Biological and molecular insights into invasive subterranean termites

(Coptotermes, Heterotermes) in the Caribbean Basin

Susan C. Jones and Tyler D. Eaton

Department of Entomology The Ohio State University

Columbus, OH

TERMITES Formerly in order Isoptera Now in order Dictyoptera

• Hemimetabolous (incomplete metamorphosis) • Immatures perform tasks in the colony (child labor)

• Diploid-diploid genetic system

• Role for females and males (sexual equality)

• Eusocial insects (highest level of sociality)

3 termite castes in the termite colony:

• Worker

• Soldier

• Reproductive

Alate

Queen

King

Neotenic reproductive

Conspicuous arboreal nests of Nasutitermes corniger are common

throughout the Caribbean.

soldiers

Termite nest types ( =very common in Caribbean)

One-piece nests: inside wood―colony size constrained by wood size Intermediate nests: inside wood, but underground gallery network allows colonization of additional wood pieces Separate nests: nest completely distinct from termites’ food

» Subterranean nests-completely under ground, but covered workings {shelter tubes} are constructed above ground

» Epigeal nests-protruding above the soil surface, i.e., mounds

» Arboreal nests-built on a tree trunk or branch, but linked with the soil by shelter tubes

conspicuous

Coptotermes species distributions: native ranges (colored areas) and invasive locations (symbols). Stripes represent overlapping native distributions.

Symbols with black center indicate likely to have been eradicated at location.

Evans et al. 2013. Biology of invasive termites: a worldwide review. Annu. Rev. Entomol. 58:455–474.

Equator

Tropic of Cancer

Tropic of Capricorn

Coptotermes spp. are spread long distances through maritime trade and boats; often first

found near harbors and sea ports.

Caribaea colleagues, Please help me monitor the arrival and inevitable spread of invasive Coptotermes so interventions can be employed.

Knowledge of the problem is a critical step.

Distribution based on: aJones et al. 2017. Formosan subterranean termite established in Grand Bahama. J. Entomol Sci. 52:445-449; Scheffrahn et al. 2015. … Coptotermes in the New World. Arthropod Syst. Phylo. 73:333–348; S.C. Jones, unpubl. data

Map template: https://www.tripsavvy.com/

• Grand Bahamaa

• New Providence • GrandTurk • Providenciales (Caicos) • Cuba • Grand & Little Cayman • Jamaica • Haiti • Puerto Rico • Virgin Gorda (BVI) • Antigua & Barbuda • St. Kitts & Nevis • Monserrat • Barbados

Coptotermes formosanus Formosan subterranean termite

Coptotermes gestroi Asian subterranean termite

Endemic species: Coptotermes testaceus

C. gestroi COII cladogram

, USA

COII gene (637 bp) Phylogenetic analyses:

• Maximum likelihood • Bayesian inference

Damage to wood in Puerto Rico structures caused by C. gestroi (Asian subterranean termite):

Damage to live trees in Puerto Rico caused by C. gestroi (Asian subterranean termite):

It attacks many tree species: • Australian pine (Casuarina spp.) • Seagrape (Coccoloba uvifera) • Bottlebrush (Callistemon spp.) • Etc.…

Behavioral characteristics of Coptotermes spp. soldiers:

• Large numbers congregate at disturbance sites • Secrete a white viscous fluid from frontal gland opening on head

Photo credit: Scheffrahn et al. 2015. A single endemic and three exotic species of the termite genus Coptotermes (Isoptera, Rhinotermitidae) in the New World. Arthropod Syst. Phylo. 73:333–348.

Coptotermes formosanus Formosan subterranean termite

Distinguishing morphological features of soldiers:

Fontanelle (external opening to frontal gland)

Coptotermes gestroi Asian subterranean termite

Characteristics of Coptotermes gestroi alates (winged reproductives):

• Fly near twilight (sunset) or night until ~midnight

• Very attracted to lights • Can fly up to 1 km; mean of ~200 m • Fly during March in Puerto Rico • Tiny hairs cover the wing surface

(quickly shed wings after flying) • Dark chestnut brown body

Background: Genus Heterotermes Froggatt

Worldwide distribution in subtropical and tropical regions

Invasive species

Economic importance:

Structural pests

Attack some field crops

Speculated to be: • a single, highly variable species OR • a complex series of several closely related species, with

intergrading species between tenuis & convexinotatus

Snyder, T. E. 1924. Descriptions of new species and hitherto unknown castes of termites from America and Hawaii. Proc. U.S. Nat. Mus. 64:1-40 +5 plates.

Heterotermes in Puerto Rico

Most likely a single, highly

variable species, but requires

further taxonomic investigation.

Grand Cayman n=1

Jamaica n=3

Cuba n=1

Puerto Rico n=3

Dominican Rep. n=2

Bahamas n=20

Bonaire n=1

Florida n=1

Turks & Caicos n=1

St. Bart n=1

St. Kitts&Nevis n=2

Aruba & Curacao n=2

Antigua n=1

Martinique n=1 Barbados

n=1

Trinidad & Tobago n=4

BVI n=1

USVI n=3

St.Lucia,St.Vincent, Grenada n=3

Geographical distribution of Heterotermes 16S haplotype groupings and species in the Caribbean Region based on

59 Caribbean samples (Szalanski et al. 2004)

27 km (17 mi)

13 km (8 mi)

61 km (41 mi)

Culebra Island 30 km2 (12 mi2)

Mona Island 9104 km2

(3515 mi2) Vieques Is. 348 km2

(134 mi2)

57 km2

(22 mi2)

65 km (40 mi) Puerto Rico

180 km (112 mi)

Three samples were H. convexinotatus (Szalanski et al. 2004)

• 75 Heterotermes samples: • 69 from Puerto Rican mainland

(subtropical dry forest) • 6 from Culebra Island • GenBank sequence entries

(126 for all Het. spp.) • Sequenced two mitochondrial loci:

• 16s rRNA gene (371 bp) • COII gene (637 bp)

• Phylogenetic analyses: • Maximum likelihood • Bayesian inference

orange: Bahama Archipelago green: Greater Antilles yellow: Lesser Antilles pink: South America blue: U.S.

H. tenuis

H. cardini

H. convexanotatus

Our proposed biogeography of Heterotermes species in the Caribbean Basin:

Evans et al. 2013. Biology of invasive termites: a worldwide review. Annu. Rev. Entomol. 58:455–474.

Equator

Tropic of Cancer

Tropic of Capricorn

Heterotermes spp. & Reticulitermes spp.

Heterotermes & Reticulitermes species distributions: native ranges (colored areas) and invasive locations (symbols). Stripes represent overlapping native distributions.

Symbols with black center indicate likely to have been eradicated at location.

Eaton et al. (2016) revealed a much more complex distribution of three Heterotermes species in the Caribbean Basin.

= H. cardini (Eaton et al. 2016)

Caribaea colleagues, Please help me monitor the arrival and inevitable spread of invasive Coptotermes spp. so interventions can be employed. Likewise, we need to more information on invasive Heterotermes spp. Please send photos, samples for identification. Contact: Prof. Susan Jones at jones.1800@osu.edu Mailing address: The Ohio State University Department of Entomology Rothenbuhler Research Lab 2501 Carmack Rd. Columbus, OH 43210-1065

Acknowledgments

• Olimpia and Hue Ferguson • Zachary Griebenow • Nicky Gallagher • Kyle Jordan • Giancarlo Lopez Martinez • Juan and Jacqueline Anguillo • U.S. Park Service • USDA Forest Service • Puerto Rican Pest Control Association • Vieques National Wildlife Refuge

Thank you!