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<document id="82C0918675CC1FFB6742D2D29A9BC0A4" ID-DOI="10.5281/zenodo.7300728" ID-ISSN="1942-1354" ID-Zenodo-Dep="7300728" ID-ZooBank="4AC56F98-6474-4AAD-A2A9-51AE2F39A1E1" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,operationResults,plazi" IM.taxonomicNames_approvedBy="carolina" IM.treatments_approvedBy="carolina" checkinTime="1667845401625" checkinUser="felipe" docAuthor="Adam, Sally, Campos, Mariana, Heron, Hugh D. C., Staines, Charles L., Westerduijn, Rob &amp; Chaboo, Caroline Simmrita" docDate="2022" docId="03EE878D6605FFF2FF542664FE25FBE3" docLanguage="en" docName="InsectaMundi.2022.0945.1-23.pdf" docOrigin="Insecta Mundi 2022 (945)" docStyle="DocumentStyle:F9976E9B7F8A094D890004515A7A0592.4:InsectaMundi.2020-.journal_article" docStyleId="F9976E9B7F8A094D890004515A7A0592" docStyleName="InsectaMundi.2020-.journal_article" docStyleVersion="4" docTitle="Cassida sphaerula Boheman 1854" docType="treatment" docVersion="1" lastPageNumber="17" masterDocId="FFD7FFF5660EFFE0FF942539FFC2FF96" masterDocTitle="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)" masterLastPageNumber="23" masterPageNumber="1" pageNumber="10" updateTime="1734616069407" updateUser="carolina" zenodo-license-document="CC-BY-NC-3.0">
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<mods:title id="8E25277B60C80BBD75B848D392C61CA2">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)</mods:title>
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<mods:namePart id="92AB930F886E5CBE9054B3E239FEB952">Adam, Sally</mods:namePart>
<mods:affiliation id="8262F1CD8418EF6436D769BF88DC9AE2">Laaiplaats 59, Mossel Bay, South Africa</mods:affiliation>
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<mods:namePart id="77B62E6978E3507C8A88A53F3192710A">Campos, Mariana</mods:namePart>
<mods:affiliation id="8AB96513B698B924A3B0E5CCFD2EC754">CSIRO Health &amp; Biosecurity 147 Underwood Avenue, Floreat 6014 Western Australia, Australia</mods:affiliation>
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<mods:namePart id="D1FA89DC811FEF27149B0ADDF920EC74">Heron, Hugh D. C.</mods:namePart>
<mods:affiliation id="F10C732DF9F1CCE5664218A9B3FEDF33">P. O. Box 39042, Escombe, Queensburgh, Natal, 4070, South Africa</mods:affiliation>
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<mods:namePart id="59CBA93BD8F43113AF66316F1F698590">Staines, Charles L.</mods:namePart>
<mods:affiliation id="F49E26211FD37E4EAF4832AAE24AEA69">Smithsonian Environmental Research Center, 647 Contees Wharf Road, Edgewater, MD 21037, U. S. A.</mods:affiliation>
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<mods:namePart id="280D2DB3153A846610479401BB6F161D">Westerduijn, Rob</mods:namePart>
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<mods:namePart id="08C38CE4F2FEDD73152D60E05A224942">Chaboo, Caroline Simmrita</mods:namePart>
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<treatment id="03EE878D6605FFF2FF542664FE25FBE3" LSID="urn:lsid:plazi:treatment:03EE878D6605FFF2FF542664FE25FBE3" httpUri="http://treatment.plazi.org/id/03EE878D6605FFF2FF542664FE25FBE3" lastPageId="18" lastPageNumber="17" pageId="11" pageNumber="10">
<subSubSection id="C35D65106605FFEBFF542664FC9DFCEC" box="[192,863,860,890]" pageId="11" pageNumber="10" type="nomenclature">
<paragraph id="8BF8369B6605FFEBFF542664FC9DFCEC" blockId="11.[192,863,860,890]" box="[192,863,860,890]" pageId="11" pageNumber="10">
<heading id="D0B081F76605FFEBFF542664FC9DFCEC" box="[192,863,860,890]" fontSize="12" level="2" pageId="11" pageNumber="10" reason="2">
<emphasis id="B933EA896605FFEBFF542664FC9DFCEC" bold="true" box="[192,863,860,890]" pageId="11" pageNumber="10">
Natural history of
<taxonomicName id="4C474D186605FFEBFE3F2665FC9DFCEC" ID-CoL="7RYXH" authority="Boheman, 1854" authorityName="Boheman" authorityYear="1854" box="[427,863,860,890]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="10" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896605FFEBFE3F2665FD50FCEF" bold="true" box="[427,658,860,889]" italics="true" pageId="11" pageNumber="10">Cassida sphaerula</emphasis>
<bibRefCitation id="EFD64B6A6605FFEBFD0D2664FC9DFCEC" author="Boheman CH" box="[665,863,861,890]" pageId="11" pageNumber="10" refId="ref11645" refString="Boheman CH. 1854. Monographia Cassididarum. Tomus secundus. Holmiae. 506 p." type="book" year="1854">Boheman, 1854</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C35D65106605FFF0FF5426B3FBC2FB78" lastPageId="16" lastPageNumber="15" pageId="11" pageNumber="10" type="description">
<paragraph id="8BF8369B6605FFEBFF5426B3FA90FBC4" blockId="11.[191,1441,906,1107]" pageId="11" pageNumber="10">
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 (
<date id="FFF9105B6605FFEBFD2F2694FC83FC51" box="[699,833,941,967]" pageId="11" pageNumber="10" value="2021-12-27">27.XII.2021</date>
) 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 ‘windows’ 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
<specimenCount id="9D41FD126605FFEBFF482101FED4FBC5" box="[220,278,1080,1107]" pageId="11" pageNumber="10" type="egg">8 egg</specimenCount>
cases have been found on a single leaf. Thus, we believe this species is an autumn/winter breeder.
</paragraph>
<paragraph id="8BF8369B6605FFEBFF54215AFB52FA68" blockId="11.[192,1441,1123,1534]" pageId="11" pageNumber="10">
<emphasis id="B933EA896605FFEBFF54215AFDD1FBEB" bold="true" box="[192,531,1123,1150]" pageId="11" pageNumber="10">
Egg cases (
<emphasis id="B933EA896605FFEBFED4215DFE8DFBE8" bold="true" box="[320,335,1124,1150]" italics="true" pageId="11" pageNumber="10">n</emphasis>
=4) (
<figureCitation id="137C2A1E6605FFEBFE12215AFDC4FBEB" box="[390,518,1123,1149]" captionStart="Figures 16–18" captionStartId="12.[192,279,1799,1825]" captionTargetBox="[246,1385,213,1769]" captionTargetId="figure-14@12.[246,1385,213,1769]" captionTargetPageId="12" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="11" pageNumber="10">Fig. 16–18</figureCitation>
).
</emphasis>
Oothecae are deposited on the venter of the leaf (
<figureCitation id="137C2A1E6605FFEBFBDE215AFB58FBEB" box="[1098,1178,1123,1149]" captionStart="Figures 16–18" captionStartId="12.[192,279,1799,1825]" captionTargetBox="[246,1385,213,1769]" captionTargetId="figure-14@12.[246,1385,213,1769]" captionTargetPageId="12" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="11" pageNumber="10">Fig. 16</figureCitation>
) 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, suspended or protuberant from the leaf surface, as in some other
<taxonomicName id="4C474D186605FFEBFC1421F5FC3EFB70" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[896,1020,1228,1254]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
.
<emphasis id="B933EA896605FFEBFB9321F4FB44FB71" box="[1031,1158,1229,1255]" italics="true" pageId="11" pageNumber="10">Oviposition</emphasis>
was not observed so it is unclear how the female may prepare a site before depositing her eggs (see
<bibRefCitation id="EFD64B6A6605FFEBFB8721D6FABBFA9F" author="Muller C &amp; Rosenberger C." box="[1043,1401,1263,1289]" pageId="11" pageNumber="10" pagination="197 - 205" refId="ref14955" refString="Muller 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." type="journal article" year="2006">Müller and Rosenberger (2006)</bibRefCitation>
for possible oviposition sequences in
<taxonomicName id="4C474D186605FFEBFDAD202BFD2BFABA" authorityName="Latreille" authorityYear="1802" box="[569,745,1298,1324]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="10" phylum="Arthropoda" rank="family">Chrysomelidae</taxonomicName>
). The ootheca, secreted by colleterial glands (
<bibRefCitation id="EFD64B6A6605FFEBFB4F202BFAA9FABA" author="Hinton HE" box="[1243,1387,1298,1324]" pageId="11" pageNumber="10" refId="ref14026" refString="Hinton HE. 1981. Biology of insect eggs. Pergamon Press; Oxford. vols. 1 - 3: 1125 p." type="book" year="1981">Hinton 1981</bibRefCitation>
;
<bibRefCitation id="EFD64B6A6605FFEBFAE2202BFEE2FAD9" author="Gillot C." pageId="11" pageNumber="10" pagination="37 - 59" refId="ref13343" refString="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." type="book chapter" year="2002">Gillot 2002</bibRefCitation>
), comprises a thin opaque outer laminate membrane that appears shiny and dark brown (
<figureCitation id="137C2A1E6605FFEBFA8C200CFA56FAD9" box="[1304,1428,1333,1359]" captionStart="Figures 16–18" captionStartId="12.[192,279,1799,1825]" captionTargetBox="[246,1385,213,1769]" captionTargetId="figure-14@12.[246,1385,213,1769]" captionTargetPageId="12" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="11" pageNumber="10">Fig. 17–18</figureCitation>
). The few enclosed eggs (less than 5) are cream-colored (
<figureCitation id="137C2A1E6605FFEBFCA72061FC6FFAE4" box="[819,941,1368,1394]" captionStart="Figures 16–18" captionStartId="12.[192,279,1799,1825]" captionTargetBox="[246,1385,213,1769]" captionTargetId="figure-14@12.[246,1385,213,1769]" captionTargetPageId="12" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="11" pageNumber="10">Fig. 17–18</figureCitation>
), dorsoventrally compressed (lying flattened on leaf), and elongate-oval shaped. The ootheca lacks any additional coverings, no fecal or chewed plant material.
<emphasis id="B933EA896605FFEBFF6720A6FEA6FA2E" box="[243,356,1438,1464]" italics="true" pageId="11" pageNumber="10">Egg hatch.</emphasis>
One ootheca was collected on
<date id="FFF9105B6605FFEBFD2A20A7FCECFA2E" box="[702,814,1438,1464]" pageId="11" pageNumber="10" value="2021-10-11">11.X.2021</date>
and
<specimenCount id="9D41FD126605FFEBFCF220A7FC28FA2E" box="[870,1002,1438,1464]" pageId="11" pageNumber="10" type="larva">three larvae</specimenCount>
hatched on
<date id="FFF9105B6605FFEBFBE720A7FB27FA2E" box="[1139,1253,1438,1464]" pageId="11" pageNumber="10" value="2021-10-18">18.X.2021</date>
, confirming that more than
<specimenCount id="9D41FD126605FFEBFEA820F8FE51FA4D" box="[316,403,1473,1499]" pageId="11" pageNumber="10" type="egg">one egg</specimenCount>
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.
