Biodiversity Data Journal 4: e6732
doi: 10.3897/BDJ.4.e6732
General Article
Macro-invertebrate Biodiversity of a Coastal Prairie
with Vernal Pool Habitat
Emile Fiesler‡, Tracy Drake§
‡ Bioveyda, Torrance, California, United States of America
§ Manager, Madrona Marsh Preserve, Torrance, California, United States of America
Corresponding author: Emile Fiesler (bioveyda@yahoo.com)
Academic editor: Benjamin Price
Received: 01 Oct 2015 | Accepted: 30 Mar 2016 | Published: 27 Apr 2016
Citation: Fiesler E, Drake T (2016) Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat.
Biodiversity Data Journal 4: e6732. doi: 10.3897/BDJ.4.e6732
Abstract
Background
The California Coastal Prairie has the highest biodiversity of North America's grasslands,
but also has the highest percentage of urbanization. The most urbanized part of the
California Coastal Prairie is its southernmost area, in Los Angeles County. This
southernmost region, known as the Los Angeles Coastal Prairie, was historically dotted
with vernal pools, and has a unique biodiverse composition. More than 99.5% of its
estimated original 95 km2 (23,475 acres), as well as almost all its vernal pool complexes,
have been lost to urbanization.
The Madrona Marsh Preserve, in Torrance, California, safeguards approximately 18
hectares (44 acres) of Los Angeles Coastal Prairie and includes a complex of vernal pools.
Its aquatic biodiversity had been studied, predominantly to genus level, but its terrestrial
macro-invertebrates were virtually unknown, aside from butterfly, dragonfly, and damselfly
observations.
© Fiesler E, Drake T. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are
credited.
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New information
In order to better understand the biodiversity at the Madrona Marsh Preserve, a minimallyinvasive macro-invertebrate inventory was conducted. The results of this inventory, with
689 invertebrate organisms recorded, covering eight phyla, 13 classes, 39 orders, and 222
families, are presented in this document.
Keywords
biological diversity, ephemeral pool, wetland, dune, upland, terrestrial, aquatic,
invertebrate, Arthropoda, Hexapoda, Insecta, Lepidoptera, Odonata, Coleoptera,
Hymenoptera, Diptera, Hemiptera, Heteroptera, Orthoptera, Neuroptera, Psocodea,
Thysanoptera, Blattodea, Embioptera, Dermaptera, Microcoryphia, Collembola, Arachnida,
Araneae, Acari, Mollusca, Gastropoda, checklist, taxonomy, classification, systematics, bioinventory, macro-photography, Madrona Marsh Preserve, Torrance
Introduction
The California Coastal Prairie has the highest biodiversity of North America's grasslands
Stromberg et al. 2002, but also has the highest proportion of urbanization Loveland and
Hutcheson 1995. The most urbanized part of the California Coastal Prairie is its
southernmost area, located in Los Angeles County. This southernmost region, known as
the Los Angeles Coastal Prairie, was historically sequined with vernal pools, and has a
unique biodiverse composition Mattoni and Longcore 1997. More than 99.5% of the
estimated 95 km2 (23,475 acres), and almost all vernal pool complexes, have been lost to
urbanization Mattoni et al. 2000. The remaining estimated thirty hectares (approximately 74
acres) has undergone considerable degradation. Notwithstanding these urbanization
pressures, the remaining fragments of the Los Angeles Coastal Prairie harbor a rich and
understudied macro-invertebrate biodiversity.
The Madrona Marsh Preserve FoMM 2015 (MMP, or "the Preserve") is a public nature
preserve located in the City of Torrance, California; see Figs 1, 2. At approximately 18
hectares (44 acres), it safeguards the largest remnant of Los Angeles Coastal Prairie, as
well as its last functional vernal pool complex.
MMP is located in the South Bay area of Los Angeles County (33.827° North; 118.341°
West) at approximately 24 meters (80 feet) elevation above sea level (see maps in Fig. 3).
The Preserve is surrounded by urban areas, including California's second largest shopping
mall on its west side. It is bounded by Madrona Avenue on the west, Sepulveda Boulevard
on the south, Maple Avenue on the east, and Plaza del Amo on the north. MMP is 4.2
kilometers (2.63 miles) inland from the Pacific Ocean coast and 4.8 km (3 miles) north of
natural open space on the Palos Verdes Peninsula, particularly the Linden H. Chandler
Preserve, managed by the Palos Verdes Peninsula Land Conservancy PVPLC 2015. It is
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
3
5.3 km (3.3 miles) west-northwest of the Bixby Marshland LACSD 2015, 6.8 km (4.2 miles)
southwest of the Gardena Willows Wetland Preserve Friends of Gardena Willows 2015,
and 13.8 km (8.6 miles) southeast of the LAX Dunes LAWA 2015. MMP is 17.1 km (10.6
miles) southeast of the Ballona Wetlands Friends of Ballona Wetlands 2015, which
includes estuarine, brackish, and freshwater marshlands, and is located scarcely outside
the northern historical boundary of the Los Angeles Coastal Prairie. The Santa Monica
Mountains National Recreation Area SMMNRA 2014 lies 30 km (19 miles) to the
northwest, and the Channel Islands National Park NPS 2015, with Santa Barbara Island,
72 km (45 miles) to the southwest.
Figure 1.
Madrona Marsh Preserve
Figure 2.
Madrona Marsh Preserve - Willow forest
Fiesler E, Drake T
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Figure 3.
Location of Madrona Marsh Preserve (MMP) in Torrance, California.
Insets depict MMP's location with respect to the United States of America and southern Los
Angeles County respectively.
MMP has a Mediterranean climate with mild, dry summers and cool, wet winters. The
annual average precipitation is 37.6 centimeters (14.8 inches), which almost all falls
between mid October and late March. August is usually the warmest month of the year,
with an average maximum temperature of 25.9 degrees Celsius (78.7 degrees Fahrenheit).
December is usually the coldest month of the year with an average minimum temperature
of 7.8 °C (46.0 °F). Temperature variations between night and day vary from an average of
9 °C (17 °F) in summer to 11 °C (20 °F) in winter U.S. Climate Data 2015.
MMP is a nature oasis in an urban setting; see map in Fig. 4. Located in the City of
Torrance, being a jewel along the Pacific Flyway Pacific_Flyway 2015, MMP is well known
for its avifauna Lyle 1999, Wikipedia 2016. Aside from its well-documented bird richness,
its vertebrate fauna, as well as its vascular flora, have also been studied to some extent in
the past City of Torrance 1976. Although its aquatic biodiversity had been studied,
predominantly down to genus level Angelos 2003, its terrestrial macro-invertebrate fauna
was virtually unknown, aside from butterflies and odonata observations. MMP butterfly
sighting records have been primarily obtained by David Moody, Tracy Drake, Jess Morton,
Martin Byhower, Kevin Larson, and Mitch Heindel. MMP odonata observations began in
2004, initiated by the second author. Summaries of the butterfly and odonata tallies have
been published periodically by the second author in the Marsh Mailing, the periodic
newsletter of the Friends of Madrona Marsh FoMM 2015.
Between November 16, 2009 and August 31st, 2011 the first author conducted a minimallyinvasive biodiversity inventory of the macro-invertebrates of the Preserve, often
accompanied by the second author during field work. This study was part of a biological
inventory that also included vertebrate animals and vascular plants. Goals of the bio
inventory included researching historical data, providing a baseline inventory of the current
MMP taxa to be able to compare future inventory date to, creating factual knowledge for
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
5
education and outreach, as well as providing habitat restoration and enhancement
recommendations for the Preserve.
Figure 4.
Map of Madrona Marsh Preserve.
Each red X-mark indicates the location of one of the ten pitfall traps. Each purple C-mark
indicates the location of one of the eight sets of three coverboards.
Vegetation is shown in shades of green and yellow, vernal pools in pink, water inundation
levels in shades of blue, the remaining upland areas in shades of light brown, and man-made
structures in darker brown. Prominent trees and shrubs are indicated by black letters in pinkrimmed white circles. The Nature Center, surrounded by native plant gardens, is located north
of Plaza del Amo, and the sump, a neighborhood runoff water collection reservoir, is located in
the southeast corner of the preserve.
Map created by Jerry Cole, 2011.
The MMP macro-invertebrate bioinventory is unique in several aspects. It is the first macroinvertebrate bioinventory of its kind and scope, uniquely based on photographic vouchers,
rather than the traditional captured specimen vouchers. It was performed with minimal
manpower, and its scope is broad, aiming to include all taxa. This novel approach is made
possible due to the advent of high-quality digital photography, which allows for taking,
processing, organizing, and archiving large amounts of detailed photos, needed for such a
project. The Internet is another technology that enables small teams to complete macroinvertebrates bioinventory projects in a limited time. BugGuide in specific is an extensive
on-line resource with an unprecedented collection of more than 820,000 arthropod photos,
taxonomically classified and organized by more than 280 editors and experts BugGuide
2014.
Related studies by others - an overview
Biodiversity, including urban biodiversity, is critical to human sustainability Kim and Byrne
2006. Most biodiversity studies include vertebrate animals or vascular plants, or both. Few
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Fiesler E, Drake T
inventories include invertebrate animals, non-vascular plants, or organisms of other
kingdoms in their scope Gullison et al. 2015Pocock et al. 2015Schuldt and Assmann 2010.
Macro-invertebrates are ubiquitous and account for the largest percentage of all metazoan
biodiversity. They also account for the largest percentage of endemic taxa and taxa of
concern (threatened, endangered, or otherwise imperiled), although most such
invertebrates have not yet been officially recognized as being imperiled. The International
Union for the Conservation of Nature’s Red List IUCN Red List 2016 estimates the global
total at about 1.3 million invertebrate species and the percentage at risk of extinction at
about thirty percent (30%) of species evaluated Center for Biological Diversity 2015.
Macro-invertebrates are primary candidates for monitoring environmental impacts, as they
respond quickly to environmental changes due to their short life cycle. Moreover, they are a
crucial building block of the food-chain, and our global biosphere in general. The famous
American myrmecologist Edward O. Wilson said: "If all mankind were to disappear, the
world would regenerate back to the rich state of equilibrium that existed ten thousand years
ago. If insects were to vanish, the environment would collapse into chaos." Jarski 2007.
Surprisingly few macro-invertebrate or all-taxa inventories have taken place in general Kim
and Byrne 2006, including in well-populated Southern California. Some prominent bioinventories performed in Southern California that focused on, or included, macroinvertebrates, are listed in Suppl. material 1. The table, which is ordered by number of taxa
found, includes studies whose results were published by the end of the MMP bioinventory
in 2011. More recent studies show more up-to-date results. The bio-inventories listed in
Suppl. material 1 are also briefly discussed below; ordered in decreasing proximity, or
relevance, to MMP.
Mark Angelos performed a high-quality study of MMP's vernal pool arthropods, gastropods,
vertebrates, and plants, between 1993 and 2003. At that time he was Research Associate
in Invertebrate Zoology at the Los Angeles County of Natural History. He predominantly
used nets and recorded 76 animal taxa, of which 67 arthropods, five mollusks, and four
vertebrates; most of which he identified to genus Angelos 2003. Between 1997 and 2003
he performed a study of MMP's ostracods, small aquatic crustaceans, of which he found
seven species in MMP’s vernal pools Angelos 2005.
W. Dwight Pierce, together with Dorothy Pool and others, performed field studies in the El
Segundo Dunes from 1938 to 1939 and published highlights of their findings in a series of
articles published between 1938 and 1947 Pierce et al. 1947. Their field notes, which
include 187 arthropods and 38 (mostly marine) mollusks, were never published Pierce and
Pool 1939.
Rudi Mattoni and his team performed a study of the flora and fauna of the 746 hectares
(302 acre) El Segundo dune remnant at the west side of Los Angeles International Airport
(LAX) from June 1987 to June 1988 Mattoni 1990. The used a combination of pitfall and
yellow-pan traps, aerial net sweeping, beat-sheets, and Malaise trapping, as well as a
mercury vapor and a ultra-violet (UV) light for night data collection. Their faunal study
included invertebrates and yielded 728 arthropod and three mollusk taxa. Sand-obligate
invertebrates restricted to Southern California coastal dunes were listed in his 1993
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
7
publication on the Los Angeles Coastal Prairie Mattoni 1993. A follow-up study was
performed from 1992 to 1993 to sample arthropods on, and adjacent to, the El Segundo
Dunes Mattoni et al. 2000. They collected 169 species in yellow pan traps and 189 species
in pitfall traps. Nine (9) arthropod species (one spider, four moths, one cricket, and three
weevils) were listed as endemic to the El Segundo dunes Mattoni et al. 2000.
Travis Longcore, who participated in the arthropod study mentioned above Mattoni et al.
2000, performed a five year terrestrial arthropod study of thirteen (13) locations on the
Palos Verdes Peninsula, in southwestern Los Angeles County, plus two locations in Crystal
Cove State Park in coastal Orange County Longcore 1999. He found 204 arthropod taxa in
his pitfall traps.
The Ballona Wetlands, located north of the northern boundary of the Los Angeles Prairie,
consisted of approximately two thousand acres of coastal wetland habitat. Most of these
wetlands have been lost to urbanization, including nine hundred acres for the construction
of a small craft harbor and adjoining community in the 1960's, now known as Marina del
Rey. The remaining six hundred acres consist of estuarine, brackish, and freshwater
marshes, seasonal wetlands and other riparian habitats, as well as uplands. From 1980 to
1981, a one year biological study was performed on the Ballona Wetlands, which included
the recording of 475 insects and mollusks Schreiber 1981. They used a combination of
aerial, sweep, and aquatic nets, baited pitfall traps, Malaise trap, yellow pan traps, UV light
traps, soil sifting, and Berlese funnel sifting. The specimens have been absorbed into the
main entomology collection at the Los Angeles County Museum of Natural History.
