continuing THREADS – LOOK AT THEM AS ULTIMATE INTERCONNECTORS to keep ecosystems resilient

CASE STUDIES: Animal species dependent upon forests for some phase of their survival, not generalist species that can live anywhere, some are keystone species important in the recovery rate of forest landscapes impacted by disturbances, they are important interconnectors of structures/functions in forests

Last week: Mammals and bats – adapted to disturbances & different phases of forest growth, utilize the entire landscape and not just a few habitats, greatly impacted by land use, some are on endangered species lists, some are part of human mythology TODAY: Coqui – Lunar cycles, hurricanes, adapted to human/natural disturbances, helps forests recover when calling for mates, loved in Puerto Rico going back to original Taino people and hated in [too noisy!!], human disturbance footprint eliminated by natural disturbances & coqui love disturbances

tomorrow: Salmon – keystone species that is also a cultural symbol, fertilizes forests with ocean nutrients Mycorrhizas – allow forests to grow in nutrient poor and toxic environments, symbionts that nitrogen enrichment kills, humans eat their fruiting bodies (mushrooms) 1

Puerto Rico & Coqui [frog]: How a Disturbance Maintains a Frog and How frogs Make these Forests Resilient

Photos: Dan Vogt

Coqui Puerto Rico

1 https://annexx51.files.wordpress.com/2010/10/puerto-rico-map-physical.jpg

Taino – indigenous people of the who valued this little frog & drew images on walls “The are the heart of the Taino,” says Domingo Abreu Collado. Shown here are the Pomier Caves in the . (Maggie Steber) http://www.smithsonianmag.com/people-places/what-became-of-the-taino- 73824867/?all

http://www.smithsonianmag.com/people-places/what-became-of-the- taino-73824867/?all

Homeland rich with cave drawings, testify to hallucinogens that fueled otherworldly visions, i.e., leader sniffs cohoba power (Maggie Steber)

Puerto Rico - On the way to Aguadilla and Rincon you see this guy- La cara del Indio en Isabela, Puerto Rico; https://www.pinterest.com/morganalafae/favorite- places-spaces/

Taino carving

2 Coqui = Recorded on petroglyphs so important for indigenous communities (Tainos)

No charismatic megafauna in Puerto Rico so this little frog has that role Note: When an indigenous community includes a http://artid.com/members/blanco/art/43974-coqui/ species in its stories, what does it tell you about the importance of that species in Many Taíno Indian myths surround the their survival?? coquí. Coquíes are found in much of the Taíno art like pictographs and pottery. http://www.elboricua.com/coqui.html

TO UNDERSTAND WHY THIS FROG IS SO IMPORTANT IN PUERTO RICO: 2 FACTS YOU NEED TO KNOW: [1] The ‘Stage’ or forest where the Coqui Lives since explains why this frog is important and humans are less important [2] How their Mating Habits and their Condo house preferences Contribute to Forest RESILIENCY

COMMENT: Coqui are interconnected to: • Leaf chemical quality controlled by lunar cycles and insect populations • Forest nutrient cycles through its urination and eating a lot of insects high in nutrients • They are not keystone species because so many other animals like to eat them. Keystone - link structures and functions of these forests recovering from disturbances

Coqui’s Live in the Tabonuco Forest (so tropical forests are its home)

3 LETS GET BACK to the Tabonuco Tree or look at it’s other name ‘turpentine trees’

The oil from this tree really smells good

Look at all the other things LETS GET BACK to the you get from this tree!! Tabonuco Tree or ‘turpentine trees’ • Wood used for building railway sleepers, heavy construction, cabinetwork, boat construction & boxes, flooring, violin bows, and billiard cues • Sap used medicinally by early settlers & for making candles, incense • Sap used by Amerindian tribes to coat torches & start fires, caulking materials for boats

What is sap and where do you get it?

