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Phylogeography, intraspecific structure and
sex-biased dispersal patterns in marine mammals: Dall's porpoise and spotted
dolphins
These two species provide contrasting systems. Dall's
porpoise are endemic to the North Pacific and predominantly pelagic. They
are divided into two morphologically distinct subspecies and are currently
subjected to large commercial catches by a hand-harpoon fishery. Spotted
dolphins are distributed globally in tropical and warmer temperate waters.
In the Eastern Tropical Pacific Ocean, they are the second most common
species affected by the tuna purse seine industry. Until recently, they were
subjected to tremendous mortality. Despite the intense human impact, little
is known about the migration patterns and separation of stocks in either
species, information critical to their effective management
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Phylogeography and the origin of species
Phylogeographic research on Dall's porpoise has provided information
concerning the history and origin of present-day populations of Dall's
porpoise. These data are used to determine the "big picture", the stage
or background on which more recent population genetic processes arise.
The estimated time to the hypothetical ancestral mtDNA haplotype of the
Dall's porpoise, from which the haplotypes of present-day Dall's
porpoises derived, existed at least between 600,000 and 1,000,000 years
ago, based on coaslescent theory. Results further suggest that Dall's
porpoise experienced a rapid divergence from an ancestral population in
the western North Pacific overlain with high levels of historical gene
flow (Escorza-Treviño and Dizon, 2000).
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Taxonomic Status. Does Morphology Predict Taxonomy?
The taxonomic status of the two morphologically distinct forms of
Dall’s porpoises (dalli-type and truei-type) is
unsettled. Different investigators have considered these forms as
separate species, separate subspecies, or morphological extremes of a
single species. To help resolve this uncertainty, mitochondrial DNA
(mtDNA) sequences were used to estimate phylogenetic relationships
between the two types and, in conjunction with microsatellite markers,
the degree of genetic differentiation between truei-type and
currently recognized dalli-type populations was tested. Results
showed two distinctive clades of mtDNA haplotypes (Escorza-Treviño et al.,
2004). However, both were composed of individuals from both types,
with shared mtDNA haplotypes between morphotypes. These results suggest
that truei- and dalli-types are forms of the same species.
The truei-type animals were also typed for six microsatellite
loci. F-statistics of mitochondrial and nuclear data, however, revealed
statistically significant genetic differentiation between truei-type
and sympatric and adjacent dalli-type populations. These results
argue that the truei-type, currently subjected to direct
commercial harvesting, has demographically insignificant exchange with
the much larger dalli-type populations and should be considered a
separate management unit.
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Population structure. What are the management units?
Successful conservation of living resources is based upon the
accurate designation of "management units". A major goal of our research
effort is to define these units so that management efforts can be
appropriately placed. In Dall's porpoise, animals from several
geographic strata were sequenced for a hypervariable region of the
mitochondrial genome and typed for variation at 6 microsatellite loci.
Analysis using FST values (based on haplotype
frequencies) and FST
values (based on frequencies and genetic distances between haplotypes)
yielded statistically significant separation among most of the stocks
currently used for management purposes by the International Whaling
Commission. The analyses also revealed two additional populations,
requiring further conservation attention (Escorza-Treviño and Dizon,
2000). Overall, mtDNA analysis supported more population subdivision
than microsatellite analysis. This initially unusual result led to the
question of sex-biased dispersal patterns (see below).
Similar markers were used to investigate genetic differentiation in
spotted dolphins (Stenella attenuata) in the Eastern Tropical
Pacific and to determine the taxonomic status of the coastal subspecies
(S. a. graffmani). Analysis of mitochondrial control region
sequences, from animals present in several coastal areas and offshore,
using FST and FST
yielded statistically significant separation among several different
coastal and the offshore strata (Escorza-Treviño et al., 2002). Analysis using FST
values for seven nuclear microsatellite loci yielded statistically
significant separation between each of the coastal strata and offshore
animals, but no coastal populations were distinguished. These results
argue for the existence of at least four distinct coastal populations
that should be treated as separate units for management purposes (Escorza-Treviño
et al., 2005) and
suggest the existence of male-biased dispersal among the coastal strata.
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Sex-biased dispersal patterns: philopatry, dispersal, and the power to
homogenize populations
In mammals, female philopatry and male dispersal is the rule. This
may be especially true among marine mammals that are widely distributed
and mobile. Yet, the extent of a bias in dispersal between the sexes, or
of its effects on population structure, is little known for any marine
mammal species. For both Dall's porpoises and spotted dolphins, highly
male-biased migration was detected for several population comparisons (Escorza-Treviño
and Dizon, 2000). The sex-biased dispersal of Dall’s porpoises explained
discrepancies in the population structure data as well. It seems
possible that site fidelity may be especially high for females of both
these species and, while males disperse and homogenize the populations,
different maternal lineages separate and define populations.
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Other
research
Our immediate plans are to continue molecular studies of the spotted
dolphin complex, investigating how sex-biased dispersal and social
structure affect population structure. Through similar molecular
methodologies, we are investigating the genetic relatedness among
individuals sampled in the same group to establish if relationships are
based on kinship, gender, and/or a combination of these. In addition, we
are comparing these relatedness results between coastal and offshore
spotted dolphins in order to establish a connection between social
structure and population differentiation.
We also intend to expand our research efforts to forensic
applications of molecular techniques. In the past, we have worked on the
identification of tissue samples suspected of being from protected
species. We are planning on establishing fast and replicable methods for
species identification to curb the illegal catch, trade, and transport
of protected species. In addition, we expect to incorporate knowledge of
population differentiation to identify samples to a greater geographic
detail as a means to enhance conservation efforts. These methodologies
are easily applicable to other protected species, be they marine or
terrestrial.
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