</paragraph>
<paragraph id="8BF8369B6605FFEBFF542336FA60F904" blockId="11.[192,1443,1550,1927]" pageId="11" pageNumber="10">
<emphasis id="B933EA896605FFEBFF542336FDA3F9BE" bold="true" box="[192,609,1550,1577]" pageId="11" pageNumber="10">
Larva (
<emphasis id="B933EA896605FFEBFE872336FEE0F9BF" bold="true" box="[275,290,1551,1577]" italics="true" pageId="11" pageNumber="10">n</emphasis>
=20;
<figureCitation id="137C2A1E6605FFEBFECF2337FE17F9BE" box="[347,469,1550,1577]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="11" pageNumber="10">Fig. 13–14</figureCitation>
,
<figureCitation id="137C2A1E6605FFEBFE752337FE3CF9BE" box="[481,510,1550,1576]" captionStart="Figures 16–18" captionStartId="12.[192,279,1799,1825]" captionTargetBox="[246,1385,213,1769]" captionTargetId="figure-14@12.[246,1385,213,1769]" captionTargetPageId="12" captionText="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)." figureDoi="http://doi.org/10.5281/zenodo.7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="11" pageNumber="10">16</figureCitation>
, 19–25).
</emphasis>
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 contact. 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
<bibRefCitation id="EFD64B6A6605FFEBFB40234EFA57F904" author="Jolivet P &amp; Vasconcellos-Neto J &amp; Weinstein P." box="[1236,1429,1655,1682]" pageId="11" pageNumber="10" pagination="133 - 141" refId="ref14155" refString="Jolivet P, Vasconcellos-Neto J, Weinstein P. 1990. Cycloalexy: a new concept in the larval defense of insects. Insecta Mundi 4: 133 - 141." type="journal article" year="1990">Jolivet et al. 1990</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8369B6605FFEBFE9423A2FBC9F810" blockId="11.[192,1443,1550,1927]" pageId="11" pageNumber="10">
Larvae are found mostly on the venter of the leaves. Instar I (
<emphasis id="B933EA896605FFEBFC5523A2FC12F922" box="[961,976,1691,1716]" italics="true" pageId="11" pageNumber="10">n</emphasis>
=3; Fig. 19–20) are tear-dropped shaped, about
<quantity id="4CBF9B7E6605FFEBFE9E2384FE8CF94E" box="[266,334,1725,1752]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="11" pageNumber="10" unit="mm" value="2.0">2 mm</quantity>
long X
<quantity id="4CBF9B7E6605FFEBFE3F2384FE2DF94E" box="[427,495,1725,1752]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="11" pageNumber="10" unit="mm" value="1.0">1 mm</quantity>
at maximum width (across pronotum). The body is tan-colored. The paired caudal processes (
<figureCitation id="137C2A1E6605FFEBFEAF23D9FE1FF96D" box="[315,477,1760,1787]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="11" pageNumber="10">Fig. 21, 24–25</figureCitation>
; = supra-anal processes, urogomphi) are almost half as long as the body. Older larvae (
<figureCitation id="137C2A1E6605FFEBFF5D223AFED5F88B" box="[201,279,1795,1821]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="11" pageNumber="10">Fig. 23</figureCitation>
) are creamy yellow and with a dark brown central area; their cuticle is almost transparent, and the internal organs are somewhat visible (internal movements are easily seen). The scoli pattern (
<figureCitation id="137C2A1E6605FFEBFB0A221FFADAF8D6" box="[1182,1304,1830,1856]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="11" pageNumber="10">Fig. 21–22</figureCitation>
) and caudal processes processes are similar between instars and fit with
<bibRefCitation id="EFD64B6A6605FFEBFCCA2272FBB1F8F5" author="Swietojanska J." box="[862,1139,1865,1895]" pageId="11" pageNumber="10" refId="ref16063" refString="Swietojanska J. 2009. The immature stages of tortoise beetles with review of all described taxa (Coleoptera: Chrysomelidae: Cassidinae). Polish Taxonomical Monographs 16: 157 p." type="book" year="2009">Świętojańska’s (2009: 74)</bibRefCitation>
generalized
<taxonomicName id="4C474D186605FFEBFA952270FA94F8F5" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[1281,1366,1865,1891]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="10" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA896605FFEBFA952270FA94F8F5" box="[1281,1366,1865,1891]" italics="true" pageId="11" pageNumber="10">Cassida</emphasis>
</taxonomicName>
larvae having an ovoid dorso-ventrally flattened body with 16 pairs of lateral scoli.
</paragraph>
<caption id="DF3866136602FFECFF54223EFBA5F810" ID-DOI="http://doi.org/10.5281/zenodo.7300742" ID-Zenodo-Dep="7300742" httpUri="https://zenodo.org/record/7300742/files/figure.png" pageId="12" pageNumber="11" startId="12.[192,279,1799,1825]" targetBox="[246,1385,213,1769]" targetPageId="12">
<paragraph id="8BF8369B6602FFECFF54223EFBA5F810" blockId="12.[191,1440,1799,1927]" pageId="12" pageNumber="11">
<emphasis id="B933EA896602FFECFF54223EFEABF8B7" bold="true" box="[192,361,1799,1825]" pageId="12" pageNumber="11">Figures 16–18.</emphasis>
Ootheca and young larvae of
<taxonomicName id="4C474D186602FFECFD24223EFCB4F8B7" authorityName="Boheman" authorityYear="1854" box="[688,886,1799,1825]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="11" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896602FFECFD24223EFCB4F8B7" box="[688,886,1799,1825]" italics="true" pageId="12" pageNumber="11">Cassida sphaerula</emphasis>
</taxonomicName>
(photos: S. Adam, September 2021).
<emphasis id="B933EA896602FFECFA9A223EFAF1F8B7" bold="true" box="[1294,1331,1799,1825]" pageId="12" pageNumber="11">16)</emphasis>
Venter of host leaf with two oothecae (arrows) and two instar III with their black exuvio-fecal shields.
<emphasis id="B933EA896602FFECFB212211FB18F8D4" bold="true" box="[1205,1242,1832,1858]" pageId="12" pageNumber="11">17)</emphasis>
Ootheca (~2 mm long).
<emphasis id="B933EA896602FFECFE932273FEEFF8F2" bold="true" box="[263,301,1866,1892]" pageId="12" pageNumber="11">18)</emphasis>
Ootheca with oval-shaped egg.
<emphasis id="B933EA896602FFECFD042273FD77F8F2" bold="true" box="[656,693,1866,1892]" pageId="12" pageNumber="11">19)</emphasis>
Two young instar 1 (~2 mm long) with tiny black shield composed entirely of its own feces.
<emphasis id="B933EA896602FFECFE452255FE35F810" bold="true" box="[465,503,1900,1926]" pageId="12" pageNumber="11">20)</emphasis>
Mature instar 1 with larger shield (reared from Fig. 19).
</paragraph>
</caption>
<caption id="DF3866136603FFEDFF5423FDFA52F811" ID-DOI="http://doi.org/10.5281/zenodo.7300744" ID-Zenodo-Dep="7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="13" pageNumber="12" startId="13.[192,279,1732,1758]" targetBox="[380,1252,213,1702]" targetPageId="13">
<paragraph id="8BF8369B6603FFEDFF5423FDFA52F811" blockId="13.[191,1441,1732,1927]" pageId="13" pageNumber="12">
<emphasis id="B933EA896603FFEDFF5423FDFEAFF948" bold="true" box="[192,365,1732,1758]" pageId="13" pageNumber="12">Figures 21–25.</emphasis>
Larva of
<taxonomicName id="4C474D186603FFEDFE4B23FDFD6AF948" authorityName="Boheman" authorityYear="1854" box="[479,680,1732,1758]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="12" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896603FFEDFE4B23FDFD6AF948" box="[479,680,1732,1758]" italics="true" pageId="13" pageNumber="12">Cassida sphaerula</emphasis>
</taxonomicName>
(photos: S. Adam, September 2021).
<emphasis id="B933EA896603FFEDFBC323FDFBBFF948" bold="true" box="[1111,1149,1732,1758]" pageId="13" pageNumber="12">21)</emphasis>
Young instar with lateral projections called scoli; shield removed to expose paired caudal processes.
<emphasis id="B933EA896603FFEDFB9B23DCFBF7F969" bold="true" box="[1039,1077,1765,1791]" pageId="13" pageNumber="12">22)</emphasis>
Older instar (frontal view) with exuvio-fecal shield attached to caudal processes; feces appear dry.
<emphasis id="B933EA896603FFEDFC02223EFC7EF8B7" bold="true" box="[918,956,1799,1825]" pageId="13" pageNumber="12">23)</emphasis>
Older instar with moist exuvio-fecal shield.
<emphasis id="B933EA896603FFEDFF2B2211FF27F8D4" bold="true" box="[191,229,1832,1858]" pageId="13" pageNumber="12">24)</emphasis>
Older instar, dorsal view, with feces removed; legs and caudal processes of exuviae of previous instar apparent.
<emphasis id="B933EA896603FFEDFF2B2273FF26F8F2" bold="true" box="[191,228,1866,1892]" pageId="13" pageNumber="12">25)</emphasis>
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.
</paragraph>
</caption>
<paragraph id="8BF8369B6600FFEEFE9425EFFCEBFD94" blockId="14.[191,1442,214,515]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFE9425EFFE14FF66" box="[256,470,214,240]" italics="true" pageId="14" pageNumber="13">Exuvio-fecal shield.</emphasis>
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 (
<figureCitation id="137C2A1E6600FFEEFF5C2404FED4FEC1" box="[200,278,317,343]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 22</figureCitation>
) or wet (
<figureCitation id="137C2A1E6600FFEEFEE12404FE00FEC1" box="[373,450,317,343]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 23</figureCitation>
) and the telescoped anus periodically applies a dark wet droplet (see
<figureCitation id="137C2A1E6600FFEEFB2D2404FAC6FEC1" box="[1209,1284,317,343]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="14" pageNumber="13">Fig. 13</figureCitation>
) to the shield. Dissections of shields reveal a fan (
<figureCitation id="137C2A1E6600FFEEFDD62466FD53FEEF" box="[578,657,351,377]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 22</figureCitation>
) or pyramidal shape (
<figureCitation id="137C2A1E6600FFEEFC102466FC13FEEF" box="[900,977,351,377]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 23</figureCitation>
) 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.,
<figureCitation id="137C2A1E6600FFEEFF54249AFECDFE28" box="[192,271,419,446]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 23</figureCitation>
). In dissected shields (
<figureCitation id="137C2A1E6600FFEEFD91249AFDBDFE28" box="[517,639,419,446]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 24–25</figureCitation>
), 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
<figureCitation id="137C2A1E6600FFEEFD5A24D1FCDFFD94" box="[718,797,488,514]" captionStart="Figures 21–25" captionStartId="13.[192,279,1732,1758]" captionTargetBox="[380,1252,213,1702]" captionTargetId="figure-112@13.[380,1252,213,1702]" captionTargetPageId="13" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300744" httpUri="https://zenodo.org/record/7300744/files/figure.png" pageId="14" pageNumber="13">Fig. 24</figureCitation>
).
</paragraph>
<paragraph id="8BF8369B6600FFEEFF54272AFA42FD20" blockId="14.[191,1442,530,1037]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFF54272AFE1CFDBA" bold="true" box="[192,478,530,557]" pageId="14" pageNumber="13">
Pupa (
<emphasis id="B933EA896600FFEEFE98272AFED9FDBB" bold="true" box="[268,283,531,557]" italics="true" pageId="14" pageNumber="13">n</emphasis>
=10;
<figureCitation id="137C2A1E6600FFEEFEC1272BFE13FDBA" box="[341,465,530,556]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 26–27</figureCitation>
).
