ZooKeys 336: 47–59, doi: 10.3897/zookeys.336.5435
First record of leaf-hole shelters used and modified by leaf beetles (Coleoptera, Chrysomelidae), with descriptions of two new Orthaltica Crotch species from southern India
Kaniyarikkal Divakaran Prathapan 1,†, Alexander S. Konstantinov 2,‡, K. M. Shameem 1,§, A. P. Balan 1,|
1 Department of Entomology, Kerala Agricultural University, Vellayani P.O., Trivandrum -695 522, Kerala, INDIA
2 Systematic Entomology Laboratory, ARS, USDA, c/o National Museum of Natural History, Smithsonian Institution P.O. Box 37012, MRC-168, Washington, DC 20013-7012, U.S.A.

Corresponding author: Alexander Konstantinov (alex.konstantinov@ars.usda.gov)

Academic editor: C. Majka

received 29 April 2013 | accepted 14 August 2013 | Published 27 September 2013


(C) 2013 Kaniyarikkal Divakaran Prathapan. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


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Citation: Prathapan KD, Konstantinov AS, Shameem KM, Balan AP (2013) First record of leaf-hole shelters used and modified by leaf beetles (Coleoptera, Chrysomelidae), with descriptions of two new Orthaltica Crotch species from southern India. ZooKeys 336: 47–59. doi: 10.3897/zookeys.336.5435

Abstract

Behavioural novelties observed in adult leaf beetles of two new Orthaltica Crotch species include: 1) the use of low cost leaf-hole shelters, either in pre-formed holes produced by larger beetles that fed on the same leaf, or artificially created holes as part of an experiment; and 2) the use of faeces to partition the hole. Two new southern Indian species of the genus Orthaltica are described and illustrated: Orthaltica syzygium and Orthaltica terminalia. Host plants are identified for both species. A key to the Indian species of Orthaltica is provided.

Keywords

Leaf-hole shelter, faeces, leaf beetles, new species, host plant

Introduction

Animal architecture reflects the biology of its builder. Three broad categories of animal constructions have been recognized: homes, traps and displays (Hansell 2005; 2007). Homes or shelters protect their builders from external physical and possible biological hostilities. They provide protection by means of their architecture, effecting avoidance of detection and the prevention of invasion after detection has occurred (Hansell 2005).

In architecture birds, spiders, termites, ants, bees and wasps are prodigies. Beetles exhibit the most numerous radiation within the animalia, but there are very few architects amongst them. In leaf beetles (Chrysomelidae) (about 50, 000 species) larval defensive structures, built of faeces and exuvial skin, are known (Chaboo et al. 2007, Chaboo et al. 2008, Prathapan and Chaboo 2011). Larvae of several leaf beetles are known to use faeces for defense, and nearly 20% of the described chrysomelid species have a form of faecal covering or faeces-associated structure at some point in their lifecycle (Vencl et al. 1999; Weiss 2006).

Two genera in the Cassidinae, namely Imatidium Fabricius (Gilbert et al. 2001) and Leptispa Baly (Maulik 1919; Voronova and Zaitsev 1982; Prathapan et al. 2009), have the only leaf beetles known to build leaf shelters. Larvae of nearly all of about 4500 species of Camptosomata (Cryptocephalinae and Lamprosomatinae) build cases from their own faeces (Chaboo et al. 2008).

Builders invest considerable resources and time making constructions. Hence, natural selection favours building behaviour which reduces the cost of building, whilst maximizing the benefits it offers. Nesting in existing cavities produced by primary cavity nesters, as in woodpeckers, is a common method of cost reduction used by birds. Low cost shelters of Leptispa pygmaea Baly larvae are formed by feeding alone—no cutting, bending or secretion of silk or glue is required (Prathapan et al. 2009). A type of low cost shelter, named a leaf-hole shelter, is here reported for the first time for two species of Orthaltica Crotch, a genus of minute leaf beetles (Figs 1, 4–8).

Figures 1–8.

