.
Figures 40A-G show the arrangement of intact conidial chains on C. oxysporum and
C. sphaerospermum. The top right and middle row show the variation in node placement
and structure (an important character for identification of C. oxysporum). And the insert
(Figure 40E) and bottom right figures (Figure 40G) show the conidia of C. oxysporum and C.
sphaerospermum, respectively. Although both Cladosporium spp. are cosmopolitan species,
they are new records for mangrove forests of Puerto Rico.
Coastal environments of Puerto Rico are prime repositories of PAHs because most
industries as well as urban centers are located on the coast. For instance, for over 20 years
GB was the location of one of the biggest concentrations of petrochemical industries in the
world until it was shut down in 1982. Petrochemical industries and oil spills are the major
source of organic pollutants which are of interest to this study. Although many studies have
been conducted in areas along Puerto Rican coasts to determine the fate of pollutants by
microbial degradation (Zaidi and Imam, 1999; Zaidi et al., 2003), few have been conducted
by using fungal isolates (Acevedo, 2001). This study was baseline information on the
conditions of GB and LM.
183
In conclusion, our particular interest was on the identifying potential bioremediation
species that may be used in natural oil spills or other contaminant clean ups, which is a
crucial part of environmental protection. The two species discussed above may play an
important role in the future. However, we must discover, define, and experiment with
additional naturally occurring species to ensure the protection of our tropical environments.
LITERATURE CITED
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Biology Inter Campus Doctoral Program, Puerto Rico, Río Piedras, Puerto Rico. 85
pp.
Atlas, R. M. and R. Bartha. .1997. Microbial ecology: fundamentals and applications, fourth
edition. Benjamin-Cummings Publishing Company, San Francisco, California. 640
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Bogardt, A. H. and B. B. Hemmingsen. 1992. Enumeration of phenanthrene-degrading
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Bragg, J. R., R. C. Prince, E. J. Harner and R. M. Atlas. 1994. Effectiveness of
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in bioremediation, pp. 136-187. Cambridge University Press, Cambridge.
Corredor, J. E., J. M. Morell and C. E. del Castillo. 1990. Persistence of spilled crude oil in a
tropical intertidal environment. Marine Pollution Bulletin 21: 385-388.
Corredor, J. E., J. M. Morell, E. J. Klekowski, Jr. and R. Lowenfeld. 1995. Mangrove
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Figures 40A-G. Characteristic microscopic features of Cladosporium oxysporum (ATCC
MYA-3068) and C. sphaerospermum (ATCC MYA-3069) from coastal seawater.
A-D. Conidiophores of C. oxysporum on MEA. E. A group of oval to ellipticalshaped conidia and cylindrical ramoconidia of C. oxysporum at high magnification.
F. Ramoconidia and conidia of C. sphaerospermum on ½ V8 agar. G.
Conidiophores, ramoconidia, and conidia of C. sphaerospermum on MEA.
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CHAPTER 6
SOOTY MOLD-PLANTHOPPER ASSOCIATION ON LEAVES OF THE BLACK
MANGROVE AVICENNIA GERMINANS IN SOUTHWESTERN PUERTO RICO
ABSTRACT
Recent attention has been given to terrestrial and marine manglicolous fungi because
of mangrove tree mortalities. However, this mycobiota in many Caribbean Islands is
practically unknown. The halotolerant fungus Asteridiella sepulta (Ascomycota,
Meliolaceae), which is one of many species that form sooty mould, has been isolated from
leaves surfaces of Avicennia germinans in southwestern Puerto Rico. In this study of A.
germinans, I found the planthopper Petrusa marginata (Homoptera, Flatidae) excretes a
sugary honeydew upon which the dematiaceous mycelium of A. sepulta grows. Although A.
sepulta has been previously collected on A. germinans, the association of the fungus and the
planthopper in black mangrove had not been noted. Asteridiella sepulta produces a flat
colony with a spongy subiculum on surfaces of leaves, twigs, and small branches of A.
germinans.
RESUMEN
Se le ha dado una reciente atención a los hongos manglícolas terrestres y marinos a
causa de las mortalidades de árboles de mangle. Sin embargo, en muchas islas Caribeñas
esta micobiota es prácticamente desconocida. El hongo halotolerante Asteridiella sepulta
(Ascomycota, Meliolaceae), considerado como una de las especies que forman la fumagina,
ha sido aislado de la superficie de las hojas de Avicennia germinans en el suroeste de Puerto
Rico. En este estudio de A. germinans, encontré que el saltahojas Petrusa marginata
(Homoptera, Flatidae) excreta una secreción azucarada sobre la cual crece el micelio
dematiáceo de A. sepulta. Aunque A. sepulta ha sido previamente colectado en A.
germinans, la asociación del hongo con los saltahojas en el mangle negro es poco conocida.
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Asteridiella sepulta produce una colonia plana con un subículo esponjoso sobre la superficie
de las hojas, vástagos, y pequeñas ramas de A. germinans.
INTRODUCTION
A mangrove forest is a dynamic ecotone, or a transition zone, between the terrestrial
and marine habitats. In its simplest sense, a mangrove is used as a generic term referring to a
group of woody, halophylic plant formations that grow along sheltered tropical and
subtropical coastlines (Tomlison, 1986). The mangroves are derived from a variety of plant
taxa and they vary in their dependence upon littoral habitats (Lugo and Snedaker, 1974).
The black mangrove, Avicennia germinans (Avicenniaceae), is a small tree or shrub
(about 3 to 12-m high) that grows in lagoons and coastal swamps in paleotropics and
neotropics. It has been recorded in continental tropical America, Bermuda, The Bahamas,
United States of America (From Florida to Texas), throughout the West Indies (except
Dominica), including Trinidad, Tobago, and Curaçao (Martorell, 1976; Little et al., 2001). In
Puerto Rico, A. germinans has been reported from the main island, Vieques, and Culebra
(Martorell, 1976; Little et al., 2001). Avicennia germinans is very widely distributed along
tropical and subtropical protected silty seashores and forming mangals in brackish water at
mouths of rivers, usually with other mangrove species, but rarely forming monotypic stands
(Jiménez and Lugo, 1985a; Tomlinson, 1986; Little et al., 2001). Except for A. germinans,
the other mangrove species that occur in Puerto Rico are the red mangrove (Rhizophora
mangle) (Rhizophoraceae), the white mangrove (Laguncularia racemosa), and the
buttonwood (Conocarpus erectus) (Combretaceae) (Little et al., 2001).
Recent attention has been given to mangrove tree mortalities, which are caused by
anthropogenic misuse and unfavorable environmental conditions (Jiménez and Lugo, 1985b;
Anderson and Lee, 1995) as well as by fungal diseases (Jiménez and Lugo, 1985a; Wier et
al., 2000), among other biotic and abiotic factors.
Although terrestrial and marine
manglicolous fungi have been extensively studied in various parts of the world (Johnson and
Sparrow, 1961; Kohlmeyer and Kohlmeyer, 1979; Rollet, 1981; Hyde and Lee, 1995), in
190
many Caribbean Islands these fungi are poorly known.
Previous mangrove fungal collections in Puerto Rico are summarized in Stevenson
(1975), Acevedo (1987, 2001), Nieves-Rivera et al. (1998), Calzada (1999), Tattar et al.
(1994), Wier et al. (1996, 2000), Minter et al. (2001), and Tattar and Wier (2002). Stevenson
(1975) gave a summary of manglicolous fungi of Puerto Rico and the U.S. Virgin Islands.
Acevedo (1987) reported 18 species of manglicolous fungi (ascomycetes and mitosporic
fungi) from R. mangle in La Parguera. Nieves-Rivera et al. (1998) reported Schizophyllum
commune and Hypoxylon Sect. Hypoxylon in Avicennia nitida (= A. germinans) and
Rhizophora mangle, respectively, from Boquerón Wildlife Refuge. Calzada (1999) studied
three phytopathogenic fungi (Pestalotiopsis disseminata), Phoma eupyrena, Pterosporidium
rhizophorae) causing foliar diseases in R. mangle of La Parguera (Phosphorescent Bay and
La Parguera Channels). Acevedo (2001) assayed marine fungi (e.g., Didymosphaeria
rhizophorae, Hydronectria tethys, Hypoxylon oceanicum, Lulworthia grandispora,
Pestalotia sp., Xylaria spp., as well as other undetermined species of ascomycetes and
mitosporic fungi) by HPLC for biotransformation of phenanthrene in algal and mangrove (R.
mangle) substrates; she found that marine fungi and notably algal endophytes (e.g., Xylaria
spp.) are potentially useful organisms for bioremediation in marine environments. Minter et
al. (2001) gave a summary of manglicolous fungi of the Caribbean, including Puerto Rico.
Tattar et al. (1994), Wier et al. (1996, 2000), and Tattar and Wier (2002) documented the
incidence of Cytospora rhizophorae as a plant pathogen in R. mangle in the southwestern
coast of Puerto Rico.
Insects also have been found associated with mangrove plants. According to
Martorell (1976), two species of insects collected on A. germinans in Puerto Rico are the
cricket Hygronemobius alleni (Orthoptera, Grillidae) and the tree termite Nasutitermes
costalis (Isoptera, Termitidae). Tattar and Wier (2002) reported the termites N. costalis,
Neotermes mona, Incisertermes nr. incisus and Procryptotermes corniceps from R. mangle in
southwestern Puerto Rico.
Twenty-two species of ants (including Azteca sp.
(Dolichoderinae), Camponotus spp. Myrmelachista spp. (Formicidae), Crematogaster spp.,
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Hylomyrma sp., Pheidole spp., Solenopsis spp. (Myrmicinae), Pachycondyla villosa
(Ponerinae), Pseudomyrmex gracilis, Pseudomyrmex sp. (Pseudomyrmecinae)) have been
recorded from the Brazilian mangrove plants Avicennia schaueriana, L. racemosa, and R.
mangle (Cortes-Lopes and Dos Santos, 1996). However, the recent reports of the gall
midges Actilasioptera spp. (Gagné and Law, 1998) and Meunieriella avicenniae (Diptera,
Cecidomyiidae) (Gagné and Etienne, 1998), the causative agents of the leaf gall in black
mangroves, show that mangrove insect fauna is rather poorly known. Therefore, there has
been no assessment on the effects insects have on these plants.
On February 5, 2001, while collecting manglicolous fungi, I observed a coating on
the leaves of the black mangrove (A. germinans), with a black soot caused by mycelia giving
the false impression of pollution caused by passing vehicles. After a careful examination
under the dissecting microscope, the soot was found to be a mycelium produced by a fungus.
This fungus was Asteridiella sepulta (Ascomycota, Meliolaceae). The type collection for A.
sepulta is contained in voucher 6416, collected from A. nitida (= A. germinans) from Cataño,
Puerto Rico, by A. A. Heller (Stevenson, 1975). The purpose of the present study is to
record details of the sooty mould-planthopper association on leaves of the black mangrove in
southwestern Puerto Rico.
MATERIALS AND METHODS
Locality
Leaves of A. germinans were collected at Los Morrillos (LM), (coordinates: 17°
57.215’ N, 67° 11.867’ W), which is part of the Boquerón Commonwealth Forest, Road 301,
km 11.4, Barrio El Corozo, Cabo Rojo, Puerto Rico (Figures 2, 41, 42A). This forest is
approximately 0 to 30 m above sea level and the general environment of the region is
classified as a subtropical dry forest (Ewel and Whitmore, 1973). The rainy season in Puerto
Rico ranges from May to June and August to September, with two rainfall peaks. Annual
average rainfalls in the weather station located at Lajas Agricultural Experiment Station of
the University of Puerto Rico-Mayagüez, are 1016 to 1270 mm (Ravalo et al., 1986).
