Rodrigusia

Print version ISSN 0370-6583On-line version ISSN 2175-7860

Rodrigusiavol.66no.4RiodeJaneiro2015

http://dx.doi.org/10.1590/2175-7860201566416

ORIGINAL PAPERS

Flora of North America North of Mexico

Nancy R. Morin1

Luc Brouillet2

Geoffrey A. Levin3

1
Flora of North America Association, P.O. Box 716, Point Arena, California 95468, USA.
nancy.morin@nau.edu
2
Institut de recherch en biologie vgtale, Universit de Montral, 4101 est, rue
Sherbrooke, Montral, Qubec H1X 2B2, Canada. luc.brouillet@umontreal.ca
3
Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816
S Oak St., Champaign, IL 61820, USA. levin1@illinois.edu
ABSTRACT

The Flora of North America north of Mexico treats all native and naturalized vascular plants
and bryophytes in Canada, Greenland, St. Pierre et Miquelon, and the continental United
States including the Florida Keys and Aleutian Islands (approximately 18 million square
kilometers). It provides accepted names, literature citations, basionyms, synonyms,
morphological descriptions, habitat, geographical distribution, conservation or weed status,
and a discussion of taxonomic issues for approximately 20,000 species. Of the total 30
volumes anticipated, 18 have been published and one is in press, treating 2021 genera and
12,393 species. For the remaining volumes, 763 genera and 5,008 species have been
submitted, and 82 of the 144 families have been submitted in full. Completion is anticipated
by the end of 2017. The project is managed by the Flora of North America Association.
Content from published volumes is available through eFloras and JSTOR and has been
provided to the World Flora informatics team.

Key words: Flora; North America; United States; Canada; Greenland

RESUMO

A Flora da Amrica do Norte ao Norte do Mxico trata todas as espcies de plantas

vasculares e brifitas ocorrente sno Canad, Groenlndia, St. Pierre et Miquelon e na parte
continental dos Estados Unidos, incluindo as Florida Keys e as Aleutian Islands
(aproximadametne 18 milhes de kilmetros quadrados). Informaes como nomes
aceitos, citaes bibliogrficas, basinimos, sinonmia, descries morfolgicas, habitat,
distribuio geogrfica, estado de conservao e informao sobre invasoras, bem como
discusses relativas taxonomia de aproximadamente 20.000 espcies. A previso final
de 30 volumes, sendo que, nos 18 j publicados e naquele que est no prelo, foram
tratados 2021 gneros e 12,393 espcies. Os restantes volumes contam com a submisso
de 763 gneros e 5.008 espcies, sendo que 82 das 144 famlias esto completas. A
finalizao da flora est prevista para 2017. O projeto gerenciado pela Flora of North
America Association. O contedo dos volumes publicados est disponibilizado atravs de e-
Floras e do JSTOR, e foi repassado para o time de informtica do projeto World Flora.

Palavras-chave: Flora; Estados Unidos da Amrica; Canad; Groenlndia

INTRODUCTION

The Flora of North America North of Mexico comprises treatments of vascular plants
and bryophytes native or naturalized in Canada, Greenland, St Pierre et Miquelon, and
the continental United States including the Florida Keys and Aleutian Islands
(approximately 18 million square kilometers). Distribution information for Mexico,
especially the northern tier of states, is provided, as well as generalized world-wide
distribution. Treatments include accepted name, basionym, place of publication,
synonyms, vernacular names, morphological descriptions, flowering time, habitat,
elevation, geographical distribution, discussions, and references. Every genus and 1/6
of the species are illustrated by professional botanical artists working closely with
authors and editors, assuring that the drawings reflect the author's concepts and
complement the technical descriptions in a way that assists the general user. Small
distribution maps are provided for all species and infraspecific taxa.

A total of 30 volumes will be published including an introductory volume and a final

volume containing a cummulative index and bibliography. As of June, 2015, eighteen
volumes had been published and one was in press (Tab. 1).

Table 1 Statistics from published volumes of Flora of North America north of Mexico.

Date Gener Specie

Volume Contents
published a s
Volume 1 1993 Introductory chapters
Ferns and fern allies;
Volume 2 1993 97 553
Gymnosperms
Magnoliaceae to
Volume 3 1997 128 741
Casuarinaceae
Phytolaccaceae to
Volume 4 2003 117 652
Molluginaceae
Caryophyllaceae to
Volume 5 2005 75 739
Plumbaginaceae
Cucurbitaceae to
Volume 6 in press 104 545
Droseraceae
Volume 7 2010 Salicaceae to Brassicaceae 125 923
Volume 8 2009 Paeoniaceae to Ericaceae 125 682
Volume 9 2014 Picramniaceae to Rosaceae 74 691
Volumes 19, 20,
2006 Asteraceae 418 2413
21
Volume 22 2000 Butomaceae to Marantaceae 89 423
Volume 23 2002 Cyperaceae 27 843
Volume 24 2007 Poaceae part I 113 695
Volume 25 2003 Poaceae part II 123 728
 Date Gener Specie
Volume Contents
published a s
Pontederiaceae to
Volume 26 2002 177 908
Orchidaceae
Volume 27 2007 Bryophytes: Mosses part I 127 704
Volume 28 2014 Bryophytes: Mosses part II 206 698
Totals 2021 12393

Content of all published volumes is made available electronically through eFloras and
JSTOR Plant Science. Preparation of all remaining volumes is projected to be
completed by the end of 2017.

Brief history of Flora of North America north of Mexico project

The scientific community has recognized the need for a continental scale flora of North
America since the early 1800s. The first accounts of plants from the area were
published in Europe by European botanists (e.g., in Robert Morison's Historiae [1680-
1699], Leonard Plukenet's Phytographia [1691-1705], and John Ray's Historia
Plantarum [1686-1704]). Fredrick Pursh's Flora Americae Septentrionalis (1814) was the first flora of
continental scope. Treatises on North American plants were written by resident
botanists in the early 1800s, and the first attempt at production of a comprehensive
flora of the continent was undertaken by John Torrey and Asa Gray the 1830s (see Reveal
& Pringle 1993
). Subsequently, botanists focused on producing regional floras, until 1905
when N.L. Britton began publishing the monographic series North American Flora at
the New York Botanical Garden, with multiple authors. The first effort at a
multiinstitutional, collaborative project started in 1966, stimulated by the completion of
the Flora SSSR in 1963 (Komarov et al. 1963) and publication of the first volume of Flora
Europaea in 1964 (Tutin et al. 1964). The project was to be headquartered at the
Smithsonian Institution and funded ultimately through a congressional appropriation. A
great deal of organizational work was done and information resources were developed
but the project was suspended in 1972 because the funding plan was not successful.
Various other attempts were made to get the project restarted, and in 1982 a group of
botanists met in St. Louis and decided to try again. A plan was developed, an editorial
committee formed, and in 1987 the first funding was awarded by the Pew Charitable
Trusts and the National Science Foundation. Until 1996, the Missouri Botanical Garden
was the organizational center through which the project was funded, after which the
Flora of North America Association was incorporated and funding, along with the work
force, was distributed among Editorial Centers at various institutions.

Underlying philosophy of Flora of North America north of Mexico

The mission of the Flora of North America Association (FNAA) is to make the best
scientific knowledge of the plants of North America north of Mexico publicly available in
consistent and easy to use forms.