Orthaltica species and their low cost leaf-hole shelters. 1 Leaf-hole shelter of Orthaltica terminalia on Terminalia paniculata. Note the feeding trenches radiating from the leaf-hole shelter 2 Feeding trenches of non-shelter forming species on an unidentified plant from the Combretaceae. Note the unoccupied leaf-hole nearby 3 Eumolpine beetle (Basilepta sp.) feeding holes on the abaxial side of the leaf of Syzygium caryophyllatum 4 Orthaltica syzygium occupying the leaf-hole made by the eumolpine beetle in Fig. 3 5 Leaf-hole shelter of Orthaltica in Fig. 3 as seen on the second day, viewed from adaxial side of leaf. Note the feeding trenches radiating from the leaf-hole 6 Triangular-shaped artificial leaf-holes on Syzygium travancoricum used as shelter by Orthaltica syzygium - note feeding trenches radiating from holes 7 Single occupancy - note the first partition 8 Multiple occupancy with multiple partitions.

The genus Orthaltica is distributed in the Afrotropical, Australian, Nearctic and Oriental Regions (Seeno and Wilcox 1982; Reid 1990) and contains 44 named species, of which the majority (32 species) are distributed in the Oriental Region. The taxonomy of Orthaltica is complex, currently including eight generic synonyms. Crioceris copalina (F.), the type species, occurs in the Nearctic Region. In India the genus is represented by at least ten species level taxa, of which only six have been formally named and described: Orthaltica assamensis (Scherer, 1971), Orthaltica bengalensis (Basu & Sengupta, 1978), Orthaltica coomani (Laboissiére, 1933 in Basu and Sengupta 1978), Orthaltica dakshina (Basu & Sengupta, 1978), Orthaltica minuta indica (Medvedev, 1998) and Orthaltica purba (Basu & Sengupta, 1985). We here describe two species that utilise leaf-hole shelters. However, a revisionary study of the genus is necessary, at least in Asia, to clarify both species boundaries and the relationships between species groups currently placed in Orthaltica.

Material and methods

Five species of Orthaltica were collected from various localities in southern India, along with their host plants, forming part of our long term study of the leaf beetles of India. Species from this genus are known to feed on plants from the Combretaceae, Melastomataceae and Myrtaceae. Leaf-hole sheltering, described below, was observed in two species feeding on Syzygium (Myrtaceae) and Terminalia (Combretaceae) species respectively. The Orthaltica species feeding on Syzygium (Gaertn.) is here described as Orthaltica syzygium new species, and the species on Terminalia L. as Orthaltica terminalia new species. Orthaltica syzygium was collected from various localities in the states of Karnataka (Kottigehara, Kudremukha) and Kerala (Arippa, Kumarakom, Vattavada) on Syzygium cumini (L.) Skeels, Syzygium caryophyllatum Alston and Syzygium travancoricum Gamble. Orthalitca terminalia was also collected in the states of Karnataka (Kudremukha) and Kerala (Calicut University Campus, Kallar, Trichur), but on Terminalia cuneata Roth and Terminalia paniculata Roth. The leaf-hole shelters on Terminalia paniculata were initially formed by a Tricliona sp. (Eumolpinae). Natural populations of all the Orthaltica species were carefully observed in the field for feeding and shelter seeking behaviour. Leaf-hole shelters of Orthaltica syzygium were observed at the Kudremukha National Park on Syzygium caryophyllatum, but were initially formed by a Basilepta sp. (Eumolpinae) (Fig. 3). Other leaf feeding beetles, but particularly Eumolpinae, were observed for their feeding activity on the hosts of Orthaltica. Artificial leaf holes were presented to the beetles on leaf laminae of Syzygium travancoricum. Holes were made with a punch, generally used for making the card points on which small leaf beetles are mounted. The punch created elongate, triangular holes of 7 mm in length and a width of 2 mm at the base. About 60 leaves were provided to adults of Orthaltica syzygium on 20th April, 2013, each with six holes punched on the lamina. Occupation of the holes by these beetles was observed on 20th May, 2013 and 20th August, 2013 and could easily be determined by checking for the presence or absence of feeding trenches radiating from the hole (Fig. 6).

