Dr. Larry McEdward was a leader of the field of ecological physiology, and evolutionary ecology of marine invertebrate larvae. His primary focus was the evolutionary transitions between feeding and non-feeding larvae in marine invertebrates (e.g., planktotrophy and lecithotrophy). In 2001 Larry unexpectedly passed away—a significant loss to the field. We are organizing this symposium to honor Larry by synthesizing recent major research advances within his field and focusing on the future directions, especially those that result from Larry’s work.

Larry received a Masters degree from the University of South Florida with John Lawerence in 1979 investigating respiratory electron transport system activity during larval development of Lytechinus variegatus. He then entered the Ph.D. program at the University of Washington with Richard Strathmann. This work focused on the ecological physiology and functional morphology of echinoderm larvae. After graduating in 1984, he was a Killam Post-Doctoral Scholar at the Department of Zoology at the University of Alberta until 1987. During his postdoc he inverstigated variation in biochemical composition of echinoderm eggs. After completing his postdoc, Larry was a visiting scholar at the University of Washington until he was hired by the Department of Zoology at the University of Florida in 1989. Larry was promoted to Full Professor (posthumous) in 2001. For further details about Larry's academic history and research, please see his CV and publications.

Tribute to the Science and Life of Larry McEdward

By Richard R. Strathmann

Larry McEdward's sudden and unexpected death on July 2, 2001 was a hard blow for family and friends and also a great loss to science.

Larry's success in research and teaching came from his grasp of general issues, his ability to reduce general questions to specific questions, and his ability to make the specific questions answerable by developing new methods. My first knowledge of Larry was that there was a graduate student in Florida known as the larval marvel. He lived up to this early reputation as he continued his graduate work at the University of Washington. He pursued his studies of evolution, form, and function of marine larvae with characteristic energy and enthusiasm throughout his career. He favored echinoderms in his studies, but (peculiar as these animals are) the implications of his studies are much broader.

In his research on evolution of life histories, he focused on key problems. One of these was the relation between parental investment per offspring and offspring performance and fitness. Another was evolutionary transitions between modes of development: feeding and non-feeding larval development and non-larval alternatives. Larry explored these relationships with new models, using his skill and experience in programming, and then tested the new hypotheses with comparative and experimental studies. With his students as collaborators he examined consequences of the continuum in facultative larval feeding among closely related species. In yet another line of research, they analyzed a spectacular change in development of body axes associated with evolution of brooding. He also developed evidence for a reversion to pelagic development, despite loss of larval features. In the year before his death, Larry and his students were developing models for a more unified theory of bentho-pelagic life-histories.

When usual methods proved inadequate, Larry invented new ones. He introduced new morphometric methods for measuring complex 3-dimensional ciliary bands to assess feeding capacity of disparate larval forms. He introduced manipulation of parental investment to test evolutionary hypotheses and new methods for measuring organic content per egg to analyze variation in parental investment. Other researchers continue to apply these methods to diverse organisms.

Larry's intelligence, skill at solving problems, willingness to help people, and sense of humor made him an excellent colleague. Many of you knew Larry through his professional associations. Larry was an active member of SICB/ASZ and the current Program Officer in the Division of Invertebrate Zoology. Larry completed his Masters Degree at the University of South Florida, Ph.D. Degree at the University of Washington, and a postdoctoral fellowship at the University of Alberta, Canada before joining the Zoology Department at the University of Florida. He was also a visiting scientist and instructor at the Friday Harbor Laboratories and at the Bamfield Marine Station. Larry was one of the originators of the "Evo-Devo" course at the Friday Harbor Laboratories. He edited numerous reviews of larval biology and echinoderm biology, including Marine Invertebrate Larvae (the leading one-stop reference book for the field), the Larval Development issue of Seminars in Cell & Developmental Biology, and the recent Echinodermata issue of Canadian Journal of Zoology. This incomplete list does not do justice to the number and diversity of his professional activities and institutional associations.

Larry's life was much too short, but he made a very good life of it. He devoted himself to family, friends, science, mountain biking, climbing, kayaking, and chocolate. To an onlooker, his priorities in these pursuits were not always obvious. I know that chocolate ranked above climbing, because he would not give it up to condition himself for still more demanding climbs.