</emphasis>
The pre-pupal stage is typically when the mature larvae ceases feeding, become sedentary 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
<date id="FFF9105B6600FFEEFEC42740FE18FD05" box="[336,474,633,659]" pageId="14" pageNumber="13" value="2021-08-28">28.VIII.2021</date>
and the first adult appeared on
<date id="FFF9105B6600FFEEFCBF2740FC49FD05" box="[811,907,633,659]" pageId="14" pageNumber="13" value="2021-10-07">7.X.2021</date>
(9 days); two of these pupae failed and
<specimenCount id="9D41FD126600FFEEFAB92740FA60FD05" box="[1325,1442,633,659]" pageId="14" pageNumber="13" type="adult">four adults</specimenCount>
were reared. Thus, pupation (n=7) ranges from 9–20 days. No parasitoids emerged from these laboratory pupae.
</paragraph>
<paragraph id="8BF8369B6600FFEEFE942787FC69FC8A" blockId="14.[191,1442,530,1037]" pageId="14" pageNumber="13">
Pupae (
<figureCitation id="137C2A1E6600FFEEFEC72787FE08FD4E" box="[339,458,702,728]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 26–27</figureCitation>
) are ~
<quantity id="4CBF9B7E6600FFEEFD9B2787FD93FD4E" box="[527,593,702,728]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" pageId="14" pageNumber="13" unit="mm" value="9.0">9 mm</quantity>
long, solitary, affixed by their abdomen to the leaf venter, never on the upper part. There is seldom more than
<specimenCount id="9D41FD126600FFEEFDA027D8FD5FFD6D" box="[564,669,737,763]" pageId="14" pageNumber="13" type="pupa">one pupa</specimenCount>
per leaf. The pupa is tan-colored, and the body is ovoid and dorsoventrally flattened. Only the abdominal segments have lateral scoli.
</paragraph>
<paragraph id="8BF8369B6600FFEEFE94261CFE42FB9B" blockId="14.[191,1442,530,1037]" pageId="14" pageNumber="13">
The pupa of
<taxonomicName id="4C474D186600FFEEFE04261CFDD9FCA9" authorityName="Boheman" authorityYear="1854" box="[400,539,805,831]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="13" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896600FFEEFE04261CFDD9FCA9" box="[400,539,805,831]" italics="true" pageId="14" pageNumber="13">C. sphaerula</emphasis>
</taxonomicName>
shows
<specimenCount id="9D41FD126600FFEEFDF9261CFD18FCA9" box="[621,730,805,831]" pageId="14" pageNumber="13" type="generic" typeStatus="types">two types</specimenCount>
of shields. It may retain the final exuviae (
<figureCitation id="137C2A1E6600FFEEFB2E261CFAC9FCA9" box="[1210,1291,805,831]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 26</figureCitation>
) and the former larval shield may be found discarded nearby or the pupa may retain the entire shield structure of the 5
<superScript id="7C329BD36600FFEEFA06267EFA62FCC3" attach="none" box="[1426,1440,839,853]" fontSize="6" pageId="14" pageNumber="13">th</superScript>
instar larva (exuviae I–IV and their fecal matter) (
<figureCitation id="137C2A1E6600FFEEFC972650FC95FC12" box="[771,855,873,900]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 27</figureCitation>
). 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.
</paragraph>
<paragraph id="8BF8369B6600FFEEFF542124FCA8FB57" blockId="14.[192,1440,1052,1354]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFF542124FE2AFBA0" bold="true" box="[192,488,1052,1079]" pageId="14" pageNumber="13">
Adult (
<emphasis id="B933EA896600FFEEFE802124FEE1FBA1" bold="true" box="[276,291,1053,1079]" italics="true" pageId="14" pageNumber="13">n</emphasis>
=30;
<figureCitation id="137C2A1E6600FFEEFEC92124FE18FBA0" box="[349,474,1052,1079]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 28–30</figureCitation>
).
</emphasis>
These are ~
<quantity id="4CBF9B7E6600FFEEFDFB2124FD70FBA1" box="[623,690,1053,1079]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" pageId="14" pageNumber="13" unit="mm" value="9.0">9 mm</quantity>
long (along midline, head to posterior margin) by
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at their maximum width (across pronotum). The dorsum is generally pale green in color but can vary from translucent 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.
</paragraph>
<paragraph id="8BF8369B6600FFEEFE9421F1FB72FADF" blockId="14.[192,1440,1052,1354]" pageId="14" pageNumber="13">
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 permanent circular blackish marks in different locations of the elytra (
<figureCitation id="137C2A1E6600FFEEFC3B2034FBC2FAB1" box="[943,1024,1293,1319]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">Fig. 30</figureCitation>
), but we did not detect marked color polymorphism as in some
<taxonomicName id="4C474D186600FFEEFE7D2016FD93FADF" authorityName="Gyllenhal" authorityYear="1813" box="[489,593,1327,1353]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Cassidini">Cassidini</taxonomicName>
(
<bibRefCitation id="EFD64B6A6600FFEEFDCB2016FC8AFADF" author="Simon Thomas RT" box="[607,840,1327,1353]" pageId="14" pageNumber="13" pagination="167 - 264" refId="ref15513" refString="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." type="journal article" year="1964">Simon Thomas 1964</bibRefCitation>
;
<bibRefCitation id="EFD64B6A6600FFEEFCC62009FB66FADF" author="Verma KK &amp; Kalaichelvan T." box="[850,1188,1327,1354]" pageId="14" pageNumber="13" pagination="213 - 224" refId="ref16431" refString="Verma KK, Kalaichelvan T. 2004. Polymorphism and microtaxonomy in Chrysomelidae. p. 213 - 224. In: Jolivet P, SantiagoBlay J, Schmitt M (eds.). New contributions in biology of the Chrysomelidae. Kugler Publications; The Hague. 803 p." type="book chapter" year="2004">Verma and Kalaichelvan 2004</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8369B6600FFEEFF542060FE80FA6B" blockId="14.[191,1440,1369,1533]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFF542060FD9BFAE5" bold="true" box="[192,601,1369,1396]" pageId="14" pageNumber="13">
Courtship and mating (
<figureCitation id="137C2A1E6600FFEEFE442060FDE3FAE5" box="[464,545,1369,1395]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="14" pageNumber="13">Fig. 13</figureCitation>
,
<figureCitation id="137C2A1E6600FFEEFDB82060FD89FAE5" box="[556,587,1369,1395]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">29</figureCitation>
).
</emphasis>
Mating pairs were first observed on
<date id="FFF9105B6600FFEEFC782063FBBEFAE2" box="[1004,1148,1370,1396]" pageId="14" pageNumber="13" value="2021-08-24">24.VIII.2021</date>
, 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
<taxonomicName id="4C474D186600FFEEFCF220F9FC1DFA4C" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[870,991,1472,1498]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
during mating or when disturbed (
<bibRefCitation id="EFD64B6A6600FFEEFAFA20F8FEF4FA6B" author="Barrows EM" pageId="14" pageNumber="13" pagination="9 - 16" refId="ref11485" refString="Barrows EM. 1979. Life cycles, mating, and color change in tortoise beetles (Coleoptera: Chrysomelidae: Cassidinae). The Coleopterists Bulletin 33: 9 - 16." type="journal article" year="1979">Barrows 1979</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8369B6600FFEEFF542334FB36F918" blockId="14.[191,1440,1549,1678]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFF542334FE80F9B1" bold="true" box="[192,322,1549,1575]" pageId="14" pageNumber="13">Dormancy.</emphasis>
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.).
</paragraph>
<paragraph id="8BF8369B6600FFEEFF5423A7FCC1F811" blockId="14.[191,1442,1694,1927]" pageId="14" pageNumber="13">
<emphasis id="B933EA896600FFEEFF5423A7FC8CF92E" bold="true" box="[192,846,1694,1721]" pageId="14" pageNumber="13">
Feeding patterns (
<figureCitation id="137C2A1E6600FFEEFE0523A7FDA3F92E" box="[401,609,1694,1720]" captionStart-0="Figures 9–15" captionStart-1="Figures 16–18" captionStartId-0="10.[192,279,1698,1724]" captionStartId-1="12.[192,279,1799,1825]" captionTargetBox-0="[371,1260,213,1668]" captionTargetBox-1="[246,1385,213,1769]" captionTargetId-0="figure-14@10.[370,1260,213,1668]" captionTargetId-1="figure-14@12.[246,1385,213,1769]" captionTargetPageId-0="10" captionTargetPageId-1="12" captionText-0="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 exuviofecal 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." captionText-1="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)." figureDoi-0="http://doi.org/10.5281/zenodo.7300738" figureDoi-1="http://doi.org/10.5281/zenodo.7300742" httpUri-0="https://zenodo.org/record/7300738/files/figure.png" httpUri-1="https://zenodo.org/record/7300742/files/figure.png" pageId="14" pageNumber="13">Fig. 11–12, 14–16</figureCitation>
,
<figureCitation id="137C2A1E6600FFEEFDF823A7FD48F92E" box="[620,650,1694,1720]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="14" pageNumber="13">28</figureCitation>
) of
<taxonomicName id="4C474D186600FFEEFD2123A7FC85F92F" authorityName="Boheman" authorityYear="1854" box="[693,839,1694,1721]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="13" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896600FFEEFD2123A7FC85F92F" bold="true" box="[693,839,1694,1721]" italics="true" pageId="14" pageNumber="13">C. sphaerula</emphasis>
</taxonomicName>
.
</emphasis>
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 (
<figureCitation id="137C2A1E6600FFEEFA8623F9FAA0F94D" box="[1298,1378,1728,1755]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="14" pageNumber="13">Fig. 15</figureCitation>
). The craters of instar 1 are small (Fig. 19–20); older larvae and adults make craters up to
<quantity id="4CBF9B7E6600FFEEFBE523DAFB76F96B" box="[1137,1204,1763,1789]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" pageId="14" pageNumber="13" unit="mm" value="4.0">4 mm</quantity>
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 (
<figureCitation id="137C2A1E6600FFEEFF5E2273FED8F8F2" box="[202,282,1866,1892]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="14" pageNumber="13">Fig. 11</figureCitation>
), 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.
</paragraph>
<caption id="DF3866136601FFEFFF54223EFEE9F810" ID-DOI="http://doi.org/10.5281/zenodo.7300748" ID-Zenodo-Dep="7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="15" pageNumber="14" startId="15.[192,279,1799,1825]" targetBox="[219,1413,213,1769]" targetPageId="15">
<paragraph id="8BF8369B6601FFEFFF54223EFEE9F810" blockId="15.[192,1441,1799,1927]" pageId="15" pageNumber="14">
<emphasis id="B933EA896601FFEFFF54223EFEA9F8B7" bold="true" box="[192,363,1799,1825]" pageId="15" pageNumber="14">Figures 26–31.</emphasis>
Pupa and adult of
<taxonomicName id="4C474D186601FFEFFDD6223EFCC8F8B7" authorityName="Boheman" authorityYear="1854" box="[578,778,1799,1825]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="14" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA896601FFEFFDD6223EFCC8F8B7" box="[578,778,1799,1825]" italics="true" pageId="15" pageNumber="14">Cassida sphaerula</emphasis>
</taxonomicName>
(photos: S. Adam, September 2021).
<emphasis id="B933EA896601FFEFFB39223EFB11F8B7" bold="true" box="[1197,1235,1799,1825]" pageId="15" pageNumber="14">26)</emphasis>
Pupa, attached by venter of leaf, with shield comprising only exuviae of 5
<superScript id="7C329BD36601FFEFFCBA2210FCFEF8A1" attach="left" box="[814,828,1833,1847]" fontSize="6" pageId="15" pageNumber="14">th</superScript>
instar.
<emphasis id="B933EA896601FFEFFC052211FC75F8D4" bold="true" box="[913,951,1832,1858]" pageId="15" pageNumber="14">27)</emphasis>
Pupa with shield of exuviae I–V and feces.
