I A journal of world insect systematicsnsecta MundI 23 0945Natural history of Cassida sphaerula Boheman, 1854 Page Count: (Coleoptera: Chrysomelidae: Cassidinae: Cassidini) on Arctotheca prostrata (Salisb.) Britten Adam et al. (Asteraceae: Arctotidinae) in South Africa, with a checklist of South African Cassidinae (leaf-mining and tortoise beetles) Sally Adam Laaiplaats 59, Mossel Bay, South Africa Mariana Campos CSIRO Health & Biosecurity 147 Underwood Avenue, Floreat 6014 Western Australia, Australia Hugh D. C. Heron P.O. Box 39042, Escombe, Queensburgh, Natal, 4070, South Africa Charles L. Staines Smithsonian Environmental Research Center, 647 Contees Wharf Road, Edgewater, MD 21037, U.S.A. Rob Westerduijn Willem Klooslaan 12, 2273TZ Voorburg, The Netherlands Caroline Simmrita Chaboo Systematics Research Collections, University of Nebraska State Museum, W436 Nebraska Hall, Lincoln, NE 68588-0514, U.S.A. Date of issue: July 29, 2022 Center for Systematic Entomology, Inc., Gainesville, FL Adam S, Campos M, Heron HDC, Staines CL, Westerduijn R, Chaboo CS. 2022. Natural history of Cassida sphaerula Boheman, 1854 (Coleoptera: Chrysomelidae: Cassidinae: Cassidini) on Arctotheca prostrata (Salisb.) Britten (Asteraceae: Arctotidinae) in South Africa, with a checklist of South African Cassidinae (leaf-mining and tortoise beetles). Insecta Mundi 0945: 1–23. Published on July 29, 2022 by Center for Systematic Entomology, Inc. P.O. Box 141874 Gainesville, FL 32614-1874 USA http://centerforsystematicentomology.org/ Insecta Mundi is a journal primarily devoted to insect systematics, but articles can be published on any non- marine arthropod. Topics considered for publication include systematics, taxonomy, nomenclature, checklists, faunal works, and natural history. Insecta Mundi will not consider works in the applied sciences (i.e. medi- cal entomology, pest control research, etc.), and no longer publishes book reviews or editorials. Insecta Mundi publishes original research or discoveries in an inexpensive and timely manner, distributing them free via open access on the internet on the date of publication. Insecta Mundi is referenced or abstracted by several sources, including the Zoological Record and CAB Abstracts. Insecta Mundi is published irregularly throughout the year, with completed manuscripts assigned an individual number. Manuscripts must be peer reviewed prior to submission, after which they are reviewed by the editorial board to ensure quality. One author of each submitted manuscript must be a current member of the Center for Systematic Entomology. Guidelines and requirements for the preparation of manuscripts are available on the Insecta Mundi website at http://centerforsystematicentomology.org/insectamundi/ Chief Editor: David Plotkin, insectamundi@gmail.com Assistant Editor: Paul E. Skelley, insectamundi@gmail.com Layout Editor: Robert G. Forsyth Editorial Board: Davide Dal Pos, Oliver Keller, M. J. Paulsen Founding Editors: Ross H. Arnett, Jr., J. H. Frank, Virendra Gupta, John B. Heppner, Lionel A. Stange, Michael C. Thomas, Robert E. Woodruff Review Editors: Listed on the Insecta Mundi webpage Printed copies (ISSN 0749-6737) annually deposited in libraries Florida Department of Agriculture and Consumer Services, Gainesville, FL, USA The Natural History Museum, London, UK National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Zoological Institute of Russian Academy of Sciences, Saint-Petersburg, Russia Electronic copies (Online ISSN 1942-1354) in PDF format Archived digitally by Portico Florida Virtual Campus: http://purl.fcla.edu/fcla/insectamundi University of Nebraska-Lincoln, Digital Commons: http://digitalcommons.unl.edu/insectamundi/ Goethe-Universität, Frankfurt am Main: http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:hebis:30:3-135240 Copyright held by the author(s). This is an open access article distributed under the terms of the Creative Commons, Attribution Non-Commer- cial License, which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited. http://creativecommons.org/licenses/by-nc/3.0/ 0945: 1–23 Insecta MundI 2022 Natural history of Cassida sphaerula Boheman, 1854 (Coleoptera: Chrysomelidae: Cassidinae: Cassidini) on Arctotheca prostrata (Salisb.) Britten (Asteraceae: Arctotidinae) in South Africa, with a checklist of South African Cassidinae (leaf-mining and tortoise beetles) Sally Adam Laaiplaats 59, Mossel Bay, South Africa sallyslak@gmail.com Mariana Campos CSIRO Health & Biosecurity 147 Underwood Avenue, Floreat 6014 Western Australia, Australia Mariana.Campos@csiro.au https://orcid.org/0000-0003-3685-473X Hugh D. C. Heron P.O. Box 39042, Escombe, Queensburgh, Natal, 4070, South Africa hughheronescombe@gmail.com Charles L. Staines Smithsonian Environmental Research Center, 647 Contees Wharf Road, Edgewater, MD 21037, U.S.A. stainesc@si.edu https://orcid.org/0000-002-7411-1024 Rob Westerduijn Willem Klooslaan 12, 2273TZ Voorburg, The Netherlands rob_westerduijn@yahoo.com Caroline Simmrita Chaboo Systematics Research Collections, University of Nebraska State Museum, W436 Nebraska Hall, Lincoln, NE 68588-0514, U.S.A. cchaboo2@unl.edu https://orcid.org/0000-0002-6983-8042 Abstract. The tortoise beetle, Cassida sphaerula Boheman, 1854 (Coleoptera: Chrysomelidae: Cassidinae: Cassidini) is endemic to South Africa. Its endemic host, Arctotheca prostrata (Salisb.) Britten (Asteraceae) has been introduced in other countries where it is becoming invasive. Cassida sphaerula could provide a potential biocontrol of Arctotheca weeds as it spends the entire life cycle on this host. An intensive field study, with rearing, photography, and short films of C. sphaerula was conducted in its native habitat to document the life cycle. A checklist of Cassidinae genera in South Africa, along with 19 new host records for Cassidini species in South Africa are presented. Oothecae are simple, with few laminate membranes enclosing fewer than five eggs. There are five larval instars. Larvae and adults feed by making a series of cuts in the ventral cuticle, forming an arc, and they consume the mesophyll as the cuticle is rolled to one side. This creates many ventral craters, thickened on one margin with the rolled cuticle; these ventral craters correspond to ‘windows’ in the dorsal leaf surface where the dorsal cuticle is left intact. This unusual feeding pattern is known in three Cas- sida species, all in South Africa. Like many tortoise beetles, instar I initiates a feces-only shield on its paired caudal processes (= urogomophi); this construction is retained, along with exuviae, by subsequent instars. The shield construction was studied by film and dissections. This revealed that the columnar or pyramidal shield in this species has an exterior of dry or moist feces that obscures the central nested stack of exuviae, 2 · July 29, 2022 Adam et al. each exuviae compressed onto the caudal processes. Pupae may retain the entire larval shield of exuviae and feces or only the 5th instar exuviae; this behavioral flexibility in pupal shield retention is novel for tortoise beetles. Behaviors of C. sphaerula are discussed in the context of phylogenetic characters that can give evolu- tionary insights into the genus, tribe, and subfamily. Key words. Invasive species, pest, weed, larva, herbivore. ZooBank registration. urn:lsid:zoobank.org:pub:4AC56F98-6474-4AAD-A2A9-51AE2F39A1E1 Introduction Cassida Linnaeus, 1758 is the type of the genus of the chrysomelid subfamily Cassidinae (Gyllenhal 1813) and of the tribe Cassidini Chapuis, 1875. As currently defined, Cassidinae comprises 37 tribes, the former ‘Hispinae’ and the tortoise beetles (Chen 1940; Staines 2002; Chaboo 2007). The tortoise beetle tribes of Cassidinae are dis- tinguished from plesiomorphic “hispines” and other cryptic-feeding tribes by the exophagous larvae possessing paired caudal processes (= urogomphi) on the 9th abdominal tergite. Furthermore, these tortoise beetle larvae exhibit a unique behavior—they build and carry a shield, held on the caudal processes, and composed of materi- als of their cast exuviae only or cast exuviae + feces (Olmstead 1994; Müller and Hilker 2003). The shields have different architectures depending on the texture, quantity, and form of the feces that are stacked unto the exuviae; the feces on these shields may be semi-liquid or dry, and loose, lumpy, or filamentous (see Świętojańska 2009 and citations therein). In many tribes, the pupae can also have paired caudal processes and retain the shield from the 5th instar (Steinhausen 1950). These complex ecological, morphological, and behavioral features subtend the crown clade of Cassidinae comprising eight tribes (Chaboo 2007): Basiprionotini, Cassidini, Eugenysini, Hemi- sphaerotini, Ischyrosonychini, Mesomphaliini, Omocerini, and Spilophorini. The online catalog of Borowiec and Świętojańska (2002–2022) treat Aspidimorphini as a tribe, but it has been synonymized in Cassidini (López-Pérez et al. 2018). Their catalog includes the tribes Delocraniini, Imati- diini, and Notosacanthini; larvae in these tribes do not carry shields (Bondar 1940; García-Robledo et al. 2010; Sekerka et al. 2013; Monteith et al. 2021) and are plesiomorphic to the crown-clade Cassidinae (Chaboo 2007). The tribe Cassidini has both Old World and New World species but all other cassidine tribes have either Old World or New World distributions (Weise 1911; Spaeth 1914). Cassidini contains 87 genera; Cassida is the most speciose with 459 recognized species distributed in both Old World and New World (Borowiec and Świętojańska 2002–2022). López-Pérez et al.’s (2018) morphology-based phylogenetic analyses of Cassidini did not recover a monophyletic Cassidini (unless it includes Ischyrosonychini) nor a monophyletic Cassida. There are many synonyms in the long history of Cassida reflecting the poor definition of the genus; still, more refining of species concepts is needed to achieve a rigorous monophyletic genus concept. Documented host plants of Cassida are from about 16 plant families (Jolivet and Hawkeswood 1995; Borowiec 1995; Borowiec and Świętojańska 2002–2022); Cassida species can be mono- or oligophagous (Bordy and Doguet 1987). Goedært (1662) is the earliest report on juvenile stages in Cassidinae, for Cassida vibex Lin- naeus, 1767 and Cassida viridis Linnaeus, 1758. Today, data on juveniles are available for 136 species of Cassidini (Świętojańska 2009) and for about 61 species of Cassida, however, most morphological descriptions are not detailed. Cassida larval shields have diverse architectures; the pupae retain the shield from the 5th instar (Heron 2008; Świętojańska 2009). The fundamental problem in the systematics of Cassida, indeed in all Cassidinae, is the gap in knowledge about life history and ecology of species and limited specimen collections of all life stages for detailed comparisons. These would generate novel characters and states to better resolve more natural monophyletic groups and relationships. This paper addresses that gap by reporting the first natural history account of Cassida sphaerula Boheman, 1854, a South African endemic. The Cassidinae of South Africa comprises 10 tribes, 32 genera, and ~205 species (Table 1; Staines 2015; Borowiec and Świętojańska 2002–2022). Cassida has 36 species documented in South Africa; six new species are being described (Borowiec and Świętojańska 2022). For those South African Cassidi- nae species using Asteraceae, host plants are listed in Table 2; we provide 20 new host records. Cassida sphaerula ranges from the Cape region to the Transvaal and KwaZulu-Natal (Borowiec and Świętojańska 2002–2022). Based on label data of specimens reared in Australia, this species was reported to use the host plant, Arctotheca Natural history of Cassida sphaerula Insecta Mundi 0945 · 3 Table 1. Checklist of tribes and genera of Cassidinae in South Africa (Chrysomelidae). ‘Hispines’ = 6 tribes; 16 genera; 103 species (see Staines 2015 for species). Tortoise beetles = 4 tribes, 16 genera; ~101 species (see Borowiec and Świętojańska 2002–2022 for species).