Lecture 3: Biogeography of Honey Bees University of Florida Department of Entomology & Nematology Dr. Jamie Ellis Created by: Pablo Herrera, Catherine Nalen, Jamie Ellis What is Biogeography? 1. Biogeography is the study of the distribution of species. This is considered spatially (geographically where the species is) and temporally (when did the species arrive where it is). 2. Biogeography relies on integral knowledge of biological systems combining geographical distribution data, ecology, genetics, fossil records, phylogeny, and many others. 3. Geology, climatology and paleontology are integrated into this field to help us understand the evolutionary history of organisms. Population Distributions This map represents the global distribution of Magnolias. When an organism is distributed in two isolated locations, this is called disjunct distribution. Magnolia distribution was not always disjunct. - We know this from fossil records. 1
So how did organisms get to different continents? Permian 225 mya Jurassic 135 mya Triassic 200 mya Cretaceous 65 mya 265 mya there was one large supercontinent called Pangea 200 mya Pangea splits into Laurasia and Gondwanaland Presently, we have 7 distinct continents Indian sub-continent Present Day What is a Species? 1. There is no easy answer to this question!!!!!! 2. Best working definition for our purposes: a group of organisms that can interbreed and produce VIABLE (or fertile) offspring. + = Speciation 1. Populations descended from a common ancestor will take different evolutionary paths because they are subjected to different environmental pressures 2. These populations will accumulate differences over time because of isolation 3. They can be isolated by either distance or biogeographic barriers 1. A region or area through which an organism cannot pass 2. The Himalayas, the Darien Swamps in Panama, oceanic currents between islands, and the Sahara desert are some examples of major biogeographic barriers 2
- The honey bee species as a group (Genus Apis) is one of the most widely studied groups in the world. - Fossil records suggest the origin of the genus Apis most likely occurred in South East Asia. Bee fossil found in northern Burma, >100 myo Origins of Honey Bees Mader et al. 2010 Honey bees as a group are thought to have diverged from sphecoid wasps around 100 million years ago, with the rise of the angiosperms (flowering plants). Honey Bee Taxonomy 1. Honey bees (Tribe Apini) are most closely related to the Meliponini (Stingless bees), Bombini (Bumble bees) and Euglossini (Orchid bees). 2. There is no clear answer on their exact relationship, conflicting results have been found with different forms of analysis. 3. Most authors say that the closest relatives of honey bees are stingless bees, but recent DNA analyses indicates that they may be more closely related to orchid bees. 3
Honey Bee Taxonomy Until recently, all known honey bee fossils came from Europe, Asia and Africa Many of the oldest known fossils are European, from the Oligocene- Eocene boundary This does not mean that they are of European Origin Science News, August 15th, 2009; Vol.176 #4 The recent discovery of a fossil honey bee, Apis nearctica, in the United States has brought the evolutionary history of Honey Bees into question Characteristics of Apini 1. The Apini are a group of highly derived bees which present corbicula (pollen baskets) 2. They are Eusocial 3. They produce honey 4. Native to Eurasia and Africa 5. There is debate on the number of honey bee species in the world. The estimate is between 7-11, divided into 3 subgenera. The Recognized Honey Bee Species Engel et al.. 2009 Subgenus: Micrapis Subgenus: Megapis Subgenus: Apis 4
Global Distribution of the Apis Species / Apis mellifera / Apis nigrocincta / Apis koschevnikovi / Apis andreniformis / Apis dorsata / Apis cerana / Apis florea Subgenus Micrapis Commonly known as the dwarf honey bees They are present in Southern and Southeast Asia Two species exist: Apis florea and A. andreniformis A. florea and A. andreniformis are sympatric, but seem to have evolved allopatrically Micrapis are the most plesiomorphic (showing the greatest amount of primitive traits) of the honey bees Apis florea Dwarf honey bees Native to S. & S.E. Asia Exposed, single combs Most primitive of the honey bees Zac Huang Kept in rural Asia for honey production Video of Apis florea: http://www.youtube.com/watc h?v=11uq_9da2k4 5