<emphasis id="B933EA896601FFEFFF562273FF25F8F2" bold="true" box="[194,231,1866,1892]" pageId="15" pageNumber="14">28)</emphasis>
Teneral adult is straw colored.
<emphasis id="B933EA896601FFEFFDDE2273FD59F8F2" bold="true" box="[586,667,1866,1892]" pageId="15" pageNumber="14">29–30)</emphasis>
Mature adults are green, in copula.
<emphasis id="B933EA896601FFEFFBA02273FB9BF8F2" bold="true" box="[1076,1113,1866,1892]" pageId="15" pageNumber="14">31)</emphasis>
Older adult with black spots on elytra.
</paragraph>
</caption>
<paragraph id="8BF8369B661EFFF0FE9425EFFCEAFE2C" blockId="16.[192,1441,214,814]" pageId="16" pageNumber="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 epicuticular 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.</paragraph>
<paragraph id="8BF8369B661EFFF0FE9424FAFB64FDF5" blockId="16.[192,1441,214,814]" pageId="16" pageNumber="15">
A single larva can spend many days feeding on the same leaf. We observed and filmed the larvae of
<taxonomicName id="4C474D18661EFFF0FA1C24FAFEE9FE68" authorityName="Boheman" authorityYear="1854" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FA1C24FAFEE9FE68" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
cutting and eating trichomes (Video 1). Plants in the tribe
<taxonomicName id="4C474D18661EFFF0FC2224DDFB86FE68" box="[950,1092,484,510]" pageId="16" pageNumber="15" rank="tribe" tribe="Arctotidinae">Arctotidinae</taxonomicName>
have mostly non-glandular trichomes, although some glandular hairs can be present in certain organs (
<bibRefCitation id="EFD64B6A661EFFF0FC79273FFB65FDB6" author="Karis PO &amp; Funk VA &amp; McKenzie RJ &amp; Barker NP &amp; Chan R." box="[1005,1191,518,544]" pageId="16" pageNumber="15" pagination="385 - 410" refId="ref14186" refString="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 Taxonomy (IAPT); Vienna, Austria. 965 p." type="book chapter" year="2009">Karis et al. 2009</bibRefCitation>
). Glandular trichomes would be more deterrent to herbivory. Trichome-eating has not been observed for any
<taxonomicName id="4C474D18661EFFF0FB1F2711FAC5FDD4" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[1163,1287,552,578]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
. No trichome fragments appear in the shields we dissected (
<emphasis id="B933EA89661EFFF0FD2A2773FD0FFDF5" box="[702,717,586,611]" italics="true" pageId="16" pageNumber="15">n</emphasis>
= 4) so we assume trichomes are digested.
</paragraph>
<paragraph id="8BF8369B661EFFF0FE942755FD5CFCB8" blockId="16.[192,1441,214,814]" pageId="16" pageNumber="15">
In
<taxonomicName id="4C474D18661EFFF0FEB42755FE68FD13" authorityName="Boheman" authorityYear="1854" box="[288,426,619,646]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FEB42755FE68FD13" box="[288,426,619,646]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
, 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.
</paragraph>
<paragraph id="8BF8369B661EFFF0FF542607FDE3FB97" blockId="16.[191,1442,830,1262]" pageId="16" pageNumber="15">
<emphasis id="B933EA89661EFFF0FF542607FDF5FCCE" bold="true" box="[192,567,830,857]" pageId="16" pageNumber="15">
Natural enemies of
<taxonomicName id="4C474D18661EFFF0FE0B2607FDEDFCCF" authorityName="Boheman" authorityYear="1854" box="[415,559,830,857]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FE0B2607FDEDFCCF" bold="true" box="[415,559,830,857]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
.
</emphasis>
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 determined),
<taxonomicName id="4C474D18661EFFF0FE8A26BBFE67FC0A" authorityName="Linnaeus" authorityYear="1758" box="[286,421,898,924]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
caterpillars), but noted few interactions that might clarify which are competitors, predators, and parasites of
<taxonomicName id="4C474D18661EFFF0FEE3269DFDC3FC2B" authorityName="Boheman" authorityYear="1854" box="[375,513,931,958]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FEE3269DFDC3FC2B" box="[375,513,931,958]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
. We observed and filmed one
<taxonomicName id="4C474D18661EFFF0FCC3269DFC23FC2B" authorityName="Boheman" authorityYear="1854" box="[855,993,931,958]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FCC3269DFC23FC2B" box="[855,993,931,958]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
larva walking over a leaf and a smallersized aphid moved out of its way (Video 3). In another instance, a smaller mite moved out of the way of an approaching
<taxonomicName id="4C474D18661EFFF0FEC626D1FE1EFB97" authorityName="Boheman" authorityYear="1854" box="[338,476,999,1026]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FEC626D1FE1EFB97" box="[338,476,999,1026]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
larva.
</paragraph>
<paragraph id="8BF8369B661EFFF0FE942130FBC2FB78" blockId="16.[191,1442,830,1262]" pageId="16" pageNumber="15">
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
<taxonomicName id="4C474D18661EFFF0FCCC2174FC20FBF1" authorityName="Boheman" authorityYear="1854" box="[856,994,1101,1127]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FCCC2174FC20FBF1" box="[856,994,1101,1127]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
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.
</paragraph>
</subSubSection>
<subSubSection id="C35D6510661EFFF2FD472017FE25FBE3" lastPageId="18" lastPageNumber="17" pageId="16" pageNumber="15" type="discussion">
<paragraph id="8BF8369B661EFFF0FD472017FC4CFAC2" blockId="16.[723,910,1326,1364]" box="[723,910,1326,1364]" pageId="16" pageNumber="15">
<heading id="D0B081F7661EFFF0FD472017FC4CFAC2" box="[723,910,1326,1364]" centered="true" fontSize="16" level="1" pageId="16" pageNumber="15" reason="1">Discussion</heading>
</paragraph>
<paragraph id="8BF8369B661EFFF0FF542048FB3AF92D" blockId="16.[191,1442,1393,1927]" pageId="16" pageNumber="15">
The behaviors and life cycle of
<taxonomicName id="4C474D18661EFFF0FDA42048FD7CFA1D" authorityName="Boheman" authorityYear="1854" box="[560,702,1393,1419]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FDA42048FD7CFA1D" box="[560,702,1393,1419]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
was studied in detail and over many months (early spring-late autumn). We confirmed the choice of host plant,
<taxonomicName id="4C474D18661EFFF0FD4920AAFC9CFA3B" box="[733,862,1427,1453]" class="Magnoliopsida" family="Asteraceae" genus="Arctotheca" kingdom="Plantae" order="Asterales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="species" species="prostrata">
<emphasis id="B933EA89661EFFF0FD4920AAFC9CFA3B" box="[733,862,1427,1453]" italics="true" pageId="16" pageNumber="15">A. prostrata</emphasis>
</taxonomicName>
, in the indigenous habitat in
<collectingCountry id="F350760B661EFFF0FB3620AAFAF0FA3B" box="[1186,1330,1427,1453]" name="South Africa" pageId="16" pageNumber="15">South Africa</collectingCountry>
;
<taxonomicName id="4C474D18661EFFF0FAAF20AAFF2CFA59" authorityName="Boheman" authorityYear="1854" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FAAF20AAFF2CFA59" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
is now known on two
<emphasis id="B933EA89661EFFF0FE7D208CFD97FA59" box="[489,597,1461,1487]" italics="true" pageId="16" pageNumber="15">Arcotheca</emphasis>
species (
<bibRefCitation id="EFD64B6A661EFFF0FD2C208CFC20FA59" author="Heron H &amp; Borowiec L." box="[696,994,1461,1487]" pageId="16" pageNumber="15" pagination="625 - 658" refId="ref13876" refString="Heron H, Borowiec L. 1997. Host plants and feeding patterns of some South African tortoise beetles (Coleoptera: Chrysomelidae: cassidoid Hispinae). Genus 8: 625 - 658." type="journal article" year="1997">Heron and Borowiec 1997</bibRefCitation>
). In
<collectingCountry id="F350760B661EFFF0FB86208CFB61FA59" box="[1042,1187,1461,1487]" name="South Africa" pageId="16" pageNumber="15">South Africa</collectingCountry>
, hosts documented for
<taxonomicName id="4C474D18661EFFF0FF5420EFFEEAFA66" authorityName="Gyllenhal" authorityYear="1813" box="[192,296,1494,1520]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Cassidini">Cassidini</taxonomicName>
are in the
<taxonomicName id="4C474D18661EFFF0FE0E20EFFD8EFA66" box="[410,588,1494,1520]" class="Magnoliopsida" family="Amaranthaceae" kingdom="Plantae" order="Caryophyllales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Amaranthaceae</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF0FDC120EEFD0FFA67" box="[597,717,1495,1521]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Asteraceae</taxonomicName>
and more infrequent hosts are in
<taxonomicName id="4C474D18661EFFF0FBD520EFFB77FA66" box="[1089,1205,1494,1520]" class="Magnoliopsida" family="Aizoaceae" kingdom="Plantae" order="Caryophyllales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Aizoaceae</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF0FB2A20EFFAE7FA66" box="[1214,1317,1494,1520]" class="Magnoliopsida" family="Fabaceae" kingdom="Plantae" order="Fabales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Fabaceae</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF0FABB20EFFF30F985" class="Magnoliopsida" family="Polygonaceae" kingdom="Plantae" order="Caryophyllales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Polygonaceae</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF0FF6820C1FE5EF984" box="[252,412,1528,1554]" class="Magnoliopsida" family="Salvadoraceae" kingdom="Plantae" order="Brassicales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Salvadoraceae</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF0FE3120C1FDE0F984" box="[421,546,1528,1554]" class="Magnoliopsida" family="Solanaceae" kingdom="Plantae" order="Solanales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Solanaceae</taxonomicName>
, and
<taxonomicName id="4C474D18661EFFF0FDCF20C1FCC5F984" box="[603,775,1528,1554]" class="Magnoliopsida" family="Zygophyllaceae" kingdom="Plantae" order="Zygophyllales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="family">Zygophyllaceae</taxonomicName>
(
<bibRefCitation id="EFD64B6A661EFFF0FC8120C1FB46F984" author="Borowiec L &amp; Swietojanska J." box="[789,1156,1528,1558]" pageId="16" pageNumber="15" refId="ref11961" refString="Borowiec L, Swietojanska J. 2002 - 2022. World Catalog of Cassidinae, Wroclaw, Poland. Available at http: // www. cassidae. uni. wroc. pl / katalog % 20 internetowy / index. htm (Last accessed 3 February 2022)." type="url" year="2002">Borowiec and Świętojańska 2002</bibRefCitation>
–
<bibRefCitation id="EFD64B6A661EFFF0FB0720C1FB09F984" author="Borowiec L &amp; Swietojanska J." box="[1171,1227,1528,1554]" pageId="16" pageNumber="15" refId="ref12011" refString="Borowiec L, Swietojanska 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)." type="book" year="2022">2022</bibRefCitation>
). This is the second publication to record the feeding habit of
<taxonomicName id="4C474D18661EFFF0FD1C2323FCD2F9A2" authorityName="Boheman" authorityYear="1854" box="[648,784,1562,1588]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FD1C2323FCD2F9A2" box="[648,784,1562,1588]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
on an
<taxonomicName id="4C474D18661EFFF0FCCD2323FBE4F9A2" box="[857,1062,1562,1588]" pageId="16" pageNumber="15">
<emphasis id="B933EA89661EFFF0FCCD2323FC0CF9A2" box="[857,974,1562,1588]" italics="true" pageId="16" pageNumber="15">Arctotheca</emphasis>
species
</taxonomicName>
; a comprehensive survey of agents against
<taxonomicName id="4C474D18661EFFF0FE832305FE61F9C0" baseAuthorityName="Scott and Way" baseAuthorityYear="1990" box="[279,419,1596,1622]" class="Magnoliopsida" family="Asteraceae" genus="Arctotheca" kingdom="Plantae" order="Asterales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="species" species="calendula">
<emphasis id="B933EA89661EFFF0FE832305FE61F9C0" box="[279,419,1596,1622]" italics="true" pageId="16" pageNumber="15">A. calendula</emphasis>
</taxonomicName>
was carried out in
<collectingCountry id="F350760B661EFFF0FD122305FCD4F9C0" box="[646,790,1596,1622]" name="South Africa" pageId="16" pageNumber="15">South Africa</collectingCountry>
1986, 1987 and 1988, where
<taxonomicName id="4C474D18661EFFF0FBF22305FB31F9C0" authorityName="Boheman" authorityYear="1854" box="[1126,1267,1596,1622]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FBF22305FB31F9C0" box="[1126,1267,1596,1622]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
was noted as a potential agent although possibly not sufficiently specific (
<bibRefCitation id="EFD64B6A661EFFF0FCD82367FBF3F9EE" author="Scott JK &amp; Way MJ" box="[844,1073,1630,1656]" pageId="16" pageNumber="15" pagination="31 - 34" refId="ref15421" refString="Scott JK, Way MJ. 1990. A survey in South Africa for potential biological control agents against capeweed, Arctotheca calendula (L.) Levyns (Asteraceae). Plant Protection Quarterly 5 (1): 31 - 34." type="journal article" year="1990">Scott and Way 1990</bibRefCitation>
). Our study shows a strong association of
<taxonomicName id="4C474D18661EFFF0FEBA23B9FE75F90F" authorityName="Boheman" authorityYear="1854" box="[302,439,1663,1690]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FEBA23B9FE75F90F" box="[302,439,1663,1690]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
with
<taxonomicName id="4C474D18661EFFF0FE6223B9FDB5F90F" box="[502,631,1664,1690]" class="Magnoliopsida" family="Asteraceae" genus="Arctotheca" kingdom="Plantae" order="Asterales" pageId="16" pageNumber="15" phylum="Tracheophyta" rank="species" species="prostrata">
<emphasis id="B933EA89661EFFF0FE6223B9FDB5F90F" box="[502,631,1664,1690]" italics="true" pageId="16" pageNumber="15">A. prostrata</emphasis>
</taxonomicName>
. Further observations and testing of the specificity of
<taxonomicName id="4C474D18661EFFF0FB5E23B9FA96F90F" authorityName="Boheman" authorityYear="1854" box="[1226,1364,1663,1690]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FB5E23B9FA96F90F" box="[1226,1364,1663,1690]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
would be necessary to determine whether it could be considered a potential agent for biological control.