http://www.fs.fed.us/wildflowers/ethnobotany/resins.shtml

Pre-1900, Tabonuco Forests almost ALL cut down

BUT post-1900 - International markets stopped buying SUGAR CANE (mostly to make rum), no one wanted Puerto Rico’s sugar or coffee SO Sugar cane fields abandoned, forests started to come back

FACTS: 1900 - >90 deforested 1936 - 40% deforested, so forests growing back

4 This shows the extensive forest cover in Puerto Rico [62.2% forested today] – our story takes place in the forest that is circled

http://www.scscb.org/programs/Program_resources/cebf-2008-caribbean-forest-maps.htm; http://rainforests.mongabay.com/deforestation/2000/Puerto_Rico.htm

Eda Melendez Colom and Ariel E. Lugo. 2006. LTER publication Closed Forests BUT in dark green FORESTS are coming back Used to be forests, which is our converted story for today to agricultural fields or pastures, houses

KEY POINT: Forest surrounded by Human land-uses but doesn’t degrade KEY POINT: Natural disturbances eliminate human land-use impacts – human footprint eliminated by nature

FACT: Nature is resilient to

Invasive exotic animal species who are not adapted to common disturbances

Most other cases, invasive species push ecosystems to tipping point and degradation

But these forests still need the little coqui frog!!

5 Look at exotic invaders!! Monkey Island (Cabo Santiago) - 10 minute boat ride from Naguabo, PR [39-acre island just off coast, cannot land on it, is a research center]

Research colony for testing AIDS vaccines (Brennan Linsley/AP)

Rhesus monkeys also used in psychological, behavioral studies; Photo: Alexander http://upload.wikimedia.org/wikipedia/en/thumb/f/ff/DangerMonkeySign.jpg/220pxMazurkevich- /Shutterstock DangerMonkeySign.jpg; http://repeatingislands.com/2014/12/30/rhesus-monkeys- in-puerto-rico-remain-vital-to-research/

Monkey Island (Cabo Santiago)

• In 1960’s and 70’s - descendants of individuals who escaped from research centers 30 years ago now running wild in PR • 1,000 - 2,000 rhesus and patas monkeys exist in ~11 separate colonies since escaped

“the escapees and their progeny are raising concern among public health and environmental officials…When an automobile in an urban area near San Juan …hit an adult rhesus monkey, a number of emergency personnel were exposed to Life couldn’t get any better the monkey’s body fluids. The except for the cat!! monkey subsequently tested positive for antibodies to B- virus (Cercopithecine herpesvirus 1)” BUT http://www.ncbi.nlm.nih.gov/p mc/articles/PMC3322806/

6 Puerto Rico Lays Traps for Marauding Monkeys, Wednesday, June 13, 2007, http://www.foxnews.com/story/0,2933,282094,00.html “SAN JUAN, Puerto Rico — Authorities in Puerto Rico are using cages and mangos to try to trap hundreds of marauding monkeys — descendants of escaped research on the island of Cavo Santiago, off Puerto Rico; Local farmers blame monkeys for devastating melon, pepper and pumpkin crops over the last decade”

OKAY What does forest resiliency have to do with coqui’s?? Or Considering HISTORICAL LAND-USES cutting and removing 90% of PRs forests, why do we even have any tropical forests on the island of Puerto Rico??

QUESTION: What do you have a lot of in Puerto Rico and you have to be adapted to if you want to live in the forests?

One of the ANSWERS:

https://soundcloud.com/user6380197/tropical-thunderstorm- with?utm_source=soundcloud&utm_campaign=share&utm_m edium=email

Play for 1 minute

7 Water everywhere

Photo: Dan Vogt; https://photos.travelblog.org/Photos/176949/55014 5/f/5671451-tropical_rainstorm-0.jpg

Signature of excess water visible in this landscape

Landslides common – whole hillslopes flow down

Rocks as big as cars move with a storm

Photos: Dan Vogt; http://upload.wikimedia.org/wikipedia/commons/8/88/VolkswagenBeetle- 001.jpg

