We analyzed COI barcode sequences from 138 over-a-century old specimens of Calinaga including 36 name-bearing type specimens stored at the Natural History Museum London. These new data, combined with previously available RPS5 sequences, divide the Calinaga samples into four well-supported mitochondrial lineages that together with a novel wing-pattern analysis, support the recognition of six species (lhatso, buddha, brahma, aborica, formosana and davidis), with all other names subsumed either as subspecies or synonyms. One new taxon is described, Calinaga aborica naima Vane-Wright, ssp. n.

This essay presents various reflections on living systems, what they are and how they evolve, prompted by editing Teleonomy in Living Systems (a special issue of the Biological Journal of the Linnean Society). Conclusions include the suggestion that the linked notions of teleonomy and agency represent fundamental properties of matter that become apparent only when organized in the way that we consider to be that of a living system. As such, they are factors that form part of the intrinsic 'a priori' of living systems, as they evolve in form through space and time. Biology, the science of life and living systems, needs to be 'biological' if it is to be anything at all. Understanding the role of teleonomy (internal, inherent goal-seeking) will always play a necessary part in this endeavour: teleonomy represents one of the fundamental properties of living systems.

An introduction, overview and discussion are provided for this special issue of the Biological Journal of the Linnean Society, entitled 'Teleonomy in Living Systems'. The introduction offers a brief account of the origin of Colin Pittendrigh's notion of teleonomy, in many ways comparable to Dobzhansky's later proposal of 'internal teleology', and a subsequent externalist interpretation of the concept by Ernst Mayr which, unfortunately, has remained influential. As part of the growing movement that places organismic purpose, goal-directedness and agency back at the centre of biology, in June 2021 a 2-day online international meeting was organized by the authors through the Linnean Society of London, under the title Evolution 'On Purpose': Teleonomy in Living Systems. Our overview provides a summary, with some commentary, for each of 15 papers presented here. Together with a complementary volume of 18 papers published by MIT Press, they form a selected and extended proceedings. These papers represent scientifically founded views of evolutionary biologists and philosophers of science who seriously question the adequacy of the neo-Darwinian Modern Synthesis to account for the purposive nature of living systems. Like any other phenomena associated with life, purposive and teleonomic behaviours demand evolutionary explanations and context. Without any recourse to supernatural or non-material processes, various approaches to trying to understand how this goal-directed, teleonomic property of life has influenced the course of evolution are explored.

This paper discusses three problems concerning the Woodland Grayling,Hipparchia fagiScopoli, 1763, with respect to the identity and application of the junior namePapilio hermioneLinnaeus, 1764. In 1977, the late Otakar Kudrna designated a specimen of the Rock Grayling,Hipparchia alcyone[Denis & Schiffermüller], 1775, to become the lectotype ofPapilio hermione– as a result of whichhermionesupplantedalcyoneas the senior epithet for this species. BecauseP. hermioneis the nominal type species ofHipparchiaFabricius, 1807, Kudrna’s action rendered this a genus based on a misidentified species. Third, while a majority of lepidopterists have ignored Kudrna’s action and continue to apply the nameH. alcyoneto the Rock Grayling, and still regardP. hermioneas a junior subjective synonym ofH. fagi, the formal nomenclature for the Rock Grayling has become unstable because a large minority have nonetheless accepted Kudrna’s lectotype designation and all that follows from it. It is demonstrated here that no syntypes ofPapilio hermione(orPapilio fagi) have survived; consequently, Kudrna’s lectotype designation forP. hermioneis invalid. By designation of a single specimen of the Woodland Grayling as neotype for bothP. fagiandP. hermione, the two names are rendered objectively synonymous, thereby restoring stability to the species name for the Rock Grayling (asHipparchia alcyone), and to the application ofPapilio hermione(=Hipparchia fagi) as nominal type species of the generic nameHipparchia.