* = 6 species newly described by Borowiec and Świętojańska (2022). Tribe Basiprionotini Hincks, 1952 Tribe Gonophorini Chapuis, 1875 Metriopepla Fairmaire, 1882: 1 sp. Agonita Strand, 1942: 4 spp. Tribe Callispini Chapuis, 1875 Tribe Hispini Gyllenhal, 1813 Callispa Baly, 1858: 8 spp. Callanispa Uhmann, 1959: 1 spp. Tribe Cassidini Gyllenhal, 1813 Chrysispa Weise, 1897: 1 spp. Aspidimorpha Hope, 1840: 19 spp. Dactylispa Weise, 1897: 30 spp. Cassida Linnaeus, 1758: 41 spp.* Dicladispa Gestro, 1897: 36 spp. Chiridopsis Spaeth, 1922: 9 spp. Dorcathispa Weise, 1900: 3 spp. Conchyloctenia Spaeth, 1902: 8 spp. Hispellinus Weise, 1897: 1 spp. Fornicocassis Spaeth, 1917: 1 sp. Platypria Guérin-Méneville, 1840: 3 spp. Hybosinota Spaeth, 1909: 1 sp. Polyconia Weise, 1905: 1 spp. Ischiocassis Spaeth, 1917: 4 spp. Pseudispella Kraatz, 1895: 1 spp. Laccoptera Boheman, 1855: 11 spp. Thoracispa Chapuis, 1875: 3 spp. Limnocassis Spaeth, 1952: 3 spp. Trichispa Chapuis, 1875: 1 spp. Orobiocassis Spaeth, 1934: 3 spp. Tribe Leptispini Fairmaire, 1868 Oxylepus Desbrochers, 1884: 6 spp. Leptispa Baly, 1858: 6 spp. Psalidoma Spaeth, 1899: 1 sp.: questionable record) Rhytidocassis Spaeth, 1941: 3 spp. Tribe Notosacanthini Hincks, 1952 Smeringaspis Spaeth, 1934: 1 sp. Herminella Spaeth, 1913: 4 spp. Trichaspis Spaeth, 1911: 3 spp. Notosacantha Chevrolat, 1837: 11 spp. Tribe Cryptonychini Chapuis, 1875 Tribe Oncocephalini Chapuis, 1875 Cryptonychus Gyllenhal, 1817: 2 spp. Oncocephala Agassiz, 1846: 2 spp. calendula (L.) Levyns (Asteraceae: Arctotidinae; Heron and Borowiec 1997) which is native to South Africa but has become an introduced invasive in Australia (Groves et al. 2003) and California (Brossard et al. 2000). Taylor (1965) reported Combretum Loefl. (Combretaceae) as a host plant, but this is possibly erroneous as Combreta- ceae is not considered a host of tortoise beetles (Borowiec and Świętojańska 2002–2022) and it is a host for a few other chrysomelids (Jolivet and Hawkeswood 1995). Little else is known about C. sphaerula in its native habitat in South Africa. Here we document the second native host, Arctotheca prostrata (Salisb.) Britten. Access to a good-sized population of C. sphaerula is allowing ongoing study and specimen collection of C. sphaerula. We note its poten- tial as a biocontrol agent for A. prostrata outside of the native range, where the plant poses a weed risk. Arctotheca prostrata (Fig. 1–5) is endemic to South Africa where it has several common names, including prostrate Cape weed and Cape dandelion. It can be regarded as a weed in South African gardens—hundreds of plants are dug and removed every year from author SA’s garden. However, seed packets and garden plants are sold in local nurseries. The genus Arctotheca Vaill. comprises five species (McKenzie et al. 2005): Arctotheca calendula (Fig. 6), Arctotheca forbesiana K. Lewin, Arctotheca populifolia (P.K. Bergius) Norl., Arctotheca prostrata (Fig. 1–5), and Arctotheca marginata Beyers (Beyers 2000). These are perennial or, occasionally, annual, trailing, or erect herbs (Ghafoor and Bean 2015). Arctotheca calendula and A. prostrata have been recognized as weeds in multiple countries (Hinojosa-Espinosa and Villaseñor 2015; Atlas of Living Australia 2021; California Invasive Plant Council 2021; GBIF 2021a, b, c; Jepson eFlora 2021) and A. populifolia is a weed in Australia. These three species became weeds after escaping from cultivation (Mahoney and McKenzie 2008; Hinojosa-Espinosa and Vil- laseñor 2015; Lucid Central Weeds of Australia 2021) or due to being contaminants of stock fodder and packing straw (Wood 1994). One UK-based gardening website (Candide 2021) lists all three weedy Arctotheca species with photos and promotes two of them: A. populifolia as a garden species that attracts bees and acts as weed suppressors and dune stabilizer; and A. calendula as a bee and butterfly attractant that can be planted as a showy ground cover or bed- ding plant for spring and early summer display (Candide 2021). These plants can cover large landscapes and have 4 · July 29, 2022 Adam et al. Table 2. Asteraceae host plants used by Cassidinae in South Africa. Host plant Cassida species Comments and references Arctotheca calendula L. Cassida sphaerula Boheman, 1854 Heron and Borowiec 1997 Arctotheca prostrata (Salisbury) Britten Cassida sphaerula Boheman, 1854 new record of S. Adam www.inaturalist.org; confirmed breeding Berkheya bipinnatifida (Harvey) Roessl Cassida guttipennis Boheman, 1862 Heron and Borowiec 1997; Heron 2003, 2011; confirmed breeding Berkheya heterophylla (Thunberg) Cassida guttipennis Boheman, 1862 Label inf., SANC, PPRI O.Hoffmann Berkheya maritima J.M. Wood & M.S. Cassida quatuordecimsignata Spaeth, new record; confirmed feeding Evans 1899 Berkheya onopordifolia (DC) O. Cassida quatuordecimsignata Spaeth, new record; nomad; confirmed feeding Hoffmann ex Burtt Davy. 1899 Berkheya pinnatifida (Thunberg) Cassida quatuordecimsignata Spaeth, new record; confirmed breeding Thellung 1899 Berkheya rhapontica (DC) Hutchinson & Cassida guttipennis Boheman, 1862 new record; confirmed breeding Burtt Davy Berkheya seminivea Harvey Cassida guttipennis Boheman, 1862 new record; confirmed breeding; Label inf., SANC, PPRI Berkheya speciosa (De Candolle) O. Cassida guttipennis Boheman, 1862 Heron and Borowiec 1997; Heron 2003; Hoffmann confirmed breeding Cassida vespertilio Boheman, 1862 at Howick, Natal; confirmed breeding Cassida sp. nov. 2 (Borowiec and at Umzinto; confirmed breeding Świętojańska 2022) Berkheya sp. #1 Cassida vespertilio Boheman, 1862 new record; observed at Ingeli; confirmed breeding Berkheya sp. #2 Cassida quatuordecimsignata Spaeth, new record; Howick, Natal; confirmed 1899 breeding Berkheya sp. #3 Cassida quatuordecimsignata Spaeth, new record; Voorkeur Siding, Natal; 1899 confirmed breeding Berkheya sp. #4 Cassida guttipennis Boheman, 1862 new record; confirmed breeding. Brachylaena discolor De Candolle Basipta stolida Boheman, 1854 Borowiec 2002; Heron and Borowiec 1997; Heron 2003; Muir and Sharp 1904; species erroneously given as Basipta glauca, Identification corrected in Borowiec 1999; confirmed breeding Cassida granulicollis Spaeth, 1905 Heron and Borowiec 1997; confirmed breeding Cassida unimaculata Boheman, 1854 Heron and Borowiec 1997; Heron 2003; Muir and Sharp 1904; confirmed breeding Brachylaena elliptica (Thunberg) De Basipta stolida Boheman, 1854 new record; confirmed breeding Candolle Cassida unimaculata Boheman, 1854 new record; confirmed breeding Brachylaena huillensis O.Hoffmann Cassida granulicollis Spaeth, 1905 new record; Label inf. SANC, PPRI (shaken from tree)   Brachylaena rotundata S. Moore Cassida granulicollis Spaeth, 1905 new record; Label inf. SANC, PPRI Natural history of Cassida sphaerula Insecta Mundi 0945 · 5 Host plant Cassida species Comments and references Brachylaena uniflora Harvey Basipta stolida Boheman, 1854 new record; Label inf. SANC, PPRI (identified as “Brachylaena sp. cf. uniflora); Heron 2018; confirmed breeding Cassida unimaculata Boheman, 1854 Heron 2018; confirmed breeding Chrysanthemoides incana (Burman f.) Cassida foveolatipennis Borowiec and Borowiec and Świętojańska 2001; Norlindh Świętojańska, 2001 confirmed breeding Chrysanthemoides monilifera (L.) Cassida chrysanthemoides Borowiec and Świętojańska 2001; Norlindh subsp. monilifera (L.) Borowiec and Świętojańska, 2001 confirmed breeding Norlindh. (= Chrysanthemoides monilifera (L.) Norlindh) Chrysanthemoides monilifera (L.) Cassida foveolatipennis Borowiec and Borowiec and Świętojańska 2001; Norlindh subsp. pisifera (L.). Norlindh Świętojańska, 2001 confirmed breeding Cassida chrysanthemoides Borowiec Borowiec and Świętojańska 2001; and Świętojańska, 2001 confirmed breeding Chrysanthemoides monilifera (L.) Cassida diversipunctata Borowiec and Borowiec and Świętojańska 2001; Norlindh. subsp. rotundata (De Candolle) Świętojańska, 2001 confirmed breeding Norlindh Chrysanthemoides monilifera (L.) Cassida spatiosiformis Borowiec and Borowiec and Świętojańska 2001; beetle Norlindh. subsp. subcanescens (De Świętojańska, 2001 erroneously given as Cassida spatiosa Candolle) Norlindh in Heron and Borowiec 1997; confirmed breeding Othonna quinquedentata Thunberg Cassida pudens Boheman, 1854 Borowiec and Świętojańska 2001; Heron and Borowiec 1997 (under Cassida   subplana Spaeth, 1928) Sonchus asper (L.) Hill var. asper Cassida sulphurago Boheman, 1854 new record; label inf., SANC, PPRI (Kuntze) O. Hoffmann Tarchonanthus littoralis P. Herman Basipta luteocincta Boheman, 1854 new record; Label inf., SANC, PPRI, under T. camphoratus L.; confirmed breeding; T. camphoratus group revised by Herman 2002 Tarchonanthus obovatus De Candolle Cassida sulphurago Boheman, 1854 new record; H. de Klerk photo inf. under T. camphoratus L.; T. camphoratus group revised by Herman 2002; confirmed feeding Tarchonanthus trilobus De Candolle Basipta stolida Boheman, 1854 new record; confirmed breeding Cassida unimaculata Boheman, 1854 new record; suspected feeding attractive yellow flowers (Fig. 