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According to ‘CNN.com/weather’, the annual average climatic conditions for
Boquerón, Cabo Rojo, Puerto Rico from 2000 to 2002) were: air temperature, 28.0°C;
relative humidity, 74.0%; wind, variable (mostly from ENE or SSE at < 5 to 30 km/hr;
sunrise, 06:05 hr, sunset, 18:49 hr. In ‘Atmos Carib— Caribbean Atmospheric Research
Center’ at University of Puerto Rico-Mayagüez Campus, the annual average climatic
conditions for Magueyes Island Marine Laboratories (MIML) in La Parguera, Lajas, Puerto
Rico from 2000 to 2001 were: air temperature, 26.6°C; atmospheric pressure, 1013.9 mb.
Glynn (1973) conducted early meteorological observations in MIML. Therefore MIML was
selected because it has the same xeric conditions and its proximity to the study site. The
geologic formations of the study site are: Holocene’s mangrove swamps (Qm) and beach
deposits (Qb) (Volckmann, 1984; Torres-Figueroa, 1993). Soils of LM are classified as tidal
flats (Tf), tidal swamps (Ts), coastal beaches (Co), and limestone rock lands (Lr) (USDA,
1993).
Isolation of Sooty Mold
The author collected individual leaves of A. germinans August 3, 2001. Most of the
leaves of A. germinans were collected close to roads and trails. These were photographed
and the identity of the sooty mould was confirmed with Stevens (1916, 1917, 1927), Ciferri
(1954), and Hughes (1976). Leaves were placed in wet chambers in Petri-dishes, following
Calzada (1999). Distilled water was used in the wet chambers. The paraffin-sealed wet
chambers were placed in light/dark at room temperature. Microscopic observations were
made with a light microscope (Nikon Labophoto-2 Microscope). Drawings were made with
a camera lucida. All voucher specimens are placed at the Center for Forest Mycology
Research, in the process of curation before being deposited in the Herbarium of the
Department of Natural Sciences, University of Puerto Rico at Río Piedras (UPRRP).
Insect Collection
The insect specimens collected in Avicennia germinans were preserved in 70%
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ethanol and identified by various specialists (Drs. Arístides Armstrong, Ángel L. González,
and Silverio Medina-Gaud of the Department of Crop Protection, University of Puerto Rico,
Mayagüez).
Planthopper identification was provided by Dr. Stuart H. McKamey
(Smithsonian Institution in Washington, D.C.).
RESULTS
Asteridiella sepulta was found on the phylloplane (surface of living leaves, sensu
Hughes, 1976) (Figure 42B) producing a flat, spongy subiculum colony of sooty mould on
the front and back of the leaves, twigs, and small branches of A. germinans (Figures 42C-F).
This sooty mould was found throughout the year on the leaves of A. germinans. It must be a
halotolerant fungus to be able to grow on black mangrove leaves, which are often covered
with salt crystals. Asteridiella sepulta also grew in the margins of the leaves and the stems,
apparently following the path of running water after a rain, moist, dew, or by condensation
(Figure 42D).
Diagnostic
Colonies circular, 1-7 mm in diameter, amphigenous, dense, easily secedent, well
defined, sub-epiphyllous spots, single or confluent, black. Hyphae brown, sinous to tortuous,
branching alternate or irregular, not opposite, at acute angles, forming a mat, densely
reticulate and becoming almost solid (Figure 43A). Capitate hyphopodia alternate, more or
less antrorse, usually straight, 23.0-24.3 µm long; stalk cell cylindric, 0.9-10.4 µm long; head
cell globose to widely piriform, entire or rarely slightly rounded-angulose. Mucronate
hyphopodia few, mixed with capitate, opposite or alternate, ampulliform with short neck.
Setae none. Perithecia in loose central group, black, globose, rough, 165 µm in diameter;
most surface cells are prolonged into translucent dark brown, obtusely conoid outgrowths,
not striate (Figure 43A). Asci not seen. Spores dark brown, cylindric, obtuse, 4 septate,
constricted, ends obtuse, smooth, thin-walled, 51.1-52.6 x 18.2-19.8 µm (Figure 43B).
194
Material Studied
Los Morrillos, Boquerón Commonwealth Forest, in coastal forest next to Parador Las
Salinas, on living leaves of A. germinans, 1.5 m alt., 3 August 2001, Á. M. Nieves-Rivera,
PR-935, 936, 937 (UPRRP); Los Morrillos, Boquerón Commonwealth Forest, in the shores
of a hypersaline lagoon, next to old bridge, on living leaves of A. germinans, 0.5 m alt., 3
August 2001, Á. M. Nieves-Rivera, PR-938 (UPRRP); Guánica Dry Forest, in coastal forest
next to Parador Copa Marina, on living leaves of A. germinans, 1.5 m alt., 7 August 2001, Á.
M. Nieves-Rivera, PR-939, 940 (UPRRP).
Distribution
Sooty mould mycelia develops in the front and back of the leaves, petioles, twigs, and
branches (Reynolds, 1976) (Figures 44A-B, 45A). The range of the area covered by sooty
moulds extends about 2.0 x 1.5 km in A. germinans of LM. However, the fungus
dissemination seems to depend on the planthopper Petrusa marginata (Homoptera, Flatidae)
and A. germinans distribution along the coastline. Previous collections of A. sepulta in
Puerto Rico were reported by Stevens (1916, 1917, 1927) as Irenina (Meliola) sepulta,
Chardón (1920) as Meliola sepulta, Toro (1925) as Irene sepulta, in the Dominican Republic
by Ciferri (1954) as Meliola (Irenina) sepulta, and Trinidad by Dennis (1970) as A. sepulta.
Stevenson (1975) summarized A. sepulta distribution to be found in A. germinans forests of
Puerto Rico, the Dominican Republic, and Sierra Leone (Africa).
Insects
The planthopper P. marginata has been previously reported in Puerto Rico by Osborn
(1935), Caldwell and Martorell (1950), and Maldonado-Capriles and Medina-Gaud (1985);
however, A. germinans has not been recorded as host plant for this insect in Puerto Rico.
Many planthopper exuviae, exocuticles, and immature (nymphs) were also detected on the
leaves of A. germinans. Petrusa marginata has become a pest in plantations of coffee Coffea
arabica, coco-plum Chrysobalanus icaco, jasmine Jasminum sp. (Maldonado-Capriles and
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Medina-Gaud, 1985), and the sea grape Coccoloba uvifera (Figures 45B).
Petrusa
marginata (Oremis (Petrusina) marginata of Osborn, 1935) has been collected on Lantana
sp. and Cordia sp. at Ensenada, Aguirre, and other points throughout the island (Osborn,
1935). Caldwell and Martorell (1950) reported P. marginata from Monserrat, B. W. I., to
Mona Island, Puerto Rico. Petrusa marginata is very common along coastal areas and also
present in suitable habitat up to 762 m especially along the south coast of Puerto Rico
(Caldwell and Martorell, loc. cit.). During our study, few other insects were collected by the
senior author on A. germinans, including the common bee Apis mellifera (Hymenoptera,
Apidae), the green lacewing Chrysopa sp. (Neuroptera, Chrysopidae), the leaf gall midge M.
avicenniae, the ants Solenopsis spp. (Hymenoptera, Formicidae), the cricket H. alleni, the
tree termite N. costalis, and three spiders of the Group Aranae.
DISCUSSION
Sooty moulds are usually associated with the liquid excrement of sucking insects,
known as ‘honeydew’ is a common occurrence on many trees (Auclair, 1963). Undigested
sucrose in honeydew makes an excellent growth medium for dark-spored fungi (Tattar,
1989). However, sooty moulds also have been found on plants not infested with insects
which produce honeydew, living and dead vegetation, on the surface of rocks, and the forest
floor (Hughes, 1976).
In Puerto Rico sooty moulds caused by Capnodium spp., Trimmatostroma sp.
(Figures 44A-B), and other undetermined fungal species are found in honeydew excretions of
aphids and scale droppings on tea (Camellia sinensis), sour orange (Citrus aurantium),
orange (C. sinensis), coffee (C. arabica), mango (Mangifera indica), sea grape (Coccoloba
uvifera), white mangrove (L. racemosa) or ornamental plants (Anthurium sp., C. icaco,
Gardenia sp., Ixora sp., Jasminum spp.) (Figure 45A). The mycelial mats of these fungi are
easily removed by peeling off the surface from the leaf where they are found, usually
revealing a clean, intact plant surface (Reynolds, 1976). Maldonado-Capriles and MedinaGaud (1985) refer to sooty mould in Spanish as “hongo de hollín”.
196
The presence of saprophytic sooty mould A. sepulta does not initially infect the
leaves of A. germinans, but covers the leaf surfaces only after the honeydew of P. marginata.
Planthoppers, like aphids, feed on the leaves of both deciduous hardwoods and evergreens
(Tattar, 1989). They excrete excess sucrose in a honeydew excrement (Auclair, 1963). We
suspect this fungus-leaf covering does not adversely affect photosynthesis because of the
healthiness in leaves examined (leaves were green, robust, and intact), and its similarity to
cases of myxomycete-grass associations (Nieves-Rivera, 2000).
However, a heavy
accumulation of sooty mould can prevent photosynthesis (Tattar, 1989).
The covering of the leaves by the fungus looked like “soot,” giving the false
impression of pollution caused by passing vehicles. The “soot” was found to be fungal
mycelia which had expanded over the foliar surface to cover, in some cases, 25 to 98% of the
leaf, similar to the percentages reported in pecan leaves by Tedders and Smith (1976), and
Wood et al. (1988). Coating by A. sepulta was not detected on other mangroves species,
such as R. mangle and C. erectus. In his study of the genus Meliola in Puerto Rico, Stevens
(1917) reported Meliola lagunculariae and M. nigra on the white mangrove L. racemosa in
Puerto Rico. However, other black mangrove populations in different locations around
Puerto Rico have been observed with sooty moulds, for example, the mangals located in
Magueyes Island, La Parguera mangrove channels, Bahía de Jobos estuary in Salinas, Las
Cabezas de San Juan Natural Reserve in Fajardo, Caño Corazones in Mayagüez, Guayanilla
and Ponce coasts (Nieves-Rivera, unpubl. data).
Sooty moulds, like myxomycetes (Nieves-Rivera, 2000) are saprobes and their
fruiting bodies may cover portions of the plant, but apparently do not infect them. The
plasmodium (in the case of myxomycetes) does not affect the leaf by reducing its
photosynthesis or respiration as true fungi do (for example, powdery mildews (Mignucci and
Boyer, 1979)) (Nieves-Rivera, 2000). Although sooty moulds are not considered of
economic importance, Maldonado-Capriles and Medina-Gaud (1985) recommended the use
of Diazinon AG-500, Cygon 2.67 or Endosulfan 50 PH to control P. marginata, thus
controlling the sooty mould. However, the use of commercial pesticides and insecticides
197
might best be avoided because of the resultant pollution, the use of entomopathogenic fungi
for planthopper and leafhopper biocontrol (Soper, 1985) might be more prudent.
In conclusion, sooty mould-planthopper occurrence on black mangrove leaves is
another example of fungus/insect interaction, that does not appear to be detriment. However,
if a black mangrove forest were to be stressed from changes in climate, attack by borers in
high incidence or other negative anthropogenic impact, a heavy incidence of sooty mould
could exacerbate the stress and lead to decline. Therefore, the continued study of sooty
mould on A. germinans is merited.
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201
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Figure 41. A. Map of the southwestern end of Puerto Rico, showing Cabo Rojo municipality
collection areas (*).
203
Figures 42A-F. A. View of Los Morrillos, Boquerón Commonwealth Forest, Cabo Rojo,
Puerto Rico. B. Growth in situ of sooty mould (Asteridiella sepulta) on an upper
surface of leaf of the black mangrove Avicennia germinans. C-D. Leaves of black
mangrove Avicennia germinans, with a clean surface (left) and Asteridiella sepulta
infested surface (right). E-F. Growth of Asteridiella sepulta on the lower surface of
the front of the leaf of A. germinans, associated with infestation of the planthopper
Petrusa marginata. Photos taken at Los Morrillos, Boquerón Commonwealth Forest.