The global increase in species richness toward the tropics across continents and taxonomic groups, referred to as the latitudinal diversity gradient, stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluate several of these hypotheses to explain spatial diversity patterns in a butterfly family, the Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal, and also the extent to which these processes differ among regions at the same latitude. We generate a time-calibrated phylogeny containing 2,866 nymphalid species (~45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the latitudinal diversity gradient among regions because temporal diversification dynamics differ greatly across longitude. The Neotropical diversity results from low extinction rates, not high speciation rates, and biotic interchanges with other regions are rare. Southeast Asia is also characterized by a low speciation rate but, unlike the Neotropics, is the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, combined with tropical niche conservatism, played a major role in generating the modern latitudinal diversity gradient of nymphalid butterflies.

The life of Father Jean Xavier Hyacinthe Montrouzier (1820-1897) is briefly reviewed, with special reference to the 5 years he spent on Woodlark Island (Muyua), Papua New Guinea, and his subsequent academic studies while briefly resident in Australia. Of the four species listed by Montrouzier as members of the genus Euploea in his 1856 paper on the Lepidoptera of Woodlark, two do undoubtedly belong to the genus as currently recognised: Euploea leucostictos and E. boisduvalii (Danaini: subtribe Euploeina). A third species, originally described by Macleay as a species of Euploea but now included in the genus Tirumala (Danaini, Danaina), was correctly identified by Montrouzier. The fourth, for which Montrouzier introduced a new species name, appears to represent either a species of Neptis (Limenitidinae) or Tellervo (Danainae, Tellervini). This name, Euploea transfixa Montrouzier, is however both a nomen dubium and a nomen oblitum.

The nominal taxon Papilio vanessa Fabricius, 1793, for long thought to be an unrecognised species of Riodinidae from the Americas, is shown to be a synonym of the West African lycaenid butterfly Liptena septistrigata (Bethune-Baker, 1903), syn. nov. Papilio vanessa is however a nomen oblitum, as it has not been applied to any recognised taxon since 1889. Thus Liptena septistrigata has priority so long as this subjective synonymy is considered valid and, within this context, L. septistrigata should be treated as a nomen protectum.

Chalk rivers and streams are of conservation importance due to their ecological diversity, historical relevance and economic value. With more than 200 chalk watercourses, England is considered unusual in having the most chalk rivers in the world. However, due to increasing anthropogenic activities, many English chalk rivers and streams are becoming badly degraded. The non-biting midges or chironomids (Diptera, Chironomidae) are considered key-stone taxa in aquatic food webs, and have been used as ecological indicators of freshwater quality and environmental stress. Here we determined the generic richness, diversity, and community structure of Chironomidae across six sites in the mid-section of the Great Stour in Kent, a chalk river for which concern has been expressed regarding both water and habitat quality. Based on the morphological identification of 1336 insect larvae from the six sites (four in Westgate Parks, Canterbury, and two at nearby locations upstream and downstream from Canterbury City), a total of 20 genera of Chironomidae were identified, including some taxa indicative of freshwater habitats with low levels of organic pollution. There were different levels of generic richness and diversity among sites, and while there was little variation in the community composition among the sites within Westgate Parks, there were noticeable generic differences among Westgate Parks sites compared with those upstream and downstream, showing the highest complementarity and Beta diversity values. Overall, the results were comparable with other studies on chironomids in chalk rivers and other river systems. Although spatially limited to a small stretch of river, this represents the first study on chironomids in the Great Stour and provides baseline information on the diversity and structure of this important insect group with aquatic larvae, useful for the objective interpretation of any future biological assessments and monitoring programmes on the Kentish Stour, and also for comparisons with other chalk rivers.