6). Arctotheca calendula is promoted on a USA-based website (Gardenia.net 2021) as a tough and easy-to-grow perennial that blooms most of the year. Such information is likely to appeal to some gardeners and encourage its propagation. Arctotheca prostrata has recently been recorded in new countries. The first herbarium record of this species in Australia dates to 1946 (Australasian Virtual Herbarium 2021), whereas the first records in Mexico are from 2015 (Hinojosa-Espinosa and Villaseñor 2015) and in Italy from 2018 (Galasso et al. 2019). In Italy it is hypoth- esized to have been introduced via stolons transported in soil (Galasso et al. 2019). In the U.S.A., the species was introduced as ground cover (Veit 2014, date unspecified) and records of its occurrence begin in 2002 (EDDMapS 2021). Records of A. prostrata have in the past been amalgamated with A. calendula, despite being two distinct species (Mahoney and McKenzie 2008). The Calflora website (Calflora 2021) lists A. prostrata as: “the infertile form. The fertile form is Arctotheca calendula”. However, Veit (2014) found that Californian A. prostrata is fertile and able to produce achenes/seeds that germinate; if the populations become more numerous and closer together, cross-pollination may allow for fertile seeds to be produced. Arctotheca prostrata is a sterile perennial that spreads aggressively by prostrate stems; invasions favor roadsides and other disturbed sites (Mahoney and McKenzie 6 · July 29, 2022 Adam et al. Figures 1–6. Arctotheca prostrata (Salisb.) Britten (Asteraceae) in its native habitat, South Africa (photos: S. Adam, September 2021). 1–4) Various sites on the farm Laaiplaats, Mossel Bay. 5) Leaves appear spotted due to beetle feeding damage, farm Laaiplaats. 6) Arctotheca calendula in Australia, showing how successfully these plants overtake bare soil (photo: Stephen D. Hopper). Natural history of Cassida sphaerula Insecta Mundi 0945 · 7 2008). In California, this plant is listed as having an overall risk score of “Moderate” (Brusati 2004). Due to the invasive and weedy nature of A. prostrata, natural history of its native herbivores is important to document as they can lead to the development of biological control agents (Harley and Forno 1992). This detailed record of feeding habits of C. sphaerula on A. prostrata raises the possibility of its development and use as a biocontrol agent. Good biocontrol agents should exhibit three fundamental qualities: (1) specificity to the host species and host genotype, (2) damage (the more damage it causes the better as a biocontrol agent); and (3) potential for establishment in the introduced range (Knutson and Coulston 1997; van Klinken and Raghu 2006; Müller-Schärer and Schaffner 2008). Cassida sphaerula has been documented as a natural enemy of A. calendula (Scott and Way 1990) and possibly without enough species-specificity. Species specificity, however, is more important in the designation of a biological control agent than in its natural range. If no other non-invasive plant species are recorded as plant hosts, the use of C. sphaerula could still be supported as a biological control agent where it is wanted. Materials and Methods Author SA studied, reared, collected, photographed, and filmed many individuals of C. sphaerula on multiple plants of A. prostrata scattered in several locations (Fig. 1–5), including her own garden: SOUTH AFRICA: Province Western Cape, Mossel Bay, Laaiplaats 59, −33.966030 22.089960, 188m elev., farm habitat, 18.VIII.2021–31.I.2022, coll. S. Adam. Another site within the same farm was at −33.967890 22.094130,  95m elev., 18.VIII.2021– 31.I.2022, coll. S. Adam; Mossel Bay, farm ‘Bosrug’, -33.961200 22.093260, 91m elev, 18.VIII.2021–3.XI.2021, coll. S. Adam; Moordkuil River, −33.967890 22.094130, 95m elev, 18.VIII.2021–3.XI.2021, coll. S. Adam; Bos- man River, −33.966030 22.089960, 188m elev, 18.VIII.2021–3.XI.2021, coll. S. Adam; Blommekloof, grassy field, −33.94271805, 22.0601465875, 280m elev., 3.VI.2022 and 10.VI.2022, coll. S. Adam At Goukamma Reserve (near Knysna), −34.06556 22.94379, 21.X.2021, on the coast about an hour from Mossel Bay, the “lawn” area around the picnic site (Fig. 8–9) and amenities was very disturbed by the activity of Cape dune mole-rat (Bathyergus suillus (Schreber, 1782): Bathyergidae, Bathyerginae)) and the Arctotheca plants were extensive and abundant. The Cape Mole-rat, Georychus capensis (Pallas, 1778) (Bathyergidae, Georychinae) also occurs in the same area. However, inspection did not turn up any C. sphaerula, only snails eating the plants. On 30.X.2021, one 50 Km reconnaissance trip was conducted by SA and a coastal trip, −34.0765, 22.1655 to −33.8732, 22.0307, was done by Wendy Wiles and Sandra Falanga, searching for additional locations of plants and beetles. These trips found plants with beetle-feeding damage and beetles in multiple areas along the routes, but not above 750 m ASL elev. Throughout spring to autumn 2021, the beetles were always present at the farm habitat (Laaiplaats 59), with tenerals (straw-colored) and mature (green) adults. Brief examinations found seven larvae (three young, four mature) on 3.XI.2021; five larvae, no pupae, two green adults, one yellow-green adult, on 4.XI.2021; and one pre-pupa and a few adult teneral adults on 14.XI.2021. The last Cassida individual was seen on 23.XII.2021; no individuals have been found on the plants in mid-summer. Beetle activity resumed in autumn (early May 2022) and was observed in several sites (maximum temperature of 82.4°F (28°C) and minimum temperature of 35.6°F (2°C)). Habitat ecology. Mossel Bay is on the south coast of South Africa. The climate is classified as semi-arid (Kop- pen climate classification, Kottek et al. 2006) and is moderated by proximity to the ocean. The area has short dry summers (late November to late March) and long, cool, windy winters. Temperatures range from 52–75°F; pre- cipitation (~420 mm/yr) is somewhat even throughout the year, as rain or snow (WeatherSpark 2021). Identifications. Photos were uploaded to the online repository, iNaturalist (2021) and the beetle species was identified initially by RW. Vouchers of juveniles and adults of the beetle were collected, deposited at South Africa National Insects Collection, South African National Biodiversity Institute (SANBI), and loaned to CSC. The beetle species was confirmed after studying collected specimens by CSC and by Elizabeth Grobbelaar, ARC Iden- tification Services, Pretoria. The type is housed in Museum für Naturkunde de Humboldt-Universität (ZMHU), Berlin, which is currently relocating their insect collection so access to types is not possible. Vouchers of the 8 · July 29, 2022 Adam et al. Figures 7–8. Coastal area with extensive growth of Arctotheca prostrata, Goukamma Reserve, South Africa. Plants intact but soil disturbed by the activity of Cape dune mole-rat (Bathyergidae: Bathyerginae: Bathyergus suillus (Schreber, 1782)). host plant are deposited with and identified by Robert McKenzie, Dept. of Botany, Rhodes University and Nicola Bergh, Compton Herbarium, SANBI. Imaging. Photos were taken with a Panasonic® DMC-FZ200 camera with a Raynox® macroscopic lens M-150. Specimens were studied with a Zeiss® stereoscopic microscope with a Dino-Lite® eyepiece digital microscope/ camera. Photo editing was done in Paint.net or Photoshop®. Specimen study. Only specimens from field collections and rearing are studied here. These are deposited at the South Africa National Insect Collection, Pretoria, and the University of Nebraska-Lincoln. Specimens of C. sphaerula are in South African institutions (mainly, South Africa National Insect Collection, Iziko Museums, Dipsong Museum, and National Museum of Bloemfontein) but we are unable to travel at present, to study and confirm identifications and add more locality information in the present study. Observations. Author SA checked the C. sphaerula population on their host daily from August 2021 to June 2022. She brought eggs, larvae of different ages, pupae, and adults indoors for observation and rearing. She suc- cessfully followed eggs to hatching, and three of these hatchlings were followed through all instars, pre-pupation, pupation, and then the emergence of teneral adults. Feeding behavior. Multiple adults and larvae were maintained in small plastic containers (at ambient tem- perature, humidity, and light) to determine the feeding pattern. Intact leaves were presented for feeding and the larva (different ages) or adult was removed at different times to view their cuts into the leaf (the expanded prono- tum obscures feeding in both larvae and adults of this species). Both larval and adult feeding were observed and filmed (Video 1–4); larvae were discovered cutting and eating trichomes (Video 2). Exuvio-fecal shield architecture and construction. In several subfamilies of Chrysomelidae, larvae retain their feces directly on the body (Criocerinae, certain Galerucinae) or use it as a construction material in cases they live in (Camptosomata) or carry as an umbrella over the body (Cassidinae: tortoise beetles) (Chaboo 2007, 2011; Chaboo et al. 2007, 2008). In tortoise beetles, the larvae retain the shield on the caudal processes, hold- ing it over the body like an umbrella, or moving it in different angles, even bringing it flat against the dorsum. The caudal processes in Cassidinae have been called “apical furci”, “supra-anal processes”, and “spines” in the cassidine literature (see Chaboo 2007: 68–74); “urogomphi” is used in the insect literature but these structures are not all homologous. Other materials may be added or become established in the fecal medium of the shield, e.g., exuviae, gut microbes, trichomes, fungi, other chemicals. Author SA followed multiple larvae of various instars in plastic dishes at ambient conditions and took photos at 2-hour time intervals to capture the initiation, Natural history of Cassida sphaerula Insecta Mundi 0945 · 9 Figures 9–15. Arctotheca prostrata with feeding damage by beetle, Cassida sphaerula Boheman, 1853 (photos: S. Adam, September 2021). 9) Intact leaf, dorsal view. 10) Intact leaf, ventral view. 11) Leaf, dorsal view, with window-pane pattern where beetles leave dorsal cuticle intact. 12) Leaf, ventral view, with craters left by beetle feeding damage. 13) Leaf with paired green adults (dorsal) and cream-colored larva showing blackish exuvio- fecal shield (held on caudal processes) and wet anal droplet to apply to shield. Note hirsute dorsal and ventral surfaces of host leaf. 14) Leaf with many feeding craters and single larva with exuvio-fecal shield; note feeding is only between veins. 