Publications

Selected publications

King, B. and Rücklin, M. (2020). a Bayesian approach to dynamic homology of morphological characters, and the ancestral phenotype of jawed vertebrates.

What exactly the ancestors of jawed vertebrates looked like has been a topic of vigorous debate. A particularly interesting question is what form the first jaws took. However, different hypotheses about the jaw bones of the placoderms complicate the picture: they are either the equivalent of the palatal bones of boney fishes, or represent maxillae and premaxillae. In this paper we introduced a method that can calculate the evolutionary tree of early jawed vertebrates while simultaneously estimating bone correspondences between placoderms and boney fishes. We found that the ancestors of jawed vertebrates most likely resembled the "maxillate placoderms", from the Silurian period of China, and had maxillae and premaxillae but no palatal bones.

Journal website

King, B. (2020). Bayesian tip-dated phylogenetics in paleontology: topological effects and stratigraphic fit. Systematic Biology 70: 283-294.

A focus of my research since my PhD is the use of a type of phylogenetic analysis (evolutionary tree building) known as tip dating. In this study, I investigated how the use of tip dating affects the placement of fossils in phylogenetic trees. In particular, I wanted to know how the use of tree models that include the ages of fossils influences results. I found that tip dating does indeed have a substantial effect, greater than the differences between other phylogenetic methods. Trees produced by tip dating analysis have a better fit to the fossil record: for example, this method will disfavour trees with a very ancient species related to very recent species. The difference in the phylogenetic relationships is most clear with highly incomplete fossils. In other words, when there is very little data to tell us the relationships of a fossil, tip dating will tend to place this fossil in a position consistent with its age. Other methods on the other hand, might suggest quite surprising trees, for example suggesting that a highly incomplete fossil is the first example of a group that otherwise does not appear until many millions of years later.

pdf Journal website

King, B. and Beck, R.M.D. (2020). Tip dating supports novel resolutions of controversial relationships among early mammals. Proceedings of the Royal Society B 287: 20200943.

During the time of the dinosaurs, several groups of early mammals possessed a distinctive type of tooth with longitudinal rows of cusps. The relationships of these mammal groups, collectively known as the ‘allotherians’, have been debated, with one hypothesis being that they all descend from a single common ancestor. We analysed their relationships using a method known as ‘tip dating’, which analyses the age and morphology of fossils simultaneously, and found instead that ‘allotherians’ from different time periods were split into three separate groups. This suggests that early mammal evolution was complex, with different groups converging on similar lifestyles.

pdf Journal website

King, B. (2019). Which morphological characters are influential in a Bayesian phylogenetic analysis? Examples from the earliest osteichthyans. Biology Letters 15: 20190288.

A major ongoing debate within palaeontology is which methods should be used to build evolutionary trees of relationships. The two major competing methods are "parsimony" which looks for the tree that explains evolution in the simplest way, and "Bayesian", which is a statistical method employing a model of evolution. In the area of vertebrate palaeontology, this argument is very important for figuring out when the osteichthyans (the bony fishes, which includes all tetrapod vertebrates as well as most fishes) first appeared. The estimated relationships of the earliest examples of bony fishes are very different under Bayesian and parsimony methods. The Bayesian result suggests a surprisingly ancient origin for the modern groups of bony fishes. However, a problem with Bayesian analysis of morphological data is the somewhat blurry connection between the final result and the anatomical data used as input. In this paper I introduced a new and simple method to figure exactly which anatomical features are driving the Bayesian analysis. I found that Bayesian analyses are overly reliant on small parts of the anatomical data, and that in the case of the earliest bony fishes, the result from parsimony may in fact be more reliable.

pdf Journal website

King, B.*, Young, G.C.* and Long, J.A. (2018). New information on Brindabellaspis stensioi, Young, 1980, highlights morphological disparity in Early Devonian placoderms. Royal Society Open Science 5:180094.

