The many friends and colleagues of Steve Le Comber were devastated to learn of his untimely and sudden death from a heart attack at the age of 53. As an evolutionary and mathematical biologist with a huge passion for teaching, outreach and public engagement, it is a great personal loss for many and leaves an irreplaceable absence in the Organismal Biology Department of Queen Mary University of London where Steve had spent most of the last twenty‐four years. Steve’s work covers a wide range of subjects within evolutionary biology, including mathematical and computer models of molecular evolution and studies of spatial patterns in biology, notably in epidemiology and invasive species biology but with much of it focusing on the mathematics of spatial patterns. Steve was very active in public engagement and outreach delivering hundreds of talks to schools at science festivals and other venues, including our local St Paul’s Way Trust School where he contributed talks every year with Brian Cox. He was a much‐loved lecturer and colleague and is missed by both students and staff.

Before moving into science, Steve had a successful career as a journalist from 1984 to 1995, first for DC Thomson in his home town of Dundee, writing for Jackie magazine (including a stint as fiction editor) and undertaking sports reporting for their local and national newspapers. Later, he moved to London and among other things was Sub‐Editor for Woman's Own and Production Editor on Prima magazine (which at the time was selling over 1 million copies), as well as freelance work writing for Best and Bella magazines and editing a young person's magazine for the Sunday Times.

By 1995, Steve had come to the conclusion that he wanted to move into science writing. A passion for science was already in the family – his father Peter Le Comber FRS was a well‐known and respected solid‐state physicist. In order to achieve his goal, Steve decided that he needed more formal training and enrolled for a degree in the School of Biological Sciences at what was then Queen Mary and Westfield College. As a highly motivated and enthusiastic student, he was immediately successful, winning the Bevan Prize for outstanding academic achievement in biological sciences in 1996 and the Elizabeth Hocart Prize in Botany and Westfield Trust Prize in 1998. Steve graduated with a first‐class degree in Ecology and Genetics in 1998, soon returning to Queen Mary to undertake a PhD, supervised by Carl Smith and Chris Faulkes on ‘Alternative male mating tactics in the three‐spined stickleback, Gasterosteus aculeatus’ (2003). This involved arduous fieldwork (which Steve soon came to love) floating in Scottish streams observing fish for many hours, to the bemusement of the locals. Equally entertaining was the work with laboratory‐based experimental stickleback populations, established in a system of around 12 paddling pools set up to observe the fish mating in controlled situations, and in the process depleting the local area of children’s inflatable pools. The first paper from his PhD was subsequently published in the Journal of Zoology (Le Comber et al., 2003). Although Steve only worked on fish for a relatively short time, a spin‐off project into a new area on polyploidy in fish is his most accessed paper, with over 150 citations at the time of writing (Le Comber & Smith, 2004).

Steve began working with Chris Faulkes as early as 1998, whilst he was still in his final year of undergraduate studies, offering invaluable proofreading and editing advice, skills honed during his time as a journalist. At this time, Steve came up with the idea of using the mathematics of fractals to measure the branching complexity and shape of mole‐rat burrows, and this was to be the subject of his first publication (Le Comber et al., 2002), and several more followed on this topic, which have been well cited – including two published in Journal of Zoology (Le Comber et al., 2004, 2006a; Sichillima et al., 2008; Thomas et al., 2009). Steve remained close to the African mole‐rat research world, making invaluable contributions to several other publications (Faulkes et al., 2010; Katandukila et al., 2014) and joining several field trips in South Africa in the company of Journal of Zoology’s Editor in Chief, Nigel Bennett. These were always great adventures and those that knew Steve will be familiar with tales of accidentally becoming involved with some hair‐raising quad biking in the Drakensberg Mountains, and digging holes under the perimeter fence of a medium‐security prison looking for mole‐rats (with the last‐minute permission of the prison governor)!