15) Feeding craters, each with marginal cuticle roll. 10 · July 29, 2022 Adam et al. expansion, and maintenance of the exuvio-fecal shield. Shields were removed, by gently prying them off the liv- ing caudal processes, and abraded with a forceps to remove small fecal pieces. This allowed determination of the shield architecture. We describe the construction process based on many days of observations, many dissections of exuvial-fecal shields, and a large archive of imagery (stills and films). Results Beetles and plants are found in several areas within our main study site, the 97-ha farm, and in many populations along the Moordkuil and Bosman rivers in the area. Some A. prostrata plant patches can be heavily infested and show considerable damage with leaves appearing spotted (Fig. 5). No nearby plants of other species were found to host C. sphaerula. During winter, A. prostrata plants show some die-back/browning of the leaves, especially in areas subject to frost; in more temperate areas, they continue to grow. From early summer (27.XII.2021) until autumn (mid-May 2022), the Arctotheca plants are quite patchy, and get burned to a crisp in sunny spots but do better in shaded areas. Other animals on the plants are rare during this hot period, with only the occasional slug found on Arctotheca. At the Goukamma Reserve on the coast, we observed the “lawn” around the picnic site and amenities (Fig. 7–8) to be very disturbed by Cape dune mole-rat activity. The A. prostrata growth is excellent, but no C. sphaerula were found eating these plants, only snails. Natural history of Cassida sphaerula Boheman, 1854 Field observations over almost one year revealed the beetle’s cycle of activity. Our observations began at the end of one breeding season. From early summer (27.XII.2021) till early autumn (mid-May 2022) no adults or larvae were seen. Then the new breeding season began in late autumn. The first sign of beetle activity is the small ‘win- dows’ chewed on the plant leaves in autumn (late May); then the larvae can be seen on the underside. Egg-laying begins in May, with much larval activity by mid-June, when minimum temperatures are around 35.6°F (2°C). Up to 8 egg cases have been found on a single leaf. Thus, we believe this species is an autumn/winter breeder. Egg cases (n=4) (Fig. 16–18). Oothecae are deposited on the venter of the leaf (Fig. 16) in apparently random areas between veins. We observed a maximum of eight oothecae per leaf. The ootheca lies flattened on the long axis, shallowly tucked into the leaf surface as there is a slight depression under each one; it is not stalked, sus- pended or protuberant from the leaf surface, as in some other Cassidinae. Oviposition was not observed so it is unclear how the female may prepare a site before depositing her eggs (see Müller and Rosenberger (2006) for possible oviposition sequences in Chrysomelidae). The ootheca, secreted by colleterial glands (Hinton 1981; Gil- lot 2002), comprises a thin opaque outer laminate membrane that appears shiny and dark brown (Fig. 17–18). The few enclosed eggs (less than 5) are cream-colored (Fig. 17–18), dorsoventrally compressed (lying flattened on leaf), and elongate-oval shaped. The ootheca lacks any additional coverings, no fecal or chewed plant mate- rial. Egg hatch. One ootheca was collected on 11.X.2021 and three larvae hatched on 18.X.2021, confirming that more than one egg is oviposited at a time. We did not observe how the larvae exited the egg case, but we found the ootheca roughly torn at one end and left behind, therefore not eaten by the neonate. Larva (n=20; Fig. 13–14, 16, 19–25). The larvae are solitary, not apparently gregarious, but may be found mixed with others of different stages in a dense situation, even feeding side by side with scoli (lateral projections) in con- tact. They do not respond to disturbance by moving into groups or with coordinated cycloalexic (ring) defense where larvae move into a tight, somewhat circular, group and all flex the shield in unison (see Jolivet et al. 1990). Larvae are found mostly on the venter of the leaves. Instar I (n=3; Fig. 19–20) are tear-dropped shaped, about 2 mm long X 1 mm at maximum width (across pronotum). The body is tan-colored. The paired caudal processes (Fig. 21, 24–25; = supra-anal processes, urogomphi) are almost half as long as the body. Older larvae (Fig. 23) are creamy yellow and with a dark brown central area; their cuticle is almost transparent, and the inter- nal organs are somewhat visible (internal movements are easily seen). The scoli pattern (Fig. 21–22) and caudal processes processes are similar between instars and fit with Świętojańska’s (2009: 74) generalized Cassida larvae having an ovoid dorso-ventrally flattened body with 16 pairs of lateral scoli. Natural history of Cassida sphaerula Insecta Mundi 0945 · 11 Figures 16–18. Ootheca and young larvae of Cassida sphaerula (photos: S. Adam, September 2021). 16) Venter of host leaf with two oothecae (arrows) and two instar III with their black exuvio-fecal shields. 17) Ootheca (~2 mm long). 18) Ootheca with oval-shaped egg. 19) Two young instar 1 (~2 mm long) with tiny black shield composed entirely of its own feces. 20) Mature instar 1 with larger shield (reared from Fig. 19). 12 · July 29, 2022 Adam et al. Figures 21–25. Larva of Cassida sphaerula (photos: S. Adam, September 2021). 21) Young instar with lateral projections called scoli; shield removed to expose paired caudal processes. 22) Older instar (frontal view) with exuvio-fecal shield attached to caudal processes; feces appear dry. 23) Older instar with moist exuvio-fecal shield. 24) Older instar, dorsal view, with feces removed; legs and caudal processes of exuviae of previous instar apparent. 25) Hind end of older larva with dry exuvio-fecal shield. Paired caudal processes of previous instar are exposed, projecting dorsad. The caudal processes of this larva is hidden, stacked within the observable caudal processes. Natural history of Cassida sphaerula Insecta Mundi 0945 · 13 Exuvio-fecal shield. The larval shield is initiated in Instar 1 (Fig. 19) shortly after it initiates feeding. This shield is comprised only of larval feces that is applied to the caudal processes by the muscular telescoped anus. The shield grows into an elongate mass on the larva’s paired caudal processes (Fig. 20). The shield can appear dry (Fig. 22) or wet (Fig. 23) and the telescoped anus periodically applies a dark wet droplet (see Fig. 13) to the shield. Dissections of shields reveal a fan (Fig. 22) or pyramidal shape (Fig. 23) with a central scaffold of stacked, nested exuviae and all entirely covered in dry or moist feces. The exuviae are not easily discerned in intact shields (e.g., Fig. 23). In dissected shields (Fig. 24–25), the feces are abraded to reveal the stack of exuviae; each exuviae can be individually teased off to show the caudal processes of older instars. The larvae continue to build, applying fresh feces and wet droplets (note wet appearance in Fig. 24). Pupa (n=10; Fig. 26–27). The pre-pupal stage is typically when the mature larvae ceases feeding, become seden- tary and fixes its abdomen to the substrate. Five young larvae were followed (three from egg hatch) to adulthood; three pupated; pupation lasted nine days, 15 days, and 20 days. Six mixed-age pupae/pre-pupae were placed in a container on 28.VIII.2021 and the first adult appeared on 7.X.2021 (9 days); two of these pupae failed and four adults were reared. Thus, pupation (n=7) ranges from 9–20 days. No parasitoids emerged from these laboratory pupae. Pupae (Fig. 26–27) are ~9 mm long, solitary, affixed by their abdomen to the leaf venter, never on the upper part. There is seldom more than one pupa per leaf. The pupa is tan-colored, and the body is ovoid and dorso- ventrally flattened. Only the abdominal segments have lateral scoli. The pupa of C. sphaerula shows two types of shields. It may retain the final exuviae (Fig. 26) and the for- mer larval shield may be found discarded nearby or the pupa may retain the entire shield structure of the 5th instar larva (exuviae I–IV and their fecal matter) (Fig. 27). As far as we know, this is the first observation of such flexibility in shield retention in cassidine pupal shields. After the adults have emerged, the pupal exuviae remains attached to the plant for a long time, with or without the fecal shield. A few adults seem to have some difficulty eclosing, taking longer and struggling to exit the exuviae, but these adults eventually became hardened and moved away. Adult (n=30; Fig. 28–30). These are ~9 mm long (along midline, head to posterior margin) by 4–5mm at their maximum width (across pronotum). The dorsum is generally pale green in color but can vary from translu- cent straw to a deep green. They were observed as early as 30 August (reared) and 23 September (wild) and are generally solitary. During the observation period, the habitat experienced a frost (late August) and the beetles remained sluggish but resumed activity as temperatures rose. Color/pattern variations. Newly eclosed or teneral adults are straw (pale-brown) colored and the mature hardened adults are green. We only observed mating pairs of green individuals. As adults age, some acquire per- manent circular blackish marks in different locations of the elytra (Fig. 30), but we did not detect marked color polymorphism as in some Cassidini (Simon Thomas 1964; Verma and Kalaichelvan 2004). Courtship and mating (Fig. 13, 29). Mating pairs were first observed on 24.VIII.2021, as the frost season ended, and the region transitioned to spring. Courtship was not observed by the many mating pairs found, but pairs in copula were noted. Mature green adults exhibited no rapid (a few seconds) color changes (with temporary black spots or to golden or straw colors) as documented for some Cassidinae during mating or when disturbed (Bar- rows 1979). Dormancy. Beetle activity ceased as the summer peaked and it is unclear where they hide. The host plants do not lose leaves in winter, suggesting that the beetles can have a steady food supply, further supporting them as a good biocontrol agent. We continue observations in 2022 but have not determined if beetles pass the winter hidden under stones or in dead vegetable matter, as they tend to do in the Natal area (Heron, pers. obs.). Feeding patterns (Fig. 11–12, 14–16, 28) of C. sphaerula. Larvae and adults feed in similar ways, which creates a distinct pattern of craters on the venter of the leaf, each crater with the cuticle rolled to one side (Fig. 15). The craters of instar 1 are small (Fig. 19–20); older larvae and adults make craters up to 4 mm long. The craters are hollowed out by feeding and are irregularly shaped (hemispherical, ovoid, rounded). They have a deep basin, with the rolled ventral cuticle forming a thickened margin on one side. The dorsal cuticle of the leaf remains intact (Fig. 11), with a window-pane pattern. The mid-rib and secondary veins are not eaten but the leaves are intact dorsally and do not have a skeletonized appearance. 