To this end, FNAA coordinates the work of 900+ botanists to provide authoritative
information on the names, circumscriptions, characteristics, and geographical and
ecological distributions of the 20,000 species of plants native or naturalized in North
America north of Mexico.
The Flora of North America north of Mexico (FNANM) editorial committee developed
policies and procedures for its completion based on recommendations made by a wide
range of interested parties at a symposium titled "Floristics for the 21 st Century" (Morin et
al. 1989
). A guide for contributors was developed setting out style and format
expectations, including that descriptions within a rank should be parallel to the extent
possible and the descriptions should match the relevant entries in the identification
key. The symposium participants recommended parallel descriptions so that FNANM
would be more than a device to identify unknown plants; it should characterize the
flora in a way useful to a wide range of audiences. Experts in each taxonomic group,
when available, would be asked to write treatments of their groups in order to assure
that the most authoritative, up-to-date information and taxonomic concepts were
employed. Scientific reliability was deemed essential if the information was to inform
management and conservation decisions. Regional reviewers (now numbering about
100) would review treatments to assure that each taxon was treated appropriately
throughout its range. The editorial committee concluded that decisions should be made
by the people doing the work and that authors and editors would mostly work on a
volunteer basis.

Content

A "flora" may be a checklist, an identification key to included taxa, a list of accepted

species with distributions, a list of accepted names with brief descriptions, literature
citations, synonyms, some with maps, some with illustrations, or various combinations
of these components.

Initially, the expectation was that the accounts in FNANM would be similar to those in
Flora Europaea, with keys, short descriptions, minimum discussion, and no maps or
illustrations, and that the entire series would comprise 14 volumes. After
consideration, small distribution maps were added to help give users a quick visual
impression of the geographical range of a taxon. The geographical area, continental
North America north of Mexico, the Florida Keys, Aleutian Islands, St. Pierre et
Miquellon, and Greenland, was selected because it forms a reasonable
phytogeographical region. Mexico was not included because there were other efforts to
treat that flora (regional floras, Flora Mesoamericana, Davidse et al. 1994), but distribution
statements in FNANM include at least the northern tier of Mexican states when
possible. Illustrations, initially for every genus and one-third of the species, were
added to make the treatments more useful to non-botanists, and to show complex
features difficult to portray with just words. The length of descriptions was expected to
be 700 characters or less, but, for larger families and genera, the descriptions must be
longer in order to make descriptions parallel and to assure that they contain features
used in the key. Many of the treatments represent a botanist's lifetime of work, work
that might otherwise never be published elsewhere. The editorial team has attempted
to balance realistic constraints of time and resources (including a goal of completing
the project within their lifetime, and physical limitations of number of pages that can
be bound in one volume) with a wish to convey in the best possible way this wealth of
knowledge.

Taxonomic Arrangement

When decisions were first being made about the content of Flora of North America
volumes, the most comprehensive published modern classification system for plants
was by Arthur Cronquist (1981), and that work was adopted to set the circumscriptions of
families and the order in which they would appear in the volumes. Beginning in 2007,
Flora of North America adopted the system outlined by the Angiosperm Phylogeny Group (2003) for
remaining volumes, to the extent practicable.

Authors are encouraged to order genera and species according to their best
understanding of relationships. If this is not possible, the taxa may be arranged
alphabetically.

Community and financial support

In-kind support in the form of facilities and staff time has been essential for the
success of the project. Editorial centers are or have been located at the The Arboretum
at Flagstaff, Arizona (Volume 4), Botanical Research Institute of Texas, Fort Worth
(Asteraceae, Volumes 19, 20, and 21), California Academy of Sciences, San Francisco
(Volume 18 - now moving), Hunt

Institute for Botanical Documentation, Pittsburgh (bibliography and Volumes 5, 6, and

14, 26, 30), Harvard University Herbaria (nomenclature), Illinois Natural History
Survey, Champaign together with Canadian Museum of Nature (Volume 12), University
of Kansas together with University of Michigan (Volumes 5, 8, 17), Missouri Botanical
Garden (Volumes 1, 2, 3, 7, 10/11, 23, and the bryophyte volumes, 27, 28, 29),
University of Montreal (Volumes 9, 13), University of Washington (Volume 15), and
Utah State University (Poaceae, Volumes 24 and 25). Most editors, authors, and
reviewers donate their time and expertise. Botanical artists and technical editors work
on a contract basis and are coordinated by the Managing Editor and Scientific Director
located at Missouri Botanical Garden. When funding has been available, a botanist has
been employed on contract to write treatments for which no specialist is available.
Botanical artists and technical editors, the Managing Editor, and half of the Scientific
Director's time is paid by grants, contributions, art sponsorships, and earned income
(royalties, etc.). Also funded are basic organizational costs such as insurance, financial
audits, meetings, etc.

The botanical community has supported work on FNANM by developing cooperative

groups to agree on taxonomic approaches and engaging the efforts of specialists. For
example, Volumes 24 and 25, which treat Poaceae, were prepared in collaboration with
a large group of grass specialists working through Utah State University and funded in
part by the U.S. Department of Agriculture. Volumes 19, 20, and 21, treating
Asteraceae, were prepared under the guidance of a team of noted synantherologists
coordinated at the Botanical Research Institute of Texas and funded by a grant from
the U. S. National Science Foundation. Volumes 27, 28, and 29, Bryophytes, are being
completed by a team of bryologists.

The initial funding for the current Flora of North America project from the Pew
Charitable Trusts was matched by a grant from the U. S. National Science Foundation,
which made additional grants through 1999. A wide range of foundations made grants
to the project from 1987 through 1999. The Chanticleer Foundation provided core
funding 2000-2008. The Andrew W. Mellon Foundation and an anonymous foundation
also provided core funding in 2008 through 2012. Funding since 2012 has been
received from the Philecology Foundation, the Franklinia Foundation, the David and
Lucile Packard Foundation and other organizations and individuals.

Information Technology
The original Flora North America project was far ahead of its time in its vision of
management and analysis of botanical data using computers. The 1967 proposal to
NSF outlined goals of developing an information retrieval system and the ability to
automatically update information continuously and retrieve it in newly combined forms.
At the time, research indices like Index Kewensis were compiled by hand. The FNA
organizers hoped they could provide the initial stimulus and technical foundation for
development of comprehensive information retrieval systems.

The current Flora of North America organizers decided to focus their efforts on
acquiring the botanical treatments rather than developing information management
tools. Nonetheless, the potential for using computers to assist with preparation of the
flora was kept in mind. (Before the World Wide Web became publicly available in 1990,
all manuscripts were duplicated and distributed using the U. S. Postal Service.)
TROPICOS, a system developed by Missouri Botanical Garden botanists Robert Magill
and Marshall Crosby in the early 1980s to manage plant nomenclature and
bibliography, was adopted as the system in which FNA nomenclature would be
maintained. A bibliographic database for FNA was developed by Robert Kiger at the
Hunt Institute for Botanical Documentation. Grants from the U. S. National Science
Foundation supported development of innovative tools for collaborative work
processes, envisioning the potential for multiple participants to work on the same
manuscript and to monitor tasks and accomplishments. An initial goal of creating a
massive relational database of characteristics has evolved into the use of semantic
markup tools developed by James Macklin and Hong Cui (Cui 2010; Morris et al.2013). Treatments
parsed into their basic components are available through eFloras
(<http://www.efloras.org>) and JSTOR (<http://www.plants.jstor.org>). Resource
materials for authors and reviewers, links to published volumes, status of volumes in
preparation, and other information about the FNANM project, are available on the FNA
website: www.floranorthamerica.org. In the past year the FNA website received about
1,371,540 successful requests for 377,356 pages - on average 26,000 successful
requests per week.