</paragraph>
<paragraph id="8BF8369B661EFFF1FE9423FAFB58FEC3" blockId="16.[191,1442,1393,1927]" lastBlockId="17.[191,1442,214,1927]" lastPageId="17" lastPageNumber="16" pageId="16" pageNumber="15">
Chrysomelid females provide several lines of physical and chemical protection of their eggs, including oothecal and excremental coverings (
<bibRefCitation id="EFD64B6A661EFFF0FDF423DCFD2EF969" author="Hilker M." box="[608,748,1765,1791]" pageId="16" pageNumber="15" pagination="263 - 276" refId="ref13911" refString="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." type="book chapter" year="1994">Hilker 1994</bibRefCitation>
). Eggs have been documented for 13
<taxonomicName id="4C474D18661EFFF0FB1B23DCFAF8F969" box="[1167,1338,1765,1791]" pageId="16" pageNumber="15">
<emphasis id="B933EA89661EFFF0FB1B23DCFB27F969" box="[1167,1253,1765,1791]" italics="true" pageId="16" pageNumber="15">Cassida</emphasis>
species
</taxonomicName>
in
<collectingCountry id="F350760B661EFFF0FACA23DCFEC5F8B7" name="South Africa" pageId="16" pageNumber="15">South Africa</collectingCountry>
and females deposit their eggs singly or in small groups to the undersides of their host leaves (often alongside a vein) in simple oothecae that lacks a stalk. The ootheca of
<taxonomicName id="4C474D18661EFFF0FC152210FBC9F8D4" authorityName="Boheman" authorityYear="1854" box="[897,1035,1832,1859]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FC152210FBC9F8D4" box="[897,1035,1832,1859]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
has a single layer enclosing the eggs, in contrast to the large complex multi-membrane oothecae with many eggs in
<emphasis id="B933EA89661EFFF0FBA72273FB12F8F2" box="[1075,1232,1866,1892]" italics="true" pageId="16" pageNumber="15">Conchylotenia</emphasis>
(
<bibRefCitation id="EFD64B6A661EFFF0FB4A2273FAA5F8F2" author="Heron HDC" box="[1246,1383,1866,1892]" pageId="16" pageNumber="15" pagination="565 - 580" refId="ref13699" refString="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." type="book chapter" year="1999">Heron 1999</bibRefCitation>
) and
<taxonomicName id="4C474D18661EFFF0FF552255FDA7F810" authority="(Muir and Sharp 1904)" baseAuthorityName="Muir and Sharp" baseAuthorityYear="1904" box="[193,613,1900,1926]" class="Insecta" family="Chrysomelidae" genus="Aspidimorpha" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661EFFF0FF552255FE9FF810" box="[193,349,1900,1926]" italics="true" pageId="16" pageNumber="15">Aspidimorpha</emphasis>
(
<bibRefCitation id="EFD64B6A661EFFF0FEFE2255FD9EF810" author="Muir F &amp; Sharp D." box="[362,604,1900,1926]" pageId="16" pageNumber="15" pagination="1 - 23" refId="ref14810" refString="Muir F, Sharp D. 1904. On the egg-cases and early stages of some Cassididae. Transactions of the Royal Entomological Society of London 1904: 1 - 23." type="journal article" year="1904">Muir and Sharp 1904</bibRefCitation>
)
</taxonomicName>
. In
<taxonomicName id="4C474D18661EFFF0FD192255FCD4F810" authorityName="Boheman" authorityYear="1854" box="[653,790,1900,1926]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661EFFF0FD192255FCD4F810" box="[653,790,1900,1926]" italics="true" pageId="16" pageNumber="15">C. sphaerula</emphasis>
</taxonomicName>
, ootheca have no fecal cover. Within the genus
<taxonomicName id="4C474D18661EFFF0FAB22255FAB9F810" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[1318,1403,1900,1926]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661EFFF0FAB22255FAB9F810" box="[1318,1403,1900,1926]" italics="true" pageId="16" pageNumber="15">Cassida</emphasis>
</taxonomicName>
,
<taxonomicName id="4C474D18661EFFF1FA1C2255FE1CFF67" authority="Boheman, 1854" authorityName="Boheman" authorityYear="1854" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" lastPageId="17" lastPageNumber="16" order="Coleoptera" pageId="16" pageNumber="15" phylum="Arthropoda" rank="species" species="coagulata">
<emphasis id="B933EA89661EFFF1FA1C2255FEE8FF66" italics="true" lastPageId="17" lastPageNumber="16" pageId="16" pageNumber="15">C. coagulata</emphasis>
<bibRefCitation id="EFD64B6A661FFFF1FEBB25EFFE1CFF67" author="Boheman CH" box="[303,478,214,241]" pageId="17" pageNumber="16" refId="ref11645" refString="Boheman CH. 1854. Monographia Cassididarum. Tomus secundus. Holmiae. 506 p." type="book" year="1854">Boheman, 1854</bibRefCitation>
</taxonomicName>
is a notable exception with a larger more elaborate oothecae generally attached to their host plant stem (
<taxonomicName id="4C474D18661FFFF1FEEE25CEFDEEFE87" box="[378,556,247,273]" class="Magnoliopsida" family="Amaranthaceae" kingdom="Plantae" order="Caryophyllales" pageId="17" pageNumber="16" phylum="Tracheophyta" rank="family">Amaranthaceae</taxonomicName>
hosts in this case, not
<taxonomicName id="4C474D18661FFFF1FCBE25C1FC67FE84" box="[810,933,248,274]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="17" pageNumber="16" phylum="Tracheophyta" rank="family">Asteraceae</taxonomicName>
; 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
<taxonomicName id="4C474D18661FFFF1FDF02402FD7BFEC3" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[612,697,315,341]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661FFFF1FDF02402FD7BFEC3" box="[612,697,315,341]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
</taxonomicName>
and suggest novel phylogenetic characters.
</paragraph>
<paragraph id="8BF8369B661FFFF1FE942464FB0FFD5E" blockId="17.[191,1442,214,1927]" pageId="17" pageNumber="16">
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
<taxonomicName id="4C474D18661FFFF1FD282499FCACFE2C" box="[700,878,416,442]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B933EA89661FFFF1FD282499FCD3FE2C" box="[700,785,416,442]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
species.
</taxonomicName>
Author Heron photographed similar patterns for
<taxonomicName id="4C474D18661FFFF1FF5624FBFD8CFE4A" authority="Boheman, 1862" authorityName="Boheman" authorityYear="1862" box="[194,590,450,476]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="guttipennis">
<emphasis id="B933EA89661FFFF1FF5624FBFE5AFE4A" box="[194,408,450,476]" italics="true" pageId="17" pageNumber="16">Cassida guttipennis</emphasis>
<bibRefCitation id="EFD64B6A661FFFF1FE0A24FBFD8CFE4A" author="Boheman CH" box="[414,590,450,476]" pageId="17" pageNumber="16" refId="ref11681" refString="Boheman CH. 1862. Monographia Cassididarum. Tomus quartus. Holmiae. 504 p." type="book" year="1862">Boheman, 1862</bibRefCitation>
</taxonomicName>
on the host,
<taxonomicName id="4C474D18661FFFF1FD7124FBFAEDFE4A" authority="(Harvey) Roessl" authorityName="Roessl" baseAuthorityName="Harvey" box="[741,1327,450,476]" class="Magnoliopsida" family="Asteraceae" genus="Berkheya" kingdom="Plantae" order="Asterales" pageId="17" pageNumber="16" phylum="Tracheophyta" rank="species" species="bipinnatifida">
<emphasis id="B933EA89661FFFF1FD7124FBFC22FE4A" box="[741,992,450,476]" italics="true" pageId="17" pageNumber="16">Berkheya bipinnatifida</emphasis>
(Harvey) Roessl (Asteraceae)
</taxonomicName>
, and
<taxonomicName id="4C474D18661FFFF1FAFB24FBFDB2FE68" authority="Spaeth, 1899" authorityName="Spaeth" authorityYear="1899" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="quatuordecimsignata">
<emphasis id="B933EA89661FFFF1FAFB24FBFE18FE6B" italics="true" pageId="17" pageNumber="16">Cassida quatuordecimsignata</emphasis>
<bibRefCitation id="EFD64B6A661FFFF1FE7524DAFDB2FE68" author="Spaeth F." box="[481,624,483,510]" pageId="17" pageNumber="16" pagination="213 - 221" refId="ref15554" refString="Spaeth F. 1899. Beschreibung einiger neuer Cassididen nebst synonymischen Bemerkungen. III. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien: 213 - 221." type="book chapter" year="1899">Spaeth, 1899</bibRefCitation>
</taxonomicName>
on the host,
<taxonomicName id="4C474D18661FFFF1FC9024DAFA5DFE68" authority="J. M. Wood and M. S. Evans" authorityName="J. M. Wood and M. S. Evans" box="[772,1439,483,510]" class="Magnoliopsida" family="Asteraceae" genus="Berkheya" kingdom="Plantae" order="Asterales" pageId="17" pageNumber="16" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis id="B933EA89661FFFF1FC9024DAFC1BFE6B" box="[772,985,483,509]" italics="true" pageId="17" pageNumber="16">Berkheya maritima</emphasis>
J.M. Wood and M.S. Evans (Asteraceae)
</taxonomicName>
(see
<bibRefCitation id="EFD64B6A661FFFF1FF67273FFD8EFD89" author="Heron H &amp; Borowiec L." box="[243,588,517,544]" pageId="17" pageNumber="16" pagination="625 - 658" refId="ref13876" refString="Heron H, Borowiec L. 1997. Host plants and feeding patterns of some South African tortoise beetles (Coleoptera: Chrysomelidae: cassidoid Hispinae). Genus 8: 625 - 658." type="journal article" year="1997">Heron and Borowiec 1997: 643</bibRefCitation>
, Fig. 19;
<bibRefCitation id="EFD64B6A661FFFF1FD39273FFCA5FD89" author="Heron HDC" box="[685,871,517,544]" pageId="17" pageNumber="16" pagination="133 - 149" refId="ref13822" refString="Heron HDC. 2011. Polymorphism in four tortoise beetles from Queensburgh, South Africa (Chrysomelidae: Cassidinae). Genus 22 (1): 133 - 149." type="journal article" year="2011">Heron 2011: 137</bibRefCitation>
,
<figureCitation id="137C2A1E661FFFF1FCE6273CFC73FD89" box="[882,945,517,543]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="17" pageNumber="16">Fig. 9</figureCitation>
;
<bibRefCitation id="EFD64B6A661FFFF1FC2F273FFBAAFD89" author="Heron HDC" box="[955,1128,517,544]" pageId="17" pageNumber="16" pagination="31 - 44" refId="ref13754" refString="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." type="journal article" year="2003">Heron 2003: 43</bibRefCitation>
, Fig. XXV) without discussing 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.