Brindabellaspis is a member of the placoderms, a group of fishes that went extinct 360 million years ago. It comes from the Burrinjuck fossil site near Canberra, which preserves a 400 million-year-old coral reef ecosystem. Fossils from Burrinjuck are known for their amazing preservation of braincase anatomy, and because of this Brindabellaspis is one of the best known, but also most controversial, placoderms. In this paper we describe the previously unknown front end of the skull. Brindabellaspis turns out to have a remarkable elongated bill, similar to a paddlefish or platypus, making it one of the most bizarre known placoderms. The bill also contains a unique adaptation of the lateral line sensory system, not seen in any fish, living or fossil.

pdf Journal website

Clement A.M.*, King, B.*, Giles, S.*, Choo, B., Ahlberg, P.E., Young, G.C. and Long, J.A. (2018). Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution. eLife 7:e34349.

In 2015, during a visit to the Burrinjuck fossil site with Gavin Young, I found a small skull of an osteichthyan (bony fish). After painstaking acid-preparation to remove limestone around the bones, the tiny specimen was revealed to belong to Ligulalepis, which was previously known from a single incomplete specimen described in the 1990s, and which has been igniting debate ever since. The new specimen preserved the front half of the skull roof, which was missing in the original specimen. The original specimen was also re-analysed using CT scanning to reveal the nerves and braincase morphology, led by Alice Clement. Combined, our study is the most detailed description of a head of a fish of this age (400 million years). Ligulalepis turns out to be have strange mixture of features from bony fishes, sharks and the extinct placoderms

pdf Journal website

King, B., Hu, Y. and Long, J.A. (2018). Electroreception in early vertebrates: survey, evidence and new information. Palaeontology 61(3):325–358.

Although often overlooked because we humans lack it, electroreception is a widespread and important sensory system in aquatic vertebrates. However, its origins remain mysterious. In this paper we presented the first in-depth review of possible electroreceptors in early vertebrate fossils, presenting a comprehensive suite of new data from micro-CT scans of exceptionally-preserved fossil material. Many of the possible electroreceptors from the extinct placoderms do not stand up to scrutiny, and we find no evidence for electroreception in this group. However, we found that even the earliest osteichthyans (bony fishes) had elaborate and specialised electrosensory systems.

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King, B., Qiao, T., Lee, M.S.Y., Zhu, M. and Long J.A. (2017). Bayesian morphological clock methods resurrect placoderm monophyly and reveal rapid early evolution in jawed vertebrates. Systematic Biology 66(4):599–516

This paper formed the centrepiece of my PhD. For several years, the extinct placoderms have been viewed as a "paraphyletic" group. This means that the modern groups of jawed vertebrates (sharks, fishes, tetrapods) evolved from within placoderms, and some placoderms are more closely related to the modern groups than others. However, various discoveries led to doubts in the minds of researchers, including the presence of claspers (mating organs) in several placoderm groups; perhaps the old view that placoderms form a coherent group of their own is true after all. To investigate this problem I compiled the largest dataset of anatomical features of early vertebrates, and analysed them with "tip-dated" methods. These methods model fossil sampling, morphological evolution and fossil ages, and I suspected they may be better able to deal with the unique problems of early vertebrates. In contrast to the currently accepted view, I found strong evidence for placoderm "monophyly" (they form a group descended from a common ancestor).

pdf Journal website

King, B. and Lee M.S.Y. (2015) Ancestral state reconstruction, rate heterogeneity and the evolution of reptile viviparity. Systematic Biology 64(3):532–544.