Progress to date

Flora of North America now has published 18 of the total 30 volumes and has one
volume in press (Volumes 1-9, 19-28; 11,200 pages), containing 253 families, 2,125
genera, and 12,949 species (Flora of North America Editorial Committee, 1993+).
These volumes also contain original illustrations of 7,200 species, subspecies, or
varieties and maps of every species and infraspecific taxon. Volume 1, published in
1993, includes introductory chapters on the history of floristics and plant collecting in
North America; climate and physiography; soils; history of the vegetation and
paleoclimates, vegetation, and floras; vegetation, and phytogeography.

Completing Flora of North America north of Mexico by 2020

The taxonomic volumes remaining to be completed: Volumes 10-18 and 29, contain
144 families, 1103 genera, and 7365 species. Of these, 763 genera and 5008 species
have been submitted, and 82 of the 144 families have been submitted in full. Volume
12, Vitaceae to Garryaceae, 29 families, 122 genera, and 760 species, is in final stages
of editing and will be in press this fall. Volume 17, Polypremaceae to Orobanchaceae,
containing 9 families, 94 genera, and 942 species, will be in press by the end of 2015
or early 2016. For Volumes 10 and 11, being prepared together and containing largely
Fabaceae and Onagraceae, all but those two families have been submitted in full and
91% of the genera and species are in hand and in process of review, revision, and
editing. For the remaining five volumes containing treatments of vascular plants, more
than half of the families and genera have been submitted and are being edited. All
other treatments have authors assigned and are in preparation. The final Bryophyte
volume, containing liverworts and hornworts, is also making good progress. Status of
individual treatments in volumes to be published can be found at www.
floranorthamerica.org.

METHODOLOGY

Each volume is assigned to an editorial center under the leadership of a lead editor (or
co-lead editors). Each family is assigned to a taxon editor who works directly with
authors, reviewers, and technical editors. Authors are selected based on their interest
and expertise and with approval of the FNAA Board. Technical editors work on a
contract basis; each one is assigned to a volume. Regional review is managed by
regional coordinators, who send draft treatments to reviewers knowledgeable in their
local flora and compile responses, which are returned to taxon editors. Authors provide
material to botanical artists so they can prepare illustrations. Every genus and about
one in six species are illustrated by these contract artists. Authors are asked to use
terminology set out in (Kiger & Porter 2001). Names accepted in major regional floras are
supplied to authors and are to be accounted for in the treatments. Robert Kiger, Hunt
Institute for Documentation, edits all bibliographic references and compiles the
cumulative bibliography. Kanchi Gandhi, Harvard University Herbaria, reviews
nomenclature and citations. Some aspects of final production, such as indexing and
page composition, are carried out by contract workers under the supervision of Heidi
Schmidt, managing editor, and James Zarucchi, editorial director, who also oversee all
manuscript flow.

Challenges for completion include editorial capacity and time available to volunteer
authors and reviewers. Botanists with broad floristic knowledge are becoming scarce,
and there are plant groups for which no specialist is available. Fortunately, FNANM has
been able to enlist the help of botanists who have recently completed their graduate
degrees or are currently graduate students.

Resources for completing the Flora of North America north of Mexico

1. Regional Floras

The Flora of North America project draws on a wealth of resources. A working checklist
has been available from several sources, including the United States Department of
Agriculture PLANTS database and The Natural Heritage Program through its
Natureserve database. Canada has a working checklist through VASCAN, part of
Canadensys.

North America north of Mexico is a very large geographic area, politically and
biogeographically diverse, so it is understandable that floristic work has focused on
states, provinces, or regions. Regional floras recently completed for the United

States include The Jepson Manual, covering all of California (Baldwin et al. 2012); the
Intermountain Flora, covering all of the Great Basin (Cronquist et al. 1972); Flora of the
Southern and MidAtlantic States (Weakley 2012); and Flora Novae Angliae (Haines 2011). Less
recent but still important are Flora of the Great Plains (Great Plains Flora Association 1986); Manual
of the Vascular Plants of Texas (Correll & Johnston 1970); and Manual of vascular plants of
northeastern United States and adjacent Canada, second edition (Gleason & Cronquist 1991).

For Canada, a basic reference is the Flora of Canada (Scoggan 1978-1979); Budd's Flora of the Canadian
Provinces (Budd et al. 1987
); The vascular plants of British Columbia (Douglas et al. 1989-1994);
Vascular plants of the continental Northwest Territories (Porsild & Cody 1980); Flore
Laurentienne (Marie-Victorin, revised by Brouillet et al. 1995); Atlas of the vascular plants of the island of
Newfoundland and of Saint Pierre et Miquelon (Rouleau and Lamoureux 1992); and The flora of
New Brunswick (Hinds 2000). In preparation is a Flora of the vascular plants of the
Canadian Arctic Archipelago, updating the Illustrated Flora of the Canadian Arcitc
Archipelago (Porsild 1964).

2. Online specimen, occurrence, and nomenclatural databases

Some regional or state specimen databases also functionally serve as checklists. They
are a rich resource for authors and reviewers. As Flora of North America volumes are
published, the databases are updated to reflect new classifications. Examples of such
databases are: Country-wide Databases

2.1 Canadensys (<http://data.canadensys.net/explorer/en/search>)

The checklist covers all vascular plants reported in Canada, Greenland (Denmark), and
Saint Pierre and Miquelon (France). The latter two regions are added because their
floras are intimately related to that of Canada. Provincial distributions are provided to
help users visualize the relationship among the floras of Canadian provinces and
territories. VASCAN does not intend to replace regional or provincial lists but to act as
a complement to them. The covered regions are, in alphabetical order: Alberta, British
Columbia, Greenland, Labrador, Manitoba, New Brunswick, Newfoundland, Northwest
Territories, Nova Scotia, Nunavut, Ontario, Prince Edward Island, Quebec, Saint Pierre
and Miquelon, Saskatchewan, and Yukon. The distribution status of the plant is
indicated per region. These can be grouped as present (native, introduced or
ephemeral), previously reported but currently considered absent (excluded,
extirpated), doubtful or not reported (absent). The latter status is not recorded in the
database (null value). Excluded taxa are those considered not currently occurring in a
region, due either to nonrecurring ephemeralness, misidentification, lack of supporting
documentation, or when specimens are old and the taxon has not been observed again
in more than 50 years. All distribution statuses are defined at
(<http://data.canadensys.net/vascan/ about/#distribution>). The VASCAN website
(<http://data.canadensys.net/vascan>) provides a distribution map for each taxon.

2.2 The PLANTS Database (<http://plants.usda. gov>)

Provides standardized information about the vascular plants, mosses, liverworts,

hornworts, and lichens of the U.S. and its territories. It includes names, plant symbols,
checklists, distributional data, species abstracts, characteristics, images, crop
information, automated tools, onward Web links, and references. It can be searched by
taxonomy, morphological and habitat characteristics, geographical distribution, and
weed, wetland, and conservation status.

3. Specimen Databases available online that also can generate checklists

Most of these online databases are collaborations among regional networks of herbaria
and provide links to the individual member organizations.

3.1 Canadensys Explorer (<http://data. canadensys.net/explorer>)

Serves 180,000 vascular plant specimen records from 13 Canadian herbaria.

3.2 Consortium of California Herbaria (<http://

ucjeps.berkeley.edu/consortium>)

The Consortium of California Herbaria provides data from over 2 million specimen
records from over 30 institutions.

3.3 Consortium of Pacific Northwest Herbaria

(<http://www.pnwherbaria.org>)

The Consortium of Pacific Northwest Herbaria hosts more than 2.4 million specimen
records from 33 herbaria in Alaska, Idaho, Montana, Oregon, Washington, Wyoming,
British Columbia, Yukon Territory, and Humboldt State University in northern
California.

