BLEED publications about bryozoans


Liow L. H.  &  Taylor, P. D.  2019 Cope’s Rule in a modular organism: Directional evolution without an overarching macroevolutionary trend. Evolution 73: 1863-1872

So much of the research on macroevolutionary trends are based on solitary evo13800-fig-0003-morganisms where body size has rather clear biological meaning. But what about colonial organisms? Studying module (zooid) sizes of cheilostome bryozoans through their long evolutionary history, we find that there is no dectectable trend in zooid size. However, ancestral species tend to give rise to descendent species that have larger zooid sizes than themselves. Sounds contradictory? Read our paper for explanations!


Kopperud B., Lidgard S. & Liow L. H. 2019 Text-mined fossil biodiversity dynamics using machine learning. 286:20190022 Proceedings of the Royal Society B 286 (1901) 20190022

pages-from-kopperud-larwood-2018-presentation.pngWe really wanted a dataset of observations of bryozoan taxa through geological time so we can estimate taxonomic richness and ask all kinds of cool questions about how bryozoans evolved. But we didn’t have a dataset we can play with… and  we wanted a dataset yesterday, so we used Natural Language Processing tools to help us text mine bryozoan data. Who cares? Well, maybe you do if you also want a automatically compiled dataset for your favorite  clade. Check out the compiled data and Bjørn Kopperud’s code here!

Ecology & Paleoecology

Di Martino E., Liow L.H., Perkins T., Portell R.W. & Taylor P.D. 2020 Sneaking up on ‘enemies’: alleviating inherent disadvantages in competitive outcomes in a nearly 3‐million‐year‐old palaeocommunity from Florida, USA. Lethaia. early online

2017-10-21 16.08.56Using a 3-million-year-old bryozoan paleocommunity encrusting jingle shells from Florida, we confirm our previous findings on overgrowth competition (Big is better!), and show how both poor and good competitors gain the upper hand by  ‘attacking’ colonies of other bryozoan species from the rear and the flank…….sneaky!

Liow L.H., Reitan R., Voje  K., Taylor P.D. & Di Martino E. 2019 Size, weapons and armor as predictors of competitive outcomes in fossil and contemporary marine communities. Ecological Monographs 89(2): e01354

TraitsWe were curious as to whether we could predict the outcomes of competitive interactions , both in space and (deep) time. Using a community of about 80 species that we know well, we modeled the relationship between phenotypic traits and competitive outcomes. Size, weapons and armor do matter, but so do so many other factors, including stochastic ones. Our newly developed model might be useful in predicting sumo matches and Pokémon games too.

Liow L.H., Di Martino E., Krzeminska M, Ramsfjell M, Rust S., Taylor P.D. & Voje  K. (2017) Relative size predicts competitive outcome through 2 million years. Ecology Letters 20: 981–988

2017-Ecology_Letters small

Using communities of encrusting marine cheilostome bryozoans spanning more than 2 million years from New Zealand, we show that zooid size is a significant determinant of overgrowth outcomes (spatial competition), where colonies with larger zooids tend to overgrow those with smaller zooids. We also (serendipitously) detected temporally coordinated changes in average zooid sizes across species.

Liow L.H., Di Martino E., Voje K., Rust S. & Taylor P. D. (2016) Interspecific interactions through 2 million years: are competitive outcomes predictable?
Proceedings of the Royal Society, B 283:20160981


Using species level data on competitive overgrowth spanning more than 2 million years, we find most winner species stay winners and loser species stay losers. Bryozoans of the same species tend to cluster spatially and when they do encounter each other spatially, they stop growing at edges of encounter much more often than they try to overgrow each other. Counter-intuitively (or perhaps not) competitive ability has no bearing on ecological dominance.

Systematics & Taxonomy

Di Martino E, Taylor PD & Gordon DP (2020) Erect bifoliate species of Microporella (Bryozoa, Cheilostomata), fossil and modern. European Journal of Taxonomy 678: 1–31


Orr RJS, Haugen MN, Berning B, Bock P, Cumming R, Florence W, Hirose M, Di Martino E, Ramsfjell MH, Sannum M, Smith AB, Vieira L, Waeschenbach A, E & Liow LH (2019) A genome-skimmed phylogeny of a wide-spread bryozoan family, Adeonidae. BMC Evolutionary Biology 19 (235)


Orr RJS, Waeschenbach A, Enevoldsen ELG., Boeve JP, Haugen MN, Voje KL, Porter J, Zágoršek K, Smith AM, Gordon DP & Liow LH (2018) Bryozoan genera Fenestrulina and Microporella no longer confamilial; multi-gene phylogeny supports separation. Zoological Journal of the Linnean Society 186(1): 190-199

Microporellidae, Fenestrulinidae

Gordon D, Voje K.L. and Taylor P.D. (2017) Living and fossil Steginoporellidae (Bryozoa: Cheilostomata) from New Zealand. Zootaxa 4350: 345–362


Di Martino E., Taylor P.D., Gordon D. & Liow L.H. (2017) New bryozoan species from the Pleistocene of the Wanganui Basin, North Island, New Zealand. European Journal of Taxonomy 345: 1–15

Buskia, Microporella, Parkermavella

Di Martino, E., Taylor P. D., Gordon D., and L.H. Liow. (2016) Powellithecidae, a new Pliocene to Recent bryozoan family endemic to New Zealand. European Journal of Taxonomy 207:1–17

Powellitheca, Emballotheca, Monoporella

Rosso A., Di Martino E., Sanfilippo R., Sciuto F., and Liow L.H. (2015) Resurrection of an old forgotten name: the case of the Pliocene to Recent bryozoan Cleidochasmidra portisi (Neviani, 1895) from the Mediterranean. Bollettino della Societa Paleontologica Italiana 54: 91–102.

Cleidochasmidra, Smittia

Non-bryozoan pubs from BLEED

For non-bryozoan related publications from BLEEDers, please check out their personal webpages