Animals adapted to high water: Less water, Plants adapted to get coquis smaller rid of the excess & less able to water: attract mates • Foliage drip lines to (common get rid of water situation along • Aerenchyma – edges of modified roots - to get forests where oxygen to roots under drier) anaerobic conditions

http://luq.lternet.edu/data/lterdb09/data/CTE-photos/coqui10.JPG

8 Exotic & Invasive plants without aerenchyma not survive in these environments with high rainfall

Aerenchyma - secondary respiratory tissue or modified periderm, found in many aquatic plants and distinguished by the large intercellular spaces

http://www.mnwetlands.umn.edu/tour/tour_images/vege1.jpg

All adapted to lots of snails water and mostly active at night time!! You become a ‘night owl’ if you research these guys!!

frogs shrimp http://luq.lternet.edu/data/lterdb09/data/CTE-photos/coqui10.JPG

shrimp All adapted to lots of water!!

live part of life in fresh water streams but then go into the ocean

According to scientists, may take several years for juveniles to crawl back up to the head water streams http://luq.lternetedu/data/lterdb09/data/CTE-photos/coqui10.JPG; http://www.scielo.cl/fbpe/img/lajar/v41n4/art02-figura01.jpg

9 Another FACT – the other part of the story to understand why coqui are important for Puerto Rico’s forests are the disturbances! Major Hurricanes - frequency of hitting Puerto Rico • Once/20 yrs • Hurricane Hugo 1989 – once/40 year return frequency

http://www.photolib.noaa. Puerto Rico gov/htmls/wea00451.htm

6 Hurricanes during Vogt’s 10 yr study in PR:

• Sept 1989 – Hugo • early-mid Sept 1995 – Luis & Marilyn

• Jul 1996 – Bertha Hurricane Georges hits Puerto Rico on September 21, 1998. Image by Dennis • Sept 1996 –Hortense Chesters, Marit Jentoft-Nilsen, Craig Mayhew, and Hal Pierce, Laboratory for Atmospheres, NASA Goddard Space Flight Center from data derived • Sept 1998 - Georges from NOAA GOES-8 satellite. Image from "http://rsd.gsfc.nasa.gov/rsd/images/Georges.html".

10 Puerto Rico Take home message: ecosystems and hurricanes adapted to lots of hurricanes (if not adapted not survive in since 1980s environment where hurricanes hit frequently)

Each number is a hurricane that went over Puerto Rico – at least 15!!

Eda Melendez Colom and Ariel E. Lugo. 2006. LTER publication

Appears totally devastated but ecosystems adapted to this

AS LONG AS THEY have a resident frog that makes these forests resilient!!

In PUERTO RICO, tropical sounds at night time mostly made by frogs that are endemic (= found no where else naturally)

Eleutherodactylus or just coqui (pronounced koo-KEE)

11 Puerto Rican folk tales - If leave Puerto Rico will never sing again

Coqui image in a cup of coffee in San Juan, PR https://d262ilb51hltx0.cloudfront.net/max/2000/1*YL5aoHkw8s07ONyCviYV0Q.jpeg

Doesn’t hurt that coqui sing so well and make you smile/happy!! It also works on female coqui!! https://s-media-cache- ak0.pinimg.com/236x/c2/fd/a4/c2fda42053d18b7b831345 84fc90bdbc.jpg

Natural History • Most abundant and widespread frog (>20,000/ha) • Nocturnally active, i.e., at night time – feed, calling (calls primarily from 1-2m [3-7‘] height on exposed perches) and mating at night • Diurnal retreats - hides in forest leaf litter during day and shuttle to elevated perches at night

12 FACT: Only live 8 months so out every night calling

Role in the Food Web

• ‘Sit and wait’ predators • Mostly consume insects that fly by foliage or leaves • Consumes tremendous amount of insects (114,000 individuals/ha/night) • No predator appears to limit populations – no animal eats coqui!!

Likes human built structures – especially WET areas and where the structure funnels their calling sound for a mate – increases success of attracting a mate

13 Or even better for a coqui is a shower!! Or even light covers in the kitchen!! Showers great because its wet and the shape of a shower causes marvelous resonance that amplifies NOT QUITE the coqui LIKE THE call - attracts MOVIE Psycho females when and its shower calling scene

What is the link between the Coqui and Hurricanes?