3.4 Intermountain Regional Herbarium Network

(<http://intermountainbiota.org/portal/index.php>)

The Intermountain Regional herbarium Network hosts collection data from herbaria in
Nevada and Utah. (also served throught SEINET:
(<http://swbiodiversity.org/seinet/index.php>)).

3.5 Southwest Environmental Information Network

(<http://swbiodiversity.org/seinet/index.php>)

The Arizona Chapter of SEINET serves collections and observation data from 14
institutions in Arizona. SEINet allows searching across many major regional herbarium
networks.

3.5 Flora of Texas Database (<http://w3.biosci. utexas.edu/prc/Tex.html>)

Serves records of occurrences in Texas based on University of Texas and Lundell

herbaria.

3.7 Rocky Mountain Regional Consortium Specimen Database

(<http://www.rmh.uwyo.edu>)

Provides searching across 14 herbaria in Colorado, Idaho, Montana, and Wyoming

(also served throught SEINET: (<http://swbiodiversity. org/seinet/index.php>)).

3.8 Consortium of Midwest Herbaria (<http:// midwestherbaria.org/portal>)

Hosts specimen data from 21 herbaria in Illinois, Indiana, Iowa, Michigan, Minnesota,
Ohio, and Wisconsin. (also served through SEINET:
(<http://swbiodiversity.org/seinet/index.php>)).
3.9 Northern Great Plains Herbaria (<http:// www.ngpherbaria.org>)

Hosts data from 19 herbaria in Illinois, Iowa, Kansas, Minnesota, Missouri, Nebraska,
North Dakota, and South Dakota, (also served through SEINET:
(<http://swbiodiversity.org/seinet/index. php>)).

3.10 Consortium of Northeastern Herbaria (<http://portal.neherbaria.org>)

Serves 776,600 records of vascular plants and bryophytes from 21 herbaria in Nova
Scotia, Quebec; Connecticut, Massachusetts, New Hampshire, New Jersey, New York,
Pennsylvania, Rhode Island, and Vermont.

3.11 Southeast Regional Network of Expertise and Collections

(<http://sernec.appstate.edu>)

Hosts records from 110 herbaria in Alabama, Arkansas, Delaware, Florida, Georgia,
Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee,
Texas, Verginia, and West Virginia (also served through SEINET:
(<http://swbiodiversity.org/seinet/index.php>)).

Atlas of Florida vascular Plants (<http:// www.florida.plantatlas.usf.edu>)

Serves records from over 110,000 herbarium specimens from 8 herbaria in Florida.

CONCLUSION

Improving the knowledge of the Flora of North America north of Mexico

Flora of North America north of Mexico has stimulated the completion and publication
of research by authors of treatments and presents the best, most current
understanding of the relationships, characteristics, and distribution of plants in the
flora. It brings together in a standard, carefully reviewed form, the results of lifetimes
of study and the findings of the most recent research. Aspects of phylogeny, biology,
or ecology that need more study are mentioned in the discussion, which stimulates
new research. Because the descriptions and discussion characterize the plants in more
detail than more abbreviated floras or checklist, they provide a rich resource for
research by ecologists, plant breeders, plant morphologists, physiologists, to name just
a few. This resource will be much enhanced by the current effort to parse all
descriptions, distributions, and habitats from published volumes.

Importance of Flora of North America for Environmental Research

Flora of North America provides a common reference for the circumscription of plant
taxa, correct application of names to those taxa, and descriptive information for each
taxon. This is critically important for every kind of research from broad ecological
studies to phenomics, genomics, and phylogenetic analysis. Although there are
checklists of the plants of North America already available, none have the same level
of scientific authority, and, since they lack descriptions, they are not sufficient to
determine what is actually included within the named concept. There are few
equivalents in terms of digital datasets in the animal or insect worlds, and none dealing
with as many species.

Importance of the World Flora

It would be highly useful for further research, for conservation, and for resource
management to have a common reference for the circumscription of plant taxa, correct
application of names to those taxa, and descriptive information for each taxon, and
worldwide distribution for all plants. Only elements of this, in various guises, are so far
available. For Flora of North America north of Mexico, based on volumes published to
date, 13% of the genera and 50% of the species are endemic to the flora area. FNANM
has provided full descriptions and abbreviated worldwide distributions for 6500 species
that also occur outside the flora area and for nearly 1900 genera that also occur
outside the flora area. Of these, 315 of the genera and 1791 of the species in
published volumes are introduced. Currently it is very difficult for authors to find the
relevant data on the non-native taxa, as it must be for researchers outside North
America to find data on the 10,000 species likely to be endemic in North America north
of Mexico, since much of that information is contained in local floras or individual
monographs. A World Flora would facilitate access to these data.

Integrating Flora of North America north of Mexico data with the World Flora

All published treatments are available in electronic form, and soon all will be fully
parsed into individual components. The electronic files for volumes published have
already been provided to the World Flora Online for incorporation.

REFERENCES

Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group

classification for the orders and families of flowering plants: APG II. Botanical Journal
of the Linnean Society 141: 399-436. [ Links ]

Baldwin, B.G.; Goldman, D.; Keil, D.J.; Patterson, R.; Rosatti, T.J. & Wilken, D. (eds.).
2012. The Jepson manual: vascular plants of California, revised and expanded.
University of California Press, Berkeley. 1568p. [ Links ]

Britton, N.L. et al (eds.). 1905. North American Flora.47 vols. New York Botanical
Garden, New York. Vols. 1-34, 1905-1957; ser. 2, parts 1-13, 1954. [ Links ]

Budd, A.C.; Looman, J. & Best, K.F. 1987. Budd's flora of the Canadian Prairie
Provinces, revised and enlarged. Publication 1662. Agriculture Canada Research
Branch, Ottawa and Hull. 863p. [ Links ]

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Received: July 01, 2015; Accepted: September 14, 2015

FNA Vol. 28 Page 477

FNA | Family List | FNA Vol. 28

65. Fabroniaceae Schimper
Terry T. McIntosh
Plants very small, often in silky mats, green, yellow-green, or gray-green. Stems often erect-ascending, sparsely and irregularly bran
radiculose in scattered tufts. Leaves ovate to oblong-lanceolate, rarely lanceolate, often somewhat concave; margins plane, denticul
acuminate, or long-acuminate, subulate to piliferous, rarely blunt; costa single, 1/3-2/3 leaf length, terminal spine sometimes present,
cells rounded-rhomboidal to rhomboidal, quadrate to transversely rectangular in few rows along margins; medial and distal cells rhom
smooth; apical cells linear. Specialized asexual reproductionapparently by foliose pseudoparaphyllia. Sexual condition autoicous
short, wrinkled when old, mouth often flared with age; exothecial cells isodiametric, walls sinuate, suboral cells transversely elonga
absent; operculum conic, umbonate to mammillate; peristome single, rarely absent, erect-spreading, sometimes recurved when dry, i
pairs, red-brown, broadly lanceolate, blunt, 0.3 mm, densely papillose-striolate. Calyptra cucullate. Spores 9-14 m, coarsely papillo
Genera 5, species ca. 17 (1 genus, 2 species in the flora): North America, Mexico, West Indies, Central America, South America, Eur
Zealand), Australia.
Lower Taxon
Fabronia Raddi

FNA Vol. 28 Page 477

FNA | Family List | FNA Vol. 28 | Fabroniaceae

1. Fabronia Raddi, Atti Accad. Sci. Siena. 9: 231, plate 1. 1808. [For Giovanni Valentino Mattia Fabbroni, 1752-1822, Italian natural
Stems prostrate. Leaves loosely appressed when dry, spreading when moist. Capsule ovoid to pyriform, [0.3-]0.5-0.8[-1] mm.
Species ca. 11 (2 in the flora): North America, Mexico, West Indies, Central America, South America, Eurasia, Africa, Atlantic Islands,
Fabronia is characterized by its very small size, ovate to oblong-lanceolate, long-acuminate, usually toothed leaves tipped with linear
However, because of the wide degree of morphological variability, especially within the F. ciliaris complex, Fabronia is taxonomically
Anderson (1981) and W. R. Buck (1994b) are enlightening with respect to taxonomic problems.