Emile Fiesler conducted a macroinvertebrate bioinventory of the Oxford Basin in 2010. The
Oxford Basin, located in Marina del Rey, is one of the last remaining areas with intertidal
mud flat habitat in Los Angeles County. He recorded 85 terrestrial and marine taxa (two
mollusks, eight spiders, 67 insects, and eight other arthropods), using macro-photography,
during three half days of field work spread over four months Fiesler 2010.
Louis LaPierre and Pamela Wright surveyed the arthropod fauna of the Kenneth Hahn
State Recreation Area during June and July of 2000 LaPierre and Wright 2000. The
Kenneth Hahn State Recreation Area is located in the Baldwin Hills, which are situated
approximately halfway between the Ballona Wetlands near the Pacific Ocean and
downtown Los Angeles. They classified the habitat as "degraded Coastal Sage Scrub,
dominated by California Sagebrush and Coyote Brush." The methods they employed were:
pitfall traps, Malaise traps, yellow pan traps, beating sheets, aerial nets, black-lights and a
mercury vapor lamp. They found 114 arthropod taxa (fourteen spiders, two isopods, 70
Lepidoptera (33 butterflies and 37 moths), 15 Coleoptera, 7 Hymenoptera, and six other
insects) plus more than 99 undetermined morphospecies.
Emile Fiesler conducted a study of the non-marine macro-invertebrates of the Santa
Monica Mountains National Recreation Area (SMMNRA) Fiesler 2009. This pilot study
consisted in estimating the number of non-marine macro-invertebrate families present in
SMMNRA, compiling a complete list of butterflies that could potentially be found in the
SMMNRA, compiling a list of invertebrates of concern (rare, endemic, threatened, or
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Fiesler E, Drake T
endangered), and creating a preliminary list of 160 non-marine macro-invertebrates
representing 160 distinct families. The family list is comprised of twenty arachnids, seven
myriapods, three crustaceans, two mollusks, one nematomorph, and the rest hexapods. He
extracted this list from 410 (pre-project) plus 530 (new) specimens that are vouchered by
approximately 6,200 photos. Based on this pilot study, the first author created an on-line
field guide with the title: "Field Guide to the Insects and other Invertebrates of the Santa
Monica Mountains" Fiesler 2016.
Peter Bryant has set-up an extensive internet website on the biodiversity of "Orange
County and nearby places" Bryant 2014. Orange County is situated directly south and
southeast of Los Angeles County. The hierarchically organized website's scope includes all
taxa: animals, plants, fungi, and more. Each taxon is usually represented by its common
name, its scientific name, and one or more photos. Each photo has a caption, which
typically contains information on the photographer and the location and date where the
photo was taken. Selected taxa are accompanied by information on biology, ecology,
taxonomy, or other topics.
Nancy Hamlett is in charge of an all-taxa biota study of the 210 ha (85 acre) Robert J.
Bernard Biological Field Station of the Claremont Colleges. The Bernard Field Station is
located in the city of Claremont, approximately 58 km (36 miles) east of downtown Los
Angeles. The ongoing study covered 352 invertebrate taxa, as well as vertebrates, plants,
and lichens Hamlett 2014.
Studies on plants, vertebrate animals, and insects of the 2,469 ha (6,100 acre) Philip L.
Boyd Deep Canyon Desert Natural Reserve have been coordinated by the University of
California, Riverside. The Reserve is located five miles south of the City of Palm Desert in
Riverside County; in the Colorado Subdivision of the Sonoran Desert. A succession of
researchers, students, and visiting scientists have surveyed insects at the Reserve over the
past 34 years Frommer 1988. The insect survey, primarily utilizing Malaise traps, has
yielded about 5,000 insect taxa, 2,600 of which have been classified. The specimens are
curated at the University of California, Riverside entomology collection.
An all-taxa bioinventory was conducted as part of the San Diego Bay Integrated Natural
Resources Management Plan TD 2011. A key finding was that climate change and invasive
species are now principal drivers of change, whereas habitat loss due to development was
the main driver in the past for the San Diego Bay. More than 650 taxa of marine, estuarine,
and salt marsh invertebrates were recorded at species-, or genus-level, as part of the
bioinventory that was primarily based on trawling.
Gordon Pratt conducted a survey of the terrestrial arthropods of Edwards Air Force Base
between January 1996 and October 1998 Pratt 2000. Edwards Air Force Base is located in
the Mojave Desert, in northeastern Los Angeles County, approximately 36 km (22 miles)
northeast of Lancaster. He encountered 1,536 species from 32 sites, using sight, nets, and
pitfall traps.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
9
The Southern California Association of Marine Invertebrate Taxonomists (SCAMIT) has
compiled a list of benthic macro- and mega-invertebrates. The data is compiled from
various infaunal and epifaunal monitoring and research programs conducted in the
Southern California Bight, mostly based on trawling. The Southern California Bight area
covered by the list extends from Point Conception in Santa Barbara County, California to
Bahía Todos Santos, Baja California Sur, Mexico, in intertidal to 1000 meter depths. This
geographical area includes the off-shore areas of Los Angeles, Orange, San Diego, and
Ventura Counties, plus the southern section of Santa Barbara County and the
northwestern-most section of Baja California Norte. The ninth edition lists 3,177 taxa
Cadien et al. 2014.
Material and methods
The following sections detail the methodologies used for the terrestrial and aquatic
invertebrates surveys. The surveys performed for this project are unique as they were
conducted with minimal invasiveness. A minimally invasive study aims at studying
organisms with minimum disturbance and without collecting them; relying primarily on highresolution digital photography. To maximize taxonomic identification success, close-up
photos are taken from various angles, where possible. Nevertheless, not having a
specimen in hand tends to increase the complexity of taxonomic identification, as certain
characteristic details, some of which could be internal, might not be visible.
Terrestrial invertebrates data collection methodology
Common terrestrial invertebrates are mostly comprised of hexapods and arachnids. Other,
less species-rich taxa include: isopods, terrestrial snails, and earthworms. Terrestrial
invertebrates can be divided into (1) herbivores and detritivores, which are the primary
consumers, and (2) predators, parasites, and parasitoids. The herbivores and detritivores
comprise the lower levels of the food chain. Terrestrial herbivores are usually associated
with certain host- or food-plants, which are predominantly plants native to the area. Native
plants are therefore the base for a healthy terrestrial ecosystem.
The minimally invasive terrestrial invertebrate survey was performed in stages, using the
following methodologies:
•
•
•
•
visual detection and photo-documenting salient organisms, as well as evidence of
their presence;
periodic coverboard inspection;
overnight pitfall trapping focused on flightless invertebrates; and
night light collection, using both ultraviolet and broad-spectrum incandescent
illumination
Each of these four data collection methodologies is explained below.
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Fiesler E, Drake T
Visual detection took place during all visits of the site and formed an important part of the
data collection. We made an effort to choose a different transect for every visit to maximize
coverage of the area. We surveyed at various parts of the day, from the early morning to
after dark, with an average of 2.5 hours per survey. Keen visual observation enables
recording taxa that are not typically obtained by other methods and would go unnoticed.
Specimens were photographed using a Canon Powershot A650 IS in macro-photography
setting. Each specimen was photographed from various angles, aiming for dorsal, lateral,
and frontal exposures; limited by the time the specimen remained present.
Eight sets of three wooden coverboards were placed throughout the preserve, mostly at
shady locations to minimize direct sun exposure. Each of the 24 square coverboards
measured three by three feet (91 centimeters) in length and 0.5" inch (1.3 cm) thick. The
map in Fig. 4 contains the locations of the coverboards. During each of the transects
followed during the visual surveys explained above, we would examine the coverboards
that we encountered en route. One person would quickly lift the coverboard in such a way
as to maximize exposure of sunlight of the area that had been covered by the board.
Specimens encountered would be photographed in a manner similar to that explained
above.
Ten independent pitfall traps, consisting of 1.75-liter white polypropylene containers with
smooth, near-vertical walls, were placed on June 17 and September 10, 2010, at various
well-spaced locations from the waterline to the higher parts of the core upland areas Fig. 4.
The containers were placed in holes dug into the soil such that their upper rims were flush
with the surrounding ground. The locations were chosen to maximize inclusion of habitats.
Some were placed under the canopy of shrubs and trees, like western sycamore (Platanus
racemosa Nutt.), coast live oak ( Quercus agrifolia Née), and Mexican elderberry (
Sambucus nigra L. subsp. caerulea (Raf.) Bolli), to catch animals that might fall down due
to the wind or other mechanisms. The next morning the pitfall traps were collected, the
holes closed, and their content documented.
The night light data collection took place on April 13, May 12, June 17, August 10,
September 9, and December 1 of 2010, and on February 23, April 6, August 23, and
November 8, 2011; mostly on dates close to a new moon. New moon assures minimum
light from the sky as well as maximizes insect, especially moth, activity, apparently due to
high light polarization levels Nowinszky and Puskás 2012. At night, we used ultraviolet light
stations, as well as broad-spectrum incandescent (“white”) light stations. Given the need
for powering the lights, we typically deployed at least one ultraviolet station in the central
area of the Preserve, where we used the available 110 Volt AC outlet and multiple
extension cords to reach places near the water or on some of the walking trails. The
stations themselves consist of a light colored sheet draped over a construction made from
ladders, poles, and wooden boards. Another ultraviolet light station was created by draping
a light colored sheet over the hood or trunk of a car. This light was powered by the 12 DC
car outlet. A “white” light station was usually a battery-powered lamp placed on a light
colored sheet on the ground. We typically chose a location on a trail due to the absence of
vegetation for spreading out the sheet, as well as to optimize our access. The location on
open trail areas is also likely to intercept walkways of ground-dwelling invertebrates. Once
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
11
a specimen settles onto one of the sheets, we recorded it photographically. We usually had
a team of assistants helping with setup of the stations as well as help the authors keeping
track of new specimen arrivals.
The combined results of all the data collection methods described above, consists of more
than 21,000 digital photos. The photos have been stored in an hierarchical data structure
ordered by year and date, based on each photo's date and time stamp. Representatives of
each specimen photographed have been copied into a separate hierarchical data structure,
sorted on taxonomic order. Each digital photo in the taxonomic data structure has been
assigned a filename that includes the organism's scientific name and common name,
together with other available data, including size and field-marks. This file naming strategy
enables fast, operating-system-level, text-based searches for retrieving data.
Aquatic invertebrates data collection methodology
In addition to the data collection methodologies described above, we also performed
dedicated sampling of MMP's aquatic habitats. Data collection of aquatic organisms was
performed several times a month between April and July 2010, using a combination of two
techniques:
1.
2.
Acute sampling without a net: The bulb of a large baster was fully squeezed
before inserting the tip into the water. The tip of the baster was moved around
slowly; slowly releasing the bulb to collect the sample. The resulting sample was
slowly squeezed out into a sieve with a mesh size of about 0.1 mm. This was
repeated several times until the sieve became relatively full. The sieve was then
turned upside down over a collection container, which usually was a small glass jar,
and some clear water was squeezed through the other side of the sieve, releasing
the trapped content into the jar. This process was repeated until a sample of
sufficient volume had been collected.
Large quantity sampling with a net: First, about four ounces of water were
collected and poured into a small container. Next, a rectangular, four-by-three inch
net, with approximately 1 mm mesh size, was swished back-and-forth through the
water, collecting whatever is in the water. Once the net had a full sample, it was
pulled out of the water. Large debris, such as pieces of Schoenoplectus californicus
(C. A. Mey.) Soják (California Tule) and other plant detritus, was removed manually
and the net subsequently turned inside out into the container. The sample in the net
would free itself into the water in the basin by moving the net through the water.
Holding the container at an angle helps to keep the water in one relatively small
location. This was repeated several times until a sample of sufficient volume had
been collected. The resulting content was poured into a collection jar.
These techniques were applied at a variety of locations, including small vernal pools and
the sump, which is a large basin that collects runoff water from the surrounding
neighborhood. The sampling locations were changed each week, and at these locations
various micro-habitats (open water, near emergent vegetation, near tree roots, and under
floating aquatic vegetation) were sampled at varying depths (from the shallowest to the
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Fiesler E, Drake T
deepest water), to maximize the chances of capturing organisms from a broad range of
habitats.
The samples were taken to the MMP Nature Center’s curation lab, where they were
examined by the first author, using a variety of microscopes and photographically recorded
using a number of digital imaging techniques. After examination, the samples were
returned to the Preserve. The results of the aquatic sampling have been integrated into
Suppl. material 2.
Arthropod taxonomy
As the number of described macro-invertebrate taxa is immense, estimated at more than
1,300,000 Current Results 2015, and its classification in constant flux, a continuously
updated on-line resource was used for taxonomic hierarchy and nomenclature. The
overwhelming majority of the macro-invertebrate taxa recorded during the MMP
bioinventory are arthropods. BugGuide BugGuide 2014 offers a taxonomy for arthropods
that is updated using the latest information by consent of the scientific community directly,
or indirectly, contributing to the site. The information is stored in a hierarchical taxonomic
way and is publicly available. The arthropod taxonomy presented in BugGuide, for the 2009
to 2012 period of the MMP bioinventory, is used in this publication.