14 · July 29, 2022 Adam et al. Figures 26–31. Pupa and adult of Cassida sphaerula (photos: S. Adam, September 2021). 26) Pupa, attached by venter of leaf, with shield comprising only exuviae of 5th instar. 27) Pupa with shield of exuviae I–V and feces. 28) Teneral adult is straw colored. 29–30) Mature adults are green, in copula. 31) Older adult with black spots on elytra. Natural history of Cassida sphaerula Insecta Mundi 0945 · 15 Larvae start feeding shortly after hatching. Their feeding exhibits a stereotyped repertoire. The site is prepared by eating most of the trichomes (Video 1). The first cut of the ventral cuticle (including the leaf ’s epicu- ticular wax layer) is made by a series of bites that create an arc-shaped cut, about the same size of the pronotum. The larva starts feeding on spongy mesophyll, and its head action pushes the cuticle layer, rolling it over and ventrad. As the larvae feed on the exposed mesophyll, a crater forms, deepens, and enlarges ventrad underneath the larvae. The rolled cuticle is pushed further ventrad, underneath the larva. When the larva finishes feeding in that crater, it moves to a different spot on the same leaf. A single larva can spend many days feeding on the same leaf. We observed and filmed the larvae of C. sphaerula cutting and eating trichomes (Video 1). Plants in the tribe Arctotidinae have mostly non-glandular tri- chomes, although some glandular hairs can be present in certain organs (Karis et al. 2009). Glandular trichomes would be more deterrent to herbivory. Trichome-eating has not been observed for any Cassidinae. No trichome fragments appear in the shields we dissected (n = 4) so we assume trichomes are digested. In C. sphaerula, adult feeding resembles larval eating. The adult also makes multiple cuts in the cuticle, in an arc-shape; as it feeds deeper into the trough, the head movements push the cuticle ventrad, under the beetle, towards the posterior margin of that feeding depression (Video 4). The depression deepens and widens, and the cuticle becomes a ridge at the margin of this feeding crater. We did not observe the adults consuming trichomes. The pattern resulting from adult feeding resembles the larval pattern, but the craters are larger. Both stages leave the dorsal cuticle intact, forming windows. Natural enemies of C. sphaerula. SA observed other animals on the host plant: snails, slugs, spiders, velvet mites, springtails, insects (wasps, aphids, stink bugs, lace bugs, other beetles including one chrysomelid (to be deter- mined), Lepidoptera caterpillars), but noted few interactions that might clarify which are competitors, predators, and parasites of C. sphaerula. We observed and filmed one C. sphaerula larva walking over a leaf and a smaller- sized aphid moved out of its way (Video 3). In another instance, a smaller mite moved out of the way of an approaching C. sphaerula larva. Observations of interactions in the field were almost impossible as the host leaves lie flat, pressed against one another and it is necessary to grasp each leaf and gently pull it up to see the underside. This tends to dislodge or scare off many of the other individuals on the plant. The C. sphaerula larvae raise their shields whenever they are disturbed, including by others of the same species. They seem to spend a great deal of time sitting still, but adults are alert—they freeze when there is any movement of the leaf. Then they scuttle to the underside of the leaf, out of the light and view. Like many cassidines, adults show a definite tendency to tumble off the leaf to the ground and then scuttle to the plant stems where they are better protected. Discussion The behaviors and life cycle of C. sphaerula was studied in detail and over many months (early spring-late autumn). We confirmed the choice of host plant, A. prostrata, in the indigenous habitat in South Africa; C. sphae- rula is now known on two Arcotheca species (Heron and Borowiec 1997). In South Africa, hosts documented for Cassidini are in the Amaranthaceae, Asteraceae and more infrequent hosts are in Aizoaceae, Fabaceae, Polygona- ceae, Salvadoraceae, Solanaceae, and Zygophyllaceae (Borowiec and Świętojańska 2002–2022). This is the second publication to record the feeding habit of C. sphaerula on an Arctotheca species; a comprehensive survey of agents against A. calendula was carried out in South Africa 1986, 1987 and 1988, where C. sphaerula was noted as a potential agent although possibly not sufficiently specific (Scott and Way 1990). Our study shows a strong asso- ciation of C. sphaerula with A. prostrata. Further observations and testing of the specificity of C. sphaerula would be necessary to determine whether it could be considered a potential agent for biological control. Chrysomelid females provide several lines of physical and chemical protection of their eggs, including oothecal and excremental coverings (Hilker 1994). Eggs have been documented for 13 Cassida species in South Africa and females deposit their eggs singly or in small groups to the undersides of their host leaves (often along- side a vein) in simple oothecae that lacks a stalk. The ootheca of C. sphaerula has a single layer enclosing the eggs, in contrast to the large complex multi-membrane oothecae with many eggs in Conchylotenia (Heron 1999) and Aspidimorpha (Muir and Sharp 1904). In C. sphaerula, ootheca have no fecal cover. Within the genus Cassida, C. 16 · July 29, 2022 Adam et al. coagulata Boheman, 1854 is a notable exception with a larger more elaborate oothecae generally attached to their host plant stem (Amaranthaceae hosts in this case, not Asteraceae; H. Heron, pers. observ.). Female oviposition behaviors, including site preparation and coverings of the ootheca, the oothecal structure, and qualities of the egg mass appear to vary within the genus Cassida and suggest novel phylogenetic characters. We observed the distinct feeding pattern that pushes the epidermis to one side and leaves craters on the dorsal surface of the leaf. Comparison with images and data for other species suggests this is a distinct pattern, now known for at least three South African Cassida species. Author Heron photographed similar patterns for Cassida guttipennis Boheman, 1862 on the host, Berkheya bipinnatifida (Harvey) Roessl (Asteraceae), and Cas- sida quatuordecimsignata Spaeth, 1899 on the host, Berkheya maritima J.M. Wood and M.S. Evans (Asteraceae) (see Heron and Borowiec 1997: 643, Fig. 19; Heron 2011: 137, Fig. 9; Heron 2003: 43, Fig. XXV) without discuss- ing how the pattern arose. These three species are the only ones where such a pattern is reported; the midrib and secondary veins are not eaten, and the craters are found in areas between veins. Bieńkowski (2010) described the more typical chewing pattern in two other Cassida species. These patterns suggest intrinsic intra-generic variations within Cassida. As more feeding patterns are recognized, novel hypotheses about their significance are emerging; for example, a masquerade strategy in some leaf beetles (Konstantinov et al. 2018). The careful observation and filming of feeding in C. sphaerula allow us to determine how the windowpane feeding pattern arises. It is unclear if the rolling over of the epidermis is related to the sheer density of trichomes (see Fig. 13)—pushing trichomatous cuticle out of the way avoids energy and time costs to cut trichomes and clear a feeding path. We observed C. sphaerula larvae consuming trichomes, which has not been reported for any Chrysomelidae before. In Chrysomelinae chrysomelids, larvae of some Platyphora species were observed to cut and throw trichomes backwards unto their fecal shields (Bernardi and Scivittaro 1991; Flinte et al. 2017: 15). In Campostomate chrysomelids, larvae trim and store trichomes into a section (“attic”) of the fecal case (Brown and Funk 2005) or incorporate trichomes and feces to make the case wall (Chaboo et al. 2008). Trichome-consumption may not be a regular part of the diet and the nutritive value is unclear. The feeding process may be flexible when trichomes are less dense. Trichome density impacts movements of cassidine larval (larvae use the tarsungulus to insert into the epidermis and “tiptoe” to move) (Medeiros et al. 2004; Medeiros and Moreira 2005). Author Her- on’s observations of C. guttipennis feeding revealed that more typical circular feeding scars without rolled cuticle margin is left on those plants with less dense pubescence, e.g., Berkheya speciosa (DC.) O. Hoffm. (Asteraceae). In C. sphaerula, all five larval instars and the pupa retain an exuvio-fecal shield. Instar I has a feces-only shield (Fig. 19–20); instars II–V retain previous exuviae in a stack, with feces applied. The pupae exhibit vari- ability, retaining either the instar V exuviae only (Fig. 26) or the entire structure of the larval stages (Fig. 27). It is unclear what the different benefits are of each shield form. Within the genus Cassida, shields vary in architecture, some with exuviae only, or with exuviae covered with fecal or with fecal filaments (Świętojańska 2009). Life history data can provide a great deal of comparative information to support species concepts and evo- lutionary relationships. Some of our findings are relevant to character hypotheses presented in two phylogenetic analyses of Cassidinae, Borowiec (1995), particularly his characters 15–19, and Chaboo (2007; 16 larval char- acters). Our findings also suggests new characters and new states to expand López-Pérez et al.’s (2018) dataset for the phylogeny of Cassidini. The similarity of feeding pattern in C. guttipennis, C. quatuordecimsignata, and C. sphaerula may be clues to shared behavior and morphology, possibly defining a sub-group within Cassida. The production and relative simplicity of the ootheca in C. sphaerula compared to the more complex one in C. coagulata indicate intra-generic variations and other potential characters, for example ootheca present or absent, size (e.g., number and arrangement