The evolution of live birth from egg laying in reptiles is a classic evolutionary case study. It has long been thought that live birth evolved many times from egg-laying, but that the reverse transition almost never occurred. However, a high-profile study published in 2013 claimed to turn the established view on its head: they suggested live birth evolved early, and there were multiple reversals to egg-laying. We re-analysed the data using multiple methods, including some newly introduced, to show that the result of the 2013 study was a statistical artifact. The problem was that their model assumed that life birth evolved at the same rate across all lizard groups. In truth some groups like Geckos (left, below) are conservative, almost all lay eggs, whereas other groups such as skinks (left, above), go crazy with closely related species differing in their reproduction. We instead found that live birth evolved 115 times, whereas egg laying evolved from live birth only 3 times. The methods we proposed are increasingly being used by evolutionary biologists, as larger and larger evolutionary trees are utilised for analysis of biological traits.

pdf

King, B. and Denholm, B. (2014). Malpighian tubule development in the red flour beetle (Tribolium castaneum). Arthropod Structure and Development 43:605–613.

Malpighian tubules are the excretory organs of insects. The tubules of the fruit fly Drosophila have been extensively studied, but other insects far less. Here we described the embryonic development of Malpighian tubules in the beetle Tribolium. This is particularly significant because this beetle has what is known as a cryptonephridial system, a powerful water-saving device. In the cryptonephridial system, the ends of the Malpighian tubules attach to the rectum and maintain a high salt concentration, drawing water out of the excreted waste.

pdf Journal website

Full publications list

Rücklin, M., King, B., Cunningham, J., Marone, F. and Donoghue, P.C.J. (2021). Acanthodian dental development and the origin of gnathostome dentitions. Nature Ecology and Evolution

King, B. and Rücklin, M. (2020). A Bayesian approach to dynamic homology of morphological characters and the ancestral phenotype of jawed vertebrates. eLife.

King, B. (2020). Bayesian tip-dated phylogenetics in paleontology: topological effects and stratigraphic fit. Systematic Biology: 70: 283-294.

King, B. and Rücklin, M. (2020). Tip dating with fossil sites and stratigraphic sequences. PeerJ 8:e9368.

King, B. and Beck, R.M.D. (2020). Tip dating supports novel resolutions of controversial relationships among early mammals. Proceedings of the Royal Society B 287: 20200943.

King, B. (2019). Which morphological characters are influential in a Bayesian phylogenetic analysis? Examples from the earliest osteichthyans. Biology Letters 15: 20190288.

King, B.*, Young, G.C.* and Long, J.A. (2018). New information on Brindabellaspis stensioi, Young, 1980, highlights morphological disparity in Early Devonian placoderms. Royal Society Open Science 5:180094.

Clement A.M.*, King, B.*, Giles, S.*, Choo, B., Ahlberg, P.E., Young, G.C. and Long, J.A. (2018). Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution. eLife 7:e34349.

King, B., Hu, Y. and Long, J.A. (2018). Electroreception in early vertebrates: survey, evidence and new information. Palaeontology 61(3):325–358.

King, B., Qiao, T., Lee, M.S.Y., Zhu, M. and Long J.A. (2017). Bayesian morphological clock methods resurrect placoderm monophyly and reveal rapid early evolution in jawed vertebrates. Systematic Biology 66(4):599–516.

Qiao, T., King, B., Long, J.A., Ahlberg, P.E., Zhu, M. (2016). Early gnathostome phylogeny revisited: multiple method consensus. PLoS ONE 11(9): e0163157.

Lee, M.S.Y., Sanders, K.L., King, B. and Palci, A. (2016). Diversification rates and phenotypic evolution in venomous snakes (Elapidae). Royal Society Open Science 3:150277.

King, B. and Lee M.S.Y. (2015) Ancestral state reconstruction, rate heterogeneity and the evolution of reptile viviparity. Systematic Biology 64(3):532–544.

King, B. and Lee M.S.Y. (2015). Epoch based likelihood models reveal no evidence for accelerated evolution of viviparity in squamate reptiles in response to Cenozoic climate change. Journal of Experimental Zoology B 324(6):525–31. Journal website . pdf

King, B. and Denholm, B. (2014). Malpighian tubule development in the red flour beetle (Tribolium castaneum). Arthropod Structure and Development 43:605–613.

*Denotes equal contribution