Results
Before presenting the overall results, the outcome of a few selected data collection
methodologies are presented, as they tend to contain method-specific data of interest.
Pitfall trap results
The pitfall trap results yielded several species not recorded by the other methodologies.
The most striking find was Ceuthophilus hesperus clunicornis Hubbell, 1936, (San Diego
camel cricket); see Fig. 5. Given the fact that they are flightless, and hence limited in their
dispersal, it is very likely that camel crickets have been present at the site for a long time.
Camel crickets are among the least studied orthopterans (order: Orthoptera, or “straightwinged” insects), which include all grasshoppers, katydids, crickets, and allies.
Other unique pitfall trap discoveries are three small native beetles:
•
A somewhat elongated beige-brown beetle with two unusually long claws on each
foreleg. This is Lepidocnemeplatia sericea (Horn, 1870), a darkling beetle, see Fig.
6, (family: Tenebrionidae); which is fairly widespread in the Southwestern United
States, but not often encountered.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
13
Figure 5.
Ceuthophilus hesperus clunicornis Hubbell, 1936 (San Diego camel cricket)
•
•
A new family for the Preserve was the discovery of Xerosaprinus sp., a clown
beetle, see Fig. 7, in the family Histeridae.
The third beetle was a snout beetle, also known as a weevil. It was in one of the
two traps near the waterline. It is Stenopelmus rufinasus Gyllenhal, 1835, also
known as Azolla weevil or water fern weevil; see Fig. 8. It's life cycle is dependent
on the presence of the native Azolla filiculoides Lam. (Pacific mosquito fern) in the
family Azollaceae.
Figure 6.
Lepidocnemeplatia sericea (Horn, 1870) (a darkling beetle)
14
Fiesler E, Drake T
Figure 7.
Xerosaprinus sp. (a clown beetle)
Figure 8.
Stenopelmus rufinasus Gyllenhal, 1835 (Azolla weevil)
The pitfall trap closest to the waterline yielded, besides two aquatic ground beetles, many
hundreds of springtails (class: Collembola); see Figs 9, 10. The data collected during the
pitfall trap sessions have been added to the taxon list in Suppl. material 2.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
Figure 9.
Collembola (Springtails) and Linepithema humile (Mayr, 1868) (Argentine Ants)
Figure 10.
Collembola (Springtails) and Linepithema humile (Mayr, 1868) (Argentine Ants) - detail
Nighttime observations
An overview of the results of our night data collection efforts are shown in Table 1.
15
Fiesler E, Drake T
16
Table 1.
Overview tally of specimens recorded during night data collection sessions, grouped by selected
higher taxa (order, superfamily, family, etc.). Related higher taxa are grouped together and sorted in
phylogenic order Wheeler et al. 2001. Higher taxon groups have alternating background coloration
to aid readability.
Note: Numbers in boldface font indicate maxima - greater than 1 - for each taxon. (Sub)totals per
taxon, and per month, are added in the margins.
Year:
Phylum Class
Order
Taxon group
Common
name
Annelida Clitellata
Haplotaxida
Haplotaxida
Earthworms
& kin
Arachnida
Arachnida
Spiders & kin
Collembola
Collembola
Springtails
Insecta Orthoptera
Orthoptera
Grasshoppers
& kin
Dermaptera
Earwigs
Aphidoidea
Aphids
Arthopoda
Dermaptera
Hemiptera
2010
Apr May Jun Aug Sep Dec Feb Apr Aug Nov Total
1
1
Coleoptera
1
Psylloidea
Psyllids
Heteroptera
Bugs
Psocoptera
Barklice
Scarabeidae
Scarabs
Hemerobiiformia
Hymenoptera "Parasitica"
3
1
3
2
2
1
2
1
2
13
1
9
1
1
1
1
2
4
1
1
1
3
6
1
1
4
5
1
3
3
4
1
1
1
2
1
1
3
2
4
11
6
16
1
Parasitoid
Wasps
1
1
1
1
3
2
1
1
11
12
Ants
Moths
3
33
24
Chironomidae
Midges
6
15
4
1
other
Nematocera
other "lower"
Flies
9
5
3
8
Brachycera
"higher" Flies
4
2
1
3
2
25
71
49
30
38
18
2
1
Lacewings
Total:
2
1
Formicidae
Lepidoptera
Diptera
1
1
other Coleoptera other Beetles
Neuroptera
2
1
1
Auchenorrhyncha Hoppers
Psocoptera
2011
2
5
16
1
4
4
2
2
4
110
28
1
3
7
2
30
3
3
1
19
14
39
9
282
The results of the night data collection show that moths are by far the most frequently
encountered, followed by chironomidae (non-biting midges) and various other Nematocera
("lower flies"). The maximum number of specimens were encountered in May, followed by
June and September. Only five individuals, and no moths at all, were recorded for
December. The species recorded by our night data collection efforts are included in Suppl.
material 2.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
17
Aquatic invertebrates
Besides macro-invertebrates, we also recorded the presence of some of the more salient
aquatic micro-invertebrates that were caught in our water samples. These taxa have been
added to Suppl. material 2, but likely represent only a small fraction of MMP's micro-fauna.
It is difficult to compare our data one-to-one with those obtained by Mark Angelos (1993–
2004; as detailed above), due to the fact that his specimens were identified to genus or
family level and those in our study (2009–2010) to various levels ranging from family to
(sub)species. Nevertheless, we attempt to compare our data by estimating the number of
genera obtained in each study, and then comparing which genera are present in both
studies. Genera recorded during 2009–2010, but not during 1993–2004, are added in the
column labeled "gained", and genera recorded during 1993–2004, but not during 2009–
2010, are added in a column labeled "lost" The combined result is shown in Table 2.
Table 2.
Comparison of aquatic invertebrate data obtained by Mark Angelos' ("1993-2004") and our study
("2009-2010").
The numbers represent estimated number of genera. Totals can be found in the bottom-most row of
the table.
Nomenclature used: "gained" = number of genera recorded in this study, but not by Mark Angelos;
"lost" = number of genera recorded by Mark Angelos, but not in this study; "vs." = versus.
Phylum
Class
Order
1993-2004
2009-2010
"gained"
"lost"
Comments
Mollusca
Gastropoda
5
2
0
3
Arthropoda
Ostracoda
6
5
0
1 difficult to
match: higher
taxa vs.
species
Arthropoda
Branchiopoda
Anostraca
3
2
0
1
Arthropoda
Insecta
Ephemeroptera
1
1
0
0 same species
Arthropoda
Insecta
Odonata
8
10
3
1 lost: Lestes
congener
Arthropoda
Insecta
Hemiptera
7
3
0
4
Arthropoda
Insecta
Coleoptera
14
6
1
9
Arthropoda
Insecta
Diptera
24
24
8
8 difficult to
match:
matures vs.
immatures
54
44
12
22 net loss = 10
taxa = 19%
Totals:
18
Fiesler E, Drake T
It is difficult to proof that an organism is not present in a given area. There are however
some salient organisms that have not been encountered during our study or reported
during the last decade. We are fairly sure that these organisms have been extirpated, or
become extinct, at MMP. The following two salient invertebrate taxa were recorded by Mark
Angelos and have not been observed during the last decade: Lestes congener Hagen,
1861 (Spotted Spreadwing) and Mesovelia sp. (Water Treader). These losses might be due
to a decrease in water quality. Also potentially extirpated are Buenoa and Nononecta spp.
(Backswimmers), Haliplus sp. (Crawling Water Beetle), Hydraenidae (Minute Moss
Beetles), and Noteridae (Burrowing Water Beetles); although more intensive aquatic
sampling over a longer time frame might reveal the continued presence of these taxa.
Mark Angelos has donated his MMP aquatic invertebrate specimen collection, and
associated documentation, to the MMP Nature Center. This enables the possibility to
identify his specimens to species level and recreate a detailed MMP aquatic invertebrate
fauna for the period 1993–2004, as well as a higher quality comparison to MMP's current
aquatic invertebrate fauna.
Environmental conditions
To put the data collection period in perspective, we add some information on the weather
during the timeframe of the project. A lot of the data was collected in the year 2010, which
was a year with unusual weather. The spring and summer of 2010 were surprisingly
breezy, and the coolest on record, temperature-wise. However, the highest temperatures
ever measured in Los Angeles were recorded on September 27, 2010, with an all-time
record of 113 degrees Fahrenheit for the downtown area. Furthermore, in late December
2010, Los Angeles received an exceptional amount of rain from one storm-system, more
than seven inches. Invertebrates are exothermic, relying on the ambient temperature to
warm their body in order to get active. The cool and breezy weather during the peak annual
activity time was therefore not optimal for invertebrate abundance. During 2011 we
collected data until August 30th. This year (2011) also had a cool summer. We
nevertheless recorded a large amount of data, and can only expect to encounter even
larger biodiversity during more 'normal' years.
Discussion
During the project period, more than 12,500 digital photos were taken, processed,
identified, and archived. Besides those, more than 9,500 digital photos had been taken and
archived during the period from 2003 until the start of the project. A small fraction of this
older data have been processed and identified and these taxa have been included in this
report. Combining all the available data, a total of 689 invertebrate organisms were
recorded, covering 8 phyla, 13 classes, 39 orders, and 222 families. A summary of the
statistics for the highest, and some of the most salient, taxa are given in Table 3. The
orders with the largest number of recorded taxa are, in decreasing order: Diptera (155
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
19
taxa), Coleoptera (106 taxa), Hymenoptera (101 taxa), Lepidoptera (84 taxa), Hemiptera
(82 taxa), and Araneae (48 taxa).
Table 3.
Summary of order-level statistics of recorded invertebrate taxa.
The data is grouped by most salient higher-level taxa. (Sub)totals per higher-level taxon are added
in the bottom and top row, highlighted in yellow.
Class
ePhylum
8
Order
Family
taxa
Common Name (MMP taxa)
16
39
222
689 = Totals
Annelida
1
2
3
5 Annelids
Arthropoda
8
29
205
669 Arthropods
3
34
73 Arachnids
Acari
14
23 Mites
Araneae
19
48 Spiders
Solifugae
1
2 Solifugids
3
4 Fairy Shrimp, Waterfleas, & kin
Cladocera
2
2 Waterfleas
Anostraca
1
2 Fairy Shrimp
Arachnida
Branchiopoda
2
Chilopoda
1
1
1 Centipedes
Collembola
2
3
6 Springtails
16
Insecta
157
574 Insects
Coleoptera
23
106 Beetles
Dermaptera
2
2 Earwigs
Diptera
Embiidina
Ephemeroptera
36
1
1
Hemiptera
25
Hymenoptera
27
Blattodea
Lepidoptera
1
20
155 Flies, Mosquitos, & kin
1 Webspinners
1 Mayflies
82 True Bugs, Hoppers, Aphids, & kin
101 Wasps, Ants, Bees, & Sawflies
1 Termites
84 Butterflies & Moths
Microcoryphia
1
1 Bristletails
Neuroptera
3
3 Lacewings & Antlions
Odonata
3
15 Dragonflies & Damselflies
Orthoptera
5
11 Grasshoppers, Crickets & kin
Psocoptera
5
7 Barklice
Thysanoptera
3
3 Thrips
Zygentoma
1
1 Silverfish
20
Fiesler E, Drake T
Malacostraca
3
5
6 Crayfish, Scuds, Isopods, & kin
Amphipoda
2
3 Scuds
Decapoda
1
1 Decapods = Crayfish
2
2 Isopods
Maxillopoda
Isopoda
1
1
1 Copepods
Ostracoda
1
1
4 Seed Shrimp
Cnidaria
1
1
1
1 Hydroids
Mollusca
1
2
7
8 Molluscs
2
Gastropoda
Nematoda
7
8 Snails & Slugs
Stylommatophora
5
6 Terrestrial Snails & Slugs
Basommatophora
2
2 Freshwater Snails
2
2
2
2 Nematodes = Roundworms
Platyhelminthes
1
1
2
2 Flatworms
Rotifera
1
1
1
1 Rotifers = Wheel Animals
Tardigrada
1
1
1
1 Tardigrades = Water Bears
16
39
222
8
689
=Totals
All the 689 taxa are listed in Suppl. material 2, sorted alphabetically in a hierarchical way,
starting at phylum level, down to subspecific level. The list contains all invertebrate taxa
recorded, including small arthropods like aphids, thrips, and mites, as well as all recorded
microscopic aquatic taxa. Some taxa above species level appear identical in Suppl.
material 2. These taxa are based on sufficiently different morphotypes.
Of particular interest is the presence of native flightless species, that have likely persisted
throughout the human-influenced history of the land that is now Madrona Marsh Preserve,
including:
•
Typhoctes peculiaris peculiaris (Cresson, 1875), a bradynobaenid wasp, see Fig.
11; whose females are flightless,
•
Pseudomethoca anthracina (Fox, 1892), a velvet ant, see Fig. 12; whose females
are also flightless,
•
Eremobates sp., a large solifugid, see Figs 13, 14, and
•
Ceuthophilus hesperus clunicornis Hubbell, 1936, San Diego Camel Cricket, see
Fig. 5.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
Figure 11.
Typhoctes peculiaris peculiaris (Cresson, 1875) (a bradynobaenid wasp) female
Figure 12.
Pseudomethoca anthracina (Fox, 1892) (a velvet ant) male
21
22
Fiesler E, Drake T
Figure 13.
Eremobates sp. (a solifugid)
Figure 14.