Descriptive terminology follows Konstantinov (1998). Holotypes of the new species are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (USNM). Paratypes will be deposited in the National Pusa Collection, Indian Agricultural Research Institute, New Delhi, India (NPC); University of Agricultural Sciences, Bangalore, India (UASB); Natural History Museum, London, United Kingdom (BMNH); and in the personal collection of the first author (PKDC). Plant vouchers with accession numbers are deposited in the Calicut University Herbarium, Calicut, India. Different labels on specimens are denoted by numbers and separated by “;”.

Results
Orthaltica syzygium Prathapan & Konstantinov, sp. n.

http://zoobank.org/B1CFE822-AA5E-498C-8000-F64A0A5791F2

http://species-id.net/wiki/Orthaltica_syzygium

Figs 415
Description.

Body length 1.30–1.60 mm, width 0.67–0.81 mm. Body dark brown (Figs 9, 10). Basal antennomeres and tarsi paler and yellowish-brown. Vertex with 8 long and more than 10 short setae. Supracallinal sulci straight; orbital sulci short, poorly developed; and suprafrontal and suprantennal sulci poorly developed. Antennal calli narrow and elongate (Fig. 11), relatively widely connected, with two large setiferous pores. Frontal ridge parallel-sided, ending before reaching anterofrontal ridge. Anterofrontal ridge of uniform height, but sloping abruptly towards clypeus. Labrum as wide as distance between outer edges of antennal sockets. Setiferous pores on dorsal surface of labrum small, not possible to count. Antennomere 3 slightly shorter than 4 and 5 separately.

Figures 9–15.

Orthaltica syzygium Prathapan & Konstantinov, sp. n. 9 Dorsal habitus 10 Lateral habitus 11 Head frontal view 12 Aedeagus, ventral and lateral view 13 Vaginal palpi 14 Spermatheca 15 Tignum.

Pronotum widening towards apex and narrowing abruptly after reaching anterolateral callosity. Lateral margins slightly uneven with a prominent submedial denticle, and two long setae (Fig. 10). Antebasal transverse impression distinct (Fig. 9). Pronotal surface densely covered with large punctures, their diameter larger than the distance between punctures.

Elytra densely and evenly pubescent (Figs 9, 10), with erect setae on interstices. Elytral punctures arranged in striae located in relatively deep grooves. Humeral calli distinct. Pro- and middle tibiae lacking spurs. Claws appendiculate.

Apex of aedeagus narrowing gradually (Fig. 12), base above basal opening as wide as medial width, and apex slightly down-curved in lateral view.

Spermatheca with short thick pump (Fig. 14), distinctly separated from receptacle. Tignum straight, widening abruptly anteriorly (Fig. 15). Apical abdominal segment attached near middle of vaginal palpi (Fig. 13).

Type material.

Holotype, male. 1) India: Kerala, Arippa, 8°50'11"N, 77°1'46"E, 30.xi.2011, 236m. D. Prathapan & K. Shameem Coll.; 2) H:396 Syzygium travancoricum; 3) Orthaltica sp. 1 KD Prathapan det 2012; 4) Holotype Orthaltica syzygium n. sp. Prathapan & Konstantinov 2012 (USNM). Paratypes: 13 specimens, the same labels as holotype; 8 specimens, the same labels as holotype except for the date 23.iii.2012. 29 specimens. 1) India Kerala, Kumarakom, 2.vii.2010 Prathapan Coll.; 2) 157 Syzygium caryophyllatum; 3) Paratype Orthaltica syzygium n. sp. Prathapan & Konstantinov 2012. 16 specimens. 1) India Karnataka, Kottigehara, 13°7'7.7"N, 75°30'37.9"E, 8.v.2011, 938m. D. Prathapan & K. Shameen Coll. 2) Host Syzygium cumini 3) Paratype Orthaltica syzygium n. sp. Prathapan & Konstantinov 2012. 19 specimens. 1) India Karnataka, Kudremukha N. P., 13°14'42"N, 75°6'44"E, 12.v.2011, 110m. D. Prathapan & K. Shameem Coll. Syzygium caryophyllatum; 2) Paratype Orthaltica syzygium n. sp. Prathapan & Konstantinov 2012 (5 – BMNH, 64 – NPC, 5 – PKDC, 5 – UASB, 9 – USNM).