<bibRefCitation id="EFD64B6A661FFFF1FBCC2770FAE9FDF5" author="Bienkowski AO" box="[1112,1323,585,611]" pageId="17" pageNumber="16" pagination="1 - 10" refId="ref11618" refString="Bienkowski AO. 2010. Feeding behavior of leaf beetles (Coleoptera, Chrysomelidae). Entomological Review 90 (1): 1 - 10." type="journal article" year="2010">Bieńkowski (2010)</bibRefCitation>
described the more typical chewing pattern in two other
<taxonomicName id="4C474D18661FFFF1FD742753FC57FD12" box="[736,917,618,644]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B933EA89661FFFF1FD742753FCF7FD12" box="[736,821,618,644]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
species.
</taxonomicName>
These patterns suggest intrinsic intra-generic variations within
<taxonomicName id="4C474D18661FFFF1FE1627B5FE15FD30" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[386,471,652,678]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661FFFF1FE1627B5FE15FD30" box="[386,471,652,678]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
</taxonomicName>
. As more feeding patterns are recognized, novel hypotheses about their significance are emerging; for example, a masquerade strategy in some leaf beetles (
<bibRefCitation id="EFD64B6A661FFFF1FC382797FB03FD5E" author="Konstantinov AS &amp; Prathapan KD &amp; Vencl FV" box="[940,1217,686,712]" pageId="17" pageNumber="16" pagination="311 - 320" refId="ref14287" refString="Konstantinov AS, Prathapan KD, Vencl FV. 2018. Hiding in plain sight: leaf beetles (Chrysomelidae: Galerucinae) use feeding damage as a masquerade decoy. Biological Journal of the Linnean Society 123: 311 - 320." type="journal article" year="2018">Konstantinov et al. 2018</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8369B661FFFF1FE9427E9FA40FBE8" blockId="17.[191,1442,214,1927]" pageId="17" pageNumber="16">
The careful observation and filming of feeding in
<taxonomicName id="4C474D18661FFFF1FCBD27E9FC70FD7C" authorityName="Boheman" authorityYear="1854" box="[809,946,720,746]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661FFFF1FCBD27E9FC70FD7C" box="[809,946,720,746]" italics="true" pageId="17" pageNumber="16">C. sphaerula</emphasis>
</taxonomicName>
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
<figureCitation id="137C2A1E661FFFF1FF61262AFE82FCBB" box="[245,320,787,813]" captionStart="Figures 9–15" captionStartId="10.[192,279,1698,1724]" captionTargetBox="[371,1260,213,1668]" captionTargetId="figure-14@10.[370,1260,213,1668]" captionTargetPageId="10" captionText="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 exuviofecal 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." figureDoi="http://doi.org/10.5281/zenodo.7300738" httpUri="https://zenodo.org/record/7300738/files/figure.png" pageId="17" pageNumber="16">Fig. 13</figureCitation>
)—pushing trichomatous cuticle out of the way avoids energy and time costs to cut trichomes and clear a feeding path. We observed
<taxonomicName id="4C474D18661FFFF1FDAE260CFD00FCD9" authorityName="Boheman" authorityYear="1854" box="[570,706,821,847]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661FFFF1FDAE260CFD00FCD9" box="[570,706,821,847]" italics="true" pageId="17" pageNumber="16">C. sphaerula</emphasis>
</taxonomicName>
larvae consuming trichomes, which has not been reported for any
<taxonomicName id="4C474D18661FFFF1FF54266FFEABFCE6" authorityName="Latreille" authorityYear="1802" box="[192,361,854,880]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="family">Chrysomelidae</taxonomicName>
before. In Chrysomelinae chrysomelids, larvae of some
<taxonomicName id="4C474D18661FFFF1FC77266FFB6DFCE6" box="[995,1199,854,880]" pageId="17" pageNumber="16">
<emphasis id="B933EA89661FFFF1FC77266FFB98FCE6" box="[995,1114,854,880]" italics="true" pageId="17" pageNumber="16">Platyphora</emphasis>
species
</taxonomicName>
were observed to cut and throw trichomes backwards unto their fecal shields (
<bibRefCitation id="EFD64B6A661FFFF1FCAA2641FB41FC04" author="Bernardi N &amp; Scivittaro A." box="[830,1155,888,914]" pageId="17" pageNumber="16" pagination="531 - 534" refId="ref11518" refString="Bernardi N, Scivittaro A. 1991. Estagios imaturos de Platyphora zonata (Germar, 1824) (Coleoptera, Chrysomelidae, Chrysomelinae). Revista Brasileira de Zoologia 7 (4): 531 - 534." type="journal article" year="1991">Bernardi and Scivittaro 1991</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661FFFF1FB1B2641FAB6FC05" author="Flinte V &amp; Abejanella A &amp; Daccordi M &amp; Monteiro RF &amp; Macedo MV" box="[1167,1396,888,915]" pageId="17" pageNumber="16" pagination="5 - 22" refId="ref12969" refString="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." type="journal article" year="2017">Flinte et al. 2017: 15</bibRefCitation>
). In Campostomate chrysomelids, larvae trim and store trichomes into a section (“attic”) of the fecal case (
<bibRefCitation id="EFD64B6A661FFFF1FAB126A3FEF4FC40" author="Brown CG &amp; Funk DJ" pageId="17" pageNumber="16" pagination="711 - 725" refId="ref12128" refString="Brown CG, Funk DJ. 2005. Aspects of the natural history of Neochlamisus (Coleoptera: Chrysomelidae): fecal-case associated 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." type="journal article" year="2005">Brown and Funk 2005</bibRefCitation>
) or incorporate trichomes and feces to make the case wall (
<bibRefCitation id="EFD64B6A661FFFF1FC232685FB4AFC40" author="Chaboo CS &amp; Brown CG &amp; Funk DJ" box="[951,1160,956,982]" pageId="17" pageNumber="16" pagination="315 - 335" refId="ref12464" refString="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." type="journal article" year="2008">Chaboo et al. 2008</bibRefCitation>
). 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) (
<bibRefCitation id="EFD64B6A661FFFF1FD4D2118FC7DFBAD" author="Medeiros L &amp; Bolignon DS &amp; Moreira GRP" box="[729,959,1057,1083]" pageId="17" pageNumber="16" pagination="291 - 303" refId="ref14638" refString="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." type="book chapter" year="2004">Medeiros et al. 2004</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661FFFF1FC5D2118FAC6FBAA" author="Medeiros L &amp; Moreira GRP" box="[969,1284,1057,1084]" pageId="17" pageNumber="16" pagination="339 - 350" refId="ref14703" refString="Medeiros L, Moreira GRP. 2005. Larval feeding behavior of Gratiana spadicea (Klug) (Coleoptera: Chrysomelidae: Cassidinae) on its host plant, Solanum sisymbriifolium Lamarck (Solanaceae): interaction with trichomes. The Coleopterists Bulletin 59 (3): 339 - 350." type="journal article" year="2005">Medeiros and Moreira 2005</bibRefCitation>
). Author Heron’s observations of
<taxonomicName id="4C474D18661FFFF1FE34217AFDFAFBCA" authorityName="Boheman" authorityYear="1862" box="[416,568,1091,1117]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="guttipennis">
<emphasis id="B933EA89661FFFF1FE34217AFDFAFBCA" box="[416,568,1091,1117]" italics="true" pageId="17" pageNumber="16">C. guttipennis</emphasis>
</taxonomicName>
feeding revealed that more typical circular feeding scars without rolled cuticle margin is left on those plants with less dense pubescence, e.g.,
<taxonomicName id="4C474D18661FFFF1FCF8215DFABCFBE8" authority="(DC.) O. Hoffm." authorityName="O. Hoffm." baseAuthorityName="DC." box="[876,1406,1124,1150]" class="Magnoliopsida" family="Asteraceae" genus="Berkheya" kingdom="Plantae" order="Asterales" pageId="17" pageNumber="16" phylum="Tracheophyta" rank="species" species="speciosa">
<emphasis id="B933EA89661FFFF1FCF8215DFBECFBE8" box="[876,1070,1124,1150]" italics="true" pageId="17" pageNumber="16">Berkheya speciosa</emphasis>
(DC.) O. Hoffm. (Asteraceae)
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8369B661FFFF1FE9421BEFAF5FAB1" blockId="17.[191,1442,214,1927]" pageId="17" pageNumber="16">
In
<emphasis id="B933EA89661FFFF1FEB421BFFE71FB36" box="[288,435,1158,1184]" italics="true" pageId="17" pageNumber="16">
<taxonomicName id="4C474D18661FFFF1FEB421BFFE6DFB36" authorityName="Boheman" authorityYear="1854" box="[288,431,1158,1184]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="sphaerula">C. sphaerula</taxonomicName>
,
</emphasis>
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 variability, retaining either the instar V exuviae only (
<figureCitation id="137C2A1E661FFFF1FD7021F0FCF1FB75" box="[740,819,1225,1251]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="17" pageNumber="16">Fig. 26</figureCitation>
) or the entire structure of the larval stages (
<figureCitation id="137C2A1E661FFFF1FA8121F0FAA7FB72" box="[1301,1381,1225,1252]" captionStart="Figures 26–31" captionStartId="15.[192,279,1799,1825]" captionTargetBox="[219,1413,213,1769]" captionTargetId="figure-71@15.[219,1413,213,1769]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.7300748" httpUri="https://zenodo.org/record/7300748/files/figure.png" pageId="17" pageNumber="16">Fig. 27</figureCitation>
). It is unclear what the different benefits are of each shield form. Within the genus
<taxonomicName id="4C474D18661FFFF1FB9921D2FBA0FA93" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[1037,1122,1259,1285]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661FFFF1FB9921D2FBA0FA93" box="[1037,1122,1259,1285]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
</taxonomicName>
, shields vary in architecture, some with exuviae only, or with exuviae covered with fecal or with fecal filaments (