Eremobates sp. (a solifugid)
Some taxa were notable for their absence, including Diplopoda (millipedes), Opiliones
(harvestmen),
Scorpiones
(scorpions),
Pseudoscorpiones
(pseudoscorpions),
cockroaches, Mantodea (mantids), Phasmida (stick insects), Nematomorpha (hairworms),
and Diplura. Diplopoda (millipedes) have been observed by Walt Wright (MMP manager
preceding Tracy Drake) in the past, but have possibly been extirpated. Similarly, only one
kind of Chilopoda (centipede) was found, and only in small numbers. Native Corydiidae
(sand roaches) are not expected at MMP due to lack of soft sand habitat, and adventive
cockroaches have been recorded in urban areas not far from the Preserve, but are closely
associated with human activity and not expected to be permanent resident on the
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
23
Preserve. Native mantids are relatively uncommon in Los Angeles County and not
expected at MMP due to its isolation within urbanization. Adventive mantids are introduced
in urban settings as pest control, and it is no surprise not to find them on the Preserve
either. Most of the remaining taxa that were not found are associated with fairly undisturbed
habitats or other habitats that are not present at the Preserve.
Some notorious and ubiquitous Los Angeles County adventives, like Ceratitis capitata
(Wiedemann, 1824) (Mediterranean Fruit Fly), Homalodisca vitripennis (Germar, 1821)
(Glassy-winged Sharpshooter), and Sophonia orientalis (Matsumura, 1912) (Oriental
Leafhopper) were not recorded from MMP. An interesting adventive species that was
recorded from the Preserve is the Outer Barklouse Ectopsocus strauchi Enderlein 1906,
see Figs 15, 16, which is native to Europe and found in eastern parts of the United States.
This finding constitutes the first west coast record for this species. Overall, relatively few
adventive taxa were identified: six introduced for bio-control, two migrants that expanded
their range, and 44 other adventive taxa, of which only a fraction might be labeled as
destructive pest. All three categories of adventive taxa combined represent less than 8 %
of MMP's invertebrate fauna.
Figure 15.
Ectopsocus strauchi Enderlein, 1906 (a barklouse)
No doubt species will be found in the future that are not on the list. These will
predominantly be new colonizers, visiting vagrants, and obscure species in under-sampled
habitats, including infauna of the soil, microscopic aquatic fauna, and ectoparasites of
vertebrates, which were outside the scope of this project. Also, as mentioned before, years
with different weather patterns are likely to reveal species that were not encountered during
the study period.
This study can nevertheless be considered quite successful, given the available time and
other resources. The list of recorded taxa will change over time, as some species come
and some go, especially winged ones. The current list constitutes a solid baseline to
Fiesler E, Drake T
24
compare future data with, and can be used as data source for educational and outreach
purposes.
Figure 16.
Ectopsocus strauchi Enderlein 1906 (a barklouse)
Acknowledgements
Gratefully acknowledged are the Friends of Madrona Marsh and the City of Torrance for
their financial support of part of this work.
References
•
•
•
•
•
•
Angelos M (2003) Madrona Marsh Fairy Shrimp Report. Prepared by Mark Angelos,
Natural History Museum, for Madrona Marsh Preserve management, City of Torrance.
unpublished: 40 pp.
Angelos M (2005) Ostracod Community Organization in Vernal Pools. Natural History
Museum of Los Angeles County unpublished presentation: September.
Bryant P (2014) Natural History of Orange County, California and nearby places. http://
nathistoc.bio.uci.edu/index.htm website: accessed December, 2014.
BugGuide (2014) BugGuide website: http://bugguide.net/. Accessed multiple times
between 2010 and 2013, and in 2014.
Cadien D, Lovell L, (editors) (2014) A Taxonomic Listing of Benthic Macro- and
Megainvertebrates from Infaunal & Epifaunal Monitoring and Research Programs in the
Southern California Bight. Edition 9. Prepared by The Southern California Association of
Marine Invertebrate Taxonomists, Natural History Museum of Los Angeles County
Research & Collections, Los Angeles, California.
Center for Biological Diversity (2015) Invertebrates. URL: http://
www.BiologicalDiversity.org/species/invertebrates/
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
•
25
City of Torrance (1976) Madrona Marsh Wildlife Habitat Study.
Current Results (2015) Number of species. URL: http://www.CurrentResults.com/
Environment-Facts/Plants-Animals/number-species.php
Fiesler E (2009) Family-level Invertebrate Inventory of the Santa Monica Mountains
National Recreation Area. Report number SMM-FII-2009-2a. Prepared for the National
Park Service. Santa Monica Mountains National Recreation Area, Thousand Oaks,
California.
Fiesler E (2010) Oxford Basin Invertebrate Study. Report number OBIS-2010-1-F.
Prepared for: Hamilton Biological, Inc., Long Beach, California.
Fiesler E (2016) Field Guide to the Insects and other Invertebrates of the Santa Monica
Mountains. version 1. National Park Service - Santa Monica Mountains National
Recreation Area. Release date: 2016 4 18. URL: http://www.nps.gov/samo/learn/
nature.htm
FoMM (2015) Friends of Madrona Marsh. URL: http://
www.FriendsofMadronaMarsh.com/
Friends of Ballona Wetlands (2015) Friends of Ballona Wetlands website. URL: http://
www.BallonaFriends.org/
Friends of Gardena Willows (2015) Friends of Gardena Willows Wetland Preserve
website. URL: http://www.GardenaWillows.org/
Frommer S (1988) A Hierarchic Listing of the Arthropods Known to Occur Within the
Deep Canyon Transect. University of California at Riverside, Entomology Research
Museum. 133 pp.
Gullison T, Hardner J, Anstee S, Meyer M (2015) Good Practices for the Collection of
Biodiversity Baseline Data. Prepared for the Multilateral Financing Institutions
Biodiversity Working Group & Cross-Sector Biodiversity Initiative.
Hamlett N (2014) The Robert J. Bernard Biological Field Station - Biota. URL: http://
www.BFS.Claremont.edu/biota/index.html
IUCN Red List (2016) International Union for Conservation of Nature and Natural
Resources Red List. http://www.iucnredlist.org/
Jarski R (2007) Words From The Wise. Skyhorse Publishing, page 269.
Kim KC, Byrne L (2006) Biodiversity loss and the taxonomic bottleneck: emerging
biodiversity science. Ecological Research 21 (6): 794‑810. DOI: 10.1007/
s11284-006-0035-7
LACSD (2015) Bixby Marshland. Sanitation Districts of Los Angeles County. URL: http://
www.LACSD.org/education/tours/bixby_marshland.asp
LaPierre L, Wright P (2000) Arthropods of the Baldwin Hills. In: Molina, KC (Ed.) "Biota
of the Baldwin Hills: An Ecological Assessment". Baldwin Hills Conservancy, p. 39-63.
URL: http://www.BHC.ca.gov/documents_1.html
LAWA (2015) Los Angeles International Airport (LAX) Dunes website. Los Angeles
World Airports. URL: www.LAWA.org/LAXDunes/
Longcore TR (1999) Terrestrial Arthropods as Indicators of Restoration Success in
Coastal Sage Scrub. Dissertation. Department of Geography, University of California,
Los Angeles, Los Angeles, 145 pp.
Loveland TR, Hutcheson HL (1995) Monitoring Changes in Landscapes from Satellite
Imagery. In: Society USG (Ed.) Our Living Resources: A Report to the Nation on the
Distribution, Abundance, and Health of U.S. Plants, Animals, and Ecosystems. U.S.
Department of Interior, National Biological Service, Washington, D.C. p. 468-473.
26
•
•
•
•
•
•
•
•
•
•
•
•
•
•
Fiesler E, Drake T
Lyle J (1999) Design for Human Ecosystems: Landscape, Land Use, and Natural
Resources. Island Press, 279 pp. [ISBN 155963720X, 9781559637206]
Mattoni R (1990) Species diversity and habitat evaluation on the El Segundo sand
dunes at LAX. Unpublished report. Los Angeles Board of Airport Commissioners.
Mattoni R (1993) Natural and Restorable Fragments of the Former El Segundo Sand
Dunes Ecosystem. In: Keeley JE (Ed.) Interface Between Ecology and Land
Development in California. Southern California Academy of Sciences, Los Angeles p.
289-294.
Mattoni R, Longcore T (1997) The Los Angeles Coastal Prairie, a vanished community.
Crossosoma 23 (2): 71‑102.
Mattoni R, Longcore T, Novotny V (2000) Environmental Auditing: Arthropod Monitoring
for Fine-Scale Habitat Analysis: A Case Study of the El Segundo Sand Dunes.
Environmental Management 25 (4): 445‑452. DOI: 10.1007/s002679910035
Nowinszky L, Puskás J (2012) Light-Trap Catch of the Harmful Moths Depending of
Moonlight in North Carolina and Nebraska States of USA. ISRN Zoology 2012: 1‑6.
DOI: 10.5402/2012/238591
NPS (2015) Channel Islands National Park website. United States National Park
Service. URL: http://www.NPS.gov/CHIS/
Pacific_Flyway (2015) Pacific Flyway. Wikipedia. URL: https://en.wikipedia.org/wiki/
Pacific_Flyway
Pierce WD, Pool D (1939) Unpublished field notes. Los Angeles County Museum of
Natural History, Los Angeles.
Pierce WD, Pool D, et.al. (1947) The Fauna and Flora of the El Segundo Sand Dunes.
part 1: Vol. 37, p. 93–97, 1938; part 2: Vol. 37, p. 98–100, 1938; part 3: Vol. 37, p. 100,
1938; part 4: Vol. 37, p. 101–104, 1938; part 5: Vol. 38, p. 39–42, 1939; part 6: Vol. 38,
p. 43–53, 1939; part 7: Vol. 38, p. 112–119, 1939; part 8: Vol. 38, p. 119, 1939; part 9:
Vol. 38, p. 191–197, 1939; part 10: Vol. 38, p. 198–200, 1939; part 11: Vol. 39, p. 6–27,
1940; part 12: Vol. 40, p. 14–28, 1941; part 13: Vol. 40, p. 29–38, 1941; part 14: Vol. 42,
p. 1–30, 1943; part 15: Vol. 42, p. 90–103, 1943; part 16: Vol. 46, p. 51–53, 1947; part
17: Vol. 46, p. 53–54, 1947. Bulletin of the Southern California Academy of Sciences.
Pocock MO, Newson S, Henderson I, Peyton J, Sutherland W, Noble D, Ball S,
Beckmann B, Biggs J, Brereton T, Bullock D, Buckland S, Edwards M, Eaton M, Harvey
M, Hill M, Horlock M, Hubble D, Julian A, Mackey E, Mann D, Marshall M, Medlock J,
O'Mahony E, Pacheco M, Porter K, Prentice S, Procter D, Roy H, Southway S, Shortall
C, Stewart AA, Wembridge D, Wright M, Roy D (2015) Developing and enhancing
biodiversity monitoring programmes: a collaborative assessment of priorities. Journal of
Applied Ecology 52 (3): 686‑695. DOI: 10.1111/1365-2664.12423
Pratt G (2000) Terrestrial Arthropods of Edwards Air Force Base, 1996–1998. U.S.
Army Engineer Research and Development Center Final Report ERDC/EL TR-00-20,
Edwards Air Force Base, Edwards, CA.
PVPLC (2015) Linden H. Chandler Preserve website. Palos Verdes Peninsula Land
Conservancy. URL: http://www.PVPLC.org/_lands/Chandler.asp
Schreiber RW (1981) Biota of the Ballona region - Los Angeles County. Marina del Rey
/ Ballona Local Coastal Plan, supplement 1. Los Angeles County Natural History
Museum Foundation.
Macro-invertebrate Biodiversity of a Coastal Prairie with Vernal Pool Habitat
•
•
•
•
•
•
•
27
Schuldt A, Assmann T (2010) Invertebrate diversity and national responsibility for
species conservation across Europe – A multi-taxon approach. Biological Conservation
143 (11): 2747‑2756. DOI: 10.1016/j.biocon.2010.07.022
SMMNRA (2014) Santa Monica Mountains National Recreation Area website. URL: htt
p://www.NPS.gov/SAMO/
Stromberg M, Kephart P, Yadon V (2002) Composition, invasability, and diversity in
coastal California grasslands. Madroño 48: 236‑252.
TD (2011) San Diego Bay Integrated Natural Resources Management Plan. Tierra Data
Inc. Prepared for U.S. Department of the Navy. Escondido, California.
U.S. Climate Data (2015) http://www.usclimatedata.com/
Wheeler W, Whiting M, Wheeler Q, Carpenter J (2001) The Phylogeny of the Extant
Hexapod Orders. Cladistics 17 (2): 113‑169. DOI: 10.1111/j.1096-0031.2001.tb00115.x
Wikipedia (2016) https://en.wikipedia.org/wiki/Madrona_Marsh
Supplementary materials
Suppl. material 1: Supplementary file 1 = Selected biological surveys and inventories,
that include macro-invertebrates, for Southern California
Authors: E. Fiesler
Data type: data spreadsheet
Brief description: The county, or counties, where the survey took place is highlighted in light
green for surveys areas including Los Angeles County.
Abbreviations and symbols used: A = arachnids & myriapods; B = Beach; F = fresh-water
invertebrates; H = hexapods (= insects and kin); ha = hectares; invert. = invertebrates; LA = Los
Angeles County; LAX = Los Angeles International Airport; "n/a" (non-applicable; highlighted in
orange) = survey range does not include this habitat; M = mollusks; MI = marine invertebrates; OC
= Orange County; PV = Palos Verdes Peninsula; SD = San Diego County; SRA = State
Recreation Area; "unk." = unknown; "~" = approximately; ">" = greater than; "+" = at least.