Etymology.

The specific epithet is a noun in apposition, based on the host plant name.

Host plants.

Syzygium cumini (L.) Skeels (Accession no. 6516), Syzygium caryophyllatum Alston (Accession no. 6626) and Syzygium travancoricum Gamble (Accession no. 6693) (Myrtaceae).

Remarks.

Orthaltica syzygium can easily be distinguished from all known Indian Orthaltica species using the key provided below. Orthaltica terminalia can be distinguished from the most similar species using the following characters: body dark brown (Fig. 9); vertex with eight long and more than 10 short setae (Fig. 11); elytra densely pubescent (Figs 9–10); apex of aedeagus narrowing gradually (Fig. 12); and spermatheca with short thick pump (Fig. 14).

Orthaltica terminalia Prathapan & Konstantinov, sp. n.

http://zoobank.org/040BDB2D-61C5-4E47-83DF-E15A898C06CC

http://species-id.net/wiki/Orthaltica_terminalia

Figs 1, 16–22
Description.

Body length 1.20 – 1.50 mm, width 0.75 – 0.84 mm. Body shiny brownish-black (Figs 16, 17). Antennomeres (except most apical) and legs (except base of some femora) yellowish-brown. Vertex with 4 long and 6 short setae. Supracallinal sulci slightly curved; orbital sulci absent; suprafrontal sulcus poorly developed; and suprantennal sulcus shallow. Antennal calli narrow and elongate (Fig. 18), relatively widely connected, with two large setiferous pores. Frontal ridge parallel-sided, ending before reaching anterofrontal ridge. Anterofrontal ridge of uniform height, but sloping abruptly towards clypeus. Labrum as wide as distance between outer edges of antennal sockets. Setiferous pores on dorsal surface of labrum small, more than two, but not possible to count. Antennomere 3 slightly shorter than 4 and 5 separately.

Figures 16–22.

Orthaltica terminalia Prathapan & Konstantinov, sp. n. 16 Dorsal habitus 17 Lateral habitus 18 Head frontal view 19 Aedeagus, ventral and lateral view 20 Vaginal palpi 21 Spermatheca 22 Tignum.

Pronotum about as wide basally as apically, with sides evenly curved. Lateral margins slightly uneven and lacking a prominent denticle; bearing two long setae (Fig. 17). Antebasal transverse impression distinct (Fig. 16). Pronotal surface sparsely covered with slightly elongate punctures, their diameter smaller than distance between punctures.

Elytra sparsely pubescent (Figs 16, 17), with erect setae mainly near apices. Elytral punctures arranged in striae located in relatively deep grooves. Humeral calli distinct. Pro- and middle tibiae lacking spurs. Claws appendiculate.

Apex of aedeagus narrowing abruptly (Fig. 19), base above basal opening narrower than medial width, and apex slightly up-curved in lateral view.

Spermatheca with long thin pump (Fig. 21) which is not distinctly separated from receptacle. Tignum straight widening gradually anteriorly (Fig. 22). Apical abdominal segment attached to basal third of vaginal palpus (Fig. 20).

Type material.