Remember what happens to these forests when a hurricane hits. Lots of new homes – habitats – produced with debris

Condo heaven!!

14 Coqui’s are in ‘habitat BLISS’ after hurricanes

It can’t be better than this!!

Karen H. Beard http://luq.lternet.edu/data/lterdb09/data/CTE-photos/coqui10.JPG

Why the Coqui is so important for Puerto Rico?

• Eats a lot of insects which eat plant leaves (reduces herbivory) • When calling for mates, deposits a lot of very high quality and readily available nutrients in their urine • Trees need these nutrients to grow especially after a hurricane

Coquís eat a lot of Insects – is this important and why??

Therefore, fewer insects flying around reduces plant leaves being eaten so able to photosynthesize

Coqui eat so many insects each night EATING INSECTS = GET LOTS NUTRIENTS http://flatrock.org.nz/static/frontpage/assets/animals/hawaiian_frogs2.jpg

15 Back to our story about our frog!!

PLUS Coquis URINATE while calling for mates Coquis increase nutrient availability through urination (deposition of excrements) Just leaves No diapers http://www.clipsahoy.com/webgraphics2/as1978.htm; http://c2.diapers.com/images/products/p/STF/STF-093_1z.jpg

Beard et What is in URINE made of?? al. Oecologia Coquí 5 No Coquí 4 3

(mg/L) 2 1

Mean Concentration Mean 0 DOC DON NH4+ NO3- Ca K Mg P Elements

Nutrients found on water found on leaf surface [X axis] & Y axis mineral concentrations Why does urine help plants?

Would you believe plant growth limiting nutrients supplied in the urine!!

See Class Reading: Lunar Influence: http://www.keepandshare.com/htm/calendars/lun Understanding Chemical Variation ar_moon_phases_calendar/moon_phases.php and Seasonal Impacts on Botanicals. by Ian Cole, Michael J. Balick ; HerbalGram. 2010;85:50-56

http://www.youtube.com/w atch?v=b9xhOQ26QYI

http://www.lostamerica.com /

16 -the homicide rate -traffic accidents LUNAR CYCLES and -crisis calls to police or fire stations HUMANS goes back -domestic violence -births of babies a long time!! -suicide -major disasters -casino payout rates -assassinations -kidnappings -aggression by professional hockey players http://en.wikipedia.org/wiki/Phases_of_the_moon -violence in prisons -psychiatric admissions -agitated behavior by nursing home residents -assaults -gunshot wounds -stabbings -emergency room admissions -behavioral outbursts of psychologically challenged rural adults -vampirism -alcoholism -sleep walking -epilepsy

Lunar cycle - NOTE: NOTE: How How LIGHT DARK

http://www.whatsthatbug.com/beetles10.html

Lunar cycle -

FULL MOON (FM) - Lots of complex carbon compounds in leaves Light at night!!

Insects are NEW MOON (NM) – hungry! Less complex carbon Not enough compounds in leaves Dark Nights!! to eat since hard to What lunar digest, no phase do you think coqui like food value!! better?? Too dark for insects to see plants http://www.whatsthatbug.com/beetles10.html

17 How does the lunar cycle interconnect structures and functions in the Puerto Rico forests?

More leaves eaten since no protection

Q?