Filename: Supp-1 [was Table 1]= CA~S biosurveys IMAGE version-2.jpg - Download file (272.80
kb)
Suppl. material 2: Supplementary file 2 = Annotated taxonomic List of MMP
Invertebrates
Authors: E. Fiesler
Data type: taxa checklist
Brief description: The following is a list of all invertebrates recorded at Madrona Marsh
Preserve for the project described in this document.
For each taxon, the columnar data in table in the Appendix consists of:
Scientific name & taxonomy: Phylum, Class, Order, Family, Tribe, Genus, Species, Subspecies /
Variant
Common Name
Whether the taxon is adventive / non-native
Whether the specimen was observed at night
Abbreviations used: A = adventive taxon; B = introduced for biocontrol; cf. = (Latin: confer) =
compare; E = migrant that expanded its range; imm = immature; N = recorded after sunset; n/a =
28
Fiesler E, Drake T
not applicable; nr. = near; S. Cal. = Southern California; s.l. = (Latin: sensu lato) = broadly; var. =
variant.
Color coding used: GREEN = high taxonomic certainty; YELLOW = intermediate taxonomic
certainty; ORANGE = low taxonomic certainty.
Filename: Supp-2 (was Table 2)= MMP Invertebrates taxa table version-2.xls - Download file
(247.50 kb)
Supplementary material 1:
Selected biological surveys and inventories, that include macro-invertebrates, for
Southern California
Author:
E. Fiesler
Description:
The county, or counties, where the survey took place is highlighted in light green
for surveys areas including Los Angeles County.
Abbreviations and symbols used:
A = arachnids & myriapods;
B = Beach;
F = fresh-water invertebrates;
H = hexapods (= insects and kin);
ha = hectares;
invert. = invertebrates;
LA = Los Angeles County;
LAX = Los Angeles International Airport;
"n/a" (non-applicable; highlighted in orange) = survey range does not include this
habitat;
M = mollusks;
MI = marine invertebrates;
OC = Orange County;
PV = Palos Verdes Peninsula;
SD = San Diego County;
SRA = State Recreation Area;
"unk." = unknown;
"~" = approximately;
">" = greater than;
"+" = at least.
Supplementary material 2: Annotated taxonomic List of MMP Invertebrates
Author: Emile Fiesler
Data type: taxa checklist
Brief description: The following is a list of all invertebrates recorded at Madrona
Marsh Preserve for the project described in this document.
For each taxon, the columnar data in table in the Appendix consists of:
 Scientific name & taxonomy: Phylum, Class, Order, Family, Tribe, Genus,
Species, Subspecies / Variant
 Common Name
 Whether the taxon is adventive / non-native
 Whether the specimen was observed at night
Abbreviations used:
A = adventive taxon;
B = introduced for biocontrol;
cf. = (Latin: confer) = compare;
E = migrant that expanded its range;
N = recorded after sunset;
n/a = not applicable;
nr. = near;
S. Cal. = Southern California;
s.l. = (Latin: sensu lato) = broadly;
var. = variant.
Color coding used:
GREEN = high taxonomic certainty;
YELLOW = intermediate taxonomic certainty;
ORANGE = low taxonomic certainty.
Higher
Family
Subfamily
Tribe
taxa
Phylum: Annelida = Annelids
Class: Clitellata = Annelid Worms
Order: Haplotaxida = Earthworms & kin
Lumbricidae
Lumbricinae
n/a
Lumbricidae
Lumbricinae
Lumbricini
Naididae
Naidinae
Derini
Naididae
Naidinae
Derini
Order: Rhynchobdellida = Jawless Leeches
Glossiphoniidae
Haementeriinae n/a
Genus
Species
Aporrectodea
Lumbricus
Dero
Aulophorus
trapezoides
terrestris
limosa
vagus
Helobdella
triserialis
ssp. / var. / A Ni
form
d g
A
A N
Phylum: Arthropoda = Arthropods
Class: Arachnida = Arachnids = Spiders, Mites, Ticks, Harvestmen, Scorpions, & kin
Order: Acari = Mites & Ticks
Acaridae
Tyrophaginae
Tyrophagini
Tyrophagus
Anystidae
Anystinae
Anystini
Anystis
Anystidae
Anystinae
Anystini
Anystis
Anystidae
Erythracarinae
Bdellidae
Bdellidae
Bdellinae
n/a
Bdella
Bdellidae
Odontoscirinae
Bdellidae
Odontoscirinae n/a
Bdellodes
Cunaxoidinae
Digamasellidae
Dendrolaelapinae Dendrolaelapini Dendrolaelaps
Eriophyidae
Eriophyinae
Aceriini
Aceria
boycei
Tetranychidae
Bryobiinae
Bryobiini
Bryobia
praetiosa
Eupodidae
Eupodinae
n/a
Linopodes
Erythraeidae
Erythraeinae
Erythraeidae
Balaustiinae
n/a
Balaustium
Erythraeidae
Balaustiinae
n/a
Balaustium
Erythraeidae
Balaustiinae
n/a
Balaustium
aonidophagus
Parasitidae
Penthaleidae
n/a
n/a
Penthaleus
major
Tetranychidae
Tetranychinae
Tetranychini
Oligonychus
platani
Trombidiidae
Trombidiidae
Tydeidae
Order: Aranea = Spiders
Agelenidae
n/a
Amaurobiidae
Amaurobiinae
Anyphaenidae
Anyphaeninae
Araneidae
Araneinae
Araneidae
Araneinae
Araneidae
Araneinae
Araneidae
Araneinae
Araneidae
Araneinae
Araneidae
Argiopinae
Corinnidae
Castianeirinae
Desidae
n/a
Dysderidae
Dysderinae
Gnaphosidae
Herpyllinae
Gnaphosidae
Drassodinae
Gnaphosidae
Drassodinae
Gnaphosidae
Herpyllinae
Gnaphosidae
Zelotinae
Gnaphosidae
Zelotinae
Linyphiidae
Linyphiidae
Erigoninae
Linyphiidae
Erigoninae
Lycosidae
Lycosinae
Lycosidae
Lycosinae
Lycosidae
Pardosinae
Lycosidae
Pardosinae
Lycosidae
Piratinae
Miturgidae
Eutichurinae
Oecobiidae
n/a
Oxyopidae
n/a
Oxyopidae
n/a
Philodromidae
Philodrominae
Pholcidae
n/a
Pholcidae
n/a
Salticidae
Heliophaninae
Salticidae
Pelleninae
Salticidae
Pelleninae
Salticidae
Pelleninae
Salticidae
Pelleninae
Salticidae
Pelleninae
Salticidae
Dendryphantinae
Salticidae
Dendryphantinae
n/a
n/a
n/a
Araneini
Mangorini
Mangorini
Araneini
Araneini
n/a
n/a
n/a
Dysderini
n/a
Drassodini
Drassodini
n/a
Zelotini
Zelotini
Hololena
Amaurobius
Anyphaena
Araneus
Neoscona
Neoscona
Araneus
Araneus
Argiope
Castianeira
Badumna
Dysdera
Scotophaeus
Drassodes
Drassyllus
Sergiolus
Trachyzelotes
Zelotes
n/a
n/a
n/a
n/a
Pardosini
Pardosini
n/a
n/a
n/a
n/a
n/a
Thanatini
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
n/a
Erigone
Spirembolus
Alopecosa
Schizocosa
Pardosa
Pardosa
Pirata
Cheiracanthium
Oecobius
Oxyopes
Peucetia
Tibellus
Psilochorus
Psilochorus
Menemerus
Habronattus
Habronattus
Habronattus
Habronattus
Habronattus
Phidippus
Sassacus
N
agastus
crebrispina
N
crucifera
oaxacensis
andrewsi
gemma
trifasciata
nr cingulata
longinqua
crocata
blackwalli
A
A
A N
gertschi
lyonneti
dentosa
kochii
mccooki
sternalis
piraticus
inclusum
navus
salticus
viridans
chamberlini
caiforniae
nr. hesperus
bivittatus
americanus
americanus
pyrrithrix
signatus
tarsalis
audax
vitis
N
N
A
A
Higher
Family
taxa
Tetragnathidae
Theridiidae
Theridiidae
Theridiidae
Theridiidae
Theridiidae
Thomsidae
Subfamily
Tetragnathinae
Latrodectinae
Latrodectinae
Latrodectinae
Theridiinae
Theridiinae
Thomisinae
Order: Solpugida = Sun Spiders
Eremobatidae
Eremobatinae
Eremobatidae
Therobatinae
Tribe
n/a
n/a
n/a
n/a
Theridiini
Theridiini
Diaeini
Genus
Tetragnatha
Crustulina
Latrodectus
Steatoda
Theridion
Theridion
Xysticus
Species
versicolor
sticta
hesperus
grossa
melanurum
murarium
californicus
n/a
n/a
Eremobates
Hemerotrecha
kraepelini
californica
Class: Branchiopoda = Fairy Shrimp, Waterfleas, & kin
Order: Anostraca = Fairy Shrimp
Branchinectidae
Branchinectinae n/a
Branchinecta
Branchinectidae
Branchinectinae n/a
Branchinecta
Order: Cladocera = Waterfleas
Chydoridae
Chydorinae
Daphniidae
Daphniinae
n/a
n/a
Chydorus
Daphnia
ssp. / var. / A Ni
form
d g
A
N
N
lindahli
lindahli x sandiegoensis
laevis
Class: Chilopoda = Centipedes
Order: Lithobiomorpha = Stone Centipedes
Henicopidae
Class: Collembola = Springtails
Order: Entomobryomorpha = Elongate-bodied Springtails
Entomobryidae
Entomobryinae
Entomobryini Entomobrya
Entomobryidae
Seirinae
Seirini
Seira
Isotomidae
Proisotominae
n/a
Ballastrura
Isotomidae
Proisotominae
n/a
Folsomia
Order: Symphypleona = Globular Springtails
Dicyrtomidae
Dicyrtominae
Dicyrtomidae
Ptenothricinae
n/a
Class: Insecta = Insects
Order: Coleoptera = Beetles
Anobiidae
Ernobiinae
Anobiidae
Ernobiinae
Anobiidae
Ptilininae
Bostrichidae
Psoinae
Cantharidae
Cantharinae
Carabidae
Carabinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Harpalinae
Carabidae
Trechinae
Cerambycidae
Cerambycinae
Chrysomelidae
Bruchinae
Chrysomelidae
Cassidinae
Chrysomelidae
Chrysomelinae
Chrysomelidae
Cryptocephalinae
Chrysomelidae
Cryptocephalinae
Chrysomelidae
Eumolpinae
Chrysomelidae
Eumolpinae
Chrysomelidae
Galerucinae
Chrysomelidae
Galerucinae
Chrysomelidae
Galerucinae
Chrysomelidae
Galerucinae
Coccinellidae
Chilocorinae
Coccinellidae
Chilocorinae
Coccinellidae
Coccinellinae
Coccinellidae
Coccinellinae
Coccinellidae
Coccinellinae
Coccinellidae
Coccinellinae
Coccinellidae
Coccinellinae
Coccinellidae
Scymninae
Coccinellidae
Scymninae
Coccinellidae
Scymninae
Corylophidae
Corylophinae
Corylophidae
Corylophidae
Corylophinae
N
Ptenothix
Ernobiini
Ernobius
Ozognathini
Ozognathus
Ptilinini
Ptilinus
Polycaonini
Polycaon
Cantharini
Pacificanthia
Carabini
Calosoma
Harpalini
Harpalus
Harpalini
in Stenolophina-subtribe
Lebiini
Apristus
Lebiini
Axinopalpus
Platynini
Agonum
Platynini
Agonum
Platynini
Calathus
Pterostichini
Pristonychus
Zabrini
Amara
Zabrini
Amara
Bembidiini
Bembidion
Phoracanthini Phoracantha
n/a
Stator
Chalepini
Stenopodius
Chrysomelini Trachymela
Cryptocephalini Diachus
Cryptocephalini Pachybrachis
Eumolpini
Chrysochus
Typophorini
Paria
Alticini
Crepidodera
Luperini
Diabrotica
Luperini
Diabrotica
Luperini
Diabrotica
Chilocorini
Axion
Chilocorini
Chilocorus
Coccinellini
Coccinella
Coccinellini
Cycloneda
Coccinellini
Harmonia
Coccinellini
Hippodamia
Psylloborini
Psyllobora
Scymnini
Cryptolaemus
Scymnini
Scymnus
Stethorini
Stethorus
Sericoderini
Sericoderus
Sericoderini
atrocincta
bipunctata
Sericoderus
N
cornutus
flavipennis
stoutii
consors
semilaeve
biplagiatus
ruficollis
complanatus
ruficollis