1 Leaf apices acute or acuminate; margins dentate or sometimes entire, not 1 Fabronia ciliaris
ciliate, teeth of 1 cell.
+ Leaf apices acute to long-acuminate; margins ciliate-dentate, teeth often of 2 Fabronia pusilla
more than 1 cell.

Lower Taxa
Fabronia ciliaris (Bridel) Bridel
Fabronia pusilla Raddi
2. Fabronia pusilla Raddi, Atti Accad. Sci. Siena. 9: 231, plate 1. 1808.
Leaves 0.4-0.8 mm; margins ciliate-dentate, teeth often of more than 1 cell; apex acute to long-acuminate;
medial laminal cells 30-45 9-12 m.
Capsules mature late spring- summer. Rock, bark at base of trees; low to high elevations; B.C.; Ariz., Calif.,
Colo., Idaho, Oreg., Wash.; Mexico (Baja California Sur); Europe; n Africa.
Fabronia pusilla is easily distinguished by its long-acuminate and ciliate-dentate leaves.
1. Fabronia ciliaris (Bridel) Bridel, Bryol. Univ. 2: 171. 1827.
Hypnum ciliare Bridel, Muscol. Recent., suppl. 2: 155. 1812; Fabronia ciliaris var. polycarpa (Hooker) W. R.
Buck; F. ciliaris var. wrightii (Sullivant) W. R. Buck; F. ravenelii Sullivant; F. wrightii Sullivant
Leaves 0.4-0.9 mm; margins dentate or sometimes entire, not ciliate, teeth of 1 cell; apex acute or acuminate;
medial laminal cells 30-45 11-12 m.
Capsules mature summer. Bark of trees, rock; low to high elevations; Ariz., Ark., Calif., Colo., Ga., Ind., Kans.,
Ky., La., Mich., Minn., Mo., N.J., N.Mex., N.C., Ohio, Okla., Oreg., Pa., S.C., S.Dak., Tenn., Tex., Va., Wis.;
Mexico; West Indies; Central America (Guatemala); South America; Europe; e Asia (Japan); Pacific Islands
(Hawaii, New Zealand); Australia.
This treatment of Fabronia ciliaris generally follows that of W. R. Buck (1994b). Although the varieties
recognized by Buck are not recognized here, his varietal keys and associated discussions are very useful. That
work underlines the high degree of morphological variability within this species. Although many species and
varieties have been described within the concept of F. ciliaris, examination of the variation among and within
populations, especially with respect to the degree of leaf margin dentition, does not support the acceptance of
most of these taxa.

Colombia forestal, Vol. 19, Nm. 1 (2016)

Olascuaga-Vargas, D., Mercado-Gmez, J. & Snchez-Montao,
L. (2016). Anlisis de la vegetacin sucesional en un fragmento de bosque
seco tropical en Toluviejo-Sucre (Colombia). Colombia Forestal, 19(1), 41-
52.

DOI: http://dx.doi.org/10.14483/udistrital.jour.colomb.for.2016.1.a03

Artculo de investigacin
 DIVERSIDAD DE BRIFITOS EN LOS MONTES DE
MARA, COLOS (SUCRE, COLOMBIA)
Bryophyte diversity in the Montes de Mara, Colos (Sucre, Colombia)
Stevens Garca Martnez1, Hermes De Jess Basilio Banqueth2, Fran Yair Herazo Vitola3, Jorge David
Mercado Gmez4 & Mara Eugenia Morales Puentes5

1
Universidad de Sucre. Direccin: Carrera 28 No. 5-267 Barrio Puerta Roja. Sincelejo,
Colombia. stevens.garcia@unisucre.edu.co. Autor para corresondencia.
2
Universidad de Sucre. Direccin: Carrera 28 No. 5-267 Barrio Puerta Roja. Sincelejo,
Colombia. basilio_05@hotmail.com
3
Universidad de Sucre. Direccin: Carrera 28 No. 5-267 Barrio Puerta Roja. Sincelejo,
Colombia. fran.herazo@unisucre.edu.co.
4
Universidad de Sucre. Direccin: Carrera 28 No. 5-267 Barrio Puerta Roja. Sincelejo,
Colombia. jorge.mercado@unisucre.edu.co
5
Universidad Pedaggica y Tecnolgica de Colombia. Direccin: Av. Central del Norte, Edificio Centro de
Laboratorios. Tunja, Colombia. maria.morales@uptc.edu.co

Recepcin: 30 de junio de 2015 / Aprobacin: 24 de agosto de 2015

RESUMEN

Se determin la diversidad (alfa y beta) y la distribucin en los sustratos de los

brifitos refilos en tres quebradas (localidades de Salto del Sereno, Pajarito y Paraso)
ubicadas en los Montes de Mara (Sucre-Colombia). Se realizaron dos transectos de 50
X 2 m en cada sitio de muestreo en poca de lluvia. La clase Bryopsida present 22
especies y la Hepaticopsida nueve. Las familias de musgos con mayor nmero de
especies fueron Bryaceae (3 especies), Neckeraceae (2), Pottiaceae (2) y en hepticas
Lejeuneaceae (6), y los gneros ms ricos fueron Fissidens (5
especies), Fabronia (2), Lejeunea (3) y Mastigolejeunea (2). El tipo de sustrato con
mayor incidencia de brifitos fue el epiltico (29 especies), seguido de epifito-cortcola
(16) y terrestre (13). Las 31 especies aqu registradas son nuevos registros para el
departamento, 24 para la regin Caribe y una (Hyophiladelphus agrarius) para
Colombia.

Palabras clave: bosque seco tropical, brifitos, diversidad, sustrato, Montes de Mara.

ABSTRACT

Alpha and beta diversity and the distribution of substrates of rheophilous bryophytes in
three streams were determinate (Salto del Sereno, Pajarito and Paraso localities),
located in Montes de Mara (Sucre, Colombia). Two transects of 50 x 2m were
conducted at each sampling site during the rainy season. The class Bryopsida
presented 22 species and Hepaticopsida presented 9. The families of mosses with the
largest number of species were Bryaceae (3 species), Neckeraceae (2), Pottiaceae (2).
In liverworts, Lejeuneaceae (6), and the richest genera were Fissidens (5 species),
Fabronia (2), Lejeunea (3) and Mastigolejeunea (2). The type of substrate with
increased incidence of bryophytes was the epilithic (29 species), followed by epiphyte-
corticolous (16) and terrestrial (13). The 31 species registered here, are new records
for the department, 24 for the Caribbean region and one for Colombia
(Hyophiladelphus agrarius).

Key words: tropical dry forest, bryophytes, diversity, substratum, Montes de Mara.