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Figures 43A-B. Asteridiella sepulta. A. Hyphae with perithecium and young ascostroma
bearing hyphal appendages, on leaves of the black mangrove Avicennia germinans
(Cabo Rojo, Puerto Rico). B. Spores.
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Figures 44A-B. Example of the sooty mold, Trimmatostroma sp. A. Perithecium or ascoma with asci and
ascospores inside. B. Dematiaceous hyphal subiculum (sooty mold) on twig of the sapodilla (níspero)
Manilkara zapota (Mayagüez, Puerto Rico).
Figure 45A-B. Sea grape Coccoloba uvifera, showing sooty mold (A) and with an infestation of Petrusa
marginata on lower surface (B). A. Dematiaceous hyphae (Dh) and young ascostroma of Asteridiella
sepulta and Trimmatostroma sp., on leaves of the black mangrove Avicennia germinans surface.
Photograph taken at Playa Jobos, Isabela, Puerto Rico.
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CHAPTER 7
MANGLICOLOUS BASIDIOMYCETES OF SOUTHWESTERN PUERTO RICO
AND SOUTHWESTERN FLORIDA (U.S.A.)
ABSTRACT
Field surveys of the Boquerón Commonwealth Forest, Boquerón Wildlife Refuge,
and Magueyes Island in southwestern Puerto Rico, and the Aquatic Preserves of
southwestern Florida from July 2001 throughout 2003, yielded 59 specimens of manglicolous
basidiomycetes that were catalogued and taxonomically examined. All the specimens were
identified to taxa by observation of morphological and microscopic characters, and compared
with specimens from the U.S. National Fungus Collections at Maryland (BPI), University of
Puerto Rico at Río Piedras (UPRRP), The New York Botanical Garden at Bronx (NY), and
University of Oslo at Blindern (O) herbaria. They represented 8 families, 12 genera, and 14
taxa. Basidiomycetes grew on dead bark and wood of Rhizophora mangle, Avicennia
germinans, and Laguncularia racemosa. Coriolopsis floccosa, Phellinus merrillii, and
Tyromyces cf. chioneus are new records for Puerto Rican mangroves and Phlebia sp. is a new
record for Florida mangroves.
RESUMEN
Los reconocimientos de campo en el Bosque Estatal de Boquerón, el Refugio de Vida
Silvestre de Boquerón e Isla Magueyes en el sudoeste de Puerto Rico, y en las Reservas
Acuáticas del sudoeste de la Florida durante julio de 2001 y a través de 2003, produjeron 59
especímenes de basidiomicetos manglícolas que fueron catalogados y examinados
taxonómicamente. Todos los especímenes fueron identificados a especies mediante la
observación de los caracteres morfológicos y microscópicos, junto con la comparación de
especímenes de los herbarios de las Colecciones Nacionales Fúngica de los EEUU en
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Maryland (BPI), Universidad de Puerto Rico en Río Piedras (UPRRP), el Jardín Botánico de
Nueva York en el Bronx (NY), y la Universidad de Oslo en Blindern (O).
Estos
representaron 8 familias, 12 géneros y 14 taxones. Los basidiomicetos crecieron en corteza
muerta y madera de Rhizophora mangle, Avicennia germinans y Laguncularia racemosa.
Coriolopsis floccosa, Phellinus merrillii y Tyromyces cf. chioneus son registros nuevos para
los mangles de Puerto Rico y Phlebia sp. fue un registro nuevo para los mangles de la
Florida.
INTRODUCTION
Until recently, there have been few mycological studies on the manglicolous
basidiomycetes of the Caribbean region, although information on their occurrence would be
of considerable value for biologists and conservationists. Previous reports summarized in
Kohlmeyer (1969) recorded eight manglicolous basidiomycetes (Fomes avicenniae,
Phellinus gilvus, Psathyrella sp., Schizophyllum commune, Trametes rhizophorae,
Tulasnella bifrons, T. pacifica, and T. violacea from Avicennia sp., Rhizophora mangle or
Hibiscus tiliaceus (Reichardt, 1870; Baccarini, 1916; Olive, 1957; Cooke 1961; Kohlmeyer,
1969). Kohlmeyer (1969) also collected two basidiomycetes, F. avicenniae and P. gilvus
from the Heeia Swamp, Oahu, Hawaii, on R. mangle. Lee and Baker (1973) reported three
basidiomycetes from R. mangle roots (Fomes sp., Polyporus cinnabarinus, and Psathyrella
sp.).
The neotropical occurrence of manglicolous basidiomycetes were reported from
Brazil (Sotão et al., 1991; Almeida Filho et al., 1993; Campos and Cavalcanti, 2000; Sotão et
al., 2002), Lesser Antilles (Pegler, 1983a; Minter et al., 2001), Panama (Gilbert and Sousa,
2002), Puerto Rico (Stevenson, 1975; Lodge, 1996a; Nieves-Rivera et al., 1998; Minter et
al., 2001) and Venezuela (Dennis, 1970). Minter et al. (2001) survey of Caribbean fungi
recorded 11 taxa of manglicolous basidiomycetes for Puerto Rico (e.g., Gloeophyllum
striatum, Pleurotus djamor, Pleurotus sp., Polyporus fulvocinereus, and Tyromyces sp. on R.
mangle; Phellinus sp. and Trametes villosa on Conocarpus erectus; Coriolopsis sp., Inonotus
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porrectus, and Phellinus sp. on Laguncularia racemosa; and S. commune on Laguncularia
sp.). More recently, the manglicolous fungi surveys by Poonyth et al. (2000) and Schmit and
Shearer (2003) summarized what is known about worldwide fungal species and their
mangrove hosts, and recorded about 30 species of manglicolous basidiomycetes (e.g.
Crepidotus krieglsteiner, P. gilvus, Psathyrella rhizophorae, Pycnoporus cinnabarius, S.
commune on R. mangle, and Dacrymyces intermedius on H. tiliaceus). We note here the
occurrence of 14 basidiomycetes from mangrove coastal forests of southwestern Puerto Rico
and 4 basidiomycetes for southwestern Florida (U.S.A.); this marks the second formal
documentation of manglicolous basidiomycetes for Puerto Rico, and the sixth neotropical
occurrence of manglicolous basidiomycetes.
MATERIALS AND METHOD
The Puerto Rican study areas were located at the Boquerón Commonwealth Forest
(BCF, 18°01’N, 67°10’W, secondary road PR-307, next the town of Boquerón, Barrio
Boquerón, Cabo Rojo), Boquerón Wildlife Refuge (BWR, 18°01’N, 67°09’W, secondary
road PR-301, near to the town of Boquerón, Cabo Rojo), and Magueyes Island Marine
Laboratory (MIML, 17°58’N, 67°02’W, secondary road PR-304, close to the town of La
Parguera, Barrio La Parguera, Lajas), all at mean sea level, southwestern Puerto Rico
(Figures 2 and 3). The general environment of the region is classified as a subtropical dry
forest (Ewel and Whitmore, 1973). Further details on climatology, geology, and edaphic
formations of BCF, BWR, and MIML are discussed in Glynn (1973), Vázquez (1983), Toro
and Colón (1986), Winter et al. (1998), and Nieves-Rivera et al. (2002). Rhizophora mangle
occasionally forms thick coastal woodland in BCF and in the rest of the southwestern coast
of Puerto Rico (Cintrón et al., 1978; Lugo, 1989; Vázquez and Kolterman, 1998).
The Aquatic Preserves of Southwest Florida (APSF) are located SE of Venice, at the
Gulf of Mexico (Figures 46A-B). The APSF have over 2,574 km of coastline, a network of
barrier islands and mangals providing more than 907 km2 of bays, lagoons, and other water
bodies sheltered from the open gulf. Southwest Florida contains the highest density of
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aquatic preserves in the state. Most of the coastal waters here are contained within aquatic
preserves. The eight aquatic preserves in this region are: (1) Lemon Bay; the Charlotte
Harbor Aquatic Preserves, which include (2) Cape Haze, (3) Charlotte Harbor/Gasparilla
Sound, (4) Matlacha Pass and (5) Pine Island Sound; (6) Estero Bay; (7) Rookey Bay; and (8)
Cape Romano/Ten Thousand Islands. The collection sites at APSF were: Punta Gorda
(26°47’N, 82°04’W), Cape Coral (26°45’N, 82°04’W), and Cape Haze (26°54’N, 82°10’W).
The Punta Gorda site was at sea level and was flooded with each tide. The Cape Coral and
Cape Haze sites were on berms approximately 0.5 m above sea level. The general
environment of the region is classified as a subtropical dry forest. APSF normally receive
1,376 mm on average of precipitation each year. The driest months are November,
December and April with precipitation is less than 50 mm. Wildfires are common in late
spring, since April is dry and hot, and sometimes-rainy season does not begin in June. In
these cases, the fire season can also extend into June (unpublished data). Further details on
climatology, geology, and edaphic formations of APSF are discussed in Davis (1940),
Craighead (1971), and Kangas and Lugo (1990). Avicennia germinans forms thick coastal
woodland in APSF and in the rest of the southwestern coast of Florida (Davis, 1940;
Craighead, 1971; Kangas and Lugo, 1990).
As a result of three field surveys to the BCF, BWR, and MIML in southwestern
Puerto Rico and the Aquatic Preserves of southwest Florida from July 2001 through 2003, 59
specimens of manglicolous basidiomycetes were catalogued and taxonomically examined,
using the works of Cooke (1961), Dennis (1970), Gilbertson and Ryvarden (1986, 1987),
Pegler (1983a, b), and Larsen and Cobb-Poulle (1990). Each taxon newly recorded for
Puerto Rico and Florida is marked with an asterisk at the head of the specific name. To study
the material, microscopical observations were made from slides mounted in 5.0% KOH,
Meltzer reagent or lactophenol, following Largent et al. (1977). All the specimens were
identified by observation of morphological and microscopic characters, along with
comparison with specimens from the U.S. National Fungus Collections at Maryland (BPI),
University of Puerto Rico at Río Piedras (UPRRP), The New York Botanical Garden at
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Bronx (NY), and University of Oslo at Blindern (O) herbaria.
RESULTS
Manglicolous basidiomycetes represented 8 families, 12 genera and 14 taxa.
Basidiomycetes occurred on host trees R. mangle (11 species), A. germinans (2 species) and
L. racemosa (2 species). Coriolopsis floccosa, Phellinus merrillii, and Tyromyces cf.
chioneus were new records for Puerto Rican mangroves, and Phlebia sp. was a new record
for Florida mangroves.
Crepidotaceae
Crepidotus uber (Berk. & M.A. Curtis) Sacc., Syll. Fung. 5: 878. 1887.
(Figures 47A-E)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Commonwealth Forest, next to
Puerto Real, boardwalk that passes through a mangrove forest and reaches the lagoon on the
Boquerón-Guaniquilla mangrove forest, on Rhizophora mangle on bark and dead wood, also
on rotting aerial roots, and upright tree trunks, position aboveground 0.3 to 1.5 m, 14-IV2003, Á. M. Nieves-Rivera (BPI 843767).
Remarks: Crepidotus uber (BPI 843767), the main character that distinguishes this form
from the type species Crepidotus mollis is the lack of encrusting-pigmented hyphae on the
cutis, an epicutis undifferentiated from the underlying context, and a gelatinized layer
reaching the surface (Pegler, 1983a). In general, C. uber appears to be widespread in the
tropics and subtropics and sequencing data suggest that it may be a complex of species;
microscopically it is indistinguishable from C. uber from Madagascar (Mary Catherine Aime,
unpublished data). Crepidotus uber has been collected from Bonin Island, Dominica, and
Guadeloupe in the Lesser Antilles (Pegler, 1983a).
Coprinaceae
Psathyrella sp.
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(Figures 48A-B)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Wildlife Refuge, on R. mangle leaf
litter, position aboveground 0.1 m, 13-III-1995, Á. M. Nieves-Rivera (UPRRP PR-641.1).