ULTIMATE NASTINESS: Initially humans get bitten Toxic leaf chemicals by lots of insects until coqui start to eat them!! Too many insects Puerto Rico immediately after a hurricane we were bitten by a lot by insects!! http://www.palmtreepassion.com/images/damaged_palm_leaf.jpg; http://flatrock.org.nz/static/frontpage/assets/animals/hawaiian_frogs2.jpg; http://www.whatsthatbug.com/beetles10.html; http://www.medicinenet.com/bad_bugs_pictures_slideshow/article.htm; https://www.google.com/search?q=images+of+insects+biting+people+in+tropics&biw=1680&bih=901&tbm=isch&tbo=u&source=univ&sa=X&ei=i5lCVfP- IoOvogT97YGgCA&ved=0CDIQ7Ak#imgrc=RqNq1uaOd4K8eM%253A%3B_SinR6aZ8JINZM%3Bhttp%253A%252F%252Fi.telegraph.co.uk%252Fmultimedia%252Farchive%252F02190%252 Fmosquito_2190958b.jpg%3Bhttp%253A%252F%252Fwww.telegraph.co.uk%252Fnews%252Fscience%252Fscience-news%252F9884790%252FMosquito-repellent-Deet-losing-its- effectiveness.html%3B620%3B387

Lunar-based harvesting Many harvesters of natural resources in tropics (, Panama, Puerto Rico) believe critical to harvest during the full moon because materials harvested at this phase are more durable or nutritious

NOTE: plants change their chemistry in response to lunar phases and need lunar phases for triggering these changes

AMBIO: A Journal of the Human Environment Article: pp. 485–490, Indigenous Knowledge Informing Management of Tropical Forests: The Link between Rhythms in Plant Secondary Chemistry and Lunar Cycles, Kristiina A. Vogt, Karen H. Beard, Shira Hammann, Jennifer O'Hara Palmiotto, Daniel J. Vogt, Frederick N. Scatena, Brooke P. Hecht

Lunar-based harvesting practice mentioned by indigenous groups to ethnobotanists/social ecologists studying use/harvest of products in parts of tropical Africa and tropical Americas. Examples: • rice in • certain wet woods in Puerto Rico • palm fronds for roofing thatch in the American tropics

Rice collection

http://dandapani.org/blog/wp-content/uploads/2013/09/central-java-rice- paddy-women.jpg Moon image taken by the Galileo Orbiter, 1998.

18 Experiments With Mystical "Full Moon" Beer. Reuters, Date: 27-Sep-10; Country: BELGIUM; Author: Emily Coleman

Full moons are often associated with tides, insanity and creatures like werewolves, but it turns out they're also good for brewing beer.

 family-owned brewery produced first batch of specialist beer brewed by A woman holds up a the light of a full autumn moon bottle of Paix-Dieu  Full moon speeds up fermentation beer in Brussels process, shortening it to five days September 24, 2010, Photo: Thierry Roge from seven, which adds extra punch to the beer without making it harsh, according to connoisseurs

The Coqui (Eleutherodactlus coqui) is only native to Puerto Rico. This map shows its distribution outside of its native range

http://fwcb.cfans.umn.edu/courses/nresexotics3002/GradPages/Coqui/Pages/distribution.htm

Spread of the coqui to southeast US and eventually …

Hawaii!!!

http://www.lonelyplanet.com/maps/north-america/usa/hawaii/map_of_hawaii.jpg

19 Why did they get to Hawaii!!!

Introduced with ornamental plants Introduced to control insect populations

Dilemma – frogs going extinct around the world but Hawaii doesn’t want them Coqui are New species in Hawaii: • Hawaii has no native amphibians so open ecological niche • Coqui compete with birds for food, reduce pollinator populations • Coqui have no native predators so nothing controls its population density

http://fwcb.cfans.umn.edu/courses/nresexotics3002/GradPages/Coqui/#Potential impacts on Hawaiian natives

Initially, coquis were mainly consuming mostly leaf litter non-native invertebrates but this has changed. TODAY they are eating native invertebrate species

They preferred prey items: ants, amphipods – sand fleas [small shrimp like crustaceans]

http://en.wikipedia.org/wiki/Amphipod

20 ~ 20,570 individuals/ha in Puerto Rico & consume 114,000 invertebrates/ha/night ~ 90,000 individuals/ha in Hawaii; eating 690,000 invertebrates/ha/night

http://luq.lternet.edu/data/lterdb09/data/CTE-photos/coqui10.JPG Karen H. Beard

Coqui frog population densities in Lava Tree State Monument (Pahoa, HI) are the highest in the state of Hawai’I