of eggs), structure (membranes, additional layers of chewed plant material or feces). The preparation of the oviposition site and the post-ovipositional behaviors of the female await compara- tive study and evolutionary analysis. The exuvio-fecal shield that diagnoses the eight derived tribes of Cassidinae is a unique morpho-behavioral complex (Chaboo 2007), an example of an extended phenotype, like a bird’s nest (Dawkins 1989). This is a signifi- cant macroevolutionary event in the evolution of Cassidinae, however, our current picture of its origin is murky. At the base of the tortoise beetle clade, Delocraniini larvae were described as “pouco encobertas pelos excre- mentos” (=barely covered by excrement) so not carrying a shield (Bondar 1940: 1 02), Hemisphaerotini larvae have caudal processes and a unique “bird-nest” shield architecture (Chaboo and Nguyen 2004), and Spilophorini larvae have caudal processes and an exuviae-only shield (Nishida et al. 2020). In contrast, the mining larvae of Natural history of Cassida sphaerula Insecta Mundi 0945 · 17 Notosacanthini lack caudal processes and lack shields (Monteith et al. 2021). Also, remarkable is the independent origin of shield retention in the distantly related ‘hispine’, Oediopalpa Baly, 1858 (Bruch 1906). The tortoise beetle shield has been considered as a protection and a defense. Réaumur (1737) hypothesized that it protected against sun and flies. Weise (1893) hypothesized its function as defense against desiccation. More observations led to the hypothesis that shields are a defense against enemies and used cheaply-available defeca- tion products and exuviae and perhaps even chemicals in exocrine glands of those exuviae (Olmstead 1994). Mechanical defense against predators has been tested experimentally, with support by several researchers (Eisner et al. 1967; Olmstead and Denno 1993; Eisner and Eisner 2000) but contradicted by others (Müller and Hilker 1999; Nogueira-de-Sá and Trigo 2002). Further studies with Cassida larvae point to more selective shield defense to certain enemies: Schenk and Bacher (2002) showed shields were effective against vespid predators only, while Bacher and Luder (2005) showed they were effective against parasitoids only and offer some protection against desiccation and wind, but not so against abiotic factors of UV-radiation. Müller (2002) also found variable effec- tiveness of shields to deter different predators. Chemical defense via enteric discharges in shields was proposed by Pasteels et al. (1988). Chemicals sequestered from host plants or by de novo synthesis can enhance shield defenses (Gómez et al. 1999; Vencl et al. 1999, 2005, 2009, 2011; Nogueira-de-Sá and Trigo 2002, 2005), however, chemicals were also found to have no impact on larval survival (Bottcher et al. 2009). This succession of ideas and continuing testing are crucial to illuminating the origin, function (i.e., cost benefit analyses), and diversity of fecal architectures. Phylogenetic studies in Cassidinae have relied largely on adult characters. In the past, a few characters and states of juvenile stages have been proposed: Borowiec (1995) tested four characters of larvae for his phylogeny of Cassidinae, Chaboo’s (2007) study included 20 from juveniles, López-Pérez et al. (2018) tested one larval character. Going forward, we anticipate more studies like López-Pérez et al. (2021) that hypothesized nine novel characters with their possible states for pupae (their shield present/absent is equal to Chaboo 2007: char. 19). Juvenile stages, behavior, and ecology offer a wealth of new characters that could strengthen systematics of Cas- sidinae (indeed, all insects), from species concepts to tribal relations. Juvenile stages of most insects are extremely underrepresented in museum collections. The research challenge is detailed field studies and collections and descriptions of specimens. Acknowledgments We are grateful to Beth Grobbelaar in South Africa for discussion and to Prof. Maureen Coetzee, University of Witwatersrand, for the loan of a stereo microscope to SA. Thanks to Robert J. McKenzie and Nicola Bergh for confirming the plant identification. We thank Cheryl Barr, Geoff Monteith, and Alexey Tishechkin for helping us locate researchers studying Arcotheca pest issues. We thank Sandra Falanga and Wendy Wiles for traveling to inspect plants for beetle activity in new areas, to John Scott for directing us to certain literature, and to Stephen D. Hopper for use of his landscape photograph. We also thank Michael Geiser, BMNH, UK and Dmitri Logunov, Manchester Museum, UK for answering some collection questions, as well as Lech Borowiec for Cassida ques- tions. We are grateful to reviewers Orlando Calcetas, Sara López-Pérez, and Divakaran Prathapan, for their time and comments that improved the final manuscript. Finally, we thank the editorial team at Insecta Mundi for their help. CSC’s effort was funded by U.S.A. National Science Foundation grant EAGER 1663680 (PI: CS Chaboo). Literature Cited Agassiz L. 1846. Nomenclatoris zoologici index universalis: Nomina systematica classium, ordinum, familiarum et generum animalium omnium. Fasciculus XI. Continens Coleoptera. Soloduri. 170 p. Atlas of Living Australia. 2021. Atlas of Living Australia – Open Access to Australia’s Biodiversity Data. National Research Infrastructure for Australia. Available at https://www.ala.org.au/ (Last accessed 31 August 2021.) Australasian Virtual Herbarium. 2021. Council of Heads of Australasian Herbaria. Available at https://avh.chah.org.au (Last accessed 11 October 2021). Bacher S, Luder S. 2005. Picky predators and the function of the faecal shield of a cassidine larva. Ecology 19: 263–272. Baly JS. 1858. Catalogue of Hispidae in the collection of the British Museum. Printed by order of the Trustees; London. 172 p. 18 · July 29, 2022 Adam et al. Barrows EM. 1979. Life cycles, mating, and color change in tortoise beetles (Coleoptera: Chrysomelidae: Cassidinae). The Coleopterists Bulletin 33: 9–16. Bernardi N, Scivittaro A. 1991. Estágios imaturos de Platyphora zonata (Germar, 1824) (Coleoptera, Chrysomelidae, Chrys- omelinae). Revista Brasileira de Zoologia 7(4): 531–534. Beyers JBP. 2000. Haplocarpha. p. 303–306. In: Goldblatt P, Manning JC (eds.). Cape plants: A conspectus of the cape flora of South Africa. Strelitzia 9. National Botanical Institute, Pretoria, and Missouri Botanical Garden; St. Louis, MO. 743 p. Bieńkowski AO. 2010. Feeding behavior of leaf beetles (Coleoptera, Chrysomelidae). Entomological Review 90(1): 1–10. Boheman CH. 1854. Monographia Cassididarum. Tomus secundus. Holmiae. 506 p. Boheman CH. 1855. Monographia Cassididarum. Tomus tertius. Holmiae. 543 p. + 1 tab. Boheman CH. 1862. Monographia Cassididarum. Tomus quartus. Holmiae. 504 p. Bondar G. 1940. Insectos nocivos e moléstias do Coqueiro (Cocos nucifera) no Brasil. Boletim Instituto Central de Fomento Economica da Bahia 8: 160 p. Bordy B, Doguet S. 1987. Contribution à la connaissance des Cassidinae de France. Étude de leur spermathèque (Coleop- tera, Chrysomelidae). Nouvelle Revue d’Entomologie (N.S.) 4(2): 161–176. Borowiec L. 1995. Tribal classification of the cassidoid Hispinae (Coleoptera: Chrysomelidae). p. 541–558. In: Pakaluk J, Ślipiński SA (eds.). Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN; Warszawa, Poland. 1092 p. Borowiec L. 1999. A world catalogue of the Cassidinae (Coleoptera: Chrysomelidae). Biologica Silesiae; Wrocław, Poland. 476 p. Borowiec L. 2002. A monograph of the Afrotropical Cassidinae (Coleoptera: Chrysomelidae). Part III. Revision of the tribe Cassidini 1, except the genera Aethiopocassis sp., Cassida L., and Chiridopsis sp. Biologica Silesiae; Wrocław, Poland. 292 p. + 17 pl. Borowiec L, Świętojańska J. 2001. Revision of the Cassida litigiosa group from southern Africa (Coleoptera: Chrysomelidae: Cassidinae). Annales Zoologici 51(2): 153–184. Borowiec L, Świętojańska J. 2002–2022. World Catalog of Cassidinae, Wrocław, Poland. Available at http://www. cassidae.uni.wroc.pl/katalog%20internetowy/index.htm (Last accessed 3 February 2022). Borowiec L, Świętojańska J. 2022 [In press]. A monograph of the Afrotropical Cassidinae (Coleoptera: Chrysomelidae). Part 6. Revision of the tribe Cassidini 3, the genus Cassida L. Zootaxa (monograph). Bottcher A, Zolin JP, Nogueira-de-Sá F, Trigo JR. 2009. Faecal shield chemical defense is not important in larvae of the tortoise beetle Chelymorpha reimoseri (Chrysomelidae: Cassidinae: Stolaini). Chemoecology 19: 63–66. Brossard CC, Randall JM, Hoshovsky MC. 2000. Invasive Plants of California’s Wildlands. University of California Press; Berkeley, CA. 360 p. Brown CG, Funk DJ. 2005. Aspects of the natural history of Neochlamisus (Coleoptera: Chrysomelidae): fecal-case associ- ated life history and behavior, with a method for studying the construction of insect defensive structures. Annals of the Entomological Society of America 98(5): 711–725. Bruch C. 1906. Metamorfosis y biológia de coleópteros argentines. II. Agasicles vittata Jac., Plectonycha correntina Lac., Amplipalpa negligens Weise. Revista del Museo La Plata 12: 207–219. Brusati E. 2004. Plant Assessment Form: Arctotheca prostrata. In: California Invasive Plant Council website. Available at https://www.cal-ipc.org/plants/paf/arctotheca-prostrata-plant-assessment-form/ (Last accessed 11 October 2021). Calflora. 2021. The Calflora Database [web application]. Available at https://www.calflora.org/. Berkeley, CA (Last accessed October 2021). Candide. 2021. “Arctotheca on Candide.” Available at https://candidegardening.com/GB/search?search=arctotheca&search_ category=website (Last accessed 11 October 2021). California Invasive Plant Council. 2021. Available at https://www.cal-ipc.org/California Invasive Plant Council (Last accessed 31 August 2021). Chaboo CS. 2007. Biology and phylogeny of the Cassidinae (tortoise and leaf-mining beetles) (Coleoptera: Chrysomelidae). Bulletin of the American Museum of Natural History 305: 1–250. Chaboo CS. 2011. Defensive behaviors in leaf beetles: from the unusual to the weird. p. 59–69. In: Vivanco JM, Weir T (eds.). Chemical Biology of the Tropics: An Interdisciplinary approach. Springer; Berlin, Germany. 