Remarks: This taxon grew on decayed plant material (leaf litter of R. mangle and Thespesia
populnea, and blades of Thalassia testudinum)
Lentinaceae
Lentinus crinitus (L.: Fr.) Fr., Syst. Orb. Veg. 77. 1825. [= Lentinus swartzii Berk. (Lodge,
1996)]
(Figure 48C)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Wildlife Refuge, on R. mangle
fallen trunk, position aboveground > 1 m, 15-III-2002, Á. M. Nieves-Rivera (BPI 863546).
Remarks: This extremely common species has a basidioma which is extremely variable and
difficulty is experienced in delimiting the species (Pegler, 1983a, b). Lentinus crinitus is
found single or gregarious on moist dead hardwoods logs or stumps.
Coriolaceae
Coriolopsis badius (Cooke) Murrill, Bull. Torrey Bot. Club 34: 466. 1907.
Material studied: UNITED STATES. Florida: Aquatic Preserves of Southwest Florida, Cape
Coral, on Avicennia germinans dead wood, position aboveground > 1 m, 5-I-2003, T. A.
Tattar (BPI 843741); Cape Haze, on A. germinans dead wood, position aboveground > 1 m,
20-X-2002, T. A. Tattar (BPI 843742).
Remarks: This species is common in the Caribbean. Coriolopsis badius had a position
aboveground from 25 cm to 2 m and also grew on Conocarpus erectus dead wood.
*Coriolopsis floccosa (Jungh.) Ryv., Genera Polyp. Syn. Fung. 5: 316. 1991.
(Figure 48D)
Material studied: PUERTO RICO. Lajas: Magueyes Island, on R. mangle bark, position
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aboveground < 2 m, 14-IV-2003, Á. M. Nieves-Rivera (O PR-979).
Remarks: Spores of this species were non-amyloid in Melzer’s reagent. Coriolopsis
floccosa cause white rot in R. mangle.
Gloeophyllum striatum (Sw.: Fr.) Murrill, Bull. Torrey Bot. Club 32: 370. 1905.
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Commonwealth Forest, next to
Puerto Real, boardwalk that passes through a mangrove forest and reaches the lagoon on the
Boquerón-Guaniquilla mangrove forest, on R. mangle bark, position aboveground 0.5 to 2.0
m, 19-IV-2003, Á. M. Nieves-Rivera (BPI 843744).
Remarks: Gloeophyllum striatum has also been found on R. mangle decay wood in the
Boquerón Wildlife Refuge in Cabo Rojo.
Hexagonia hydnoides (Fr.: Sw.) M. Fidalgo, Mem. N. Y. Bot. Gdn. 17: 69. 1968.
(Figure 48E)
Material studied: UNITED STATES. Florida: Aquatic Preserves of Southwest Florida, Punta
Gorda, on Laguncularia racemosa dead wood, position aboveground 50 cm, 20-X-2002, T.
A. Tattar (BPI 843745); position aboveground 1 m, 20-X-2002, T. A. Tattar (BPI 843746).
Remarks: Specimen (BPI 843746) is young. This species is common in the Caribbean and is
found especially in xeric localities. It has been found growing on R. mangle dead wood in
Boquerón Wildlife Refuge in Cabo Rojo.
Pycnoporus sanguineus (L.: Fr.) Murrill, Bull. Torrey Bot. Club 31: 421. 1904.
(Figure 48F)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Commonwealth Forest, next to
Puerto Real, boardwalk that passes through a mangrove forest and reaches the lagoon on the
Boquerón-Guaniquilla mangrove forest, on R. mangle dead wood, position aboveground 50
cm, 19-IV-2003, Á. M. Nieves-Rivera (UPRRP PR-984.2).
Remarks: Pycnoporus sanguineus is the one of causative agents of the white rot of dead
213
hardwoods. This species has hyphal contents in some areas of tramal tissue strongly
dextrinoid in Melzer’s reagent. Pycnoporus sanguineus is widely distributed throughout the
subtropical and tropical regions of the world.
*Tyromyces cf. chioneus (Fr.: Fr.) Karst., Rev. Mycol. 3: 17. 1881.
(Figure 48G)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Commonwealth Forest, next to
Puerto Real, boardwalk that passes through a mangrove forest and reaches the lagoon on the
Boquerón-Guaniquilla mangrove forest, on R. mangle bark, position aboveground 60 cm, 19IV-2003, Á. M. Nieves-Rivera (UPRRP PR-984.3).
Remarks: This cosmopolitan species is found single or gregarious on dead hardwoods and
occasionally on conifers. Tyromyces sp. has been previously collected from Rhizophora sp.
(Sotão et al., 2002).
Hymenochaetaceae
Phellinus cf. gilvus (Schw.) Pat., Essai Taxon. Hyménon., p. 97. 1900.
Material studied: UNITED STATES. Florida: Aquatic Preserves of Southwest Florida, Punta
Gorda, on L. racemosa dead wood, position aboveground 50 m, 20-X-2002, T. A. Tattar
(BPI 843747).
Remarks: Basidioma badly eaten by insects and shows almost no pores left. This species
also has been collected on A. germinans.
*Phellinus merrillii (Murrill) Ryv., Norw. J. Bot. 19: 234. 1972.
Material studied: PUERTO RICO. Lajas: Magueyes Island, on R. mangle bark and dead
wood, position aboveground 1.2 to 3.0 m, 8-V-2003, Á. M. Nieves-Rivera (O PR-985).
Remarks: Phellinus merrillii was originally described from the Philippine Islands. The
globose, pigmented spores, lustrous context, and lack of setae characterized P. merrillii.
214
Meruliaceae
*Phlebia sp.
(Figures 18A-B)
Material studied: UNITED STATES. Florida: Aquatic Preserves of Southwest Florida, Cape
Haze, on R. mangle bark on dead aerial roots, position aboveground 25 cm, April 2003, T. A.
Tattar (O FL-9).
Remarks: This species has a very striking color (bright orange towards center with white
margins) and very large spores; it was sent to Dr. Kurt Hjortstam for further study. A similar
specimen (UPRRP PR-982) also growing on R. mangle dead bark and decay wood was
collected from Boquerón Commonwealth Forest in Cabo Rojo. Phlebia acanthocystis was
reported from mangroves Bruguiera gymnorrhiza and Rhizophora mucronata in Japan
(Maekawa et al., 2003).
Schizophyllaceae
Schizophyllum commune Fr.: Fr., Syst. Mycol. 1: 330. 1831.
Material studied: PUERTO RICO. Lajas: Magueyes Island, on R. mangle bark, position
aboveground 0.5 to 3.0 m, 5-XI-1998, Á. M. Nieves-Rivera (BPI 843748).
Remarks: This species is commopolitan. Schizophyllum commune has been seen on a liana
in R. mangle canopy at 5.0 m in height in Magueyes Island. Causes wood rot and is
pathogenic to humans.
Dacrymycetaceae
Dacryopinax spathularia (Schw.) G. W. Martin, Lloydia 11: 116. 1948.
(Figure 48H)
Material studied: PUERTO RICO. Cabo Rojo: Boquerón Commonwealth Forest, on
Rhizophora mangle decay wood, sometimes in bark, position aboveground 0 to 1.2 m, 13-III1995, Á. M. Nieves-Rivera (BPI 843743).
Remarks: BWR specimens of D. spathularia were also found on R. mangle wood subjected
215
once or twice a year to brackish water, with an annual surface salinity range of 7 to 31 g/L
(unpublished data).
DISCUSSION
When compared with the assemblage of taxa reported from the Caribbean Islands, the
Florida records were lower in basidiomycete diversity (e.g., Coriolopsis badius, Hexagonia
hydnoides, Phellinus cf. gilvus, Phlebia sp.). In general, all basidiomycetes collected showed
a tendency of dominant species adapted to coastal surroundings and a few limited unreported
species demonstrate an interesting local distribution of basidiomycetes in both collection
sites. Although many of these basidiomycetes have been reported previously from Puerto
Rico (Stevenson, 1975; Lodge, 1996a; Nieves-Rivera et al., 1999; Lodge et al., 2000; Minter
et al., 2001), their presence on R. mangle, A. germinans, and L. racemosa suggests that
mangroves, although usually neglected as a potential substrate for basidiomycetes, may
support a larger assemblage of species than indicated by the previous published records.
The basidiomycetes of the forests in Puerto Rico, especially the Agaricales are poorly
known, and 15 to 25% is undescribed taxa (Lodge, 1996b). For instance, Lodge (1996b)
mentioned that 30 taxa of Entolomataceae found by Dr. Timothy J. Baroni (State University
of New York at Cortland) in Puerto Rico were not previously listed by Stevenson (1975).
Examples such as these shows that much remains to be learned and it will be a monumental
task to produce a reasonably complete mycobiota for Puerto Rico. Therefore, it would seem
important to continue studying such habitats, in order to contribute to the conservation and
knowledge of the biodiversity of basidiomycetes of Puerto Rico.
LITERATURE CITED
Almeida Filho, O. M., R. Bueno and V.L. R. Bononi. 1993. Algumas espécies de fungos
basidiomicetos dos manguezais do Estado de São Paulo. Hoehnea 20: 87-92.
Baccarini, P. 1916. Eumycetes. In Chiovenda, E. (ed.), Le collezioni botaniche della
missione Stefanini-Paoli nella Somalia italiana, pp. 1-241. Pubblicazioni del Regio
216
Istituto di Studi Superiori e di Perfezionamiento di Firenze, L’Erbario Tropicale di
Firenze, Rome, Italy.
Campos, E. L. and M. A. Q. Cavalcanti. 2000. Primeira ocorrência de Phellinus mangrovicus
(Imaz.) Imaz. para o Brasil. Acta Botanica Brasilica 14: 263-265.
Cintrón, G., C. Goenaga and J. González-Liboy. 1978. Ecología del manglar en una zona
árida: exposición al oleaje y estructura del manglar. In Departamento de Recursos
Naturales de Puerto Rico (ed.), Quinto Simposio de los Recursos Naturales, pp. 5786. Estación Experimental Agrícola de la Universidad de Puerto Rico, Río Piedras,
Puerto Rico.
Cooke, W. B. 1961. The genus Schizophyllum. Mycologia 53: 575-599.
Craighead, F. C. 1971. The trees of South Florida. Volume 1. University of Miami Press,
Coral Gables, 212 pp.
Davis, J. H., Jr. 1940. The ecology and geologic role of mangroves in Florida. Carnegie
Institute of Washington, D.C. Publication 517: 305-412.
Dennis, R. W. G. 1970. Fungus flora of Venezuela and adjacent countries. Kew Bulletin
Additional Series II. Her Majesty's Stationery Office, London, 531 pp.
Ewel, J. J. and J. L. Whitmore. 1973. The ecological life zones of Puerto Rico and the U. S.
Virgin Islands. Forest Service Research Paper ITF-18. U. S. Forest Service, Institute
of Tropical Forestry, Río Piedras, Puerto Rico. 72 pp.
Gilbert, G. S. and W. P. Sousa. 2002. Host specialization among wood-decay polypore fungi
in a Caribbean mangrove forest. Biotropica 34: 396-404.
Gilbertson, R. L. and L. Ryvarden. 1986. North American polypores. Volume 1. Fungiflora,
Oslo, pp. 1-433.
Gilbertson, R. L. and L. Ryvarden. 1987. North American polypores. Volume 2. Fungiflora,
Oslo, pp. 434-885.
Glynn, P. W. 1973. Ecology of a Caribbean coral reef. The Porites reef-flat biotope: Part I.
Meteorology and hydrology. Marine Biology 20: 297-318.
Kangas, P. C. and A. E. Lugo. 1990. The distribution of mangroves and saltmarshes in
217
Florida. Tropical Ecology 31: 32-39.