INTRODUCCIN

En Colombia, los estudios sobre la diversidad, distribucin y ecologa de la brioflora

(musgos y hepticas) en diferentes regiones geogrficas se han incrementado
considerablemente en los ltimos aos (Sastre et al., 1986; Parra et al., 2002; Ramrez & Churchill,
2002; Vasco et al., 2002; Pinzn et al., 2003; Pinzn & Linares, 2006; lvaro et al., 2007; Avendao &
Aguirre, 2007; Barbosa et al., 2007; Lagos et al., 2008; Santos & Aguirre, 2010). Sin embargo, estos
estudios se han enfocado principalmente en las regiones de vida andina, paramuna y
amaznica, dejando a un lado las tierras bajas de las llanuras, valles interandinos y el
Caribe colombiano, donde solo se han realizado los estudios de Sipman (1984), Van
Reenen et al. (1984), Aguirre & Ruiz (2001), Ruiz & Aguirre (2003), Avendao & Aguirre (2007), Aguirre &
Avendao (2008a, b), Avendao & Aguirre (2009a, b), Santos & Aguirre (2010), entre otros;
tales estudios describen aspectos ecolgicos y taxonmicos de brifitos y lquenes en
las zonas montaosas de la Sierra Nevada de Santa Marta (Sipman, 1984; Van Reenen et al.,
1984) y la Serrana del Perij (Aguirre & Ruiz, 2001; Ruiz & Aguirre, 2003; Aguirre &
Avendao, 2008a, 2008b; Avendao & Aguirre, 2009a, 2009b).

No obstante, en localidades consideradas como bosque seco son pocos los trabajos que
analizan la diversidad de este grupo de plantas, por lo que en la actualidad se
desconoce el nmero de especies de briofitos que alimentan la diversidad del bosque
seco. Teniendo en cuenta que este grupo de organismos necesitan de una alta
concentracin de agua para su supervivencia, se ha conllevado a que diferentes
autores estimen la diversidad en este tipo de ecosistemas como muy baja para el pas
(Uribe & Gradstein, 1999; Aguirre & Avendao, 2008a). Sin embargo, el departamento de Sucre
que se ubica sobre la subregin Montes de Mara, posee un paisaje constituido por
colinas onduladas, cubiertas parcialmente por bancos de niebla durante gran parte del
ao (en las horas de la maana y al atardecer, debido al efecto lluvia generado por la
humedad proveniente del golfo); estas caractersticas permiten sugerir condiciones
ambientales necesarias para una alta diversidad de brifitos, por lo tanto, realizar un
anlisis sobre aspectos ecolgicos de estos taxones en el departamento incrementara
nuestro conocimiento sobre los brifitos en las zonas bajas del pas.

Este trabajo es el inicio de una serie de estudios de los brifitos de las zonas secas del
Caribe y valles interandinos. Es as como en este caso, se realiz un anlisis de
diversidad en las comunidades de brifitos refilos y su distribucin por sustrato en el
rea de estudio.

MATERIALES Y MTODOS

rea de estudio
Los Montes de Mara, Serrana de Coraza, se localizan al norte del departamento de
Sucre a los 935 N y 75 22 E (Galvn & De La Ossa, 2009), constituidos por una franja de
colinas, entre los 0-1000 m, temperatura entre 25-28C, precipitacin de 896 a 1233
mm anual y la humedad relativa es de 83.5% anual (Aguilera, 2005; IGAC, 1969).
Segn Galvn et al. (2009), los Montes de Mara son una formacin de bosque seco tropical,
conformados por factores climticos de sequa, con suelos calcreos, y comprenden
entre 40-45% del rea del municipio de Colos (Figura 1).

Para determinar la diversidad alfa y beta de brifitos refilos en el municipio de Colos,

se escogieron tres puntos de muestreo: Salto del Sereno (Sitio1), Pajarito (Sitio 2) y
Paraso (Sitio 3). Estas localidades fueron escogidas por sus caractersticas
ambientales e importancia ecolgica; por ejemplo, en el sitio 1 se encuentra la
Estacin Ecolgica Primates, la cual es considerada como el centro de la Reserva
Coraza; y en los sitios 2 y 3 se encuentran quebradas como fuentes hdricas, que
abastecen de agua al municipio.

Mtodos

Para determinar la diversidad y la distribucin (sustrato) de los brifitos en los puntos

de muestreo, se llevaron a cabo dos transectos durante los meses de agosto,
septiembre y octubre (2013), dentro de los cuales, se increment considerablemente
la precipitacin. Cada transecto fue de 50 x 2 m, separados a una distancia 100 m en
lnea recta, entre cada uno para evitar sobre muestreos, teniendo en cuenta que en las
quebradas se generan meandros y sobre estos se realizaron recolectas de musgos y
hepticas; registrando informacin sobre localidad, coordenadas, elevacin, hora,
fecha, entre otros, desde el nivel del suelo hasta dos metros de altura del rbol. As
mismo, y con el fin de establecer la cobertura y abundancia se tomaron datos con una
plantilla cuadriculada de acetato transparente de 20 x 20 cm (modificado de Iwatsuki,
1960) sobre cada uno de los sustratos evaluados.

Posterior, a cada uno de los ejemplares recolectados, y con el fin de brindar

informacin ecolgica sobre los taxones, se les registraron datos de forma de
crecimiento, cobertura y tipo de sustrato, teniendo en cuenta la clasificacin para
sustrato propuesta por Aguirre (2008) en: epilticos, epifito-cortcolas, terrestres (suelos)
y materia orgnica en descomposicin (maordes).

La identificacin de las especies de musgos y hepticas se realiz a travs de claves

como las de Churchill & Linares (1995), Bello (1997), Uribe & Aguirre (1997) y Gradstein et al., (2001), y
con el apoyo del especialista colombiano, el profesor Edgar Linares (Universidad
Nacional de Colombia).

Por ltimo, para evaluar la diversidad alfa, se determin la riqueza de especies con el
ndice de Margalef y la estructura con los ndices de Simpson (dominancia) y Shannon-
Wiener (uniformidad). Tambin se evalu la diversidad beta (distribucin) con el uso
del ndice de Whittaker y la similaridad presente en los sitios de muestreo con el ndice
de Jaccard. Para estos ndices, se tom como referencia a Moreno (2001) y fueron
calculados por medio de una matriz de presencia/ausencia a travs del programa
PAST-Palaeontological Statistics ver. 3.06 (Hammer et al., 2001).

RESULTADOS

Se registran 31 especies distribuidas en 24 gneros y 18 familias, a partir de 144

levantamientos (Tabla 1).

Se encontraron 22 taxones de la clase Bryopsida y nueve de Hepaticopsida. Las

familias con mayor nmero de especies fueron: Bryaceae (3 especies), Neckeraceae
(2) y Pottiaceae (2); y los gneros con mayor nmero de especies son Fissidens (5)
y Fabronia (2). En Hepaticopsida, Lejeuneaceae con seis especies y los
gneros Lejeunea con tres especies y Mastigolejeunea con dos. Se resalta, que las 31
especies encontradas, corresponden a nuevos registros para el departamento de Sucre
24 para la regin Caribe y uno para Colombia (Hyophiladelphus agrarius).
Diversidad

En trminos de riqueza (ndice de Margalef), se establece que la diversidad de brifitos

(musgos y hepticas) est relacionada con su ambiente, siendo el Sitio 2 el ms rico
en familias, gneros y especies, en comparacin con los Sitios 1 y 3, en donde la
riqueza fue menor (Tabla 2). De modo similar, y segn el ndice de Shannon-Wiener, el
Sitio 2 fue el ms uniforme con un valor de 1.485, seguido del Sitio 3; mientras que el
Sitio 1 present menor valor; lo que otorga al Sitio 2, el ambiente ms propicio para el
crecimiento y desarrollo de los brifitos. Respecto a la dominancia (ndice de Simpson),
el Sitio 1 es el ms rico en especie y con un ndice de dominancia menor, en relacin
con el Sitio 2 y 3 ; por lo tanto, el Sitio 1 es el punto de muestreo que presenta menor
diversidad de especie (Tabla 2).