As a teacher and researcher, Larry leaves a model for his students to follow. There is much research and teaching that Larry would have done that will now never be done, but even so, he leaves discoveries and insights that will stimulate young biologists around the world. In addition to his scientific legacy, he leaves many good memories. Larry's humor joined insight with generosity. Being remembered with a smile is a great legacy.

Biking with Larry, told from a student's prespective

By Ben Miner

The first two paragraphs are excerpts from Larry's webpage and lead into the third, which is a recount of a ride I did with Larry.

"Some recent good rides: Smugglers Mountatin & Hunter Creek Valley in Aspen, Colorado; Kaches Ridge, Devil's Gulch, Naches Ridge, and the Northway Trail at Crystal Mtn in the Cascade Mountains of Washington; Mt Constitution, Orcas Island, Washington (39 min); and the Olympic Mountains (6200 ft of climbing). Every now and then it seems that my bike and I don't quite see eye to eye on details such as which side of that tree to go around, whether or not we have the speed and height to clear that rock or log, or even who is supposed to be on top and who is supposed to be rolling along in the dirt. Casualty list to date for Lar: bruised/cracked ribs (three times), an interesting racing stripe up the middle of my abdomen (skewered by a broken seatpost), dislocated thumb, and a few assorted cuts and bruises. Casualty list to date for bike: 5 broken rims (& unknown # of spokes), twice mangled rear derailleur, blown shifter, worn out chainrings (4 sets), chains (10), cassettes (3), brakes & many sets of pads, hubs, fork damping cartridge, handlebars, tires (lots), pedals, shoes (4 pairs!), a bent derailleur hanger, and some curious dents & scrapes (of unknown origin) on the bars ends, and one incredible OCLV carbon frame (lifetime warranty is a good thing!). There are a few original parts still on the bike and they deserve mention for their courageous service: seat, stem, shifter, um ..., I guess that is all, not even the original paint job has survived.. Credit for repairs and upgrades goes to the guys at Gator Cycle (Gainesville, FL) and Island Bicycles (Friday Harbor, WA).

Check out the article in the February 1997 issue of Bike magazine entitled "Snow Biking: its not as dumb as it looks". Actually it is a lot of fun, if you are not overly concerned with control details (e.g., turning and braking). Unfortunately it is rare that we get to snow bike in Florida. Luckily, it is more common on the homestead in New Hampshire (even in mid-April!)."

I would now like to recount a ride that I did with Larry. Convinced by Larry that riding with him was safe, I decided to go for a ride with Larry up and down Jolly mountain in the WA Cascades. With a name like Jolly mountain, I thought I had a pretty good chance of (1) surviving and (2) having a good time. We assended up a fire trail to the single track we planned on riding. However, the single track was covered with snow. I am pretty certain that Larry knew about the snow, but he played dumb. So Larry proposed that we could carry our bikes down the first part of the trail and pick it up at the bottom. As I soon found out, walking down a steep snowy hill while carrying a bike is extremely difficult, and I slipped shortly after starting and slid to the bottom (Larry was kind enough to the capture the moment on film, shown to the right). We then rode a bit through the snow during which time I managed to eat snow 3 to 4 times. However, it soon became clear that we had made a wrong turn somewhere because of all the snow. We searched for about 2 hours before giving up and debated our options. It was getting dark and Larry convinced me that we should retrace our steps and hike back up to where fire road and take that back down. So, off we went. I soon realized that hiking down the snow covered trail, althought difficult, was much much easier than hiking up the snow covered trail. We finally made it to the top (after Larry had to rescue me from a little dirt outcrop I became stranded on) and headed back down the hill. Although I always end the story with "and that was the worst ride I have ever been on", Larry always end the story with, "and that was one of the best rides I have ever been on".

Now, how and the hell he can consider snow biking "a lot of fun" is beyond me, but one of the reasons he was special. I will miss him very much.

Publications of Larry R. McEdward:

Reitzel, A.M., B.G. Miner, and L.R. McEdward. In press. Relationships between Spawning Date and Larval Development Time for Benthic Marine Invertebrates: A Modeling Approach. Marine Ecology Progress Series.

McEdward, L.R., and B.G. Miner. 2003. Fecundity-time models of reproductive strategies in marine benthic invertebrates: fitness differences under fluctuating environmental conditions. Marine Ecology Progress Series 256: 111-121.