immunda
recurva
pruininus
insularis
sloanei
auratus
turbidus
cobaltinus
N
N
A
A
opulenta
balteata
(blue variety)
undecimpunctatahowardi
undecimpunctataundecimpunctata
plagiatum
bipustulatus
septempunctata
B
sanguinea
axyridis
B
convergens
vigintimaculata
montrouzieri
B
loewii
punctum
picipes
debilis
quadratus
Higher
Family
taxa
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Curculionidae
Cryptophagidae
Cryptophagidae
Dermestidae
Dytiscidae
Dytiscidae
Dytiscidae
Dytiscidae
Dytiscidae
Heteroceridae
Histeridae
Hydrophilidae
Latridiidae
Latridiidae
Latridiidae
Meloidae
Melyridae
Melyridae
Mycetophagidae
Monotomidae
Scarabaeidae
Scarabaeidae
Scarabaeidae
Scarabaeidae
Scarabaeidae
Scarabaeidae
Scarabaeidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Staphylinidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Tenebrionidae
Subfamily
Cossoninae
Curculioninae
Curculioninae
Curculioninae
Curculioninae
Dryophthorinae
Dryophthorinae
Dryophthorinae
Dryophthorinae
Entiminae
Entiminae
Erirhininae
Erirhininae
Erirhininae
Atomariinae
Cryptophaginae
Dermestinae
Agabinae
Colymbetinae
Hydroporinae
Hydroporinae
Colymbetinae
Tribe
Rhyncolini
Anthonomini
Erirhinini
Erirhinini
Tychiini
Sphenophorini
Sphenophorini
Sphenophorini
Sphenophorini
Naupactini
Peritelini
Sitonini
Stenopelmini
Stenopelmini
Atomariini
Cryptophagini
Dermestini
Agabini
Genus
Species
Epimechus
Anthonomus
Smicronyx
Tychius
Sphenophorus
Sphenophorus
Sphenophorus
Sphenophorus
Naupactus
Thinoxenus
Sitona
Notiodes
Stenopelmus
Atomaria
Cryptophagus
Dermestes
Agabus
mobilis
nr tenuis
sordidus
lineelus
australis
phoeniciensis
simplex
venatus
cervinus
squalens
cylindricollis
punctatus
rufinasus
puella
Bidessini
Hydroporini
Colymbetini
Liodessus
Hygrotus
Rhantus
Lanternarius
Saprininae
n/a
Xerosaprinus
Hydrophilinae
Hydrophilini
Tropisternus
Corticariinae
Corticariini
Corticaria
Corticariinae
Corticariini
Corticarina
Latridiinae
Latridiini
Aridius
Meloinae
Epicautini
Epicauta
Malachiinae
Malachiini
Attalus
Malachiinae
Malachiini
Collops
Mycetophaginae Typhaeini
Typhaea
Monotominae
Monotomini
Monotoma
Aphodiinae
Aphodiini
Aphodius
Aphodiinae
Aphodiini
Aphodius
Aphodiinae
Aphodiini
Dellacasiellus
Cetoniinae
Gymnetini
Cotinis
Dynastinae
Cyclocephalini Cyclocephala
Melolonthinae
Sericini
Serica
Melolonthinae
Sericini
Serica
Aleocharinae
Aleocharinae
Aleocharinae
Aleocharinae
Athetini
Philhygra
Aleocharinae
Lomechusini Meronera
Oxytelinae
Oxytelinae
Thinobiini
Carpelimus
Paederinae
Paederini
Paederus
Paederinae
Pinophilini
Pinophilus
Staphylininae
Staphylinini
Dinothenarus
Staphylininae
Staphylinini
Ocypus
Staphylininae
Staphylinini
Platydracus
Opatrinae
Opatrini
Blapstinus
Pimeliinae
Cnemeplatiini Lepidocnemeplatia
Pimeliinae
Coniontini
Coniontis
Pimeliinae
Coniontini
Coniontis
Pimeliinae
Edrotini
Pimeliinae
Edrotini
Pimeliinae
Nyctoporini
Nyctoporis
Tenebrioninae
Amphidorini
Eleodes
Tenebrioninae
Amphidorini
Eleodes
Tenebrioninae
Amphidorini
Eleodus
Tenebrioninae
Ulomini
Tenebrioninae
Ulomini
Uloma
Order: Dermaptera = Earwigs
Anisolabididae
Carcinophorinae n/a
Forficulidae
Forficulinae
n/a
Order: Diptera = Flies, Mosquitos, & kin
Agromyzidae
Agromyzinae
Agromyzini
Agromyzidae
Agromyzinae
Ophiomyiini
Agromyzidae
Agromyzinae
Ophiomyiini
Agromyzidae
Phytomyzinae
n/a
Agromyzidae
Phytomyzinae
n/a
Agromyzidae
Phytomyzinae
n/a
Antomyiidae
Antomyiidae
Antomyiidae
Antomyiidae
Anthomyiinae
Anthomiini
Euborellia
Forficula
Agromyza
Melanagromyza
Melanagromyza
Cerodontha
Liriomyza
Phytobia
ssp. / var. / A Ni
form
d g
A
australis
glyceriae
A
N
A
N
talpinus
gutticollis
brunneus
N
N
lateralis
serrata
herbivagans
nodifer
unicolor
vittatus
stercorea
picipes
fimetarius
militaris
pseudofucosus
mutabilis
longula
N
A
N
N
N
N
compotens
saphyrinus
olens
tarsalis
brevicollis
sericea
N
N
carinata
acuticaudus
littoralis
osculans
longula
annulipes
auricularia
A N
minimoides
virides
pictella
N
N
Higher
Family
taxa
Antomyiidae
Antomyiidae
Asilidae
Asilidae
Asilidae
Asilidae
Asilidae
Bibionidae
Bibionidae
Bibionidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Bombyliidae
Calliphoridae
Calliphoridae
Calliphoridae
Cecidomyiidae
Cecidomyiidae
Chamaemyiidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chironomidae
Chloropidae
Chloropidae
Chloropidae
Chloropidae
Chloropidae
Culicidae
Culicidae
Culicidae
Dolichopodidae
Dolichopodidae
Dolichopodidae
Dolichopodidae
Dolichopodidae
Dolichopodidae
Dolichopodidae
Drosophilidae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Ephydridae
Fanniidae
Fanniidae
Heleomyzidae
Heleomyzidae
Heleomyzidae
Heleomyzidae
Lauxaniidae
Lauxaniidae
Lauxaniidae
Limoniidae
Limoniidae
Limoniidae
Lonchaeidae
Muscidae
Muscidae
Muscidae
Muscidae
Subfamily
Anthomyiinae
Anthomyiinae
Asilinae
Asilinae
Asilinae
Asilinae
Dasypogoninae
Bibioninae
Bibioninae
Bibioninae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Anthracinae
Toxophorinae
Calliphorinae
Calliphorinae
Polleniinae
Cecidomyiinae
Cecidomyiinae
Chamaemyiinae
Tribe
Anthomiini
Anthomiini
Asilini
Asilini
Asilini
Asilini
Dasypogonini
n/a
n/a
n/a
Exoprosopini
Villini
Villini
Villini
Villini
Villini
Villini
Villini
Villini
Gerontini
Luciliini
Luciliini
Polleniini
Cecidomyiidi
Lasiopteridi
Leucopini
Genus
Chironominae
Chironominae
Chironominae
Chironominae
Chironominae
Chironominae
Chironominae
Chironominae
Tanypodinae
Chloropinae
Chloropinae
Chloropinae
Oscinellinae
Siphonellopsinae
Culicinae
Culicinae
Culicinae
Diaphorinae
Dolichopodinae
Dolichopodinae
Hydrophorinae
Medeterinae
Medeterinae
Sciapodinae
Drosophilinae
Ephydrinae
Ephydrinae
Ephydrinae
Ephydrinae
Hydrelliinae
Hydrelliinae
Hydrelliinae
Hydrelliinae
Parydrinae
n/a
n/a
Heleomyzinae
Suilliinae
Trixoscelidinae
Trixoscelidinae
Homoneurinae
Lauxaniinae
Lauxaniinae
Chioneinae
Chioneinae
Chioneinae
Lonchaeinae
Chironomini
Chironomini
Chironomini
Chironomini
Chironomini
Chironomini
Chironomini
Chironomini
Procladiini
n/a
n/a
n/a
Oscinellini
n/a
Culicini
Culicini
Culicini
n/a
Dolichopodini
Dolichopodini
Hydrophorini
Medeterini
Medeterini
n/a
Drosophilini
Ephydrini
Ephydrini
Scatellini
Scatellini
Hydrelliini
Hydrelliini
Ilytheini
Notiphilini
n/a
n/a
n/a
Heleomyzini
Suilliini
Trixoscelidini
Trixoscelidini
n/a
n/a
n/a
Eriopterini
Eriopterini
Eriopterini
Lonchaeini
Axarus
Chironomus
Chironomus
Chironomus
Chironomus
Chironomus
Chironomus
Procladius
Chlorops
Thaumatomyia
Thaumatomyia
nr Oscinella
nr Apotropina
Culex
Culex
Culiseta
Achradocera
Dolichopus
Pelastoneurus
Hydrophorus
Medetera
Medetera
Condylostylus
Drosophila
Ephydra
Ephydra
Paracoenia
Scatella
Hydrellia
Hydrellia
Ilythea
Notiphila
Brachydeutera
Fannia
Fannia
Pseudoleria
Suillia
Trixoscelis
Trixoscelis
Homoneura
Meiosimyza
Minettia
Erioptera
Erioptera
Symplecta
Lonchaea
Azeliinae
Coenosiinae
Coenosiinae
Reinwardtiini
Coenosiini
Coenosiini
Muscina
Coenosia
Coenosia
cf. Anthomyia
Machimus
Mallophora
Negasilus
Stenopogon
Saropogon
Dilophus
Dilophus
Dilophus
Ligyra
Hemipenthes
Thyridanthrax
Thyridanthrax
Villa
Villa
Villa
Villa
Villa
Geron
Lucilia
Lucilia
Pollenia
Rhopalomyia
Leucopis
Species
ssp. / var. / A Ni
form
d g
fautrix
astutus
luteus
orbatus
strigilatus
tibialis
gazophylax
sinuosa
atratus
nugator
nr. agrippina
agrippina
atrata
nr. inops
lateralis
nigripes
mexicana
sericata
rudis
americana
N
N
N
N
N
N
N
N
N
N
cf. Holotanypus
pipiens
tarsalis
incidens
arcuata
N
N
quinquefasciatusN
N
N
occidentalis
similis
californiensis
N
N
gracilis
niveiceps
stagnalis
nr caudata
argentata
N
N
nuda
occidentalis
flaveola
caliptera
pilipes
cana
striatifrons
N
N
dorsilinea
N
N
Higher
Family
taxa
Muscidae
Muscidae
Muscidae
Muscidae
Muscidae
Muscidae
Mycetophilidae
Mycetophilidae
Mycetophilidae
Mycetophilidae
Phoridae
Platypezidae
Psychodidae
Rhagionidae
Sarcophagidae
Sarcophagidae
Sarcophagidae
Sarcophagidae
Sarcophagidae
Scatopsidae
Sciaridae
Sciaridae
Sciaridae
Sciomyzidae
Sciomyzidae
Sciomyzidae
Sepsidae
Stratiomyidae
Stratiomyidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Syrphidae
Tachinidae
Tachinidae
Tachinidae
Tachinidae
Tachinidae
Tachinidae
Tachinidae
Tachinidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Tephritidae
Therevidae
Therevidae
Therevidae
Tipulidae
Tipulidae
Tipulidae
Subfamily
Coenosiinae
Coenosiinae
Coenosiinae
Coenosiinae
Coenosiinae
Muscinae
Tribe
Coenosiini
Coenosiini
Coenosiini
Coenosiini
Coenosiini
Reinwardtiini
Genus
Coenosia
Coenosia
Coenosia
Coenosia
Coenosia
Synthesiomyia
Species
Mycetophilinae
Sciophilinae
Sciophilinae
Metopininae
Platypezidae
Psychodinae
Rhagioninae
Miltogramminae
Sarcophaginae
Sarcophaginae
Sarcophaginae
Sarcophaginae
Scatopsinae
Mycetophilini
Leiini
Leiini
Metopinini
n/a
Psychodini
n/a
Miltogrammini
Mycetophila
Leia
Leia
Megaselia
Plesioclythia
Psychoda
Chrysopilus
Senotainia
perita
striata
Sciarinae
Sciarinae
Sciomyzinae
Sciomyzinae
Tetanocerinae
Sepsinae
Sciarini
Bradysiini
Sciomyzini
Tetanocerini
n/a
Sepsini
Sciara
Bradysia
Pherbellia
Sepedon
Dictya
Themira
Hermetiinae
Eristalinae
Eristalinae
Eristalinae
Eristalinae
Eristalinae
Eristalinae
Syrphinae
Syrphinae
Syrphinae
Syrphinae
Syrphinae
Syrphinae
Syrphinae
Dexiinae
Dexiinae
Dexiinae
Exoristinae
Phasiinae
Phasiinae
Tachininae
Tachininae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Tephritinae
Therevinae
Therevinae
Therevinae
Tipulinae
Tipulinae
Tipulinae
Hermetiini
Eristalini
Eristalini
Eristalini
Eristalini
Eristalini
Xylotini
Paragini
Syrphini
Syrphini
Syrphini
Syrphini
Toxomerini
Toxomerini
Dexiini
Voriini
Zeliini
Hermetia
Eristalis
Eristalis
Lejops
Palpada
Palpada
Syritta
Paragus
Allograpta
Eupeodes
Scaeva