El ndice de Whittaker (diversidad beta), mostr que el Sitio 2 tiene el valor ms alto
de diversidad (0.4), debido al reemplazo de las especies encontradas en las
comunidades presentes en los transectos, y no se evidencia en los Sitios 1 y 3 (con los
valores ms bajos, 0.27 y 0.33 respectivamente), causado por la variacin de
especies. Finalmente, y segn los resultados obtenidos, las comparaciones por
similaridad (ndice de Jaccard) entre los puntos de muestreos (Figura 2), los Sitios 2 y 1,
tienen cinco especies en comn (Leucomium strumosum, Neckeropsis
undulata, Porotrichum substriatum, Splachnobryum obtusum y Lejeunea flava);
mientras que el sitio tres presenta tres especies compartidas con el Sitio 1 (Fissidens
mollis, Fissidens prionodes y Lejeunea maxonii) y una con el Sitio 2 (Lejeunea
trinitensis); y las especies en comn entre localidades fueron: Fabronia
ciliaris, Fissidens dissitifolius, Fissidens steerei, Hyophila involuta y Thuidium
tomentosum (Tabla 1).

Distribucin por sustrato

Los brifitos ms abundantes fueron los epilticos con 18 especies; 11 son epifito-
cortcolas, nueve terrestres y sobre la materia orgnica en descomposicin se
encontraron siete especies.

En trminos de familias, fueron encontradas 11 epilticas y ocho epifito-corticcolas,

siguen en orden, siete terrestres y seis sobre materia orgnica en descomposicin
(Figura 3). No obstante, al analizar la distribucin de los brifitos en cada uno de los
puntos de muestreo hallaron, por ejemplo, en el Sitio 1, 10 familias epilticas, siete
epifito-cortcolas, cuatro terrestres (suelo) y tres en materia orgnica en
descomposicin. Respecto a las epilticas fueron halladas Fissidentaceae (3 especies) y
Pottiaceae (2), las epifitas-cortcolas como Calymperaceae, Stereophyllaceae,
Fabroniaceae, Leucomiaceae, Neckeraceae y Thuidiaceae presentan una especie cada
una, mientras que en las terrestres (suelo), Fissidentaceae con dos especies y en
materia orgnica en descomposicin se registraron Leucomiaceae y Thuidiaceae (una
especie cada una). Por su parte, Lejeuneaceae fue la nica heptica que se encontr
en el suelo y maordes, y Plagiochilaceae en sustrato epifito-cortcola.

Las especies de T. tomentosum (19.44%), L. strumosum (16.67%) y F.

prionodes (13.89%) fueron los taxones epilticos ms frecuentes. Entre los epifito-
cortcolas, T. tomentosum con el 25.00% (musgos), en terrestres L. flava con 33.33%
(hepticas), H. involuta y P. substriatum con el 22.22% (musgos). Para terminar, T.
tomentosum sobre materia orgnica en descomposicin fue la ms abundante.

En el sitio dos, se registraron ocho familias epilticas como las ms abundantes, siete
en suelo, cinco epfitos-cortcolas y cuatro en materia orgnica en descomposicin.
Fissidentaceae con dos especies es la familia epiltica ms abundante, en el suelo
Bryaceae (2 especies), epfitas-cortcolas con una especie cada una y son:
Fabroniaceae, Neckeraceae, Sematophyllaceae y Thuidiaceae; en materia orgnica en
descomposicin Fabroniaceae, Leucomiaceae Pottiaceae y Thuidiaceae presentaron una
especie respectivamente. En hepticas, solo se registran Lejeuneaceae y
Marchantiaceae en los sustratos epiltico y suelo con una especie cada una (L.
flava y Marchanta chenopoda), para el epfito-cortcola se registra Lejeuneaceae con
cuatro especies (L. trinitensis, Mastigolejeunea auriculata, M.
Plicatiflora y Stictolejeunea squamata) y en materia orgnica en descomposicin no se
encontraron hepticas.

En musgos, las especies epilticas ms frecuentes fueron L.strumosum con 33.33%,

seguida de Fissidens steerei (musgos) y L. flava (hepticas) con 13.33% cada una,
mientras que en el suelo L. flava (35.71 %), S.obtusum (21.43%) y Rhodobryum sp.
(14.29%) fueron las ms representativas. Entre las epifto-cortcolas Neckeropsis
undulata ocupa el 29.41%, al igual que T. tomentosum, seguido de Mastigolejeunea
auriculata (11.76%); finalmente, en materia orgnica en descomposicin, se
registra Fabronia ciliaris (57.14%).
En el sitio tres, se registraron ocho familias epilticas y en materia orgnica en
descomposicin, as como, un taxn terrestre y uno epifto-cortcola. La familia epiltica
ms frecuente fue Fissidentaceae (4 especies) y entre las terrestres solo se registra
Pottiaceae; en materia orgnica en descomposicin, Fabroniaceae y Fissidentaceae
(2); musgos epiftos-cortcolas no se registraron y hepticas solo Lejeuneaceae como
epifito-cortcola y en maordes. Las especies epilticas ms frecuentes
fueron Hyophiladelphus agrarius (33.33%), F. dissitifolius (20%), F. steerei y T.
tomentosum (13.33%). En materia orgnica en descomposicin, Fabronia
ciliaris (38.46%), Lejeunea maxonii y T. tomentosum (23.08% cada una) y en el
suelo, H. involuta y epfito-cortcola Lejeunea trinitensis.

DISCUSIN

Las especies de musgos y hepticas registradas en el Bosque seco Tropical de los

Montes de Mara (Colos), son evidencia de la diversidad que caracteriza un pequeo
fragmento de bosque en el departamento de Sucre, y donde se han realizado
estimaciones de la diversidad, y as considerada como un rea de baja diversidad
(Churchill, 1989) para brifitos.

Los resultados obtenidos a partir de riqueza y estructura para cada sitio de estudio,
parece tener una explicacin en la resistencia de brifitos con adaptaciones
morfolgicas y fisiolgicas, frente a las condiciones generadas por la fragmentacin y
transformacin del bosque, debido a las acciones antrpicas (tala de rboles, la
ganadera y la agricultura).

El Sitio 2 present el mayor nmero de familias, gneros y especies, probablemente

por sus condiciones geomorfolgicas y ambientales al interior del bosque, ya que este
presenta una vegetacin de ladera y galera menos afectada por acciones humanas, la
presencia de meandros, condiciones favorables para el crecimiento y desarrollo de
brifitos (Galvn et al., 2009).

En esta localidad se encontraron familias de musgos, como Bryaceae, Fissidentaceae y

Neckeraceae, taxones tambin registrados en otros ecosistemas del interior del pas
(Churchill & Linares, 1995; Pinzn & Linares, 2006).

Estos resultados muestran que estas especies, que normalmente han sido halladas en
zonas altas, tambin pueden ser tolerantes a ecosistemas donde las condiciones
ambientales (exposicin directa a la luz del sol, fluctuaciones de humedad, entre otros)
no son totalmente favorables para su desarrollo. Tal es el caso del
genero Fissidens taxn registrado por Morales et al. (2008), el cual demostr ser el grupo
con el mayor nmero de especies (F. dissitifolius, F. mollis, F. prionodes y F. steerei)
registradas en este estudio, lo cual puede derivar de las adaptaciones morfolgicas que
este taxn posee, ya que, sus hojas se disponen dsticamente y una lmina
evaginante, que le permiten desarrollarse en ambientes con variaciones hdricas, e
incluso, en estos lugares donde el agua es limitada por pocas (Pinzn y Linares, 2006).