Miner, B.G., J.C. Cowart, and L.R. McEdward. 2002. Egg energetics for the facultative planktotroph Clypeaster rosaceus (Echinodermata: Echinoidea), revisited. Biological Bulletin 202: 97-99.

McEdward, L.R., W.B. Jaeckle, and M. Komatsu. 2001. Phylum Echinodermata: Asteroidea. Invited chapter in An Atlas of Invertebrate Larvae, C.M. Young, M.E. Rice, and M. Sewell (eds.), Academic Press.

McEdward, L.R. and B.G. Miner. 2001. Echinoid larval ecology, pp: 59-78. The Biology and Ecology of Edible Sea Urchins, J.M. Lawrence (ed.), Elsevier, Amsterdam. (Invited review).

McEdward, L.R. and B.G. Miner. 2001. Larval and life cycle patterns in echinoderms. Canadian Journal of Zoology 79:1125-1170. (Invited review).

McEdward, L.R. and P.Y Qian. 2001. Effects of the duration and timing of starvation during larval life on the metamorphosis and juvenile size of the polychaete Hydroides elegans. Journal of Experimental Marine Biology and Ecology 261:185-197.

McEdward, L.R. and K.H. Morgan. 2001. Interspecific relationship between egg size and the level of parental investment per offspring in echinoderms. Biological Bulletin 200:33-50.

McEdward, L.R. 2000. Introduction; larval evolution. Seminars in Cell and Developmental Biology 11:383-384. (Invited editorial).

McEdward, L.R. 2000. Adaptive evolution of larvae and life cycles. Seminars in Cell and Developmental Biology 11:403-409. (Invited review).

McEdward, L.R. 2000. The origin and evolution of marine invertebrate larvae. Quarterly Review of Biology 75:191-192. Book Review.

McEdward, L.R. and J.C. Herrera. 1999. Body form and skeletal morphometrics during larval development of the sea urchin Lytechinus variegatus. Journal of Experimental Marine Biology and Ecology 232:151-176.

McEdward, L.R.. 1997. Reproductive strategies of marine benthic invertebrates revisited: facultative feeding by planktotrophic larvae. Amer. Nat. 150:48-72.

McEdward, L.R. and D.A. Janies. 1997. Relationships among development, ecology, and morphology in the evolution of echinoderm larvae and life cycles. Biological Journal of the Linnean Society 60:381-400.

McEdward, L.R. 1996. Experimental manipulation of parental investment in echinoid echinoderms. American Zoologist. 36:169-179.

Herrera, J.C., S. McWeeney, and L.R. McEdward. 1996. Diversity of energetic strategies among echinoid larvae and the transition from feeding to nonfeeding development. Oceanologica Acta 19:313-321.

McEdward, L.R. & M.G. Hadfield. 1996. Flexibility in life cycles. Oceanologica Acta 19:468. (invited conference proceedings paper).

Nichols, D., L.R. McEdward, & A.B. Smith. 1996. Evolution of life history traits. Oceanologica Acta 19:469-470. (invited conference proceedings paper).

McEdward, L.R. (ed.) 1995. Ecology of Marine Invertebrate Larvae. CRC Press, Boca Raton.

McEdward, L.R. 1995. Evolution of pelagic direct development in the starfish Pteraster tesselatus (Asteroidea: Velatida). Biological Journal of the Linnean Society. 154:299-327.

Janies, D.A. and L.R. McEdward. 1994. A hypothesis for the evolution of the concentricycloid water-vascular system, pp: 246-257, in Reproduction and Development of Marine Invertebrates, W.H. Wilson jr, S.A. Stricker, and G.L. Shinn eds., Johns Hopkins Press

Janies, D.A. and L.R. McEdward. 1994. Heterotopy, pelagic direct development, and new body plans in velatid asteroids, pp:319-324, in Echinoderms through Time (proceedings of the 8th International Echinoderms Conference, Dijon, France), B. David et al., eds, Balkema Press, Rotterdam, Netherlands.

McEdward, L. R. and D. A. Janies. 1993. Life cycle evolution in asteroids: What is a larva? Biological Bulletin 184:255-268.

Janies, D. A. and L. R. McEdward. 1993. Highly derived coelomic and water-vascular morphogenesis in a starfish with pelagic direct development. Biological Bulletin 185:56-76.

