THE
UNIQUE ARTHROPOD BIODIVERSITY OF THE KLAMATH/SISKIYOU MOUNTAIN REGION IN NW
USA
CHALLENGES
FOR INVENTORY AND CONSERVATION
A. R. Moldenke1, B. G. Marcot2, D. DellaSala3 & N. Rappaport4
1Dept of Entomology, 2046 Cordley Hall, Oregon State University,
Corvallis, OR 97331-2907; 2US Forest Service, 1221 SW Yamhill St., PO
Box 3890, Portland, OR 97208-3894; 3World Wildlife Fund, 116 Lithia Way – Suite 7, Ashland, OR 97520; 4Forest & Range Experiment Laboratory, USDA Forest Service, Albany,
CA.
E mail:
moldenka@bcc.orst.edu
ABSTRACT:
This region has long
been recognized as of extreme significance botanically. Escape from
glaciation, diversity of geologic substrates combined with varied relief, and
location at the interface of mesic vs. xeric biomes should have rendered this
region as unique entomologically as it is botanically. Though scattered
literature supports this hypothesis, a careful documentation of the endemic
levels of arthropod biodiversity needs to be undertaken.
The World Wildlife
Fund, the Forest Service (land owners) and entomologists from Oregon State
University are collaborating in a unique attempt to define and protect the
ecological processes in which arthropods participate and to preserve
biodiversity hot-spots and premier taxa, while permitting economic utilization
of the forest resources as a sustainable long-term resource.
Relict ancient taxa
representative of the transcontinental forests before the Miocene mountain
orogeny split the continent in two characterize this region (i.e., Tricholepidion gertschi (Microcoryphia) endemic to CA; Cryptocercus punctulatus (communal Blattaria)
amphi-continental distribution).
Beta-diversity is
driven by habitat heterogeneity and recent adaptive radiation (i.e.,
Amaurobiidae (Araneae)). Additionally, taxa with limited mobility have spawned
localized geographic isolates as they recolonized the Pacific Northwest after
glaciation (i.e., Pleocominae (Scarabaeidae), Caseyidae (Diplopoda)).
Though the vast
majority of taxa inhabiting the region is widely distributed geographically,
endemicity is likely to be relatively high.
With adequate
inventory a system of protection can be implemented before this ecosystem is
more heavily impaired.
Who Cares?
Under the current and new “ecosystem management” paradigm for forested
lands of the US Federal Government (Forest Service)
biodiversity and ecosystem processes are of paramount importance. Thus, one
management requirement is knowledge of the pattern of biodiversity across the
landscape.
Are there regions of higher species diversity (hotspots)?
Are there regions characterized by higher species endemicity?
Arthropod species are also important for what they do. Who
are the prime pollinators, nutrient cyclers, herbivore controls?
Examples:
Harpaphe haydeniana (millipede) – this
one millipede species feeds
upon 33-50% of all the dead deciduous leaves and coniferous
needles
that fall upon the forest floor. By crushing up the plant cells it
allows the
process of microbial nutrient recycling to begin.
Eulonchus tristis (acrocerid fly) –
this species is the prime pollinator of several dozen
genera of plants on the old-growth forest floor, and also the prime
population regulator
of the invertebrate alpha-predator of the forest floor (trap-door
spiders).
Background:
Long ago in the Miocene period, the transcontinental mesic forests of North America were split,
leaving a small forest “island” on the West Coast. (Klamath Mountains long predate the
Cascades, Sierras and Coast Ranges.)
-During this period,
the global climate cooled and the warm subtropical flora receded to the South.
-The Rocky Mountains
continued to build and the Cascades and Sierras began to uplift,
resulting
in the drying and formation of the Great
Basin and the Great Plains.
-The arid-adapted
flora and fauna of the Southwest US and Mexico spread into the region.
CURRENT EXAMPLES
RESULTING FROM FOREST ISLAND ON WEST COAST :
• Ancient relict “Palaeoendemics” in western North America
Sequoia (conifer); Ascaphis (frog); Tricholepidion (Microcoryphia)
• Taxa with a Split-continental distribution
Torreya (conifer); Plethodontidae (salamanders); Cryptocercus (Blattaria)
Most of the basic floral relationships of today were established in
this region by the end of the Miocene/Pliocene. Later, during the Pleistocene, warm ocean-current circulation spared much of the Klamath/Siskiyou
Region from glaciation – followed by a rapid
colonization and radiation throughout adjacent areas. Alternating cooler and warmer periods left isolated mesic plant stocks on north-facing higher
altitudes and dry-adapted species on open-canopy xeric serpentine habitats.