On some nights, the frogs’ chorus in the park reaches 70 decibels – about as loud as a vacuum cleaner

Hawaii" (Honolulu Star- Bulletin, 07JAN2007) "The coqui frog is the worst Impacts “We must not let tiny shrieking monsters destroy threat ever to tourism and the state's overall economic health...”" (excerpted from article)

http://www.hear.org/species/eleutherodactylus_coqui/; https://www.toovia.com/top/7-issues-about-hawaii-s-coqui-frog-conflict; http://2.bp.blogspot.com/_qkGEo7smLnY/St63pRYkP-I/AAAAAAAAAAM/Ob7sd7mJUQQ/S226/keepkonaquiet.jpg t

21 Pest alerts

Pest Alert: Stop the spread of Caribbean frogs Pest Alert poster to educate the public and prevent further spread of Caribbean frogs in Hawaii. A letter to the horticulture industry A letter to the horticulture industry from the Hawaii Department of Agriculture (HDOA) requesting help in stopping the spread of coqui and greenhouse frogs within the state. Caribbean frogs (Eleutherodactylus coqui and Eleutherodactylus planirostris): MISC target species Caribbean frogs (Eleutherodactylus coqui and Eleutherodactylus planirostris) are targeted by the Maui Invasive Species Committee (MISC).

http://www.hear.org/species/eleutherodactylus_coqui/

http://mauisierraclub.org/wp-content/uploads/2013/05/coquipest-624x441.png

http://ecx.images- amazon.com/images/I/515DY J9AK0L._SY344_BO1,204,2 03,200_.jpg

http://honolulumagazine- images.dashdigital.com/images/2012/Jun1 2/coquiFrogs/coquifrogs1.jpg?ver=1338848 990

Coqui killed with caffeine – overdose kills coqui – but not allowed to use today https://faculty.washington.edu/chudler/gif/ffff1.gif Spraying water with lime to kill coqui http://archives.starbulletin.com/2002/07/07/news/arti.jpg

Christmas trees are cooled down in the refrigerated containers and later opened and pounded on the ground to shake any pests loose http://hdoa.hawaii.gov/wp-content/uploads/2013/11/Xmas2013- treereloading.jpg

22 Control methods

Heat being used to kill coqui frog "A local [ed.: Waimanalo, Hawaii] nursery is using heat to kill the pesky coqui frog in potted plants. In a recent test done at Leilani Nursery, nine out of ten frogs died after being exposed to temperatures of more than 113 degrees

for five minutes." (AP, as reported by KPUA.net 30 January 2006)

Citric acid is the only EPA approved chemical for use in controlling coqui. Frogs breathe through their skin so they are highly sensitive to chemicals contacting their skin. Other products have been reported to be effective such as baking soda, concentrated Simple Green Soap, and hydrated lime. Caffeine, at about 10 times the concentration found in coffee, was studied for

use but not approved. http://www.lehuanet.com/coquicontrol/indexBioCtl.html

If coqui are well-established in your area:

REPLACE thick understory vegetation with thinner vegetation. SPRAY all landscape and potted plants with citric acid or hot water. MOW a buffer zone around your house to help reduce noise. What does managing vegetation do to coqui’s??

BEFORE - dead leaves - leaf litter

AFTER - leaves trimmed - debris cleared

If these do not work, do what a student from Spring 2013 did on the Big Island, Hawaii

Photo Credit Isaac Shozuya

23 Invasive in Hawaii but not Puerto Rico but frogs are endangered around the world so hard to kill them

Endangered amphibians worldwide Amphibians have existed on earth for about 300 Check the laws in your area. In many parts of million years, yet within the world, it's illegal to kill non-invasive species, the last several decades but people are encouraged to manage invasive more than 120 species are thought to have species. http://www.wikihow.com/Get-Rid-of-Frogs disappeared for ever because of human activities. http://www.endangeredspeciesinternational.org/amphibians 4.html

24