<bibRefCitation id="EFD64B6A661FFFF1FBC32036FAEEFAB1" author="Swietojanska J." box="[1111,1324,1293,1323]" pageId="17" pageNumber="16" refId="ref16063" refString="Swietojanska J. 2009. The immature stages of tortoise beetles with review of all described taxa (Coleoptera: Chrysomelidae: Cassidinae). Polish Taxonomical Monographs 16: 157 p." type="book" year="2009">Świętojańska 2009</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8369B661FFFF1FE942016FD97F90C" blockId="17.[191,1442,214,1927]" pageId="17" pageNumber="16">
Life history data can provide a great deal of comparative information to support species concepts and evolutionary relationships. Some of our findings are relevant to character hypotheses presented in two phylogenetic analyses of
<taxonomicName id="4C474D18661FFFF1FED0204BFE7DFA1A" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[324,447,1394,1420]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
,
<bibRefCitation id="EFD64B6A661FFFF1FE5F204BFD4AFA1B" author="Borowiec L." box="[459,648,1394,1421]" pageId="17" pageNumber="16" pagination="541 - 558" refId="ref11773" refString="Borowiec L. 1995. Tribal classification of the cassidoid Hispinae (Coleoptera: Chrysomelidae). p. 541 - 558. In: Pakaluk J, Slipinski SA (eds.). Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80 th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN; Warszawa, Poland. 1092 p." type="book chapter" year="1995">Borowiec (1995)</bibRefCitation>
, particularly his characters 15–19, and Chaboo (2007; 16 larval characters). Our findings also suggests new characters and new states to expand López-Pérez et al.’s (2018) dataset for the phylogeny of
<taxonomicName id="4C474D18661FFFF1FE3B208FFDD8FA46" authorityName="Gyllenhal" authorityYear="1813" box="[431,538,1462,1488]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Cassidini">Cassidini</taxonomicName>
. The similarity of feeding pattern in
<taxonomicName id="4C474D18661FFFF1FC2E208FFB91FA59" authorityName="Boheman" authorityYear="1862" box="[954,1107,1462,1488]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="guttipennis">
<emphasis id="B933EA89661FFFF1FC2E208FFB91FA59" box="[954,1107,1462,1488]" italics="true" pageId="17" pageNumber="16">C. guttipennis</emphasis>
</taxonomicName>
,
<emphasis id="B933EA89661FFFF1FBF5208FFAACFA46" box="[1121,1390,1462,1488]" italics="true" pageId="17" pageNumber="16">
<taxonomicName id="4C474D18661FFFF1FBF5208FFAA8FA46" authorityName="Spaeth" authorityYear="1899" box="[1121,1386,1462,1488]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="quatuordecimsignata">C. quatuordecimsignata</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C474D18661FFFF1FF5420E1FE8EFA67" authorityName="Boheman" authorityYear="1854" box="[192,332,1495,1522]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661FFFF1FF5420E1FE8EFA67" box="[192,332,1495,1522]" italics="true" pageId="17" pageNumber="16">C. sphaerula</emphasis>
</taxonomicName>
may be clues to shared behavior and morphology, possibly defining a sub-group within
<taxonomicName id="4C474D18661FFFF1FAD020EEFA5BFA67" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[1348,1433,1495,1521]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661FFFF1FAD020EEFA5BFA67" box="[1348,1433,1495,1521]" italics="true" pageId="17" pageNumber="16">Cassida</emphasis>
</taxonomicName>
. The production and relative simplicity of the ootheca in
<taxonomicName id="4C474D18661FFFF1FCAA20C0FC0BF985" authorityName="Boheman" authorityYear="1854" box="[830,969,1529,1555]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="sphaerula">
<emphasis id="B933EA89661FFFF1FCAA20C0FC0BF985" box="[830,969,1529,1555]" italics="true" pageId="17" pageNumber="16">C. sphaerula</emphasis>
</taxonomicName>
compared to the more complex one in
<taxonomicName id="4C474D18661FFFF1FA1D20C0FEE8F9A3" authorityName="Boheman" authorityYear="1854" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="16" phylum="Arthropoda" rank="species" species="coagulata">
<emphasis id="B933EA89661FFFF1FA1D20C0FEE8F9A3" italics="true" pageId="17" pageNumber="16">C. coagulata</emphasis>
</taxonomicName>
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 comparative study and evolutionary analysis.
</paragraph>
<paragraph id="8BF8369B661FFFF2FE94239BFB1AFE84" blockId="17.[191,1442,214,1927]" lastBlockId="18.[191,1441,214,1141]" lastPageId="18" lastPageNumber="17" pageId="17" pageNumber="16">
The exuvio-fecal shield that diagnoses the eight derived tribes of
<taxonomicName id="4C474D18661FFFF1FC5E239BFB81F92A" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[970,1091,1698,1724]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
is a unique morpho-behavioral complex (Chaboo 2007), an example of an extended phenotype, like a bird’s nest (
<bibRefCitation id="EFD64B6A661FFFF1FBAA23FAFB20F948" author="Dawkins R." box="[1086,1250,1731,1758]" pageId="17" pageNumber="16" refId="ref12757" refString="Dawkins R. 1989. The extended phenotype. Oxford University Press; Oxford, UK. 440 p." type="book" year="1989">Dawkins 1989</bibRefCitation>
). This is a significant macroevolutionary event in the evolution of
<taxonomicName id="4C474D18661FFFF1FD7323DCFCA0F969" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[743,866,1765,1791]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
, 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 excrementos” (=barely covered by excrement) so
<emphasis id="B933EA89661FFFF1FD212210FD1BF8D4" box="[693,729,1833,1858]" italics="true" pageId="17" pageNumber="16">not</emphasis>
carrying a shield (
<bibRefCitation id="EFD64B6A661FFFF1FC382210FB99F8D5" author="Bondar G." box="[940,1115,1833,1859]" pageId="17" pageNumber="16" refId="ref11697" refString="Bondar G. 1940. Insectos nocivos e molestias do Coqueiro (Cocos nucifera) no Brasil. Boletim Instituto Central de Fomento Economica da Bahia 8: 160 p." type="book" year="1940">Bondar 1940: 1</bibRefCitation>
02),
<taxonomicName id="4C474D18661FFFF1FB032210FA97F8D5" authorityName="Monros &amp; Viana" authorityYear="1951" box="[1175,1365,1833,1859]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="tribe" tribe="Hemisphaerotini">Hemisphaerotini</taxonomicName>
larvae have caudal processes and a unique “bird-nest” shield architecture (
<bibRefCitation id="EFD64B6A661FFFF1FC382273FB16F8F2" author="Chaboo CS &amp; Nguyen T." box="[940,1236,1866,1893]" pageId="17" pageNumber="16" pagination="171 - 184" refId="ref12558" refString="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, Schmitt M (eds.). New contributions in biology of the Chrysomelidae. Kugler Publications; The Hague, Netherlands. 803 p." type="book chapter" year="2004">Chaboo and Nguyen 2004</bibRefCitation>
), and Spilophorini larvae have caudal processes and an exuviae-only shield (
<bibRefCitation id="EFD64B6A661FFFF1FCD12255FBDDF810" author="Nishida K &amp; Ferrufino-Acosta L &amp; Chaboo CS" box="[837,1055,1900,1927]" pageId="17" pageNumber="16" pagination="263 - 267" refId="ref15020" refString="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." type="journal article" year="2020">Nishida et al. 2020</bibRefCitation>
). In contrast, the mining larvae of
<taxonomicName id="4C474D18661CFFF2FF5425EFFEAEFF66" authorityName="Hincks" authorityYear="1952" box="[192,364,214,240]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="3" phylum="Arthropoda" rank="tribe" tribe="Notosacanthini">Notosacanthini</taxonomicName>
lack caudal processes and lack shields (
<bibRefCitation id="EFD64B6A661CFFF2FC8925EFFBC2FF66" author="Monteith GB &amp; Sandoval Gomez VE &amp; Chaboo CS" box="[797,1024,214,240]" pageId="18" pageNumber="17" pagination="329 - 354" refId="ref14755" refString="Monteith GB, Sandoval Gomez VE, Chaboo CS. 2021. Natural history of the Australian tortoise beetle Notosacantha dorsalis (Waterhouse, 1877) (Coleoptera: Chrysomelidae: Cassidinae: Notosacanthini) with summary of the genus in Australia. The Australian Entomologist 48 (4): 329 - 354." type="journal article" year="2021">Monteith et al. 2021</bibRefCitation>
). Also, remarkable is the independent origin of shield retention in the distantly related ‘hispine’,
<taxonomicName id="4C474D18661CFFF2FCD525C1FB16FE84" authority="Baly, 1858" authorityName="Baly" authorityYear="1858" box="[833,1236,248,275]" class="Insecta" family="Chrysomelidae" genus="Oediopalpa" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661CFFF2FCD525C1FC7DFE84" box="[833,959,248,274]" italics="true" pageId="18" pageNumber="17">Oediopalpa</emphasis>
<bibRefCitation id="EFD64B6A661CFFF2FC5125C1FBF5FE85" author="Baly JS" box="[965,1079,248,275]" pageId="18" pageNumber="17" refId="ref11457" refString="Baly JS. 1858. Catalogue of Hispidae in the collection of the British Museum. Printed by order of the Trustees; London. 172 p." type="book" year="1858">Baly, 1858</bibRefCitation>
(
<bibRefCitation id="EFD64B6A661CFFF2FBD125C1FB0EFE84" author="Bruch C." box="[1093,1228,248,274]" pageId="18" pageNumber="17" pagination="207 - 219" refId="ref12184" refString="Bruch C. 1906. Metamorfosis y biologia de coleopteros argentines. II. Agasicles vittata Jac., Plectonycha correntina Lac., Amplipalpa negligens Weise. Revista del Museo La Plata 12: 207 - 219." type="journal article" year="1906">Bruch 1906</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8369B661CFFF2FE942422FE52FCAB" blockId="18.[191,1441,214,1141]" pageId="18" pageNumber="17">
The tortoise beetle shield has been considered as a protection and a defense.
<bibRefCitation id="EFD64B6A661CFFF2FBDB2422FAC4FEA3" author="Reaumur RAF" box="[1103,1286,283,309]" pageId="18" pageNumber="17" refId="ref15318" refString="Reaumur RAF de. 1737. Memoirs pour servir a l'histoire des insects. Tome 3. Paris Imprimie Royale; Paris. 47 pl. + 532 p." type="book" year="1737">Réaumur (1737)</bibRefCitation>
hypothesized that it protected against sun and flies.