Kohlmeyer, J. 1969. Ecological notes on fungi in mangrove forests. Transactions of the
British Mycological Society 53: 237-250.
Largent, D. L., D. Johnson, D. and R. Watling. 1977. How to identify mushrooms to genus.
III. Microscopic features. Mad River Press, Eureka. 148 pp.
Larsen, M. and L. A. Cobb-Poulle. 1990. Phellinus (Hymenochaetaceae): a survey of the
world taxa. Synopsis Fungorum 3. Fungiflora, Oslo. 206 pp.
Lee, B. K. H. and G. E. Baker. 1973. Fungi associated with the roots of red mangrove,
Rhizophora mangle. Mycologia 65: 894-906.
Lodge, D. J. 1996a. Microorganisms. In Reagan, D.P. and R.W. Waide (eds.), The food web
of a tropical rain forest, pp. 53-108. University of Chicago Press, Chicago.
Lodge, D. J. 1996b. Fungi of Puerto Rico and the United States Virgin Islands: a history of
previous surveys, current status, and the future. In Figueroa Colón, J. C. (ed.), The
scientific survey of Puerto Rico and the Virgin Islands: an eighty-year reassessment
of the island’s natural history, Vol. 776, pp. 123-129. Annals of the New York
Academy of Science, New York.
Lodge, D. J., T. J. Baroni, K. K. Nakasone, L. Ryvarden, E. K. Horak, R. J. Vilgalys, N.
Legon, K.-H. Larsson, R. Halling, O. K. Miller, Jr., S. Cantrell, J. Carranza, P. J.
Roberts and A. Ferrer. 2000. Basidiomycetes of the Greater Antilles.
http://www.cortland.edu/nsf/ga.html (Accessed from World Wide Web on 4 October
2004).
Lugo, A. E. 1989. Los manglares de La Parguera. Acta Científica 3: 135-140.
Maekawa, N., H. Suhara, K. Kinjo and R. Kondo. 2003. Corticioid fungi (Basidiomycota) in
mangrove forests of the islands Iriomote and Okinawa, Japan. Mycoscience 44: 403409.
Minter, D. W., M. Rodríguez Hernández and J. Mena Portales. 2001. Fungi of the
Caribbean: an annotated checklist. Published by the senior author and PDMS
Publishing, Middlesex, United Kingdom. 946 pp.
218
Nieves-Rivera, Á. M., D. J. Lodge and O. K. Miller, Jr. 1998. Contributions to the study of
gasteromycetes of Puerto Rico. McIlvainea 13: 50-58.
Nieves-Rivera, Á. M., C. Betancourt and J. S. Mignucci. 1999. Hymenomycetes and
gasteromycetes (Basidiomycotina) of Mona Island Commonwealth Reserve, Puerto
Rico. University of Puerto Rico Agriculture Experiment Station Bulletin 298: 1-91.
Nieves-Rivera, Á. M., T. A. Tattar and E. H. Williams Jr. 2002. Sooty mould-planthopper
association on leaves of the black mangrove Avicennia germinans (L.) Stearn in
southwestern Puerto Rico. Arboricultural Journal 26: 141-155.
Olive, L. S. 1957. Tulasnellaceae of Tahiti. A revision of the family. Mycologia 49: 663-679.
Pegler, D. N. 1983a. Agaric flora of the Lesser Antilles. Her Majesty's Stationery Office,
London, 668 p.
Pegler, D. N. 1983b. The genus Lentinus: a world monograph. Her Majesty's Stationery
Office, London, 281 pp.
Poonyth, A. D., K. D. Hyde, A. Aptroot and A. Peerally. 2000. Mauritania rhizophorae gen.
et sp. nov. (Ascomycetes, Requienellaceae), with a list of terrestrial saprobic
mangrove fungi. Fungal Diversity 4: 101-116.
Reichardt, H.W. 1870. Fungi, Hepaticae et Musci frondosi. In Fenzl, E. (ed.). Reise der
österreichischen Fregatte Novara um die Erde in den Jahren. Botanischer Teil. Band
1, Sporenpflanzen., pp. 131-196. Kaiserlich-Königlichen Hof- und Staatsdruckerei,
Wien.
Schmit, J. P. and C. A. Shearer. 2003. A checklist of mangrove-associated fungi, their
geographical distribution and known host plant. Mycotaxon 85: 423-477.
Sotão, H. M. P., V. L. R. Bononi and T. S. Figueiredo. 1991. Basidiomycetes de manguezais
da Ilha de Maracá, Amapá, Brasil. Boletim do Museu Paraense Emílio Goeldi Série
Botânica 7: 109-114.
Sotão, H. M. P., E. L. Campos, S. P. S. E. Costa, O. A. Melo and J. C. Azevedo. 2002.
Basidiomycetes macroscópicos de manguezais de Bragança, Pará, Brasil. Hoehnea
29: 215-224.
219
Stevenson, J. A. 1975. Fungi of Puerto Rico and the American Virgin Islands. Contribution
of Reed Herbarium No. 23. Braun-Brumfield, Ann Arbor, 743 pp.
Toro, J. A. and J. A. Colón. 1986. Suplemento de información técnica para el plan de manejo
del área de planificación especial del suroeste—Segmento de Boquerón. Oficina de
Zona Costanera, Área de Investigaciones Científicas, Departamento de Recursos
Naturales de Puerto Rico, San Juan, 83 pp.
Vázquez, M.A. 1983. The effects of impounding on a mangrove forest. Master of Science
Thesis, University of Florida, Gainesville, 117 p.
Vázquez, O. J. and D. A. Kolterman. 1998. Floristic composition and vegetation types of the
Punta Guaniquilla Natural Reserve— Cabo Rojo, Puerto Rico. Caribbean Journal of
Science 34: 265-279.
Winter, A., R. S. Appeldoorn, A. Bruckner, E. H. Williams, Jr. and C. Goenaga. 1998. Sea
surface temperatures and coral reef bleaching off La Parguera, Puerto Rico
(northeastern Caribbean Sea). Coral Reefs 17: 377-383.
220
Figures 46A–B. Forests of Avicennia germinans and Rhizophora mangle of the Aquatic
Preserves of Southwest Florida, located southwestern of Venice, at the Gulf of
Mexico, Florida (U.S.A.). A. Pneumatophores and trees of Avicennia germinans. B.
Aerial roots, buttresses, and trees of Rhizophora mangle.
221
Figures 47A–E. Manglicolous basidiomycete collected on southwestern Puerto Rico. A–B.
Crepidotus uber on dead upright Rhizophora mangle tree trunk (A–B, young
basidioma) and branch (C–E, mature basidioma).
222
Figures 48A–H. Further manglicolous basidiomycetes collected on southwestern Puerto Rico and Florida
(U.S.A.). A–B. Psathyrella sp. (Pr = primordium; Bs = basidioma) on Rhizophora mangle leaf litter.
C. Lentinus crinitus. D. Coriolopsis floccosa. E. Hexagonia hydnoides. F. Pycnoporus sanguineus.
G. Tyromyces cf. chioneus on R. mangle wood. H. Dacryopinax spathularia on R. mangle wood.
223
CHAPTER 8
THE OCCURRENCE OF STEMONITIS SPLENDENS (MYXOMYCOTA:
STEMONITALES) ON RHIZOPHORA MANGLE
ABSTRACT
The myxomycete Stemonitis splendens is reported on the red mangrove, Rhizophora
mangle. This is the first report of this myxomycete on mangroves in the Caribbean region
and the fifth report of a myxomycete from R. mangle. Although S. splendens was reported
previously from Puerto Rico, its presence on R. mangle suggests that mangroves, although
usually neglected as a potential substrate for myxomycetes, may support a larger assemblage
of species than indicated by the few published records.
RESUMEN
Se informa el mixomiceto Stemonitis splendens en el mangle rojo, Rhizophora
mangle. Este es el primer informe de este mixomiceto en mangles en la región del Caribe y
el quinto informe de un mixomiceto en R. mangle. Aunque S. splendens se informó
previamente para Puerto Rico, su presencia en R. mangle sugiere que los mangles,
usualmente despreciados como sustrato potencial para los mixomicetos, pueden sostener una
gran asociación de especies que la indicada por los pocos registros publicados.
INTRODUCTION
The myxomycetes in Puerto Rico are still poorly known, although a number of
reports exist (a summary of previous work was included in Novozhilov et al., 2001). The
genus Stemonitis (Myxomycota, Stemonitales) includes a number of common and
widespread species, some of which are fairly distinctive. The single most important
distinguishing feature of this genus is the presence of a reticulum (also known as surface net),
a structure lacking in the morphologically similar genus Comatricha. This reticulum, which
224
develops just beneath the peridium and is connected to the tips of the branches of the
capillitium, forms a well-defined netlike covering over the surface of the spore mass
(Stephenson and Stempen, 1994). Other features used to distinguish Stemonitis from other
members of the Stemonitales are differences in stipe structure and ontogeny, as well as the
origin of the capillitium (Stephenson and Stempen, 1994). The type species of the genus is
Stemonitis fusca.
We found Stemonitis splendens Rostafinski on a red mangrove (Rhizophora mangle)
in a forest site at Cabo Rojo, Puerto Rico (Nieves-Rivera and Darrah, 2002). This marks the
first Caribbean record and the fifth published report of the occurrence of a myxomycete on
red mangrove. The first species reported was Arcyria cinerea, which was collected on June
4, 1968 in the Heeia Swamp, Oahu, Hawaii, on a branch of R. mangle (Kohlmeyer, 1969).
Jan Kohlmeyer gave the material to Constantine J. Alexopoulos who identified the specimen
and deposited it in the University of Texas Myxomycete Collection (UTMC), under
herbarium number 1922 (J. Kohlmeyer, personal communication, 2003). Later, Lee and
Baker (1973) reported Arcyria virescens, Ceratiomyxa sp., and Physarum sp. from living
roots of R. mangle above the tidal line at the same locality. Arcyria cinerea and A. virescens
were the only myxomycetes included in the Schmit and Shearer (2003) survey of mangroveassociated fungi and fungal-like organisms. The occurrence of myxomycetes on mangroves
appears to be rare.
MATERIALS AND METHOD
The study area is located at the Boquerón Commonwealth Forest (BCF), adjacent to
the town of Boquerón, Barrio Boquerón, Cabo Rojo, southwestern Puerto Rico, in a
secondary road (PR-307) at km 17.9, at 18°01’68.5”N, 67°10’49.8”W, at mean sea level, on
29 May 2002 (Figure 2). The region is classified as a subtropical dry forest (Ewel and
Whitmore, 1973). Further details on climatology, ecology, geology, and edaphic formations
of BCF are discussed in Vázquez (1983), Toro and Colón (1986), and Nieves-Rivera et al.
(2002). Rhizophora mangle occasionally forms thick coastal woodland in BCF and in the
225
rest of the southwestern coast of Puerto Rico (Cintrón et al., 1978; Lugo, 1989; Vázquez and
Kolterman, 1998).
RESULTS AND DISCUSSION
Stemonitis splendens Rostafinski, Sluzowce (Mycetozoa) Monografia 195, 1874.
(Figure 49A-B)
Description.–Sporangia 12-15 mm high, clustered and usually forming a large colony,
densely crowded or agglutinated, stipitate, cylindric, rigid and more or less erect, or flexuous,
obtuse to acuminate at the apex, dark purplish brown (Figures 495A-B). Sporotheca
cylindrical, deep purplish brown, obtuse, more or less erect but flexuose towards the apex,
0.5 mm in diameter. Stipe 3-4 x 0.1-0.2 mm, slender, lustrous, conspisuously flared at the
base, arising from a widely expanded, silvery to somewhat purplish hypothallus. Columella
reaching nearly to the sporangial apex, attenuate, often coiled and tortuous toward the tip,
dark reddish brown. Capillitium open-meshed and arising from the columella by relatively
few major branches, sometimes with membranous junctions; surface reticulum fairly robust,
smooth, with irregular, rounded to polygonal meshes, mostly 30-90 µm in diameter,
incomplete or absent in agglutinated fruitings, brown. Spores 7-9 µm in diameter, globose,
thin-walled, faintly verrucose (warted), lilaceous brown by transmitted light, dark purplish
black in mass.