1. The chemical defences of monarch butterflies involve two kinds of secondary plant metabolites: cardiac glycosides (CGs) that are obtained from larval hostplants, and pyrrolizidine alkaloids (PAs) that are gathered by adults usually independent from feeding behaviour. While monarchs and CGs have received great attention, monarchs and PAs appear largely ignored in science as well as by the public. 2. We review and discuss PA‐pharmacophagy specifically for Danaus plexippus and its closest relatives and aim to encourage research and future consideration of this peculiar type of insect‐plant relationship, which represents an intrinsic but surprisingly understudied aspect of the lives of adult monarchs.

The latitudinal diversity gradient (LDG) is arguably one of the most striking patterns in nature. The global increase in species richness toward the tropics across continents and taxonomic groups stimulated the formulation of many hypotheses to explain the underlying mechanisms of this pattern. We evaluated several of these hypotheses to explain spatial diversity patterns in the butterfly family, Nymphalidae, by assessing the contributions of speciation, extinction, and dispersal to the LDG, and also the extent to which these processes differ among regions at the same latitude. We generated a new, time-calibrated phylogeny of Nymphalidae based on 10 gene fragments and containing ca . 2,800 species (∼45% of extant diversity). Neither speciation nor extinction rate variations consistently explain the LDG among regions because temporal diversification dynamics differ greatly across longitude. For example, we found that Neotropical nymphalid diversity results from low extinction rates, not high speciation rates, and that biotic interchanges with other regions were rare. Southeast Asia was also characterized by a low speciation rate but, unlike the Neotropics, was the main source of dispersal events through time. Our results suggest that global climate change throughout the Cenozoic, particularly during the Eocene-Oligocene transition, combined with the conserved ancestral tropical niches, played a major role in generating the modern LDG of butterflies. ### Competing Interest Statement The authors have declared no competing interest.

The involvement of the diverticula, a synapomorphy for Itunina, in protrusion and expansion of hairpencils by male Lycorea halia (Hübner, 1816) is demonstrated for the first time. They facilitate maintaining the haemolymph pressure necessary to keep the hairpencils everted. The diverticula are curved hook-like lobes, open to the body cavity and densely filled with tracheae and threads made by units of two staggered cells surrounding a central extracellular fibril bundle. Such complex structures, apparently metabolically active, have not been reported for insects previously and might indicate additional functions, but their functional role(s) remains a puzzle. When a male emerges from pupa, the diverticula are not yet formed; this happens only during the first protrusion of the hairpencils.

The British Swallowtail, Papilio machaon britannicus, is an iconic flagship for its unique but now restricted and fragmented fenland ecosystem in the UK's Norfolk Broads. Occurrence in 1 km2 recording squares fell by 56% over the period of 1974–2014 and by 13% in 2005–2014, but the breeding populations, mainly confined to reserves, have increased in size. Climate change‐induced sea‐level rise and consequent seawater incursion into the Norfolk Broads represent a significant existential threat to the butterfly. Translocation to more secure fenland sites is recommended; several are being restored or recreated through stakeholder partnerships in East Anglia and Somerset. Well‐researched introduction and management of the foodplant, Milk‐parsley, is essential for such translocations to succeed. A better understanding is needed of the genetic structuring of the British Swallowtail populations using modern sequencing technologies, in particular to elucidate the significance of gene flow in relation to the viability of introductions to small or isolated sites. In a species in which hybridisation is commonplace across the Holarctic, the continued influx and future spread in Britain of the continental subspecies Papilio machaon gorganus may present a threat to the genetic integrity of subspecies britannicus, despite their differences in habitat and larval foodplants.

Although the contributions of James Petiver to the early development of systematic natural history are widely acknowledged, he is often criticized for scientific, curatorial and even social shortcomings. This rather dubious reputation is at odds with his standing among entomologists as 'the father of British butterflies'. Shortly before his death in 1718, Petiver published a densely packed eight-page pamphlet entitled Papilionum Britanniae. Analysis of this work, which at first sight makes an apparently exaggerated claim of accounting for 'above eighty English butterflies', reveals that Petiver was an original, perceptive and truly systematic entomologist, in several important respects ahead of his time.