Sphaerophoria
Toxomerus
Toxomerus
Ptilodexia
Raviniini
Ravinia
Raviniini
Ravinia
Rhegmoclematini
Rhegmoclemina
ssp. / var. / A Ni
form
d g
tigrina
nr. tigrina
nr. tigrina
nr. tigrina
nudiseta
N
N
var. a
scalaris
agarici
N
nana
N
stimulans
melanderi
N
N
vitalis
pacifica
montana
putris
S. Cal. clinal form
A
illucens
nemorum
polygrammus
alhambra
mexicana
pipiens
haemorrhous
obliqua
volucris
pyrastri
contigua
marginatus
occidentalis
A
Zelia
Gymnosomatini Gymnosoma
Phasiini
Phasia
Dejeaniini
Archytas
Tachinini
Peleteria
Eutretini
Paracantha
Tephrellini
Trupanea
Tephrellini
Trupanea
Tephritini
Euaresta
Tephritini
Euaresta
Tephritini
Neotephritis
Tephritini
Dioxyna
Tephritini
Tephritis
Terelliini
Neaspilota
Cyclotelini
Ozodiceromyia
Therevini
Thereva
Therevini
Thereva
Tipulini
Nephrotoma
Tipulini
Tipula
Tipulini
Tipula
occidentale
aldrichii
apicifer
Order: Embiidina = Webspinners
Oligotomidae
n/a
n/a
Haploembia
solieri
Order: Ephemeroptera = Mayflies
Baetidae
Baetinae
Baetini
Callibaetis
pictus
Order: Hemiptera = True Bugs, Cicadas, Hoppers, Aphids, & kin
Aleyrodidae
Aleyrodinae
Aleyrodini
Aleyrodes
Aleyrodidae
Aleyrodinae
Aleyrodini
Pealius
Alydidae
Alydinae
n/a
Alydus
Alydidae
Alydinae
n/a
Alydus
Anthocoridae
Anthocorinae
Anthocorini
Anthocoris
Anthocoridae
Anthocorinae
Oriini
Orius
Aphididae
cultaris
femoralis
wheeleri
aequalis
stigmatica
finalis
picciola
baccharis
brunneostigmata
cf. signatipennis
comata
suturalis
oleracea
planicornis
spiroeoides
kelloggi
conspersus
eurinus
antevolens
tristicolor
wulpiana
A N
A
B
Higher
Family
taxa
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aphididae
Aradidae
Berytidae
Cercopidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Cicadellidae
Coreidae
Corixidae
Delphacidae
Delphacidae
Delphacidae
Geocoridae
Geocoridae
Gerridae
Lygaeidae
Lygaeidae
Membracidae
Miridae
Miridae
Miridae
Miridae
Miridae
Miridae
Miridae
Nabidae
Pentatomidae
Pentatomidae
Pentatomidae
Pentatomidae
Psyllidae
Psyllidae
Psyllidae
Psyllidae
Reduviidae
Reduviidae
Reduviidae
Rhopalidae
Rhopalidae
Rhopalidae
Rhopalidae
Rhyparochromidae
Rhyparochromidae
Saldidae
Tingidae
Triozidae
Triozidae
Subfamily
Tribe
Aphidinae
Macrosiphini
Aphidinae
Aphidini
Aphidinae
Aphidini
Aphidinae
Aphidini
Aphidinae
Macrosiphini
Aphidinae
Macrosiphini
Aphidinae
Macrosiphini
Aphidinae
Macrosiphini
Aphidinae
Macrosiphini
Aphidinae
Macrosiphini
Chaitophorinae Siphini
Eriosomatinae
Aradinae
n/a
Metacanthinae
Metacanthini
Clastopterinae
Clastopterini
Cicadellinae
Cicadellini
Cicadellinae
Cicadellini
Deltocephalinae
Deltocephalinae Acinopterini
Deltocephalinae Athysanini
Deltocephalinae Athysanini
Deltocephalinae Deltocephalini
Deltocephalinae Deltocephalini
Deltocephalinae Macrostelini
Deltocephalinae Macrostelini
Deltocephalinae Pendarini
Deltocephalinae Scaphoideini
Deltocephalinae Scaphytopiini
Idiocerinae
Idiocerini
Ledrinae
Xerophloeini
Megophthalminae Agalliini
Typhlocybinae
Dikraneurini
Typhlocybinae
Dikraneurini
Typhlocybinae
Empoascini
Typhlocybinae
Empoascini
Coreinae
n/a
Corixinae
n/a
Delphacinae
Delphacini
Delphacinae
Delphacini
Delphacinae
Delphacini
Geocorinae
n/a
Geocorinae
n/a
Gerrinae
Gerrini
Lygaeinae
n/a
Orsillinae
Nysiini
Smiliinae
Smiliini
Bryocorinae
Dicyphini
Mirinae
Mirini
Mirinae
Mirini
Mirinae
Mirini
Mirinae
Mirini
Philinae
Philini
Philinae
Philini
Nabinae
Nabini
Pentatominae
Antestiini
Pentatominae
Pentatomini
Pentatominae
Pentatomini
Pentatominae
Strachiini
Genus
Uroleucon
Aphis
Aphis
Aphis
Species
Acyrthosiphon
Sitobion
Uroleucon
Uroleucon
Sipha
pisum
ssp. / var. / A Ni
form
d g
cornifoliae
gossypii
cf. lambersius
sonchellum
maydis
N
A
N
Aradus
Jalysus
Clastoptera
Draeculacephala
Giprus
wickhami
juniperina
minerva
Acinopterus
Colladonus
Texananus
Deltocephalus
Polyamia
Macrosteles
Macrosteles
Paraphlepsius
Osbornellus
Scaphytopius
Idiocerus
Xerophloea
Ceretagallia
Alconeura
Dikraneura
Empoasca
Empoasca
Leptoglossus
Corisella
Delphacodes
Javesella
Stobaera
Geocoris
Isthmocoris
Gerris
Melanopleurus
Nysius
Telamona
Tupiocoris
Closterotomus
Lygus
Lygus
Taylorilygus
angulatus
reductus
latipex
grex
Spanagonicus
Nabis
Thyanta
Chlorochroa
Murgantia
Bagrada
albofasciatus
alternatus
pallidovirens
sayi
histrionica
hilaris
N
N
N
N
quadrilineatus
cf. severini
nr. borealis
irroratus
nr. pallidus
N
quadrimaculata
N
N
fabae
elongella
clypealis
decolor
nr. puella
N
concinna
uliginosus
imperialis
belfragei
raphanus
californicus
norvegicus
A
elisus
apicalis
A
N
Spondyliaspidinae SpondyliaspidiniGlycaspis
Harpactorinae
n/a
Rhynocoris
Harpactorinae
n/a
Sinea
Harpactorinae
n/a
Zelus
Rhopalinae
Harmostini
Harmostes
Rhopalinae
Niesthrini
Arhyssus
Rhopalinae
Rhopalini
Liorhyssus
Rhopalinae
Rhopalini
Stictopleurus
RhyparochrominaeMegalonotini Megalonotus
RhyparochrominaeRhyparochrominiXanthochilus
Saldinae
Saldoidini
Saldula
Tinginae
Tingini
Corythucha
Triozinae
Triozini
Trioza
Triozinae
Triozini
Trioza
Order: Hymenoptera = Wasps, Ants, Bees, Sawflies, & kin
Andrenidae
Panurginae
Calliopsini
Calliopsis
Anthophoridae
Xylocopinae
Xylocopini
Xylocopa
Apidae
Apinae
Anthophorini Anthophora
spinosa
A
A
brimblecombei
A
ventralis
var. femoralis
diadema
tetracanthus
reflexulus
hyalinus
punctiventris
sabulicola
saturnius
N
A
A
marmorata
N
Nomadopsis s.l.
varipuncta
californica
Higher
Family
taxa
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Apidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Braconidae
Bradynobaenidae
Colletidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Crabronidae
Cynipidae
Cynipidae
Cynipidae
Diapriidae
Encyrtidae
Encyrtidae
Eulophidae
Eurytomidae
Figitidae
Formicidae
Formicidae
Formicidae
Halictidae
Halictidae
Halictidae
Halictidae
Halictidae
Halictidae
Halictidae
Halictidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Ichneumonidae
Megachillidae
Megaspilidae
Mutillidae
Mutillidae
Mutillidae
Perilampidae
Platygastridae
Pompilidae
Pteromalidae
Pteromalidae
Pteromalidae
Pteromalidae
Scoliidae
Pompilidae
Subfamily
Apinae
Apinae
Apinae
Apinae
Apinae
Apinae
Apinae
Bombinae
Bombinae
Nomadinae
Nomadinae
Tribe
Anthophorini
Apini
Emphorini
Eucerini
Eucerini
Eucerini
Eucerini
Bombini
Bombini
Epeolini
Epeolini
Genus
Anthophora
Apis
Diadasia
Eumelissodes
Melissodes
Melissodes
Melissodes
Bombus
Bombus
Epeolus
Triepeolus
Agathidini
Aphiidini
Meteorini
Bassus
Diaeretiella
Meteorus
Cotesini
Cotesia
Typhoctini
Colletini
Astatini
Crabronini
Larrini
Larrini
Larrini
Larrini
Pemphredonini
Pemphredonini
Philanthini
Typhoctes
Colletes
Astata
Ectemnius
Liris
Tachysphex
Tachysphex
Tachysphex
Diodontus
Pemphredon
Philanthus
peculiaris
Cynipinae
Cynipinae
Diapriinae
Cynipini
Cynipini
Diapriini
Andricus
Dryocosmus
Trichopria
quercussuttoni
minusculus
Encyrtinae
Eulophinae
Trechnitini
Psyllaephagus
bliteus
Tapinomini
Formicoxenini
Myrmicini
Augochlorini
Halictini
Halictini
Halictini
Halictini
Halictini
Halictini
Halictini
Linepithema
Cardiocondyla
Pogonomyrmex
Augochlorella
humilis
mauritanica
anzensis
pomoniella
Agapostemon
Halictus
Halictus
Lasioglossum
Lasioglossum
Lasioglossum
virescens
Anomalonini
Anomalon
Lathrostizus
Cremastini
Cryptini
Temelucha
Idiolispa
Mesochorini
Ophionini
Ophionini
Pimplini
Pimplini
Phytodietini
Mesochorus
Enicospilus
Ophion
Pimpla
Pimpla
Netelia
Agathidinae
Agathidinae
Aphidiinae
Meteorinae
Microgastrinae
Microgastrinae
Opiinae
Typhoctinae
Colletinae
Astatinae
Crabroninae
Crabroninae
Crabroninae
Crabroninae
Crabroninae
Pemphredoninae
Pemphredoninae
Philanthinae
Eucoilinae
Dolichoderinae
Myrmicinae
Myrmicinae
Halictinae
Halictinae
Halictinae
Halictinae
Halictinae
Halictinae
Halictinae
Halictinae
Anomaloninae
Campopleginae
Campopleginae
Cremastinae
Cryptinae
Ichneumoninae
Mesochorinae
Ophioninae
Ophioninae
Pimplinae
Pimplinae
Triphoninae
Megachillinae
ssp. / var. / A Ni
form
d g
Species
squammulosa
mellifera
enavata
A
pallidisignata
sonorus
vosnesenskii
helianthi
N
peculiaris
argentatus
terminatus-group
multimaculatus
B N
(ectopia)
A
A
rubicundus
actinosum
incompletum
cf. grapholithae
analis
ignea
merdarius
SphaeropthalminaePseudomethocini
Pseudomethoca
SphaeropthalminaePseudomethocini
Pseudomethoca
SphaeropthalminaeDasymutillini Dasymutilla
Perilampinae
Perilampini
Perilampus
Platygastrinae
Platygastrini
Synopeas
Pepsinae
Ageniellini
Ageniella
anthracina
anthracina
Campsomerinae Campsomerini Campsomeris
Pepsinae
Pepsini
Pepsis
pilipes
thisbe
hyalinus
coronata
N
N
N
Higher
Family
taxa
Pompilidae
Pompilidae
Sphecidae
Sphecidae
Sphecidae
Sphecidae
Sphecidae
Sphecidae
Sphecidae
Tenthredinidae
Tenthredinidae
Vespidae
Vespidae
Vespidae
Vespidae
Vespidae
Vespidae
Vespidae
ssp. / var. / A Ni
Species
form
d g
conterminus
posterus
phoenix
nigricans
luctuosa
californicum
californicum
caementarium
thomae
ichneumoneus
Subfamily
Pompilinae
Pompilinae
Ammophilinae
Ammophilinae
Sphecinae
Sphecinae
Sphecinae
Sphecinae
Sphecinae
Selandriinae
Selandriinae
Eumeninae
Eumeninae
Eumeninae
Eumeninae
Eumeninae
Polistinae
Polistinae
Tribe
Pompilini
Pompilini
Ammophilini
Ammophilini
Sceliphrini
Sceliphrini
Sceliphrini
Sphecini
Sphecini
Nematini
Nematini
Alticini
Alticini
Eumenini
Eumenini
Eumenini
Polistini
Polistini
Genus
Episyron
Perissopompilus
Ammophila
Podalonia
Chalybion
Chalybion
Sceliphron
Prionyx
Sphex
Euura
Pontania
Ancistrocerus
Ancistrocerus
Eumenes
Stenodynerus
Euodynerus
Polistes
Polistes
Order: Isoptera = Termites
Kalotermitidae
Incisitermitinae
Incisitermitini
Incisitermes
minor
Coleophora
Coleophora
Crambus
Euchromius
Evergestis
Dicymolomia
Stegea
Stegea
Hellula
Uresiphita
Lygropia
Nomophila
Glyphodes
Elachista
Estigmene
Catocala