115 p. Chaboo CS, Brown CG, Funk DJ. 2008. Faecal case architecture in the gibbosus species group of Neochlamisus Karren, 1972 (Coleoptera: Chrysomelidae: Cryptocephalinae: Chlamisini). Zoological Journal of the Linnean Society 152: 315–335. Chaboo CS, Grobbelaar E, Larsen A. 2007. Fecal ecology in leaf beetles: novel records in the African arrow-poison beetles, Diamphidia Gerstaecker and Polyclada Chevrolat (Chrysomelidae: Galerucinae). The Coleopterists Bulletin 61(2): 297–309. Chaboo CS, Nguyen T. 2004. Immatures of Hemisphaerota palmarum (Boheman), with a discussion of the caudal process and shield architecture in the tribe Hemisphaerotini (Chrysomelidae: Cassidinae). p. 171–184. In: Jolivet P, Santiago-Blay J, Natural history of Cassida sphaerula Insecta Mundi 0945 · 19 Schmitt M (eds.). New contributions in biology of the Chrysomelidae. Kugler Publications; The Hague, Netherlands. 803 p. Chapuis F. 1875. Groupe XI. Cassidites. p. 383–392. In: Lacordaire JT. Histoire naturelle des insectes. Genera des Coléop- tères, Vol. 11, Famille des Phytophages. Encylopédique de Roret; Paris, France. 420 p. Chen SH. 1940. Attempt at a new classification of the leaf beetles. Sinensia 11: 451–481. Chevrolat LAA. 1837. Notosacantha Chevrolat. p. 391. In: Dejean PFMA. Catalogue des Coléoptères de la collection de M. le comte Dejean. Troisième edition, revue, corrigée et augmentée, livr. 5. Mequignon-Marvis; Paris, France. 503 p. Dawkins R. 1989. The extended phenotype. Oxford University Press; Oxford, UK. 440 p. Desbrochers J. 1884. (Description of Oxylepus). Bulletin de l’Académie d’Hippone 19: 100. EDDMapS. 2021. Early detection and distribution mapping system. The University of Georgia - Center for Invasive Species and Ecosystem Health. Available at http://www.eddmaps.org/ (Last accessed 12 October 2021.) Eisner T, Eisner M. 2000. Defensive use of a fecal thatch by a beetle larva (Hemisphaerota cyanea). Proceedings of the National Academy of Sciences 97: 2632–2636. Eisner T, van Tassel E, Carrel JE. 1967. Defensive use of a ‘fecal shield’ by a beetle larva. Science 158: 1471–1473. Fairmaire L. 1868. Famille des Chrysomelides. p. 205–268. In: Du Val J (ed.). Genera des Coléoptères d’Europe. Chez Dey- rolle fils; Paris, France. 292 p. Fairmaire L. 1882. Rapports, lectures, communications. Annales de la Société Entomologique de Belgique 26: 57. Flinte V, Abejanella A, Daccordi M, Monteiro RF, Macedo MV. 2017. Chrysomelinae species (Coleoptera, Chrysomelidae) and new biological data from Rio de Janeiro, Brazil. ZooKeys 720: 5–22. Galasso G, Domina G, Ardenghi NM, Aristarchi C, Bacchetta G, Bartolucci F, Bonari G, Bouvet D, Brundu G, Buono S, Caldarella O. 2019. Notulae to the Italian alien vascular flora: 7. Italian Botanist 7: 157–182. García-Robledo C, Horvitz CC, Staines CL. 2010. Larval morphology, development, and notes on the natural history of Cephaloleia “rolled-leaf ” beetles (Coleoptera: Chrysomelidae: Cassidinae). Zootaxa 2610: 50–68. Gardenia.net. 2021. Arctotheca calendula (cape dandelion). Available at https://www.gardenia.net/plant/arctotheca- calendula (Last accessed 11 October 2021). Gestro R. 1897. Materiali per lo studio delle Hispidae. Annali del Museo Civico di Storia Naturale di Genova (2)18(38): 37–138. GBIF. 2021a. GBIF.org. GBIF Occurrence Download –Arctotheca calendula. Available at https://doi.org/10.15468/dl.zjqxxx (Last accessed 01 October 2021.) GBIF. 2001b. GBIF.org. GBIF Occurrence Download –Arctotheca populifolia. Available at https://doi.org/10.15468/ dl.hkah5n, (Last accessed 01 October 2021.) GBIF. 2021c. GBIF.org. GBIF Occurrence Download –Arctotheca prostrata. Available at https://doi.org/10.15468/dl.sp3tp3, (Last accessed 01 October 2021.) Ghafoor A, Bean T. 2015. The tribe Arctotideae. p. 159–171. In: Wilson A (ed.). Flora of Australia. Volume 37. Asteraceae 1. CSIRO Publishing; Canberra, Australia. 638 p. Gillot C. 2002. Insect accessory reproductive glands: key players in production and protection of eggs. p. 37–59. In: Hilker M, Meiners T (eds.). Chemoecology of insect eggs and egg deposition. Blackwell Publishing; Berlin, Germany. 390 p. Goedært J. 1662. Metamorphosis et Historia Naturalis Insectorum. Jacques Fierens; Middelburg, The Netherlands. 494 p. Gómez NE, Witte L, Hartmann T. 1999. Chemical defense in larval tortoise beetles: essential oil composition of fecal shields of Eurypedus nigrosignata and foliage of its host plant, Cordia curassavica. Journal of Chemical Ecology 25: 1007–1027. Groves RH, Hosking JR, Batianoff GN, Cooke DA, Cowie ID, Johnson RW, Keighery GJ, Lepschi BJ, Mitchell AA, Moerkerk M, Randall RP, Rozefelds C, Walsh NG, Waterhouse BM. 2003. Weed categories for natural and agricul- tural ecosystem management. Bureau of Rural Sciences; Canberra, Australia. 194 p. Guérin-Méneville FE. 1840. Description de cinq espèces d’Hispes, formant une division distincte dans ce genre. Revue zoologique 3: 139–142. Gyllenhal L. 1813. Insecta Suecica. Classis 1. Coleoptera sive Eleuterata, Tomus I, pars III. Scaris. 734 p. Gyllenhal L. 1817. Appendix. Descriptiones Novarum specierum. Insectorum. Hispa. p. 3–7. In: Schönherr CJ. Synonymia insectorum, oder: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet. Vol. 1, pt. 3. Stockholm. 506 p. Harley KLS, Forno IW. 1992. Biological control of weeds: a handbook for practitioners and students. Inkata Press; Canberra, Australia. 74 p. Herman PPJ. 2002. Revision of Tarchonanthus camphoratus complex (Asteraceae - Tarchonantheae) in Southern Africa. Bothalia 32: 21–28. Heron HDC. 1999. The biology of Conchylotenia punctata (Fabricius) – a cycloalexic cassid (Chrysomelidae: Cassidinae). p. 565–580. In: Cox ML (ed.). Advances in Chrysomelidae Biology 1. Backhuys Publishers; Leiden, The Netherlands. 691 p. 20 · July 29, 2022 Adam et al. Heron HDC. 2003. Tortoise beetles (Chrysomelidae: Cassidinae) and their feeding patterns from the North Park Nature Reserve, Durban, KwaZulu-Natal, South Africa. Durban Museum Novitates 28: 31–44. Heron H. 2008. Novel trophic behaviour in two South African tortoise beetles (Chrysomelidae, Cassidinae). Chrysomela 50–51: 13, 26. Heron HDC. 2011. Polymorphism in four tortoise beetles from Queensburgh, South Africa (Chrysomelidae: Cassidinae). Genus 22(1): 133–149. Heron HDC. 2018. Tortoise beetle-host plant relationships at North Park Nature Reserve, Queensburgh. PlantLife SA 46: 8. Heron H, Borowiec L. 1997. Host plants and feeding patterns of some South African tortoise beetles (Coleoptera: Chryso- melidae: cassidoid Hispinae). Genus 8: 625–658. Hilker M. 1994. Egg deposition and protection of eggs in Chrysomelidae. In: Jolivet PH, Cox ML, Petitpierre E (eds.). Novel aspects of the biology of Chrysomelidae. Series Entomologica 50: 263–276. Kluwer Academic Publishers; Dordrecht. 582 p. Hincks WD. 1952. The genera of the Cassidinae. Transactions of the Royal Entomological Society of London 103: 327–358. Hinojosa-Espinosa O, Villaseñor JL. 2015. Arctotheca prostrata (Asteraceae: Arctotideae), a South African species now present in Mexico. Botanical Sciences 93(4): 877–880. Hinton HE. 1981. Biology of insect eggs. Pergamon Press; Oxford. vols. 1–3: 1125 p. Hope FW. 1840. The Coleopterist’s Manual. Part 3. J. C. Bridgewater; London. 191 p. iNaturalist. 2021. A Communtiy for Naturalists – iNaturalist. Available at https://www.inaturalist.org/(Last accessed 1 Feb- ruary 2022.) Jepson eFlora. 2021. University and Jepson Herbaria Home Page. Available at https://ucjeps.berkeley.edu (Last accessed 31 August 2021.) Jolivet P, Hawkeswood TJ. 1995. Host-plants of Chrysomelidae of the world. Backhuys; Leiden. 281 p. Jolivet P, Vasconcellos-Neto J, Weinstein P. 1990. Cycloalexy: a new concept in the larval defense of insects. Insecta Mundi 4: 133–141. Karis PO, Funk VA, McKenzie RJ, Barker NP, Chan R. 2009. Arctotideae. 385–410. In: Funk VA, Susana A, Stuessy TF, Bayer RJ (eds.). Systematics, Evolution, and Biogeography of Compositae. International Association for Plant Tax- onomy (IAPT); Vienna, Austria. 965 p. Knutson L, Coulston JR. 1997. Procedures and policies in the USA regarding precautions in the introduction of classical biocontrol agents. Bulletin OEPP/EPPO Bulletin 27: 133–142. Konstantinov AS, Prathapan KD, Vencl FV. 2018. Hiding in plain sight: leaf beetles (Chrysomelidae: Galerucinae) use feed- ing damage as a masquerade decoy. Biological Journal of the Linnean Society 123: 311–320. Kottek M, Grieser J, Beck C, Rudolf B, Rubel F. 2006. World Map of the Köppen-Geiger climate classification updated. Meteorologische Zeitschrift 15(3): 259–263. Kraatz G. 1895. Hispinae von Togo. Deutsche Entomologische Zeitschrift 1895: 189–200. Linnaeus C. 1758. Systema Naturae, sive regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima, reformata. I. Holmiae. IV + 824 p. López-Pérez S, Rodríguez-Mirón GM, Chaboo CS. 2021. Pupal morphology of Physonota humilis Boheman, 1856 and Physonota stigmatilis Boheman 1854 (Chrysomelidae: Cassidinae: Ischyrosonychini). Zootaxa 5027(1): 107–119. López-Pérez S, Zaragoza-Caballero S, Ochoterena H, Morrone JJ. 2018. A phylogenetic study of the worldwide tribe Cas- sidini Gyllenhal, 1813 (Coleoptera: Chrysomelidae: Cassidinae) based on morphological data. Systematic Entomology 43: 372–386. Lucid Central Weeds of Australia. 2021. Arctotheca calendula. Available at https://keyserver.lucidcentral.org/weeds/data/ media/Html/arctotheca_calendula.htm. (Last accessed 18 November 2021.) Mahoney AM, McKenzie RJ. 2008. Notes on two Southern African Arctotis species (Arctotideae: Asteraceae) growing in California. Madroño 55(3): 244–247. McKenzie RJ, Samuel J, Muller EM, Skinner AK, Barker NP. 2005. Morphology of cypselae in subtribe Arctotidinae (Com- positae-Arctotideae) and its taxonomic implications. Annals of the Missouri Botanical Garden 92: 569–594. Medeiros L, Bolignon DS, Moreira GRP. 2004. Morphological and behavioral adaptations to movement on different leaf surfaces: studies with Cassidinae larvae. p. 291–303. In: Jolivet P, Santiago-Blay J, Schmitt M (eds.). New contributions in biology of the Chrysomelidae. Kugler Publications; The Hague. 803 p. Medeiros L, Moreira GRP. 2005. Larval feeding behavior of Gratiana spadicea (Klug) (Coleoptera: Chrysomelidae: Cassidi- nae) on its host plant, Solanum sisymbriifolium Lamarck (Solanaceae): interaction with trichomes. The Coleopterists Bulletin 59(3): 339–350. Monteith GB, Sandoval Gomez VE, Chaboo CS. 2021. Natural history of the Australian tortoise beetle Notosacantha dorsa- lis (Waterhouse, 1877) (Coleoptera: Chrysomelidae: Cassidinae: Notosacanthini) with summary of the genus in Aus- tralia. The Australian Entomologist 48(4): 329–354. Natural history of Cassida sphaerula Insecta Mundi 0945 · 21 Muir F, Sharp D. 1904. On the egg-cases and early stages of some Cassididae. Transactions of the Royal Entomological Soci- ety of London 1904: 1–23. Müller C. 2002. Variation in the effectiveness of abdominal shields of cassidine larvae against predators. Entomologia Exper- imentalis et Applicata 102: 191–198. Müller C, Hilker M. 1999. Unexpected reactions of a generalist predator towards defensive devices of cassidine larva (Cole- optera: Chrysomelidae). Oecologia 118: 166–172. Müller C, Hilker M. 2003. The advantages and disadvantages of larval abdominal shields on the Chrysomelidae: a mini- review. p. 243–259. In: Furth DG (ed.). Special topics in leaf beetle biology. Pensoft; Sofia. 