McEdward, L. R. 1992. Morphology and development of a unique type of pelagic larva in the starfish Pteraster tesselatus. Biological Bulletin 182:177-187.

McEdward, L. R. & F.S. Chia. 1991. Size and energy content of eggs from echinoderms with pelagic lecithotrophic development. Journal of Experimental Marine Biology and Ecology 147:95-102.

Paulay, G. & L.R. McEdward. 1990. A simulation model of island reef morphology: The effects of sea level fluctuations, growth, subsidence, and erosion. Coral Reefs 9:51-62. Received best paper award from the journal Coral Reefs for 1990.

Qian, P.Y., L.R. McEdward, & F.S. Chia. 1990. Effects of delayed settlement on survival, growth, and reproduction in the spionid polychaete, Polydora ligni. International Journal of Invertebrate Reproduction and Development 18:147-152.

McEdward, L.R., S.F. Carson & F.S. Chia. 1988.. Energetic content of eggs, larvae, and juveniles of Florometra serratissima and the implications for the evolution of crinoid life histories. International Journal of Invertebrate Reproduction and Development 13:9-21.

McEdward, L.R. 1988. Experimental embryology as a tool for studying the evolution of echinoderm life histories. In: Echinoderm Phylogeny and Evolutionary Biology, A. Smith & C.R.C. Paul (eds.). Ch. 15, pp:189-196.

Sinervo, B. & L.R. McEdward. 1988. Developmental consequences of an evolutionary change in egg size: An experimental test. Evolution 42:885-900.

McEdward, L.R. & R.R. Strathmann. 1987. The body plan of the cyphonautes larva of bryozoans prevents high clearance rates: Comparison with the pluteus and a growth model. Biological Bulletin 172:30-45.

Emlet, R.B., L.R. McEdward & R.R. Strathmann. 1987. Echinoderm larval ecology viewed from the egg. in: Echinoderm Studies, Vol. 2, pp:55-136, edited by M. Jangoux & J.M. Lawrence. A.A. Balkema.

McEdward, L.R., & S.F. Carson. 1987. Variation in organic content and its relationship with egg size in the starfish Solaster stimpsoni. Marine Ecology Progress Series 37:159-169.

McEdward, L.R. & L.K. Coulter. 1987. Egg volume and energetic content are not correlated among sibling offspring of starfish: Implications for life history theory. Evolution 41:914-917.

McEdward, L.R. 1986. Comparative morphometrics of echinoderm larvae. I. Some relationships between egg size and initial larval form in echinoids. Journal of Experimental Marine Biology and Ecology 96:251-265.

McEdward, L.R. 1986. Comparative morphometrics of echinoderm larvae. II. Larval size, shape, growth, and the scaling of feeding and metabolism in echinoplutei. Journal of Experimental Marine Biology and Ecology 96:267-286.

Strathmann. R.R. & L.R. McEdward. 1986. Cyphonautes' ciliary sieve breaks a rule of biological inference. Biological Bulletin 171:694-700.

McEdward, L.R. 1985. An apparatus for measuring and recording the depth dimension of microscopic organisms. Transactions of the American Microscopical Society 104:194-200.

McEdward, L.R. 1985. Effects of temperature on the body form, growth, ETS activity, and development rate of an echinopluteus. Journal of Experimental Marine Biology and Ecology 93:169-181.

Day, R.L. & L.R. McEdward. 1984. Aspects of the physiology and ecology of pelagic larvae of marine benthic invertebrates. pp: 93-120. in: Marine Plankton Life Cycle Strategies, edited by K.A. Steidinger & L.M. Walker. CRC Press.

McEdward, L.R. 1984. Morphometric and metabolic analysis of the growth and form of an echinopluteus. Journal of Experimental Marine Biology and Ecology 82:259-287.

McEdward, L.R. & J.M. Lawrence. 1981. Respiratory electron transport system activity during larval development of Lytechinus variegatus Lamarck (Echinodermata: Echinoidea). Comparative Biochemistry and Physiology 70B:653-655.

Diehl, W.J., L.R. McEdward, E. Proffitt, V. Rosenberg & J.M. Lawrence. 1979. The response of Luidia clathrata (Echinodermata: Asteroidea) to hypoxia. Comparative Biochemistry and Physiology 62A:669-67