“Neoendemics” in Klamath/Siskiyou region:
• Plant radiation/speciation on serpentine
soils
• Rasenkreise of Ensatina eschscholtzii (salamander)
• Parapatric swarms of poorly dispersed arthropods such as
millipedes
in the family
Caseyidae and flightless pleocomine Scarabaeidae.
Survey Design: Field
Protocols
Alpha-diversity
& beta-diversity
Experimental Units:
Adjacent pairs of 40-80 acre
watersheds (15-30 hect.)
(standard US Forest Service management units)
Biodiversity Drivers (established by prior studies):
1o determinant – season
Hypothesis: Within the Pacific
Northwest forest, the turnover in both species
presence and activity levels between wet and dry
seasons increases with decreasing latitude. Most summer-active taxa are derived from “Madro-Tertiary” stock which have
colonized from the arid SW and Mexico.
_____________________________________________________________________
2o determinant – successional stage
Hypothesis: Arthropod biomass should
increase throughout the year in early successional stages in the North (WA) due to the large annual photosynthetic
production in clear-cuts; however, moisture
limitation in the Klamath mountains should force most summer arthropod
activity into the forest shade. In the summer and fall insectivorous
vertebrate activity should be greater in clear-cuts in the North, but mature forests in the South.
typical pattern of successional comparisons:
a) whole groups in
one stage only
ex:
Apoidea in early succession (200+ spp)
b) individual
species in one stage only
ex:
Taracus (Phalangid) in mature forest
c) sibling species
in alternate successional stages
ex:
Araneus mariposa in early succession
Araneus nordmanni in mature forest
_____________________________________________________________________
3o determinant – exposure or slope-face
Hypothesis: In the North (WA) there
should be higher species turnover with differing slope-face in the early succession than in the mature forest; in the South
(Klamath/Siskiyou) species turnover dependent upon
slope-face should be greater in the mature forest. In the North forest canopy
is usually dense enough to significantly insulate the
understory. In the South the clearcuts are so dry that it makes little difference what the slope-face is; under the forest canopy the
moisture gradient is prominent.
_____________________________________________________________________
4o determinant – edaphic substrate specialization
In
Klamath Region the dominant plant genera on serpentine soils are the same as
those on
other soil types but the species are all
restricted to serpentine (i.e., trees: Pinus, Quercus; shrubs: Arctostaphylos, Ceanothus). Many plant genera are restricted only to serpentine (ex: Darlingtonia).
QUESTION:
Are the insect faunas different as well?
Goals of Field Research Component:
Description
of how alpha-diversity of a number of arthropod groups differ:
(1) within a site throughout the seasons
(2) & how species are replaced (beta-diversity) between adjacent sites
differing in either successional stage,
slope-face, or soil type.
Library Research: Reference
monographs
Gamma-diversity
& endemism
There are 5 major patterns, generally speaking for arthropods as a whole, of geographic
distributions characteristic of
species inhabiting the Klamath/Siskiyou Region. (Individual taxonomic groups
differ widely from the norm.)
montane transcontinental Cascade-
w. N.Am sub-boreal Sierras
(30%) (20%)
(20%)
Xeric endemic others
Great
Basin Klamath (variable)
(15%) (10%) (5%)
Classic
examples of Klamath/Siskiyou endemism:
a) strong
Klamath/Siskiyou richness hotspot
Klamath
Region = 45 spp of Amaurobiidae all
endemic
Rest of N
Amer = 38 spp nearly all very widespread
b) edaphic
endemism
(1) limestone endemism
subterranean blind cavernicolous millipede –
Aprosphylosoma
(monotypic genus)
(2)
serpentine endemism
(associated
with nearly ALL of the plant endemicity)
Question: are there more examples from arthropods??
Forest management practices can be adapted to
insure continued alpha-diversity and forest ecosystem health. Once we know the pattern of diversity these can be maintained
with preserves (if any are needed in addition to
those established for rare plants or vertebrates), or adequate buffers.
However, the scientific community needs to detail
which arthropod taxa are really unique to the Pacific Northwest and occur nowhere else.
Which bugs are analogous to the unique redwood?
This is not as much a question of conserving
biodiversity through management protocols as it is better understanding the evolutionary uniqueness of the region, and developing
an appreciation for the life-forms we have that exist nowhere else.
Punchline:
The World Wildlife Fund has designated the Klamath/Siskiyou region
as one of the areas of most critical
concern for “biodiversity conservation” worldwide.
Botanists and herpetologists appreciate the area’s unique taxa.
Entomologists now need to learn and carefully document just how
significant this region is !
Hexura
Agulla
Zootermopsis
Edith’s checkerspot:
serpentine endemic
Eulonchus tristis
Tanypteryx
Grylloblatta
Bequaert’s fly
Eschscholtz’s
ground
beetle
Long-jawed
millipede
munching
ground beetle
Mountain midge
ancient
spider family
Amaurobiidae