332 p. Müller C, Rosenberger C. 2006. Different oviposition behavior in Chrysomelid beetles: Characterization of the interface between oviposition secretion and the plant surface. Arthropod Structure and Development 35: 197–205. Müller-Schärer H, Schaffner U. 2008. Classical biological control: exploiting enemy escape to manage plant invasions. Bio- logical Invasions 10: 859–874. Nishida K, Ferrufino-Acosta L, Chaboo CS. 2020. A new host plant family for Cassidinae s.l.: Calyptocephala attenuata (Spaeth, 1919) (Coleoptera: Chrysomelidae: Cassidinae: Spilophorini) on Smilax (Smilacaceae) in Costa Rica. Pan- Pacific Entomologist 96(4): 263–267. Nogueira-de-Sá F, Trigo JR. 2002. Do fecal shields provide physical protection to larvae of the tortoise beetles Plagiome- triona flavescens and Stolas chalybea against natural enemies? Entomologia Experimentalis et Applicata 104: 203–206. Nogueira-de-Sá F, Trigo JR. 2005. Faecal shield of the tortoise beetle Plagiometriona aff. flavescens (Chrysomelidae: Cas- sidinae) as chemically mediated defense against predators. Journal of Tropical Ecology 21: 189–194. Olmstead KL. 1994. Waste products as chrysomelid defenses. p. 311–318. In: Jolivet PH, Cox ML, Petitpierre E (eds.). Novel aspects of the biology of Chrysomelidae. Series Entomologica 50. Kluwer Academic Publishers; Dordrecht. 582 p. Olmstead K, Denno RF. 1993. Effectiveness of tortoise beetle larval shields against different predator species. Ecology 74: 1394–1405. Pallas PS. 1778. Novae species quadrupedum e glirium ordine cum illustrationibus variis complurium ex hoc ordine anima- lium. Wolfgang Walther; Erlangen, Germany. 388 p. Pasteels JM, Braekman JC, Daloze D. 1988. Chemical defense in Chrysomelidae. p. 233–252. In: Jolivet PH, Cox ML, Petit- pierre E (eds.). Biology of Chrysomelidae. Kluwer Academic Publishers; Dordrecht. 615 p. Réaumur RAF de. 1737. Mémoirs pour servir à l'histoire des insects. Tome 3. Paris Imprimie Royale; Paris. 47 pl. + 532 p. Schenk D, Bacher S. 2002. Functional response of a generalist insect predator to one of its prey species in the field. Journal of Animal Ecology 71: 524–531. Schreber JCD. 1782. Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen. Supplementband III [Dritte Ubtheilung: Die Beutelthiere und Rage]. Wolfgang Walther; Erlangen, Germany. 628 p. Scott JK, Way MJ. 1990. A survey in South Africa for potential biological control agents against capeweed, Arctotheca calen- dula (L.) Levyns (Asteraceae). Plant Protection Quarterly 5(1): 31–34. Sekerka L, Windsor D, Staines CL. 2013. A new species of Cephaloleia from Panama with description of larva and first record of orchid-feeding in Cephaloleiini (Coleoptera: Chrysomelidae: Cassidinae). Acta Entomologica Musei Natio- nalis Pragae 53: 303–314. Simon Thomas RT. 1964. Some aspects of life history, genetics, distribution, and taxonomy of Aspidimorpha adhaerens (Weber, 1801) (Cassidinae, Coleoptera). Tijdschrift voor Entomologie 107: 167–264. Spaeth F. 1899. Beschreibung einiger neuer Cassididen nebst synonymischen Bemerkungen. III. Verhandlungen der Zoolo- gisch-Botanischen Gesellschaft in Wien: 213–221. Spaeth F. 1902. Eine neue Casside aus Birma. Entomologisk Tidskrift 24: 111–112. Spaeth F. 1909. 7. Coleoptera. 13. Cassidae. In: Sjöstedt I. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massai-steppen Deutsch-Ostafrikas 1905–1906 unter Leitung von Prof. Ingve Sjöstedt 7(13): 267–287. Spaeth F. 1911. Beschreibung neuer Cassididen nebst synonymischen Bemerkungen VIII. Verhandlungen der Kaiserlich- Koeniglichen Zoologisch-Botanischen Gesellschaft in Wien 61: 239–277. Spaeth F. 1913. Studien über die Gattung Hoplionota Hope und Beschreibung einer verwandten neuen Gattung. Verhand- lungen der Kaiserlich-Koeniglichen Zoologisch-Botanischen Gesellschaft in Wien 63: 381–534. Spaeth F. 1914. Coleopterorum Catalogus. Pars 62. Chrysomelidae: 16. Cassidinae. W. Junk; Berlin. 182 p. Spaeth F. 1917. Neuer Beitrag zur Kenntnis der Ost- und Zentralafrikanischen Cassidinen. Annales Historico-Naturales Musei Nationalis Hungarici 15: 422–444. Spaeth F. 1922. Cassidinae. p. 275–363. In: Alluaud C, Jeannel R. Voyage de Ch. Alluaud et R. Jeannel en Afrique orientale (1911–1912). Résultats scientifiques. Coleoptera XVIII. A. Schultz; Paris. 575 p. Spaeth F. 1934. Neue Beiträge zur Kenntnis der Afrikanischen Cassidinen (Col. Chrys.). Revue de Zoologie et de Botanique Africaines 24(4): 380–393. 22 · July 29, 2022 Adam et al. Spaeth F. 1941. Neue Cassidinen aus Italienish-Ost-Afrika des Mueso Civico di Storia Naturale-Trieste. Atti del Museo Civico di Storia Naturale di Trieste 14: 315–318. Spaeth F. 1952. Limnocassis. p. 346. In: Hincks WD. The genera of the Cassidinae. Transactions of the Royal Entomological Society of London 103: 327–358. Staines CL. 2002. The New World tribes and genera of hispines (Coleoptera: Chrysomelidae: Cassidinae). Proceedings of the Entomological Society of Washington 104: 721–784. Staines CL. 2015. Catalog of the Hispines of the World (Coleoptera: Chrysomelidae: Cassidinae). Available at https:// naturalhistory.si.edu/research/entomology/collections-overview/coleoptera/catalog-hispines-world (Last accessed 14 November 2021.) Steinhausen W. 1950. Vergleichende Morphologie, Biologie und Ökologie der Entwicklungstadien der in Niedersachen hemischen Schildkäfer (Cassidinae, Chrysom. Col.) und deren Bedeutung für die Landwirtschaft. Dissertation Tech- nische Hoschsch: 5–69. Strand E. 1942. Miscellanea nomenclatorica zoologica et palaeontologica X. Folia Zoologica et Hydrobiologica 11: 386–402. Świętojańska J. 2009. The immature stages of tortoise beetles with review of all described taxa (Coleoptera: Chrysomelidae: Cassidinae). Polish Taxonomical Monographs 16: 157 p. Taylor JS. 1965. Notes on some South African tortoise beetles (Cassidinae: Chrysomelidae). The Entomologist’s Record 77: 187–190. Uhmann E. 1959. Callanispa rasa gen. nov., spec. nov. aus Südafrika. 195. Beitrag zur Kenntnis der Hispinae (Coleopt.: Chrysomelidae). Journal of the Entomological Society of South Africa 22: 229–232. van Klinken RD, Raghu S. 2006. A scientific approach to agent selection. Australian Journal of Entomology 45: 253–258. Veit J. 2014. Testing the fertility and allelopathic abilities of Arctotheca prostrata (Salisb.) Britten (Asteraceae, Arctotideae), a South African plant species that has naturalized in California. Unpublished Master’s thesis, Minnesota State University, Mankato. Cornerstone: A Collection of Scholarly and Creative Works for Minnesota State University, Mankato. Avail- able at https://cornerstone.lib.mnsu.edu/etds/323/ (Last accessed 11 October 2021.) Vencl FV, Gómez NE, Ploss K, Boland W. 2009. The chlorophyll catabolite, pheophorbide a, confers predation resistance in a larval tortoise beetle shield defense. Journal of Chemical Ecology 35: 281–288. Vencl FV, Nogueira-de-Sá F, Allen BJ, Windsor DM, Futuyma DJ. 2005. Dietary specialization influences the efficacy of larval tortoise beetle shield defenses. Oecologia 145(3): 409–414. Vencl FV, Schultz JC, Mumma RC, Morton TC. 1999. The shield defense of a larval tortoise beetle. Journal of Chemical Ecology 25: 549–566. Vencl FV, Trillo PA, Geeta R. 2011. Functional interactions among tortoise beetle larval defenses reveal trait suites and escalation. Behavioral Ecology and Sociobiology 65: 227–239. Verma KK, Kalaichelvan T. 2004. Polymorphism and microtaxonomy in Chrysomelidae. p. 213–224. In: Jolivet P, Santiago- Blay J, Schmitt M (eds.). New contributions in biology of the Chrysomelidae. Kugler Publications; The Hague. 803 p. Weather Spark. 2021. The Weather Year Round Anywhere on Earth - Weather Spark. Available at https://weatherspark.com/ (Last accessed 24 October 2021.) Weise J. 1893. Naturgeschichte der Insecten Deutschland. Erste Abtheilung Coleoptera. Sechster Band. Nicolaische Verlags- Buchhandlung R. Stricker; Berlin. xiv + 1161 p. Weise J. 1897. Kritisches Verzeichnifs der von Mr. Andrews eingesandten Cassidinen und Hispinen aus Indien. Deutsche Entomologische Zeitschrift 1897: 97–150. Weise J. 1900. Breschreibungen africanischer Chrysomeliden nebst synonymischen Bemerkungen. Deutsche Entomologi- sche Zeitschrift 1900: 446–459. Weise J. 1905. Bemerkungen über Hispinen. Deutsche Entomologische Zeitschrift 1905: 317–320. Weise J. 1911. Coleopterorum Catalogus. Pars 35. Chrysomelidae: Hispinae. W. Junk; Berlin. 94 p. Wood H. 1994. The introduction and spread of capeweed, Arctotheca calendula (L.) Levyns (Asteraceae) in Australia. Plant Protection Quarterly (Australia) 9(3): 94–100. Received April 27, 2022; accepted June 24, 2022. Review editor Angélico Asenjo. Natural history of Cassida sphaerula Insecta Mundi 0945 · 23 Supplementary Materials Video 1. Cassida sphaerula larva feeding, site preparation, trichome-eating, first cut. The beetle larva clears the site by chewing trichomes, then making an arc-shaped cut in the epicuticle that is rolled underneath the body. The larva then feeds on the spongy mesophyll. Filmed by Sally Adam, Cape Town, South Africa (2021). Link: https://youtu.be/rlyAl_jj3VA (1.5 minutes). Video 2. Larva of Cassida sphaerula (Chrysomelidae) eating trichomes. Trichomes are considered an anti-predatory innovation of plants yet here is a beetle larva cutting and consuming trichomes of its host plant, of Arctotheca prostrata (Asteraceae). The black structure at the hind end of the larvae is made of its own feces. Filmed by Sally Adam, Cape Town, South Africa (2021). Link: https:// youtu.be/Ea-kbpM2qU4 (1.4 minutes). Video 3. Cassida sphaerula larva walking over a leaf with aphid. The larva ignores the aphid on the host, Arctotheca prostrata (Asteraceae). Filmed by Sally Adam, Cape Town South Africa (2021). Link: https://youtu.be/yoJzpn9FtXg (20 seconds). Video 4. Cassida sphaerula beetle adult feeding. The adult beetle feeds by cutting and rolling the epicuticle out of the way as it con- sumes the spongy mesophyll of its host, Arctotheca prostrata (Asteraceae) in South Africa. Filmed by Sally Adam, Cape Town, South Africa (2021). Link: https://youtu.be/8RZ3VWtkTRk (3 minutes).