Para el caso de las hepticas, Lejeuneaceae fue el grupo con ms taxones reportados,
y al igual que las especies del gnero Fissidens esta familia se encontr en lugares con
alta exposicin a la radiacin solar (>80%), condicin que da lugar a gneros
caractersticos de regiones xerofticas y subxerofticas como Lejeunea (Pinzn & Linares,
2006; Vilas & Passos, 1998) y Mastigolejeunea con la ocurrencia de un mayor nmero de
especies (M. auriculata y M. plicatiflora) en dicho ambiente. No obstante, la presencia
de Marchantiaceae y Plagiochilaceae marcan una gran diferencia en el Sitio 2, debido a
que no fueron registradas en los otros puntos de muestreos (Tabla 1), destacndose
as Marchantia con relacin a franjas hmedas y conservadas (Lisboa & Ilkiu, 1995).

En el Sitio 3, se observ una baja diversidad en comparacin con el sitio dos, esta
diferencia puede resultar debido a la ausencia de meandros, los cuales limitan la
formacin de microhbitats en los alrededores y al interior del arroyo, dificultando la
supervivencia y el desarrollo de los brifitos; adems, este lugar present un mayor
impacto antrpico que el observado en el Sitio 2, dicha observacin es confirmada por
la alta presencia de especies de Pottiaceae y Lejeuneaceae, las cuales, han sido
registradas en varios trabajos como indicadores de ambientes perturbados (Pinzn &
Linares, 2006; Vilas & Passos, 1998).

En el sitio uno, fue posible encontrar una diversidad intermedia, es decir, este sitio
comparte un nmero de especies con las otras reas de estudio (Tabla 2), aspecto
atribuido quizs a la similitud (Leucomiaceae y Neckeraceae) que este posee con el
Sitio 2, al presentar ambos bosques de galera y suelos calcreos; con respecto al Sitio
3, las semejanza puede deberse a que estos puntos presentan cierto grado de
antropicidad que pueden alojar iguales comunidades de brifitos.

Con respecto a la diversidad evidenciada en los tipos de sustratos, es claro que los
evaluados (epiltico, epifito-cortcola, suelo y materia orgnica en descomposicin)
presentan comunidades de brifitos. No obstante, el epiltico fue el sustrato con mayor
nmero de individuos, lo cual es coherente con los espacios donde fueron realizados
los levantamientos de vegetacin, ya que estos eran zonas rodeadas por fuentes de
aguas en donde predomina el sustrato rocoso, cuyas caractersticas morfolgicas
incluan fisuras y agujeros que pueden acumular polvo, materia orgnica y
temporalmente agua, favoreciendo el desarrollo de un buen nmero de especies de
musgos de crecimiento acrocrpico (Pinzn & Linares, 2006) como F. steerei, H.
involuta, S. obtusum, Anomobryum conicum, Mielichhoferia megalocarpa, entre otros.

En cuanto a los sustratos con respecto a los sitios de muestreo, la diversidad de

especies vara considerablemente entre los sustratos en cada uno de los sitios; por
ejemplo, en los Sitios 1 y 2, se recolectaron un mayor nmero de especies de musgos
y hepticas epifito-cortcola, en comparacin con el suelo y materia orgnica en
descomposicin. T. tomentosum, Neckeropsis undulata (musgos) y Mastigolejeunea
auriculata (hepticas), son epfito-cortcolas y son representativos, debido a las
coberturas frondosas que proporcionan ambientes hmedos y sombros para el
crecimiento y desarrollo sobre los rboles de estos epifitos (Barbosa et al., 2007).
Adems, Santos & Aguirre (2010)postularon que la colonizacin de cortezas por brifitos
epifitos es una adaptacin hacia la utilizacin del agua como recurso generalmente
sobrante en las plantas hospederas, siendo las familias Fabroniaceae, Neckeraceae,
Sematophyllaceae, Thuidiaceae (musgos) y Lejeuneaceae (hepticas) las ms
dominantes sobre este tipo de sustrato (epfito-cortcola).

En suelo y materia orgnica en descomposicin hay sustratos que estn ms

expuestos a las fluctuaciones ambientales (oscilacin de temperatura, luz y humedad a
lo largo del da-noche), condicin que restringen la colonizacin. Los taxones
encontrados fueron S. obtusum, P. substriatum, H. involuta, T. tomentosum, Fabronia
ciliaris (musgos) y L. flava (hepticas).

El sitio tres se caracteriza por presentar menor nmero de especie sobre el suelo y
epfito-cortcola, tanto en musgos (H. involuta) como en hepticas (Lejeunea maxonii),
debido a la cobertura arbrea rala, a una mayor intervencin antrpica que genera
claros dentro del bosque, lo cual implica una mayor incidencia de la luz (Barbosa et al.,
2007) que genera una reduccin de la humedad y un incremento en la radiacin, factor
restrictivo para el crecimiento y desarrollo de brifitos en estos sustratos (Equihua et al.,
2001). Adems, dichos factores segn Aguirre & Avendao (2008) son influyentes en la
presencia y dominancia de especies como es el caso de hepticas (Lejeuneaceae) al
borde del camino, las cuales estn adaptadas a la tolerancia de la desecacin y de la
radiacin solar directa (Barbosa et al., 2007).

Es as como, se evidencia que Lejeuneaceae se encontr en todos los sustratos

estudiados, como lo demuestran tambin Lisboa & Ilkiu (1995), quienes encontraron que
Lejeuneaceae crece sobre una variedad de sustratos como, troncos (vivos y en
descomposicin), hojas y a veces sobre rocas y suelo, probablemente por sus
caractersticas morfo-fisiolgicas (no evaluadas), las cuales les permiten poder
colonizar y adaptarse a los diferentes sustratos y microambientes que se encuentran
en los bosques de tierras bajas. Por ejemplo, los lbulos, se interpretan como una
adaptacin al crecimiento epfito y su tamao est relacionado con la humedad del
ambiente (Castroviejo & Ibez, 2005).

Por ltimo, la diversidad y la distribucin de los brifitos, se encuentra determinada

por la complejidad del ambiente y el grado de perturbacin en los sitios de muestreos.
Adems, los brifitos son considerados un grupo indicador de reas conservadas, ya
que, brindan informacin valiosa para conocer las condiciones en un rea de estudio,
dadas por sus caractersticas morfolgicas que implican amplios requerimientos de
humedad, disponibilidad de agua, sombra, entre otros.

CONCLUSIONES

Este estudio permiti aportar informacin al conocimiento de la flora de bosques

neotropicales de tierras bajas (Departamento de Sucre), registrando un total de 18
familias, 24 gneros y 31 especies de brifitos.
31 especies son nuevos registros para el departamento de Sucre, 24 para la regin
Caribe colombiana y uno (Hyophiladelphus agrarius) para Colombia.

La diversidad y distribucin de los brifitos, se encuentra determinada por la

complejidad del ambiente y el grado de perturbacin en los sitios de muestreos.

Las familias, gneros y especies de brifitos del municipio de Colos, se distribuyen

sobre el sustrato epiltico, seguido por epifito-cortcola. Sin embargo, esta distribucin
disminuye para los sustratos suelos y materia orgnica en descomposicin; siendo las
familias Fissidentaceae y Pottiaceae con mayor distribucin sobre el sustrato epiltico
en los tres sitios de muestreos.

AGRADECIMIENTOS

A los herbarios COL y UPTC, al Instituto de Ciencias Naturales de la Universidad

Nacional de Colombia por las facilidades y soportes brindados durante la ejecucin de
este trabajo. A dgar Linares y Jorge Gil por su colaboracin en la identificacin de las
especies.
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