<bibRefCitation id="EFD64B6A661CFFF2FDCA2407FD2DFECE" author="Weise J." box="[606,751,318,344]" pageId="18" pageNumber="17" refId="ref16516" refString="Weise J. 1893. Naturgeschichte der Insecten Deutschland. Erste Abtheilung Coleoptera. Sechster Band. Nicolaische Verlags- Buchhandlung R. Stricker; Berlin. xiv + 1161 p." type="book" year="1893">Weise (1893)</bibRefCitation>
hypothesized its function as defense against desiccation. More observations led to the hypothesis that shields are a defense against enemies and used cheaply-available defecation products and exuviae and perhaps even chemicals in exocrine glands of those exuviae (
<bibRefCitation id="EFD64B6A661CFFF2FB7724BAFA56FE0B" author="Olmstead KL" box="[1251,1428,387,413]" pageId="18" pageNumber="17" pagination="311 - 318" refId="ref15159" refString="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." type="book chapter" year="1994">Olmstead 1994</bibRefCitation>
). Mechanical defense against predators has been tested experimentally, with support by several researchers (
<bibRefCitation id="EFD64B6A661CFFF2FAC3249FFEF7FE74" author="Eisner T &amp; van Tassel E &amp; Carrel JE" pageId="18" pageNumber="17" pagination="1471 - 1473" refId="ref12875" refString="Eisner T, van Tassel E, Carrel JE. 1967. Defensive use of a ' fecal shield' by a beetle larva. Science 158: 1471 - 1473." type="journal article" year="1967">Eisner et al. 1967</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661CFFF2FED524F1FDB6FE75" author="Olmstead K &amp; Denno RF" box="[321,628,456,483]" pageId="18" pageNumber="17" pagination="1394 - 1405" refId="ref15212" refString="Olmstead K, Denno RF. 1993. Effectiveness of tortoise beetle larval shields against different predator species. Ecology 74: 1394 - 1405." type="journal article" year="1993">Olmstead and Denno 1993</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661CFFF2FD1424F1FC45FE75" author="Eisner T &amp; Eisner M." box="[640,903,456,483]" pageId="18" pageNumber="17" pagination="2632 - 2636" refId="ref12839" refString="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." type="journal article" year="2000">Eisner and Eisner 2000</bibRefCitation>
) but contradicted by others (
<bibRefCitation id="EFD64B6A661CFFF2FB4424F1FF3BFD93" author="Muller C &amp; Hilker M." pageId="18" pageNumber="17" pagination="166 - 172" refId="ref14870" refString="Muller C, Hilker M. 1999. Unexpected reactions of a generalist predator towards defensive devices of cassidine larva (Coleoptera: Chrysomelidae). Oecologia 118: 166 - 172." type="journal article" year="1999">Müller and Hilker 1999</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661CFFF2FE9724D2FDA5FD93" author="Nogueira-de-Sa F &amp; Trigo JR" box="[259,615,491,517]" pageId="18" pageNumber="17" pagination="203 - 206" refId="ref15080" refString="Nogueira-de-Sa F, Trigo JR. 2002. Do fecal shields provide physical protection to larvae of the tortoise beetles Plagiometriona flavescens and Stolas chalybea against natural enemies? Entomologia Experimentalis et Applicata 104: 203 - 206." type="journal article" year="2002">Nogueira-de-Sá and Trigo 2002</bibRefCitation>
). Further studies with
<taxonomicName id="4C474D18661CFFF2FCC824D2FC73FD93" authorityName="Adam &amp; Campos &amp; Heron &amp; Staines &amp; Westerduijn &amp; Chaboo" authorityYear="2022" box="[860,945,491,517]" class="Insecta" family="Chrysomelidae" genus="Cassida" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="B933EA89661CFFF2FCC824D2FC73FD93" box="[860,945,491,517]" italics="true" pageId="18" pageNumber="17">Cassida</emphasis>
</taxonomicName>
larvae point to more selective shield defense to certain enemies:
<bibRefCitation id="EFD64B6A661CFFF2FE0E2737FD02FDBE" author="Schenk D &amp; Bacher S." box="[410,704,526,552]" pageId="18" pageNumber="17" pagination="524 - 531" refId="ref15348" refString="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." type="journal article" year="2002">Schenk and Bacher (2002)</bibRefCitation>
showed shields were effective against vespid predators only, while
<bibRefCitation id="EFD64B6A661CFFF2FF542709FE19FDDD" author="Bacher S &amp; Luder S." box="[192,475,560,587]" pageId="18" pageNumber="17" pagination="263 - 272" refId="ref11428" refString="Bacher S, Luder S. 2005. Picky predators and the function of the faecal shield of a cassidine larva. Ecology 19: 263 - 272." type="journal article" year="2005">Bacher and Luder (2005)</bibRefCitation>
showed they were effective against parasitoids only and offer some protection against desiccation and wind, but not so against abiotic factors of UV-radiation.
<bibRefCitation id="EFD64B6A661CFFF2FC7D276AFB46FDFB" author="Muller C." box="[1001,1156,595,621]" pageId="18" pageNumber="17" pagination="191 - 198" refId="ref14842" refString="Muller C. 2002. Variation in the effectiveness of abdominal shields of cassidine larvae against predators. Entomologia Experimentalis et Applicata 102: 191 - 198." type="journal article" year="2002">Müller (2002)</bibRefCitation>
also found variable effectiveness of shields to deter different predators. Chemical defense via enteric discharges in shields was proposed by
<bibRefCitation id="EFD64B6A661CFFF2FF7127A1FE13FD25" author="Pasteels JM &amp; Braekman JC &amp; Daloze D." box="[229,465,664,691]" pageId="18" pageNumber="17" pagination="233 - 252" refId="ref15268" refString="Pasteels JM, Braekman JC, Daloze D. 1988. Chemical defense in Chrysomelidae. p. 233 - 252. In: Jolivet PH, Cox ML, Petitpierre E (eds.). Biology of Chrysomelidae. Kluwer Academic Publishers; Dordrecht. 615 p." type="book chapter" year="1988">Pasteels et al. (1988)</bibRefCitation>
. Chemicals sequestered from host plants or by de novo synthesis can enhance shield defenses (
<bibRefCitation id="EFD64B6A661CFFF2FEB82782FE34FD43" author="Gomez NE &amp; Witte L &amp; Hartmann T." box="[300,502,699,726]" pageId="18" pageNumber="17" pagination="1007 - 1027" refId="ref13420" refString="Gomez 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." type="journal article" year="1999">Gómez et al. 1999</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661CFFF2FD942782FD75FD43" author="Vencl FV &amp; Schultz JC &amp; Mumma RC &amp; Morton TC" box="[512,695,699,726]" pageId="18" pageNumber="17" pagination="549 - 566" refId="ref16364" refString="Vencl FV, Schultz JC, Mumma RC, Morton TC. 1999. The shield defense of a larval tortoise beetle. Journal of Chemical Ecology 25: 549 - 566." type="journal article" year="1999">Vencl et al. 1999</bibRefCitation>
,
<bibRefCitation id="EFD64B6A661CFFF2FD552782FD3BFD40" author="Vencl FV &amp; Nogueira-de-Sa F &amp; Allen BJ &amp; Windsor DM &amp; Futuyma DJ" box="[705,761,699,726]" pageId="18" pageNumber="17" pagination="409 - 414" refId="ref16325" refString="Vencl FV, Nogueira-de-Sa F, Allen BJ, Windsor DM, Futuyma DJ. 2005. Dietary specialization influences the efficacy of larval tortoise beetle shield defenses. Oecologia 145 (3): 409 - 414." type="journal article" year="2005">2005</bibRefCitation>
,
<bibRefCitation id="EFD64B6A661CFFF2FC972782FCF9FD43" author="Vencl FV &amp; Gomez NE &amp; Ploss K &amp; Boland W." box="[771,827,699,725]" pageId="18" pageNumber="17" pagination="281 - 288" refId="ref16283" refString="Vencl FV, Gomez 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." type="journal article" year="2009">2009</bibRefCitation>
,
<bibRefCitation id="EFD64B6A661CFFF2FCD12782FCBFFD43" author="Vencl FV &amp; Trillo PA &amp; Geeta R." box="[837,893,699,725]" pageId="18" pageNumber="17" pagination="227 - 239" refId="ref16397" refString="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." type="journal article" year="2011">2011</bibRefCitation>
;
<bibRefCitation id="EFD64B6A661CFFF2FC132782FB25FD43" author="Nogueira-de-Sa F &amp; Trigo JR" box="[903,1255,699,725]" pageId="18" pageNumber="17" pagination="203 - 206" refId="ref15080" refString="Nogueira-de-Sa F, Trigo JR. 2002. Do fecal shields provide physical protection to larvae of the tortoise beetles Plagiometriona flavescens and Stolas chalybea against natural enemies? Entomologia Experimentalis et Applicata 104: 203 - 206." type="journal article" year="2002">Nogueira-de-Sá and Trigo 2002</bibRefCitation>
, 2005), however, chemicals were also found to have no impact on larval survival (
<bibRefCitation id="EFD64B6A661CFFF2FC0827E7FB42FD6E" author="Bottcher A &amp; Zolin JP &amp; Nogueira-de-Sa F &amp; Trigo JR" box="[924,1152,734,760]" pageId="18" pageNumber="17" pagination="63 - 66" refId="ref12055" refString="Bottcher A, Zolin JP, Nogueira-de-Sa 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." type="journal article" year="2009">Bottcher et al. 2009</bibRefCitation>
). This succession of ideas and continuing testing are crucial to illuminating the origin, function (i.e., cost benefit analyses), and diversity of fecal architectures.
</paragraph>
<paragraph id="8BF8369B661CFFF2FE94267FFE25FBE3" blockId="18.[191,1441,214,1141]" pageId="18" pageNumber="17">
Phylogenetic studies in
<taxonomicName id="4C474D18661CFFF2FD9C267FFD43FCF6" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[520,641,838,864]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
have relied largely on adult characters. In the past, a few characters and states of juvenile stages have been proposed:
<bibRefCitation id="EFD64B6A661CFFF2FD2E2651FCB6FC15" author="Borowiec L." box="[698,884,872,899]" pageId="18" pageNumber="17" pagination="541 - 558" refId="ref11773" refString="Borowiec L. 1995. Tribal classification of the cassidoid Hispinae (Coleoptera: Chrysomelidae). p. 541 - 558. In: Pakaluk J, Slipinski SA (eds.). Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80 th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN; Warszawa, Poland. 1092 p." type="book chapter" year="1995">Borowiec (1995)</bibRefCitation>
tested four characters of larvae for his phylogeny of
<taxonomicName id="4C474D18661CFFF2FF7526B2FE9EFC33" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" box="[225,348,907,933]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
, Chaboo’s (2007) study included 20 from juveniles,
<bibRefCitation id="EFD64B6A661CFFF2FC2D26B2FB1BFC33" author="Lopez-Perez S &amp; Zaragoza-Caballero S &amp; Ochoterena H &amp; Morrone JJ" box="[953,1241,907,934]" pageId="18" pageNumber="17" pagination="372 - 386" refId="ref14472" refString="Lopez-Perez S, Zaragoza-Caballero S, Ochoterena H, Morrone JJ. 2018. A phylogenetic study of the worldwide tribe Cassidini Gyllenhal, 1813 (Coleoptera: Chrysomelidae: Cassidinae) based on morphological data. Systematic Entomology 43: 372 - 386." type="journal article" year="2018">López-Pérez et al. (2018)</bibRefCitation>
tested one larval character. Going forward, we anticipate more studies like
<bibRefCitation id="EFD64B6A661CFFF2FCD62697FB9AFC5E" author="Lopez-Perez S &amp; Rodriguez-Miron GM &amp; Chaboo CS" box="[834,1112,942,968]" pageId="18" pageNumber="17" pagination="107 - 119" refId="ref14430" refString="Lopez-Perez S, Rodriguez-Miron GM, Chaboo CS. 2021. Pupal morphology of Physonota humilis Boheman, 1856 and Physonota stigmatilis Boheman 1854 (Chrysomelidae: Cassidinae: Ischyrosonychini). Zootaxa 5027 (1): 107 - 119." type="journal article" year="2021">López-Pérez et al. (2021)</bibRefCitation>
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
<taxonomicName id="4C474D18661CFFF2FAF926CAFECDFBA6" baseAuthorityName="Gyllenhal" baseAuthorityYear="1813" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="2" phylum="Arthropoda" rank="subFamily" subFamily="Cassidinae">Cassidinae</taxonomicName>
(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.
</paragraph>
</subSubSection>
</treatment>
</document>