Apis andreniformis Zac Huang Zac Huang Identified in Thailand, Malaysia and the Southern Chinese peninsula Small in size like A. florea Nests on a single comb Behavior and life history similar to A. florea Subgenus Megapis The Megapis are commonly known as the giant honey bees Some authors consider these to be subspecies, while others consider them separate species. -Apis dorsata dorsata occurs primarily in India. -Apis dorsata laboriosa is the largest species. It nests in cliffs throughout the Himalayas. (Apis laboriosa) -Apis dorsata binghami occurs primarily in Indonesia and Malaysia -Apis dorsata brevigulia is found in the Philippines Apis dorsata Exposed, single combs -Nests can be over a meter wide! -Extremely defensive Pseudo-Migratory -Nests in jungles during rainy season -Move to edge habitat at the beginning of summer -Honey hunters know their migration routes Concern in Asia over population depletion by bee hunters Apis dorsata video a must see! 6
Subgenus: Apis The subgenus Apis consists of the cavity nesting bees, with 4 commonly recognized species: Apis koshevnikovi commonly known as the reddish Koshevnikov bee, which is found in Borneo Apis cerana the Asian honey bee, has two subspecies, is found in Eastern and Southern Asia & is kept similarly to A. mellifera Apis nigrocincta similar to Apis cerana but inhabits the Philippines and Indonesian islands Apis mellifera the Western honey bee is widely distributed throughout the Old World and is the only honey bee with a natural distribution outside of Asia Zac Huang Apis cerana Oriental Honey Bee Found throughout Asia Zac Huang Build multiple layers of comb in protected locations -Caves, hollow trees, rock cavities Have been kept in man-made hives for thousands of years Excellent honey producers Main species managed in Asia Natural host of varroa mites Apis koschevnikovi Only identified from Malaysia Commonly called Red bees Builds multiple parallel combs J. Woyke 1995 Behavior and life history similar to A. cerana Not usually managed 7
Apis nigrocinta Inhabits Philippine island of Mindanao and some Indonesian islands Very similar to A. cerana (only minor differences are noticable) Builds nests in cavities Apis mellifera Previously considered to originate in Asia, but recent fossil evidence may modify this theory - A. mellifera probably originated in Africa. Its ability to thermoregulate has allowed it to survive in hot and cold places where other honey bees have failed It has been transported by humans around the globe Origins of Apis mellifera Subspecies Apis mellifera populations moved out of Africa and into Eurasia at least three times, resulting in populations that are close geographically but not genetically The various subspecies resulted from these geological emigrations 8
Apis mellifera These subspecies have come from several geographic regions, and are categorized by regions of origin Near East Subspecies Tropical African Subspecies Apis mellifera anatolica Apis mellifera adami Apis mellifera cypria Apis mellifera syriaca Apis mellifera medea Apis mellifera caucasica Apis mellifera armenica Apis mellifera taurica Apis mellifera lamarckii Apis mellifera yemenitica Apis mellifera litorea Apis mellifera scutellata Apis mellifera adansonii Apis mellifera monticola Apis mellifera capensis Apis mellifera unicolor Apis mellifera These subspecies have come from several geographic regions, and are categorized by regions of origin Western Mediterranean Subspecies Apis mellifera saharensis Apis mellifera intermissa Apis mellifera iberica Apis mellifera mellifera Southeastern European Subspecies Apis mellifera sicula Apis mellifera ligustica Apis mellifera cecropia Apis mellifera macedonica Apis mellifera carnica Important Apis mellifera races in the U.S. Apis mellifera ligustica: - This is a light colored, golden bee, known commonly as the Italian Honey Bee. - It is valued as a laborious and mild tempered bee. - It produces a surplus of honey. - The main drawback to it is that it may require much feeding during the winter 9
Important Apis mellifera races in the U.S. Apis mellifera caucasica: - A subspecies from the Caucasus mountains - Often used by beekeepers who consider it both mild tempered and industrious - It is a darker bee (somewhat gray) - Uses considerable propolis Important Apis mellifera races in the U.S. Apis mellifera carnica: - It is often called the Carniolan Honey Bee, due to its presence in the Carniolan region of Slovenia. - It is a bee subspecies of the Alps and does very well in cold climates. - It is considered to be one of the least aggressive subspecies of Honey Bees. - Possibly original source of the Russian honey bee Additional Reading: 1. Arias, M. and W. Sheppard. 2005. Phylogenetic relationships of honey bees (Hymenoptera: Apidae: Apini) inferred from nuclear and mitochondrial DNA sequence data. Molecular Phylogenetics and Evolution. 37: 25-35. 2. Engel M., Hinojosa-Díaz I. and A. Rasnitsyn. 2009. A honey bee from the Miocene of Nevada and the biogeography of Apis. Proceedings of the California Academy of Sciences. 60: 23-38. 3. Michener, C. 1979. Biogeography of the bees. Annals of the Missouri Botanical Garden. 66: 277-347. 4. Michener, C.D. 2000. The Bees of the World. John Hopkins University Press, Baltimore, MD. 913 pp. 5. Ruttner, F. 1988. Taxonomy and Biogeography of Honey Bees. Springer-Verlag, Berlin, Germany. 284 pp. 10