Keeping with the theme of shape analysis, Steve went on to undertake postdoctoral work on bats and the evolution of their tooth shape with Queen Mary lecturer and Palaeontologist David Polly (Polly et al., 2005), and analysis of bat pedigrees and mating systems (Rossiter et al., 2005). With a rapidly expanding publication record and range of collaborations and project ideas, Steve was successful in becoming appointed as a lecturer in Anatomy at Queen Mary in 2006 and senior lecturer in 2013. He was fully immersed in teaching and outreach, and for many years was hugely successful as the SBCS senior admissions tutor. He thoroughly enjoyed teaching on field trips, which he viewed as holidays! Between 2008 and 2016, he also served on the editorial board of the Journal of Zoology.

Once appointed as a lecturer, Steve pioneered the introduction of geographic profiling – a statistical technique originally developed to prioritize the investigation of serial murders – to a wide range of topics in biology. This came about from a chance acquaintance arising from a seminar invitation to Canadian ex‐policeman Kim Rossmo, who originally developed the technique of geographic profiling in criminology. Steve further advanced the mathematics underlying the model, introducing a Bayesian Dirichlet process mixture (DPM) suitable for cases with large, unknown numbers of sources. In other studies, he used geographic profiling to investigate the source of invasive species, disease outbreaks and wildlife crime (Le Comber et al., 2006b; Le Comber et al., 2011; Le Comber & Stevenson, 2012; Verity et al., 2014; Faulkner et al., 2015, 2017). These wide‐ranging and varied applications really became the keystone of Steve’s research, but his ingenuity and an eye for an interesting story meant that often, these cases highlighted his ability to think outside the box. He used the same underlying mathematics in some surprising applications – from a Second World War British Secret Service operation designed to deceive the Germans about the accuracy of the V‐2 rocket bombardment on London (Aygin et al., 2019), to helping to identify Banksy (Hauge et al., 2016) or understanding the origins of London’s ring‐necked parakeets (Heald et al., 2019).

史蒂夫·勒·康伯（Steve Le Comber）的许多朋友和同事都为得知他53岁因心脏病发作而过早和突然死亡而感到震惊。作为一名进化论和数学生物学家，他对教学，推广和公众参与充满热情，对于许多人而言，这是巨大的个人损失，并且在伦敦女王玛丽大学的生物生物学系留下了无可替代的缺席，史蒂夫在过去的二十四年中大部分时间都在那里度过。史蒂夫（Steve）的工作涵盖了进化生物学的广泛领域，包括分子进化的数学和计算机模型以及生物学中空间模式的研究，尤其是流行病学和入侵物种生物学，但其中很多都侧重于空间模式的数学。史蒂夫（Steve）积极参与公众参与和外展活动，在科学节和其他场所为学校提供了数百场演讲，包括我们当地的圣保罗之路信托学校（St Paul's Way Trust School），他每年与布莱恩·考克斯（Brian Cox）进行演讲。他是一位备受推崇的讲师和同事，深受学生和员工的喜爱。

在涉足科学领域之前，史蒂夫（Steve）在1984年至1995年期间曾是一名成功的新闻工作者，首先是在其家乡邓迪（Dundee）的DC汤姆森（DC Thomson），为《杰基》杂志（包括担任小说编辑的工作）撰写文章，并为当地和国家进行体育报道报纸。后来，他移居伦敦，除其他事项外，还担任《妇女》杂志的副编辑和《Prima》杂志（当时销量超过100万册）的制作编辑，以及为Best和Bella杂志撰写自由作家并编辑《星期日泰晤士报》的年轻人杂志。