Habitat.–Gregarious or solitary on decayed fallen trunk (25 cm in diameter) of R.
mangle (0.5 m above ground).
Material Studied.–PUERTO RICO: Boquerón Commonwealth Forest, adjacent to
Puerto Real, boardwalk that passes through a mangrove forest and reaches the lagoon on the
Boquerón-Guaniquilla mangrove forest, < 1 m alt., 29-V-2002, UARK #17308.
Remarks.–Stemonitis splendens (UARK #17308) is a common species, often occurs
in large fruitings (as in Figure 49B) and has sporangia with conspicuous large-meshed
surface net and nonreticulate spores (Stephenson and Stempen, 1994). The specimen was
deposited in the herbarium (UARK) of the University of Arkansas in Fayetteville, Arkansas.
226
Although Stemonitis splendens Rostafinski was reported previously from Puerto Rico
(Novozhilov et al., 2001) and the Caribbean in general (Stevenson, 1975; Minter et al., 2001;
Camino et al., 2003), its presence on R. mangle suggests that mangroves, although usually
neglected as a potential substrate for myxomycetes, may support a larger assemblage of
species than indicated by the few published records. Therefore, it would seem important to
continue studying such habitats, in order to contribute to the conservation and knowledge of
the biodiversity of myxomycetes of Puerto Rico.
LITERATURE CITED
Camino, M., G. Moreno, A. Castillo and C. Illana. 2003. Revision of the family
Stemonitaceae in Cuba. Mycotaxon 88: 315-331.
Cintrón, G., C. Goenaga and J. González-Liboy. 1978. Ecología del manglar en una zona
árida: Exposición al oleaje y estructura del manglar. In Quinto Simposio de los
Recursos Naturales (ed.), Departamento de Recursos Naturales de Puerto Rico, pp.
57-86. Estación Experimental Agrícola, Río Piedras, Puerto Rico.
Ewel, J. J. and J. L. Whitmore. 1973. The ecological life zones of Puerto Rico and the U. S.
Virgin Islands. Forest Service Research Paper ITF-18. U. S. Forest Service, Institute
of Tropical Forestry, Río Piedras, Puerto Rico. 72 pp., 1 map.
Kohlmeyer, J. 1969. Ecological notes on fungi in mangrove forests. Transactions of the
British Mycological Society 53: 237-250.
Lee, B. K. H. and G. E. Baker. 1973. Fungi associated with the roots of red mangrove,
Rhizophora mangle. Mycologia 65: 894-906.
Lugo, A. E. 1989. Los manglares de La Parguera. Acta Científica 3: 135-140.
Minter, D. W., M. Rodríguez Hernández and J. Mena Portales. 2001. Fungi of the
Caribbean: an annotated checklist. Published by the senior author and PDMS
Publishing, Middlesex, United Kingdom. 946 pp.
Nieves-Rivera, Á. M. and R. G. Darrah. 2002. Further studies of slime molds in Puerto Rico.
Inoculum 53(5): 2-5.
227
Nieves-Rivera, Á. M., T. A. Tattar and E. H. Williams, Jr. 2002. Sooty mould-planthopper
association on leaves of the black mangrove Avicennia germinans (L.) Stearn in
southwestern Puerto Rico. Arboricultural Journal 26: 141-155.
Novozhilov, Y. K., M. Schnittler, A. W. Rollins and S. L. Stephenson. 2001. Myxomycetes
from different forest types in Puerto Rico. Mycotaxon 77: 285-299.
Schmit, J. P. and C. A. Shearer. 2003. A checklist of mangrove-associated fungi, their
geographical distribution and known host plants. Mycotaxon 85: 423-477.
Stephenson, S. L. and H. Stempen. 1994. Myxomycetes: A handbook of slime molds.
Timber Press, Oregon. 183 pp., 16 pl.
Stevenson, J. A. 1975. Fungi of Puerto Rico and the American Virgin Islands. BraunBrumfield, Inc., Michigan. 743 pp.
Toro, J. A. and J. A. Colón. 1986. Suplemento de información técnica para el plan de manejo
del área de planificación especial del suroeste—Segmento de Boquerón. Oficina de
Zona Costanera, Área de Investigaciones Científicas, Departamento de Recursos
Naturales de Puerto Rico, San Juan, Puerto Rico. 83 pp., 1 map.
Vázquez, M. A. 1983. The effects of impounding on a mangrove forest. M.Sc. Thesis,
University of Florida, Gainesville. 117 pp.
Vázquez, O. J. and D. A. Kolterman. 1998. Floristic composition and vegetation types of the
Punta Guaniquilla Natural Reserve– Cabo Rojo, Puerto Rico. Caribbean Journal of
Science 34: 265-279.
228
Figures. 49A-B. A. Sporangia of Stemonitis splendens (UARK #17308; a = side view, b =
bottom view) collected on Rhizophora mangle, Boquerón Commonwealth Forest,
Cabo Rojo, Puerto Rico, West Indies. B. Fruiting of Stemonitis splendens in New
Zealand.
229
CHAPTER 9. GENERAL DISCUSSION
In this study, I have provided historical background and added some new records to
the body of information that is available for the study of coastal fungi, including marine,
estuarine, and mangrove-associated fungi in Puerto Rico. The techniques used in this study,
although basic or simple, are currently used in taxonomic and physiological studies. In this
part of this study, I would like to identify gaps in our current knowledge where future
research should focus and propose novel areas for research in marine mycology in Puerto
Rico.
From previous studies, the methodology employed for the isolation and culturing of
fungi from marine and estuarine habitats has progressed little since the 1960’s. Also,
conclusions regarding the biodiversity and distribution of coastal fungi are often made
without the aid of statistical analysis. Most beginning mycologists have the general tendency
to identify large numbers of taxa or to force collections into existing species concepts.
However, over time, Puerto Rican mycologists have developed greater sophistication.
The substrates surveyed in Puerto Rico included vascular plant material, sand grains,
sandy soils, algae, and few terrestrial and aquatic animals, which tend to support a rich fungal
diversity within coastal environments. However, many more substrates deserve further study
such as marine flora (e.g., phytoplankton. algae, seaweeds, cryptogams) and fauna (e.g.,
zooplankton, calcareous mollusks, crustacean exoskeletons, sea birds, marine mammals).
Other habitats that deserve more attention are hypersaline lagoons and brines, estuaries,
pelagic waters, deep sea; such habitats might support physiologically-adapted fungi which
may be of interest to biotechnology. Another applied research field is the pollution of the
seas and the mutations in fungal populations, and the capability of such fungal isolates to
withstand and biotransform harmful chemicals (e.g., PAHs) into less detrimental ones. Fungi
in coastal environment such as parasites, commensals, and endophytes warrant further
attention, as do understudied groups as the lower fungi and other fungal-like organisms (e.g.,
myxomycetes, oomycetes).
230
The oomycete Halophytophthora sp. has been isolated from leaf samples. This
earliest group of fungal-like organisms decomposes senescent leaves and are quickly
replaced by marine higher fungi. Frequent inhabitants of leaves are the mitosporic fungi and
it is common to find geophilic species at early stages of incubation, for instance,
Cladosporium spp. and Pestalotiopsis spp. Species of Penicillium and Trichoderma spp.
occurred at advanced stages of leaf decomposition.
Most of the species coastal fungi found in this research are cosmopolitan. They are
common saprophytes in aquatic habitats (marine, estuarine and freshwater habitats), forest
timber, grow on sandy soil, and form mycorrhizal associations with both exotic and native
plants. In general, they also are well represented in xerophytic and non-xerophytic areas.
This study documents the large numbers of taxa found in the coasts of Puerto Rico, taking
into account all surveys available (published and unpublished) of coastal fungi. For example,
the terrestrial coast mycobiota (e.g., basidiomycetes) shows adaptability to xeric
environments because they tend to form epigeous basidiocarps avoiding sun exposure,
minimizing loss of water content. Moisture is a limiting factor in their collection; therefore,
rainy seasons are the best time for collecting them. The climate of southwestern Puerto Rico
has been described as subtropical dry, with scanty rain and strong winds from the southeast.
In general, there are two distinct rainy seasons in Puerto Rico, one in May-June, the other in
August-November. Summer and autumn months tend to be more or less rainy, while winter
and spring months are usually drier.
The evaluation of biodiversity is becoming an extremely important part in the
conservation of the environment and the naming of biological species depends on good
systematics. The southwestern coast of Puerto Rico includes endemic and extirpated species
of flora and fauna, interesting geological features, important archaeological sites, and
spectacular landscapes unlike those found in many other coasts of Puerto Rico. These
features are the product of the unique situation of this island-small size, belongs to the
Antillean island arc, volcano-oceanic origin, relative isolation, and the degree of protection
caused by remoteness from populated lands.
231
Islands located at a considerable distance from large land masses such as continents
will be reached by a relatively small number of organisms that are well adapted to dispersal
over long distances (e.g., seabirds, insects, beach plants, algae, pollen, fungal spores).
However, organisms less suited for such dispersal will reach islands less frequently, perhaps
blown during a hurricane, transported on the feet or in the digestive tract of a bird, or on an
ocean-going raft of vegetation.
A database of coastal fungi would be needed for comparison of all known species,
including aquatic and terrestrial fungi. This database would be integrated into an internet or
compact disk-read only memory (CD-ROM) medium where data can be conveniently
updated and accessed. The consideration of the use of DNA sequence analysis and
morphology characterization of sexual and asexual structures are nowadays the tools of
choice for the study of fungal identification.
In conclusion, there is no doubt that data on the existing biodiversity must be
recorded prior to the evaluation of any changes in the mycobiota. Without at least a
preliminary inventory of the fungi in Puerto Rico, it is nearly impossible to assay what is
novel (e.g., new species or records). It seems important to continue studying such habitats, in
order to contribute to the conservation and knowledge of the biodiversity of fungi and fungallike organisms of Puerto Rico.
232
CHAPTER 10. GENERAL CONCLUSIONS
1.
Most fungi and fungal-like organisms collected or isolated in mangroves of
southwestern Puerto Rico are cosmopolitan.
They are common saprophytes,
parasites, endophytes, and filoplanes in coastal forest timber or grow on leaves and
wood.
2.
A high number of fungal taxa (604 spp.) was found among coastal fungi of Puerto
Rico, when compared with previous reports from other coastal forests (i.e., Belize,
Cuba, Mexico), but similar to that recorded in the Lesser Antilles (i.e., Guadeloupe,
St. Croix). However, one must keep in mind that coastal mycology surveys are novel
in the Caribbean.
3.
Abrupt changes in the ecosystems of southwestern Puerto Rico, particularly
deforestation of coastal forests (mangroves), may cause the decrease of native
organims that depend on naturally occurring associations, such as fungi.
4.
Evaluation of the existing biodiversity must be recorded prior to the evaluation of any
changes in mycobiota. Mangroves, usually neglected as a potential substrate for
fungi, supported a larger assemblage of species than indicated by the literature.
233
CHAPTER 11. RECOMMENDATIONS
1.
To continue to study the biodegradation and bioremediation qualities of the
Cladosporium species (C. sphaerospermum and C. oxysporum) and native strains of
marine and terrigenous fungi by using HPLC or any other biochemical techniques to
determine if these species in fact metabolize PAHs.
2.
To survey other mangrove-associated plants (e.g., Acrostichum spp., Batis maritima,
etc.) for fungi and fungal-like organisms and compare their biodiversity, taxonomic
composition, population ecology, molecular biology, and phylogeny.
3.
To more exhaustively survey manglicolous fungal-like organisms, endophytes,
mycosis of marine fauna, and marine phytopathology of Puerto Rico, which are by no
means exhausted. Future collections may produce some new records.