The type material of the available name Euploea dorippus Klug, 1845, originated from northern Sudan, an area that lies beyond the core zone of the semispecies or subspecies currently known as Danaus chrysippus dorippus, and the description did not include examples of the phenotype currently referred to as Danaus chrysippus f. ‘dorippus’. Possible consequences for nomenclature of the infraspecific species group taxa and form names of Danaus chrysippus are discussed.

The challenging issue of animal welfare has focused mainly on furred and feathered vertebrates. However, unnoticed by most people, literally billions of insects are kept in captivity, in increasing numbers, and traded for a great variety of purposes. Arguably the most successful animals on Earth, insects are ignored or actively disliked by most people. Not just the different appreciation of insects by humans but the diversity of insects, and the diversity of their ecosystem services, shows that a discussion of insect welfare requires different criteria than vertebrate welfare. Their biology is very different, and insects are far less tolerant of suboptimal conditions. As a result, successful insect breeding programmes must necessarily fulfil basic welfare requirements. Insect natural history illustrates the complexity of practical welfare, even without fundamental consideration of insects as animals that have intrinsic value and their own agency, and the extent to which they are conscious or not and may or may not suffer pain. The great variety of insect lifestyles and lack of accessible information about industrial breeding mean that it is impossible to set general standards for insect welfare or provide meaningful evaluations of current practices. The best guidance that can be offered is to ‘keep insects under as natural conditions as possible’. However, even this cannot be adhered to. Conditions in live butterfly exhibits involve compromises. Insects released in billions as biocontrol agents often involve x-ray sterilisation or transgenic procedures and pose environmental risks. For insects bred for human food and animal feed, euthanasia is a pressing issue. Numerous questions and ethical and welfare dilemmas are raised. Despite this, formulation of an Insect Welfare Charter based on respect, and the need to pay more attention to insects, is encouraged, preferably also addressing insects living in the wild.

In this paper, we seek to predicate the status of Papilio aegyptius Schreber, 1759, as a junior subjective synonym of Danaus chrysippus chrysippus (Linnaeus, 1758). As early as 1764, these taxa were synonymised by Linnaeus, but the name aegyptius was later revived by Hans Fruhstorfer in 'Seitz' to represent a 'race' of D. chrysippus found in the Eastern Mediterranean, including Egypt and also the Sudan. George Talbot subsequently used the name for one of seven subspecies of Danaus chrysippus he recognised. In the 1970s, work by Jacques Pierre persuaded other authors to regard ssp. aegyptius as the pan-Afrotropical subspecies, contrary to Talbot's view of ssp. aegyptius being limited to only part of Africa and the Eastern Mediterranean. The 'Pierre classification' was quite widely followed for about 20 years. There are now two different, rival systems for African D. chrysippus : the 'Larsen classification', which includes all populations in a greatly expanded ssp. chrysippus, and the 'Smith classification', which divides African D. chrysippus into four named semispecies together with a large 'hybrid zone'. Both classifications agree, however, in placing both Egyptian and Eastern Mediterranean populations in the nominotypical taxon. In 'Butterflies of Turkey', Gerhard Hesselbarth, Harry van Oorschot & Sigbert Wagener placed Papilio aegyptius as a junior subjective synonym of Danaus chrysippus chrysippus, an action since independently supported by David Smith and his co-workers. A brief account of events leading to the return to synonymy of D. chrysippus aegyptius under the nominotypical subspecies is presented. In this history, differing interpretations of the type locality of Danaus chrysippus play a key role.

Campbell Smith was employed as an entomologist at London’s Natural History Museum for 35 years, during which time he worked on various catalogues, curated parts of the museum collections, and contributed to the output of several research groups. In 2009 he took early retirement to look after his widowed father. His sudden and unexpected death on 11th March 2019 came as a shock to former museum staff and various entomologists around the world, acknowledging his loss with sorrow mixed with good memories of this most amiable colleague and friend.