INSECTS OF GRAND CANYON: AN OVERVIEW FOR HIKING GUIDES

Larry Stevens Museum of Northern Arizona [email protected] Giants on Whose Shoulders We Stand

Nothing in biology makes sense except in the lightof evolution Theodore Dobzhansky Basin and Range Rocky Mtn. Geologic Prov. Geol. Prov.

Virgin R.

Rkm 0 Grand Canyon

Rkm Rkm 400 Rkm 100 Rkm 200 300

Basin and Range Geologic Prov. Grand Canyon Rims Map (Billingsley and Hampton 1999)

Eastern Basin Western Basin

Muav Gorge THE BIOGEOGRAPHY OF LARGE, DEEP CANYONS (STEVENS 2012)

Barrier/Filter Refugia North Rim Barrier/ Corridor Barrier/ Filter Flow Direction Filter South Rim

Barrier/Filter ARTHROPODA: Half a Billion Years of World Dominance

Segmented worm ancestor Chelicerata

Mites Bark Scorpion (Buthidae: Centruroides) Scorpiones

Silurian (430 m.y. ago)

AZ Giant Hairy Scorpion (Vaejovidae: Hadrurus) Scolopendron viridis Centipede

Hyallela Amphipod

Orthoporus ornatus Millepede

AQUATIC AND RIPARIAN INSECTS Belostoma ~fluminium Mayflies: ♂ Stoneflies: Plecoptera Ephemeroptera

Naucoridae Ranatra

Masked Clubskimmer Rhagovelia (Brechmorhoga pertinax) distincta Aquarius remigis Odonata – 89 spp, dominated by neotropical fauna, with ~equivalent boreal, range-centered, and Pacific coast elements; 78% of fauna is exogenous 13 5

18 20 3 25 endemic

1 29

ANT LIONS: MYRMELEONTIDAE Thermonectus Chrysina

Dryopoid Beetles

Prionus

Tiger Beetles Ground Beetles DIPTERA Nematocera: “Two Winged Insects” Ancestral flies with aquatic larvae (e.g., Culicidae, Simuliidae, Chironomidae) Brachycera: Tabanomorpha Asilimorpha Muscomorpha

Efferia HYMENOPTERA: BEES, ANTS, WASPS Origin in Triassic Period, with social wasps in Cretaceous 150,000 described, with many undescribed spp (e.g., parasitoids)

Aculeate Apocrita Wasps

Mymarid wasp, Giant Cicada Killer wasp, 0.1 mm long

45 mm long

Vespoidea

Apoidea Chrysidoidea

Vespoidea: Formicidae (Ants) Vespidae (Paper & Meat Wasps Triassic Period >200,00 million years agoPompilidae (Spider Wasps) Sphecidae (Thread-waisted Wasps) HYMENOPTERA: Bees, Ants, and Wasps Membrane -winged Insects Formicidae: Ants 175 species on southern Colorado Plateau Desert diurnal S is low Reduced by California Harvester Ant? Large Aculeate Wasps: Paper/Hornet, Spider, Pompilidae and Thread-waisted Wasps

Vespidae Sphecidae Apoidea - Bees: Six families, mostly solitary Honey Bee is only “kamakazi” bee

Melittidae Apidae Megachilidae

Honey Bee

Andrenidae Halictidae

Colletidae The Hymenopteran Stinger: A Girl’s Best Weapon

Many Uses of Stinger and Venom * Oviposition – egg laying * Self defense * Kill or paralyze prey * Hive defense * Self-protection by coating their bodies to deter attackers * Sterilization of pathogens * Alarm pheromones * Sex-attractant pheromones JUSTIN SCHMIDT HYMENOPTERA STING PAIN SCALE: * Pogonomyrmex barbatus Harvester Ant “Bold and unrelenting. Somebody using a drill to excavate your ingrown toenail”

* Paper Wasp – “Caustic and burning. Distinctly bitter aftertaste. Like spilling a beaker of hydrochloric acid on a paper cut”

Justin Schmidt * Fire Ant “Sharp, sudden, mildly alarming. Like walking across a shag carpet and reaching for the light switch.” Highest level of anaphylactic response. http://www.realmonstrosities.com/2012/08/schmidt-sting-pain-index-categorising.html Tarantula Hawks (Pompilidae: Pepsis and Hemipepsis

Schmidt Scale Sting Pain: Taratula Hawk - Blinding, fierce, shockingly electric. A running hair drier has been dropped into your bubble bath.”

The most painful sting in the Southwest, but short duration pain. Grand Canyon Region Butterflies: >140 Species in Six Families

Sleepy Dogface Orange Sulfur Black Swallowtail Many-tailed Swallowtail Western Swallowtail

Dainty Sulfur Mourning Cloak Buckeye Acastus Patch Red-spotted Purple

Queen Painted Lady AZ Sister Grand Canyon Brown Monarch

Great Purple Hairstreak American Snout Common White Pygmy Blue Common Hairstreak

© L.E. Stevens 2004 Arizona State Butterfly

Many-tailed Swallowtail (Papilio multicaudata) Plant Defenses: Larvae Eat Leaves Hairy leaves Cope with hairy leaves Latex compounds in sap Store toxins in skin Cardenolides (~digitalis) in sap. Shed skin 5 times Fast growth Store toxins in pupal integument

Adults - wings contain stored plant toxins deter predators APOSEMATIC COLORATION

+ apo sēmat-, sēma (above, highest) (sign)

Oncopeltus Tetraopes

J. Friedrich Müller

Mullerian Mimicry: Bad Guys Look Alike

Danaus plexippus VELVET ANTS (MUTILLIDAE): BEING BAD, LOOKING BAD - TOGETHER (Wilson et al. 2012 – Females only!)

Wilson, J.S., K.A. Williams, M.L. Forister, C.D. von Dohlen, and J.P. Pitts. 2012. Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications 3:1272, UAES 8399. Batesian Mimicry: Cheaters That Look Bad, but Aren’t

Monarch (Danaus plexippus) Viceroy (Limintis archippus) Bad Taste, Poisonous Faking It - No Bad Taste “Model” “Batesian Mimic” Henry Bates Grand Canyon region invertebrates are diverse (~20K spp) Requiring lifetimes to learn them Many new species, many grand processes

Each species we encounter has a remarkable story to tell, if we can listen carefully and ask the right questions ENDEMIC, SMALL POPULATIONS, NO MONITORING, MANY RARE SPECIES, STATUS UNCERTAIN

“Masked Clubskimmer Wetsalts Tiger Beetle (LIBELLULIDAE:Brechmorhoga pertinax)” (CICINDELIDAE: Cicindela haemorrhagica arizonae)

Viceroy Butterfly

Vulcans Well Giant Waterbug Limenitis archippus (Belostoma nr. flumineum )

OUT HERE IT’S SO HOT THAT….