4.
To conduct mycological research in Puerto Rico in the pelagic, shelf, benthic, and
deep sea areas (e.g., Puerto Rico’s Trench).
234
APPENDIXES
235
APPENDIX 1
AQUATIC FUNGI FROM ESTUARIES IN PUERTO RICO. I. MOUTH OF THE
MANATÍ RIVER
ABSTRACT
Aquatic fungi were isolated from sea foam, leaf litter, beach sand, and drift wood
from an estuary known as “La Boca” (river mouth) of the Manatí River in Barceloneta,
northern Puerto Rico. Observations of the baits (sterilized balsa wood) and incubated
organic debris revealed the presence of 28 species of aquatic fungi, 13 of which belong to the
ascomycetes and 15 to the mitosporic fungi. The species Arenariomyces triseptatus,
Astrosphaeriella aff. mangrovei, Corollospora cf. colossa, C. filiformis, Halosphaeria sp.,
Kirschsteiniothelia sp., Torpedospora radiata (Ascomycota), Brachiosphaera tropicalis,
Campylospora sp., and Clavatospora bulbosa (Mitosporic fungi) were the most common in
the samples. Eleven species are new records for Puerto Rico and six fungal isolates could
not be identified.
RESUMEN
Se aislaron hongos acuáticos de la espuma de mar, hojarasca, arena playera y madera
a la deriva de un estuario conocido como “La Boca” (boca de río) del Río Manatí en
Barceloneta, al norte de Puerto Rico. Las observaciones de los cebos (madera de balsa
esterilizada) y material orgánico incubado revelaron la presencia de 28 especies de hongos
acuáticos, de los cuales 13 pertenecen a los ascomicetos y 15 a los hongos mitospóricos. Las
especies Arenariomyces triseptatus, Astrosphaeriella aff. mangrovei, Corollospora cf.
colossa, C. filiformis, Halosphaeria sp., Kirschsteiniothelia sp., Torpedospora radiata
(Ascomycota), Brachiosphaera tropicalis, Campylospora sp. y Clavatospora bulbosa
(Hongos Mitospóricos) fueron las más comunes en las muestras. Once especies son nuevos
registros para Puerto Rico y se aislaron seis hongos que no pudieron ser identificados.
236
INTRODUCTION
Fungi occur widely in the oceans, seas, and estuaries as parasites in marine plants or
animals or as saprobes on timber, algae, sea grasses, protozoans, corals, sea foam, and other
substrata (Kirk et al., 2001). The distribution of fungi in the seas and estuaries has been
studied in several Caribbean localities, including (1) Belize (Kohlmeyer and VolkmannKohlmeyer, 1987a; Minter et al., 2001), (2) Cuba (Capó-de Paz, 1986a, b; Minter et al.,
2001; González et al., 2003), (3) Lesser Antilles (Kohlmeyer and Kohlmeyer, 1971; Fell and
Master, 1975; Stevenson, 1975; Kohlmeyer, 1981; Kohlmeyer and Volkmann-Kohlmeyer,
1987b, 1988; Minter et al., 2001), (4) Mexico (González et al., 1998, 2000, 2001; Minter et
al., 2001), (5) Puerto Rico (Rossy-Valderrama, 1956; Meyers, 1957; Kohlmeyer, 1968;
Carvajal-Zamora, 1971; Hernández-Vera, 1972, 1975, 1980; Stevenson, 1975; Galler-Rimm,
1982; Hernández-Vera and Almodóvar, 1983, 1984; Acevedo, 1987, 2001; Valdéz-Collazo
et al., 1987; Kohlmeyer and Volkmann-Kohlmeyer, 1987c; Calzada, 1988; Lodge, 1996;
Minter et al., 2001; Nieves-Rivera et al., 2002), and (6) Venezuela (Dennis, 1970; Minter et
al., 2001). Although aquatic fungi have been extensively studied worldwide, in many
Caribbean islands the estuarine and marine mycobiota is poorly known. Most estuarine and
marine collections are in some cases sporadic and interrupted by many years. The purpose of
this study was to report the incidence of aquatic fungi in marine foam from a river mouth
(estuarine conditions) in Puerto Rico, a subtropical island located between 18°00’–18°30’ N,
65°35’–67°15’W, in the northeastern Caribbean Sea. Fifteen of the samples were Mitosporic
fungi, the most common group in the samples, and 13 were ascomycetes, eleven species are
new records for Puerto Rico.
MATERIALS AND METHOD
On 25 November 1998, intertidal foam and sand were collected from the Manatí River
(also known as “Río Grande de Manatí”). Samples were baited in the laboratory with pieces
237
of sterilized balsa wood 10 cm long. Washed-up beach debris (leaf litter and drift wood) also
was collected for incubation, following the methods described by Kohlmeyer and Kohlmeyer
(1979).
A sandy beach of the Manatí River mouth, also known as “La Boca” or “Boca” (Spanish
for mouth), located next to road PR-684 in the Barceloneta municipality, about 36 km west
(18°28’81.8” N and 66°32’09.2” W) of San Juan, was selected as study site (Figure 50). The
Manatí River is about 64 km long, rises in Cordillera Central just north of Barranquitas,
flows northwest, past Ciales to the Atlantic Ocean 6.4 km northwest of Manatí. Mean annual
precipitation is less than 1,650 mm and mean annual temperature is 25.3° C (Ravalo et al.,
1986, Anonymous, 2003). The climate is described as that of the subtropical dry life zone
(Ewel and Whitmore, 1973). Surface seawater salinity ranges from 19 to 31 g/L, with
temperatures of 27 to 31°C, a pH of 7.4 to 8.7, a total alkalinity of 87 to 116 mg/L, and a
dissolved oxygen (DO) value of 7.2 to 8.6 mg/L (Nieves-Rivera, unpublished data, 2003).
At the collection site, the river mouth beach sand composition is dominated by dark minerals,
volcanic rock fragments, quartz, and feldspar (Morelock et al., 1985). The geomorphology
of La Boca was studied by Lobeck (1922), Wood et al. (1975), Morelock et al. (1985), and
Barreto-Orta (1997).
The Manatí River estuary has been found to be highly stratigraphic, with little or no
mixture in the interphase between fresh- and seawater (Carvajal-Zamora, 1977). The sea
wedge extends five to six km up river. The intrusion of seawater into the river usually causes
a decrease in DO, which is related to a high microbiological activity and chemical demand in
the sea wedge and in sediments; this decrease in DO creates anoxic conditions during the
year (Carvajal-Zamora, 1977). Slightly high phosphate (PO43– = 0.08 to 0.12 µM) and nitrate
(NO32– = 16.72 µM) values have been detected in near shore surface waters, especially near
the Manatí River mouth, probably because of agricultural runoff (Wood et al., 1975: Station
PMA-3A at 0 m depth). Carvajal-Zamora (1977) found that the concentrations of heavy
metals and nutrients in the sediments of the Manatí River were greater in the estuarine part
238
than in the riverine portion.
The procedure used in the present study for collection and preservation of marine
foam on the sandy beach follows Kohlmeyer and Kohlmeyer (1979); their illustrated keys are
described elsewhere (Kohlmeyer and Volkmann-Kohlmeyer, 1991; Hyde and Sarma, 2000).
For the isolation and identification of aquatic hyphomycetes, we followed Santos-Flores and
Betancourt-López (1997). Voucher specimens (slides) of all species were deposited at the
University of Puerto Rico at Río Piedras herbarium (UPRRP).
RESULTS AND DISCUSSION
A total of 28 fungi were found, 13 of which belong to the ascomycetes and 15 to the
mitosporic fungi (Table VIII; Figures 51A-H). The species Arenariomyces triseptatus
(Figure 51A), Astrosphaeriella aff. mangrovei (Figure 517B), Corollospora cf. colossa
(Figure 51C), C. filiformis (Figure 51D), Halosphaeria sp., Kirschsteiniothelia sp. (Figure
517E), Torpedospora radiata (Figure 51F) (Ascomycota), Brachiosphaera tropicalis (Figure
51G), Campylospora sp., Clavatospora bulbosa (Figure 51H) (Mitosporic fungi) were the
most common in the samples. Few marine ascomycetes were isolated; this scarcity was
probably due to the selectivity of the isolation method employed and the time of sampling.
Ascomycetes and mitosporic fungi (mostly aquatic hyphomycetes) were found mixed in
foam, leaf litter, and sand of the La Boca beach. Eleven species are new records for Puerto
Rico, and six fungal isolates could not be identified (Table VIII).
Along with ascospores of marine fungi, conidia of 10 species of aquatic
hyphomycetes were detected: Anguillospora cf. longissima, Articulospora tetracladia,
Brachiosphaera tropicalis, Camposporidium sp., Campylospora sp., Diplocladiella
scalaroides, Lemonniera pseudofloscula, Triscelophorus acuminatus, Triscelophorus sp.,
and Tubeufia cylindrothecia (Santos-Flores and Betancourt-López, 1997) (Table VIII).
Camposporidium sp. conidia resembled Camposporidium spp. reported from Río Sonadora
and Quebrada Jiménez nears the El Verde LTER Field Station, Puerto Rico (Hamilton,
1973). Some specimens of B. tropicalis showed a translucent halo around the central cell
239
(Figure 51G); while others B. tropicalis were typical. Aquatic hyphomycetes are not
uncommon in estuarine habitats (Johnson and Sparrow, 1961; Kohlmeyer and Kohlmeyer,
1979).
For example, Kirk (1969) reported two lignicolous aquatic hyphomycetes
(Clavatospora stellatacula and Tetraploa aristata) adapted to seawater conditions in the
Chesapeake Bay.
Conidia of six terrigenous mitosporic fungi also were isolated from sea foam
samples. These were: Curvularia sp., Fusarium sp., and conidia of four unknown species, as
listed in Table VIII. The unknown spores (also referred to as Ignotus by Acevedo, 1987,
2001) are described as dematiaceous, aseptate or septate, ranging in sizes, aleuriospore,
dictyospore, phragmospore, lenticular with or without apical pore. These terrigenous species
are apparently common to marine and estuarine habitats. Acevedo (1987) found 25 species
of mitosporic fungi in sand, from the reefs and an offshore island (Isla Cuevas) from La
Parguera, southwestern Puerto Rico.
These mitosporic fungi were: Alternaria sp.,
Aspergillus spp., Cephalosporium sp., Cladosporium spp., Curvularia spp., Diplodia spp.,
Fusarium spp., Helminthosporium spp., Penicillium spp., Scopulariopsis sp., Trichoderma
spp., and Mycelia Sterilia (Acevedo, 1987).
In this survey, we did not recover any species that were potentially pathogenic to
humans, a finding different to that of González et al. (2000) (Table IX). The assemblage of
species reported by González et al. (2000) also differs greatly from that recorded in the
present study due to methodology and bait used. González et al. (2000) isolated a total of 17
keratinophilic fungi, 13 of which were hyphomycetes and four ascomycetes. González et al.
(2001) reported a larger collection throughout Mexico; therefore, their checklist contains 47
ascomyetes, 14 mitosporic fungi, and one basidiomycete (Table IX). In the Cuban survey,
González et al. (2003) reported 29 marine fungi (25 ascomycetes and four mitosporic fungi),
19 of which were new records (Table IX). In contrast, our collection from a single locality
produced 15 mitosporic fungi and 13 ascomycetes (Table IX).
Most of the species collected are arenicolous (Arenariomyces cf. majusculus, A.
triseptatus, Corollosopora cf. colossa, C. filiformis, C. cf. pseudopulchella; see Kohlmeyer
240
and Kohlmeyer, 1979). Marine ascomycetes (Arenariomyces triseptatus, Lulworthia sp.,
Torpedospora radiata) have been recorded previously from Belize (Kohlmeyer and
Volkmann-Kohlmeyer, 1987a), Cuba (González et al., 2003), Mexico (González et al., 2000,
2001), Puerto Rico (Nieves-Rivera and Santos-Flores, unpublished data, 2004), and St. Croix
(Kohlmeyer and Volkmann-Kohlmeyer, 1988) (Table IX).