到1995年，史蒂夫已经得出结论，他想从事科学写作。家庭中已经充满了对科学的热情-他的父亲Peter Le Comber FRS是一位著名且受人尊敬的固态物理学家。为了实现他的目标，史蒂夫决定他需要更多的正规培训，并在当时的玛丽皇后大学和韦斯特菲尔德学院就读了生物科学学院的学位。作为一个充满激情和热情的学生，他立即获得了成功，1996年因其在生物科学方面的杰出学术成就而获得了Bevan奖，并于1998年获得了伊丽莎白·霍卡特的植物学奖和Westfield Trust奖。Steve毕业于生态学一等学位。 1998年获得遗传学和遗传学博士学位，不久后返回玛丽女王大学攻读博士学位，Gasterosteus aculeatus（2003）。这涉及艰苦的实地考察（史蒂夫很快就爱上了它）在苏格兰的溪流中漂浮着，观察了许多小时的鱼，这使当地人感到困惑。同样具有娱乐性的是基于实验室的实验性棘背类种群的工作，该种群建立在由大约12个戏水池组成的系统中，以观察受控情况下的鱼类交配，并在此过程中耗尽儿童充气水池的局部区域。他的博士学位的第一篇论文随后发表在《动物学杂志》上（Le Comber等人，2003年）。尽管史蒂夫仅在相对较短的时间内从事鱼类研究，但将其分解为鱼类多倍体的新领域是他访问最多的论文，撰写本文时被引用了150多次（Le Comber＆Smith，2004年）。

史蒂夫（Steve）早在1998年就开始与克里斯·福克斯（Chris Faulkes）合作，当时他仍处于本科学习的最后一年，提供了宝贵的校对和编辑建议，并在他成为记者的过程中磨练了技巧。这时，史蒂夫（Steve）提出了使用分形数学来测量mole鼠洞穴的分支复杂性和形状的想法，这是他的第一篇出版物的主题（Le Comber等，2002），还有几个跟着就这个话题，已经得到了很好的引用-包括出版了两动物学杂志（乐精梳机等，2004年，2006年a ; Sichillima等，2008 ;托马斯等。，2009年）。史蒂夫（Steve）与非洲mole鼠研究世界保持着密切的联系，为其他几本出版物（Faulkes等，2010； Katandukila等，2014）做出了宝贵贡献，并在《动物学杂志》的编辑的陪同下参加了南非的几次实地考察。总负责人奈杰尔·贝内特（Nigel Bennett）。这些历来都是伟大的冒险，而那些知道史蒂夫会熟悉偶然地参与德拉肯斯堡山上一些养毛四轮摩托的故事，以及在中等安全监狱的围墙下挖洞寻找痣的故事（经监狱长的最后许可）！

遵循形状分析的主题，Steve继续进行蝙蝠的博士后研究，并与玛丽皇后讲师和古生物学家David Polly进行了牙齿形状的演变（Polly等人，2005年），并分析了蝙蝠的血统和交配系统（ Rossiter等，2005）。凭借迅速增长的出版记录以及广泛的合作和项目构想，Steve于2006年成功被任命为玛丽皇后号解剖学讲师，并于2013年被任命为高级讲师。他全神贯注于教学和外展工作，多年来作为SBCS高级招生导师，取得了巨大的成功。他非常喜欢野外旅行教学，他认为这是假期！在2008年至2016年之间，他还担任过《动物学杂志》的编辑。

一旦被任命为讲师，史蒂夫（Steve）率先将地理分布图（一种最初用于优先研究连环谋杀案的统计技术）引入生物学的广泛主题。这是由于向加拿大前警官金·罗斯莫（Kim Rossmo）发出研讨会邀请而偶然认识的，他最初开发了犯罪学中的地理分布图技术。史蒂夫（Steve）进一步推进了该模型的数学基础，引入了贝叶斯Dirichlet过程混合物（DPM），适用于来源数量众多，来源不明的情况。在其他研究中，他使用地理分布图调查了入侵物种，疾病暴发和野生动植物犯罪的来源（Le Comber等，2006b； Le Comber等，2011年；Le Comber和Stevenson，2012年；Verity等。，2014 ; 福克纳等。，2015年，2017年）。这些广泛而多样的应用程序确实成为了史蒂夫研究的基石，但是他的独创性和对有趣故事的关注意味着，这些案例常常突显出他的创新思维能力。他在一些令人惊讶的应用中使用了相同的基础数学-从第二次世界大战英国特勤局的一项旨在欺骗德国人有关伦敦V-2火箭轰击的准确性的骗局（Aygin等人，2019年），以帮助识别班克斯（Hauge等，2016）或了解伦敦环颈长尾小鹦鹉的起源（Heald et al。，2019）。