Catocala
Caenurgia
Tetanolita
Aristotelia
Aroga
Chionodes
Prochoerodes
Digrammia
Digrammia
Eupithecia
Cyclophora
Coptodisca
Hylephila
Poanes
Polites
Lerodea
Erynnis
Heliopetes
Pyrgus
Leptotes
Brephidium
Echinargus
Everes
Plebejus
Strymon
Helicoverpa
Schinia
Lacinipolia
Protorthodes
Mythimna
Abagrotis
Spodoptera
Spodoptera
Autographa
Megalographa
Danaus
Danaus
Agraulis
Nymphalis
Vanessa
Vanessa
accordella
atriplicivora
leachellus
ocelleus
rimosalis
metalliferalis
Order: Lepidoptera = Butterflies & Moths
Coleophoridae
Coleophorinae
Coleophorini
Coleophoridae
Coleophorinae
Coleophorini
Crambidae
Crambinae
Crambini
Crambidae
Crambinae
Crambini
Crambidae
Evergestinae
Evergestini
Crambidae
Glaphyriinae
n/a
Crambidae
Glaphyriinae
n/a
Crambidae
Glaphyriinae
n/a
Crambidae
Nymphulinae
Nymphulini
Crambidae
Pyraustinae
Pyraustini
Crambidae
Pyraustinae
Spilomelini
Crambidae
Pyraustinae
Spilomelini
Crambidae
Pyraustinae
Spilomelini
Elachistidae
Elachistinae
Elachistini
Erebidae
Arctiinae
Arctiini
Erebidae
Erebinae
Catocalini
Erebidae
Erebinae
Catocalini
Erebidae
Erebinae
Euclidiini
Erebidae
Herminiinae
n/a
Gelechiidae
Gelechiinae
Gelechiini
Gelechiidae
Gelechiinae
Gelechiini
Gelechiidae
Gelechiinae
Gelechiini
Geometridae
Ennominae
Ourapterygini
Geometridae
Ennominae
Semiothisini
Geometridae
Ennominae
Semiothisini
Geometridae
Larentiinae
Eupitheciini
Geometridae
Sterrhinae
Cosymbiini
Heliozelidae
Heliozelinae
Heliozelini
Hesperiidae
Hesperiinae
Hesperiini
Hesperiidae
Hesperiinae
Hesperiini
Hesperiidae
Hesperiinae
Hesperiini
Hesperiidae
Hesperiinae
Moncini
Hesperiidae
Pyrginae
Erynnini
Hesperiidae
Pyrginae
Pyrgini
Hesperiidae
Pyrginae
Pyrgini
Lycaenidae
Polyommatinae Lampidini
Lycaenidae
Polyommatinae Polyommatini
Lycaenidae
Polyommatinae Polyommatini
Lycaenidae
Polyommatinae Polyommatini
Lycaenidae
Polyommatinae Polyommatini
Lycaenidae
Theclinae
Eumaeini
Noctuidae
Heliothinae
n/a
Noctuidae
Heliothinae
n/a
Noctuidae
Noctuinae
Eriopygini
Noctuidae
Noctuinae
Eriopygini
Noctuidae
Noctuinae
Leucaniini
Noctuidae
Noctuinae
Noctuini
Noctuidae
Noctuinae
Prodeniini
Noctuidae
Noctuinae
Prodeniini
Noctuidae
Plusiinae
Plusiini
Noctuidae
Plusiinae
Plusiini
Nymphalidae
Danainae
Danaini
Nymphalidae
Danainae
Danaini
Nymphalidae
Heliconiinae
Heliconiini
Nymphalidae
Nymphalinae
Nymphalini
Nymphalidae
Nymphalinae
Nymphalini
Nymphalidae
Nymphalinae
Nymphalini
pacifica
albophaleratus
cf. spilopterus
crucifera
crucifera
hidalgo
apachus
dominula
hidalgo
A
N
N
A
salutalis
rogatalis
reversalis
octonalis
nearctica
onychinalis
riparialis
acrea
californica
faustina
togataria
palligera
adenostomae
unifasciella
popa
forficaria
californiaria
irrorata
miserulata
nanaria
powellella
phyleus
melane
sabuleti
eufala
funeralis
ericetorum
albescens
marina
exile
isola
amyntula
acmon
melinus
zea
pulchripennis
circumcincta
rufula
unipuncta
forbesi
exigua
ornithogalli
californica
biloba
gilippus
plexippus
vanillae
antiopa
annabella
atalanta
acrea
N
N
N
N
A
cleopatra
N
N
N
forficaria
N
N
N
N
N
N
E
E
Higher
Family
taxa
Nymphalidae
Nymphalidae
Papilionidae
Papilionidae
Pieridae
Pieridae
Pieridae
Plutellidae
Plutellidae
Pterophoridae
Pyralidae
Pyralidae
Pyralidae
Sesiidae
Sphingidae
Tineidae
Tineidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Tortricidae
Subfamily
Nymphalinae
Nymphalinae
Papilioninae
Papilioninae
Coliadinae
Coliadinae
Pierinae
Plutellinae
Plutellinae
Pterophorinae
Phycitinae
Phycitinae
Phycitinae
Tinthiinae
Macroglossinae
Acrolophinae
Erechthiinae
Olethreutinae
Olethreutinae
Olethreutinae
Olethreutinae
Olethreutinae
Olethreutinae
Olethreutinae
Tortricinae
Tortricinae
Tortricinae
Order: Microcoryphia = Bristletails
Meinertellidae
Meinertellinae
Tribe
Genus
Nymphalini
Vanessa
Nymphalini
Vanessa
Papilionini
Papilio
Papilionini
Papilio
Coliadini
Nathalis
Coliadini
Colias
Pierini
Pieris
Plutellini
Plutella
Plutellini
Plutella
Pterophorini
Adaina
Phycitini
Elasmopalpus
Phycitini
Ephestiodes
Phycitini
Homoeosoma
Paranthrenini Paranthrene
Macroglossini Hyles
n/a
Ptilopsaltis
Erechthiini
Erechthias
Eucosmini
Epiblema
Eucosmini
Epinotia
Eucosmini
Eucosma
Eucosmini
Gypsonoma
Eucosmini
Notocelia
Eucosmini
Suleima
Eucosmini
Suleima
Archipini
Clepsis
Cochylini
Platphalonidia
Sparganothidini Platynota
Species
cardui
virginiensis
rutulus
zelicaon
iole
eurytheme
rapae
dammersi
xylostella
ambrosiae
lignosellus
gilvescentella
electella
robiniae
lineata
confusella
minuscula
nr strenuana
subviridis
atomosana
aceriana
illotana
helianthana
lagopana
peritana
felix
sultana
n/a
aurantiacus
Machilinus
Order: Neuroptera = Lacewings, Antlions, & kin
Chrysopidae
Chrysopinae
n/a
Chrysoperla
Hemerobiidae
Hemerobiinae
n/a
Wesmaelius
Myrmeleontidae
Myrmeleontinae Myrmeleontini Myrmeleon
Order: Odonata = Dragonflies & Damselflies
Aeshnidae
Aeshninae
Anactini
Aeshnidae
Aeshninae
Aeschnini
Libellulidae
Libellulinae
Libellulini
Libellulidae
Palpopleurinae
Palpopleurini
Libellulidae
Sympetrinae
Sympetrini
Libellulidae
Sympetrinae
Sympetrini
Libellulidae
Trameinae
Trameini
Libellulidae
Trameinae
Trameini
Libellulidae
Trameinae
Trameini
Coenagrionidae
Ischnurinae
n/a
Coenagrionidae
Ischnurinae
n/a
Coenagrionidae
Ischnurinae
n/a
Coenagrionidae
Ischnurinae
n/a
Coenagrionidae
Ischnurinae
n/a
Coenagrionidae
Ischnurinae
n/a
Anax
Rhionaeschna
Libellula
Perithemis
Pachydiplax
Sympetrum
Pantala
Pantala
Tramea
Enallagma
Enallagma
Enallagma
Ischnura
Ischnura
Ischnura
Order: Orthoptera = Grasshoppers, Crickets, Katydids, & kin
Acrididae
Cyrtacanthacridinae
Melanoplini
Melanoplus
Acrididae
Cyrtacanthacridinae
Cyrtacanthacridini
Schistocerca
Acrididae
Gomphocerinae Aulocarini
Eupnigodes
Acrididae
Gomphocerinae Aulocarini
Psoloessa
Acrididae
Oedipodinae
Chortophagini Chimarocephala
Acrididae
Oedipodinae
Trimerotropini Conozoa
Acrididae
Oedipodinae
Trimerotropini Dissosteira
Gryllidae
Gryllinae
Gryllini
Gryllus
Myrmecophilidae Myrmecophilinae Myrmecophilini Myrmecophilus
Rhaphidophoridae Ceuthophilinae
Ceuthophilini Ceuthophilus
Tettigoniidae
Phaneropterinae Phaneropterini Scudderia
Order: Psocoptera = Booklice & Barklice
Elipsocidae
Propsocinae
n/a
Ectopsocidae
Ectopsocinae
n/a
Ectopsocidae
Ectopsocinae
n/a
Ectopsocidae
Ectopsocinae
n/a
Lachesillidae
Lachesillinae
Lachesillini
Myopsocidae
Myopsocinae
Myopsocini
Psocidae
Amphigerontiinae n/a
Order: Thysanoptera = Thrips
Aeolothripidae
Aeolothripinae
Thripidae
Thripinae
Phlaeothripidae
Aeolothripini
Thripini
Propsocus
Ectopsocus
Ectopsocus
Ectopsocus
Lachesilla
Myopsocus
Blastopsocus
Aeolothrips
Thrips
ssp. / var. / A Ni
form
d g
A
N
A
N
palescens
N
N
N
N
N
A
N
N
N
N
adamsi
N
junius
multicolor
saturata
intensa
longipennis
corruptum
flavescens
hymenaea
onusta
carunculatum
civile
cyathigerum
cervula
denticollis
perparva
sanguinipes
nitens
sierranus
texana
pacifica
sulcifrons
carolina
assimilis
oregonensis
hesperus
mexicana
pulchripennis
briggsi
strauchi
vachoni
nitens
incisa
sulcifrons
N
N
A
N
N
Higher
Family
Subfamily
taxa
Order: Zygentoma = Silverfish
Lepismatidae
Lepismatinae
Tribe
Genus
Species
ssp. / var. / A Ni
form
d g
n/a
Ctenolepisma
lineata
pilifera
Class: Malacostraca = Malacostracans = Crayfish, Amphipods, Isopods, & kin
Order: Amphipoda = Scuds & Sideswimmers
Gammaridae
Gammarinae
Gammarini
Gammarus
Gammaridae
Gammarinae
Gammarini
Gammarus
Dogielinotidae
Hyalellinae
n/a
Hyalella
A
azteca
Order: Decapoda = Decapods = Crayfish, Crabs, Lobsters, Prawns, Shrimp, & kin
Cambaridae
Cambarinae
Cambarini
Procambarus
clarkii
A
Order: Isopoda = Isopods
Armadillidiidae
Armadillidiinae
Porcellionidae
Porcellioninae
A
A
n/a
n/a
Armadillidium
Porcellio
vulgare
scaber
Class: Maxillopoda = Copepods, Barnacles, & kin
Order: Eucopepoda = freeliving Copepods
Cyclopidae
Cyclopinae
n/a
Cyclops
Class: Ostracoda = Seed Shrimp
Order: Podocopida = (ten-legged) Seed Shrimp
Cyprididae
Cypridinae
n/a
Cyprididae
Cyprinotinae
n/a
Cyprididae
Eucypridinae
n/a
Cyprididae
Herpetocypridinae n/a
Cypris
Heterocypris
Eucypris
Herpetocypris
pubera
incongruens
virens
reptans
Phylum: Cnidaria = Jellies, Corals, Hydroids, & kin
Class: Hydrozoa = Hydroids
Order: Anthomedusae = Athecate Hydroids
Hydridae
n/a
n/a
Hydra
americana
Phylum: Mollusca = Molluscs
Class: Gastropoda = Snails & Slugs
Order: Stylommatophora = Terrestrial Snails & Slugs
Agriolimacidae
Agriolimacinae
n/a
Agriolimacidae
Agriolimacinae
n/a
Subulinidae
Rumininae
Ruminini
Succineidae
Catinellinae
n/a
Valloniidae
Valloniinae
n/a
Vertiginidae
Vertigininae
Vertiginini
Deroceras
Lehmannia
Rumina
Catinella
Vallonia
Nearctula
panormitanum
valentiana
decollata
rehderi
excentrica
hemphilli
Order: Basommatophora = Air-breathing Freshwater Snails
Physidae
Physinae
Physellini
Physella
Planorbidae
Planorbinae
Planorbini
Planorbella
virgata
tenuis
Phylum: Nematoda = Roundworms
Class: Adenophorea = Roundworms
Order: Gordea = (Soil) Roundworms
Mononchidae
Class: Secernentea = Caudal Papillae Roundworms
Order: Rhabditida = (Freeliving Soil) Roundworms [placement uncertain]
Rhabditidae
Phylum: Platyhelminthes = Flatworms
Class: Turbellaria = (free-living) Flatworms
Order: Tricladida = (freshwater+terrestrial) Flatworms
Dugesiidae
n/a
n/a
Dugesia
Planariidae
n/a
n/a
Phagocata
Phylum: Rotifera = Rotifers = Wheel Animals
Class: Monogononta = (single-gonad) Wheel Animals
Order: Plioma = Freshwater Wheel Animals + kin
Brachionidae
Brachioninae
n/a
Brachionus
Phylum: Tardigrada = Tardigrades = Water Bears
Class: Heterotardigrada = Water Bears (with head appendages)
Order: Echiniscoidea = (Clawed) Water Bears
Echiniscidae
n/a
n/a
Echiniscus
tigrina
morgani
A
A
B
A