Halorosellinia oceanica,
Lulworthia sp., and Torpedospora radiata are lignicolous or algicolous species (Kohlmeyer
and Kohlmeyer, 1979; Acevedo, 2001). All species isolated in this survey are saprobes
living in parts of angiosperms, plant debris, or in the blades of seagrasses.
Fifteen of the samples were Mitosporic fungi, the most common group in the
samples, and 13 were ascomycetes. Eleven species are new records for Puerto Rico. The
species we found are most similar to those from previous studies in Puerto Rico (CarvajalZamora, 1971; Hernández-Vera, 1972, 1975, 1980; Stevenson, 1975; Hernández-Vera and
Almodóvar, 1983, 1984; Acevedo, 1987, 2001; Calzada, 1988; Lodge, 1996; Minter et al.,
2001) (Table VIII).
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245
Table VIII. Summary of the aquatic fungi (obligate and facultative marine fungi sensu
Kohlmeyer, 1974) recovered from samples of sea foam, leaf litter, and wood in the
mouth of the Manatí River, northern Puerto Rico.
__________________________________________________________________________
Fungus
Substrate1
__________________________________________________________________________
ASCOMYCOTA
Arenariomyces cf. majusculus Kohlm. & Volkm.-Kohlm.*
F, S
A. triseptatus Kohlm.
F, S
Astrosphaeriella aff. mangrovei (Kohlm. & Vittal) Aptroot & K.D. Hyde*
F, L, W
Chaetomastia cf. typhicola (Karst.) Barr*
F, S
Corollospora cf. colossa Nakagiri & Tokura *
F
C. filiformis Nakagiri, in Nakagiri & Tokura*
F
C. cf. pseudopulchella Nakagiri & Tokura*
F
Coronopapilla aff. mangrovei (K.D. Hyde) Kohlm. & Volkm.-Kohlm.* F, W
Halosphaeria cucullata (Kohlm.) Kohlm.*
F, S
Halosphaeria sp.
F, S
Kirschsteiniothelia sp.*
F
Kretzschmariella culmorum (Cooke) Y.M. Ju & J.D. Rogers
F
Lindra sp.
F, S
Lulworthia sp.
F, S
Torpedospora radiata Meyers
F
Unknown sp. 1
F, S
Unknown sp. 2
F
MITOSPORIC FUNGI
Anguillospora cf. longissima (Sacc. & P. Syd.) Ingold
Articulospora tetracladia Ingold
Brachiosphaera tropicalis Nawawi, in Descals
Camposporidium sp.
Campylospora sp. (s. str. Santos-Flores & Betancourt-López, 1997)
Clavatospora bulbosa (Anastasiou) Nakagiri & Tubaki*
Curvularia sp.
Diplocladiella scalaroides G. Arnaud
Fusarium sp.
Lemonniera pseudofloscula Dyko, in Descals, J. Webster & Dyko
Triscelophorus acuminatus Nawawi
Triscelophorus sp.
Tubeufia cylindrothecia (Seaver) Höhn.*
246
F, L, S
F
F
F
F, L, W
F, S
F, L, W
F, L
F, L, W
F, L
F, S
F
F, S
Unknown sp. 1
Unknown sp. 2
Unknown sp. 3
Unknown sp. 4
F, W
F, L
F
F, W
1
Substrate: F = sea foam; L = leaf littter; S = beach sand; W = wood.
* = New record for Puerto Rico.
__________________________________________________________________________
247
Table IX. Summary of the aquatic fungi (obligate and facultative marine fungi sensu
Kohlmeyer, 1974) recovered from the Caribbean.
__________________________________________________________________________
Kohlmeyer & Volckmann-Kohlmeyer
(1987a) [Belize]
ASCOMYCOTA
Aigialus grandis
Aniptodera chesapeakensis
*Arenariomyces triseptatus
Ascocratera manglicola
Belizeana tuberculata
Biatriospora marina
Caryosporella rhizophorae
Chadefaudia corallinarum
Corollospora maritima
C. pulchella
Dactylospora haliotrepha
Didymosphaeria enalia
D. rhizophorae
Halosarpheia abonnis
H. fibrosa
Halosphaeria cucullata
H. quadricornuta
H. salina
Hydronectria tethys
Keissleriella blepharospora
Leptosphaeria australiensis
L. avicenniae
Lignicola laevis
L. tropica
Lindra marinera
L. thalassiae
L. thalassiae var. crassa
Lophiostoma mangrovis
Lulworthia grandispora
L. kniepii var. curalii
*Lulworthia sp.
Massarina thalassiae
Mycosphaerella pneumatophorae
M. salicorniae
Swampomyces armeniacus
Kohlmeyer & Volckmann-Kohlmeyer
(1988) [St. Croix]
ASCOMYCOTA
Aigialus grandis
A. parvus
Arenariomyces parvulus
*A. triseptatus
Ascocratera manglicola
Belizeana tuberculata
Caryosporella rhizophorae
Ceriosporopsis halima
Dactylospora haliotrepha
Didymosphaeria enalia
D. rhizophorae
Halosphaeria quadricornuta
H. salina
Hydronectria tethys
Keissleriella blepharospora
Leptosphaeria australiensis
Lignicola laevis
L. tropica
Lindra marinera
L. thalassiae
Lophiostoma mangrovis
Lulworthia grandispora
*Lulworthia sp.
Massarina thalassiae
M. velatospora
Ophiodeira monosemeia
Passeriniella savoryellopsis
Swampomyces armeniacus
*Torpedospora radiata
Trematosphaeria lignatilis
BASIDIOMYCOTA
Halocyphina villosa
Nia vibrissa
MITOSPORIC FUNGI
Cytospora rhizophorae
248
*Torpedospora radiata
Trematosphaeria lignatilis
BASIDIOMYCOTA
Halocyphina villosa
Nia vibrissa
MITOSPORIC FUNGI
Cytospora rhizophorae
Humicola alopallonella
Periconia prolifica
Rhabdospora avicenniae
Trichocladium achrasporum
Varicosporina ramulosa
Zalerion cf. varium
Humicola alopallonella
González et al.
(2000) [Mexico]
MITOSPORIC FUNGI
Aspergillus auricomus
A. fumigatus
A. terreus
Fusarium semitectum
F. solani
Gymnascella dankaliensis
Gymnoascus sp.
Scopulariopsis brumptii
González et al.
(2001) [Mexico]
ASCOMYCOTA
Antennospora quadriconuta
A. salina
Arenariomyces parvulus
A. trifurcatus
*A. triseptatus
Ceriosporopsis halima
Corollospora angusta
C. gracilis
C. maritima
C. pseudopulchella
C. pulchella
Etheirophora blepharospora
Halosphaeriopsis mediosetigera
Hydronectria tethys
Leptosphaeria australiensis
L. avicenniae
Lignicola tropica
Lindra crassa
L. marinera
L. thalassiae
*Lulworthia sp.
Mycosphaerella pneumatophorae
M. salicorniae
Paraliomyces lentiferus
Remispora galerita
González et al.
(2003) [Cuba]
ASCOMYCOTA
Antennospora quadriconuta
Arenariomyces parvulus
A. trifurcatus
*A. triseptatus
Ceriosporopsis halima
Corollospora armoricana
C. cinnamomea
C. gracilis
C. maritima
C. pseudopulchella
C. quinqueseptata
Falcatispora cincinnatula
Lignicola leavis
L. tropica
Lindra marinera
L. obtusa
L. thalassiae
Lulworthia grandispora
L. fucicola
L. kniepii
*Lulworthia sp.
Neptunella longirostris
Pontogeneia cubensis
Savoryella lignicola
*Torpedospora radiata
249
*Torpedospora radiata
Verruculina enalia
BASIDIOMYCOTA
Nia vibrissa
MITOSPORIC FUNGI
Cirrenalia pseudomacrocephala
Cytospora rhizophorae
Periconia prolifica
Tricocladium achrasporum
T. alopallonellum
Varicosporina ramulosa
MITOSPORIC FUNGI
Cirrenalia basiminuta
Tricocladium alopallonellum
Varicosporina ramulosa
Zalerion varium
Nieves-Rivera and Santos-Flores
(present study) [Puerto Rico]
ASCOMYCOTA
Arenariomyces cf. majusculus
*A. triseptatus
Astrosphaeriella aff. mangrovei
Chaetomastia cf. typhicola
Corollospora cf. colossa
C. filiformis
C. cf. pseudopulchella
Coronopapilla aff. mangrovei
Halosphaeria cucullata
Halosphaeria sp.
Kirschsteiniothelia sp.
Kretzschmariella culmorum
Lindra sp.
*Lulworthia sp.
*Torpedospora radiata
MITOSPORIC FUNGI
Anguillospora cf. longissima
Articulospora tetracladia
Brachiosphaera tropicalis
Camposporidium sp.
Campylospora sp.
Clavatospora bulbosa
Curvularia sp.
Diplocladiella scalaroides
Fusarium sp.
Lemonniera pseudofloscula
Triscelophorus acuminatus
250
Triscelophorus sp.
Tubeufia cylindrothecia
* = Marine fungi isolated in common from the Caribbean.
__________________________________________________________________________
251
Figure 50. Collection of sea foam samples from the mouth of Manatí River, north central
coast of Puerto Rico.
252
Figures 51A-H. Aquatic fungi isolated from samples collected of the mouth of Manatí
River, northern Puerto Rico. Ascospores: A. Arenariomyces triseptatus. B.
Astrosphaeriella aff. mangrovei. C. Corollospora cf. colossa. D. Corollospora
filiformis. E. Kirschsteiniothelia sp. F. Torpedospora radiata. Conidia: G.
Brachiosphaera tropicalis. H. Clavatospora bulbosa.
253
APPENDIX 2
BIBLIOGRAPHY ON COASTAL AND MARINE BIOLOGY IN PUERTO RICO,
WITH SPECIAL EMPHASIS ON MARINE MYCOLOGY
INTRODUCTION
This bibliography on coastal and marine biology is primarily mycological in
orientation. A bibliography on marine mycology in Puerto Rico, with special emphasis on
manglicolous fungi is updated to 31 December 2004. A total of 1,085 citations or references
are included. Due to recent interest in marine mycology, coastal mycology (including
terrestrial and estuarine fungi, and lichenized fungi), paleomycology, and general
oceanography (biological, geological, chemical, and physical oceanographies), it is our
intention to present this bibliographic recompilation for the benefit of future generations of
marine mycologists and oceanographers (professional and non-professional) alike. Although
many of the references included in this bibliography might give the impression of not being
related to Puerto Rico, in some cases the references reported one or more collections or
isolations from the island. This bibliography also includes handbooks and key papers (e.g.,
Matsushima’s works, 1971 to 2001) that are very helpful for initial studies on aquatic
hyphomycetes, higher marine fungi, filamentous fungi or fungal like organisms (e.g.,
myxomycetes, oomycetes). Also included are a few studies of marine biology, geology,
chemistry, and physics, including several marine pollution and marine zoologically-oriented
studies. Although literature searches for these latter subjects have not been exhaustive, we
have tried to add relevant papers with extensive bibliographies whenever possible. We also
have made an effort to include marine mycology-based unpublished theses for which we
have searched diligently in the various libraries of the University of Puerto Rico (basically
the Mayagüez and Río Piedras Campuses) and other unpublished reports such as those by the
Puerto Rico Department of Natural and Environmental Resources.
254
MARINE AND COASTAL MYCOLOGY
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Puerto Rican freshwater and marine sport fishes. Puerto Rico Department of Natural
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Puerto Rican freshwater sport fishes. Puerto Rico Department of Natural &
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Puerto Rican marine sport fishes. Puerto Rico Department of Natural &
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256
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de Puerto Rico, Mayagüez, Puerto Rico. 64 pp.
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