ELSEVIER
Biological Conservation 70 (1994) 19-24
© 1994 Elsevier Science Limited
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0006-3207/94/$07.00
EVOLUTION OF HIGH GENETIC VARIABILITY IN
THE RARE HAWAIIAN FERN Adenophorusperiens AND
IMPLICATIONS FOR CONSERVATION MANAGEMENT
Tom A. Ranker
University Museum and Department of EPO Biology, University of Colorado, Campus Box 350,
Boulder, Colorado 80309, USA
(Received 19 January 1993; revised version received 9 November 1993; accepted 1 December 1993)
Abstract
Adenophorus periens is a rain forest-dwelling epiphytic
fern and is known only from one population on the island
of Hawaii (Kahaualea) and from a few scattered individ-
uals on the islands of Kauai and Molokai. A genetic
survey was conducted of the Kahaualea population
employing protein electrophoresis. The individuals sam-
pied expressed extremely high levels of allozymic vari-
ability compared to two more widespread congeneric
species (A. tamariscinus and A. tripinnatifidus). All in-
dividuals examined were produced via sexual recombina-
tion. Given the extremely young volcanic substrate upon
which the Kahaualea forest has developed, the historical
factors leading to the high genetic variability of A.
periens are somewhat obscure. The outcrossed mating
system of this species may play a key role in the main-
tenance of genetic diversity in combination with its
perennial life cycle. Because the lack of neighboring
populations precludes the influx of new genetic variability
via gene flow, the population must rely solely on the
appearance and incorporation of new mutations as
sources of new variability.
The data suggest that this genetically diverse popula-
tion is not likely to suffer genetically from minor reduc-
tions in population size over the short term. The most
obvious potential threats to A. periens are those affecting
the demography of the population and include habitat de-
struction by lava flows, death of host trees through stand-
level dieback, chance loss of individuals by a variety of
mechanisms, and the adverse effects of gaseous emissions
from volcanic activity and geothermal development. The
State of Hawaii shouM undertake an artificial propaga-
tion and transplantation program to establish populations
in less threatened, suitable habitats.
Keywords: Hawaii, Adenophorus, genetics, evolution.
INTRODUCTION
Low levels of genetic variability have been documented
for a variety of geographically restricted plant species
(e.g. Ledig & Conkle, 1983; Lesica et al., 1988; Hickey
et al., 1991). Although these data are consistent with
the theoretically expected relationship between genetic
diversity and population size (Wright, 1931), several
19
exceptions to this relationship have been described
(discussions and references in Karron, 1987, 1991 and
Hamrick & Godt, 1990). Since the extent of genetic
variability is critical for the long-term survival and con-
tinued evolution of a population or species (Lande &
Barrowclough, 1987), genetic analyses of threatened
and endangered species are useful to aid conservation
decisions. Such studies may provide crucial data for the
assessment of present and future population viability
and for unraveling the historical factors accounting for
existing levels and patterns of genetic diversity.
Ranker (1992a) surveyed allozymic variability in two
widespread species of the endemic Hawaiian fern genus
Adenophorus Gaud. (Grammitidaceae). Populations of
A. tamariscinus (Kaulf.) Hook. &Grev. were found to
have higher levels of genetic variability than has been
reported for a variety of mainland homosporous ferns
(Soltis & Soltis, 1990; Ranker, 1992b). Populations of
A. tripinnatifidus Gaud. exhibited slightly lower levels
of genetic diversity, intermediate between historically
outcrossing and inbreeding continental populations.
Although these two species both occur on all of the
main Hawaiian Islands (i.e. Kauai, Oahu, Molokai,
Lanai, Maui, and Hawaii; Bishop, 1974), A. tripin-
natifidus is usually ecologically restricted to extremely
wet forests whereas A. tamariscinus occupies a greater
variety of montane habitats (Bishop, 1974; personal
observations). The greater ecological amplitude of the
latter species may be directly linked to its ability to
maintain higher levels of genetic variability. The slight
difference in levels of variability between these two
species along with the possible relationship between
genetic diversity and ecological amplitude suggests that
the extent of geographical distribution alone (i.e. in
terms of the number of islands on which a species is
found) may not be a good predictor of populational
genetic variability in species of Adenophorus.
In light of the information gathered on these two
widespread species of Adenophorus (Ranker, 1992a),
the present study was undertaken to explore the popu-
lation genetics of one of the rarest Hawaiian ferns, A.
periens L. E. Bishop. Although this epiphyte has been
collected historically on all the Hawaiian Islands that
support rain forests (Bishop, 1974), it is presently
known from only a few locations in middle-elevation