Holotype, male. 1) India Karnataka, Kudremukha N. P., 13°14'42"N, 75°6'44"E, 12.v.2011, 110m. D. Prathapan & K. Shameem Coll. Terminalia paniculata; 2) Holotype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012 (USNM). Paratypes: 45 specimens, the same labels as holotype. 1 specimen 1) India Karnataka, Guddayanadoddi, 12°43.233'N, 077°33.576'E, 29.vii.2011 905m; 2) Terminalia arjuna; 3) Orthaltica sp. 2 K D Prathapan det. 2012; 4) Paratype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012. 12 specimens 1) India Kerala, Calicut Univ. Campus, 3.vii.2012, Shameem K. Coll. Ex Terminalia paniculata; 2) Paratype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012. 8 specimens 1) India Kerala, Trichur VellaniPacha, 27.x.2010, Prathapan K. D. Coll; 2) 230 Terminalia arjuna; 3) Orthaltica sp. 2 K D Prathapan det. 2012; 4) Paratype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012. 1 specimen 1) India Kerala, Aathirappilly, 10°18'44.7"N, 076°42'19.6"E, 26.x.2010, 570m; 3) Orthaltica sp. 2 K D Prathapan det. 2012;4) Paratype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012. 9 specimens 1) India Kerala, Kallar, 17.viii.2010, Prathapan Coll.; 2) Terminalia paniculata; 3) Orthaltica sp. 2 K D Prathapan det. 2012; 4) Paratype Orthaltica terminalia n. sp. Prathapan & Konstantinov 2012. (5 – BMNH, 53 – NPC, 5 – PKDC, 5 – UASB, 9 – USNM).

Etymology.

The specific epithet is a noun in apposition, based on the host plant name.

Host plants.

Terminalia cuneata Roth (Accession no. 6576) and Terminalia paniculata Roth (Accession no. 6484) (Combretaceae).

Remarks.

Orthaltica terminalia can be easily distinguished from all known Indian Orthaltica species using the key below. Orthaltica syzygium can be distinguished from the most similar species using the following characters: body shiny black (Fig. 16); vertex with four long and six short setae (Fig. 18); elytra sparsely pubescent (Figs 16 & 17); apex of aedeagus narrowing abruptly (Fig. 19); spermatheca with thin elongate pump (Fig. 21). Orthaltica minuta indica (Medevedev) is not included in the key as there is no character to separate it from Orthaltica dakshina (Basu & Sengupta). Both species are from the same locality (Doddabetta in Tamil Nadu) and further research may prove them to be synonymous.

Key to the Indian species of Orthaltica
1 Length 2.4 mm; vertex with a pair of long widely separated setae at posterior Orthaltica purba Basu & Sengupta
Length 1.6  mm or less; vertex with a row of at least four long setae at posterior 2
2(1) Elytral interstices, between the eighth and ninth row of punctures, sharply carinated; transverse antebasal impression on pronotum sinuate Orthaltica assamensis (Scherer)
Elytral interstices, between eighth and ninth row of punctures, not carinate; transverse antebasal impression on pronotum straight 3
3(2) Anterofrontal ridge low, merging gradually with clypeus Orthaltica bengalensis (Basu & Sengupta)
Anterofrontal ridge high; sloping abruptly towards clypeus 4
4(3) General body colour reddish-brown Orthaltica dakshina (Basu & Sengupta)
General body colour dark brown to black 5
5(4) Frontal ridge merging gradually with anterofrontal ridge, together forming a triangular ridge Orthaltica coomani (Laboissiere)
Anterofrontal ridge evenly raised and transverse, abruptly joining frontal ridge to form an inverted T-shaped ridge 6
6(5) Body shiny brownish-black (Fig. 16). Vertex with 4 long and 6 short setae (Fig. 18). Elytra sparsely pubescent (Fig. 16 & 17). Apex of aedeagus narrowing abruptly (Fig. 19). Spermatheca with thin elongate pump (Fig. 21) Orthaltica terminalia sp. n.
Body dark brown (Fig. 9). Vertex with 8 long and more than 10 short setae (Fig. 11). Elytra densely pubescent (Fig. 9 & 10). Apex of aedeagus narrowing gradually (Fig. 12). Spermatheca with short thick pump (Fig. 14) Orthaltica syzygium sp. n.
Discussion

Besides Orthaltica, species of Terminalia and Syzygium also support other leaf feeding beetles belonging to the family Chrysomelidae, as well as to the family Curculionidae. These beetles, being much larger (length 2.2 – 4.5 mm) than Orthaltica (length 1.2 – 1.6 mm), feed by cutting holes in the laminae. The adults of Orthaltica are extremely small and incapable of cutting holes in the laminae, but rather produce elongate feeding trenches on their adaxial surface. Adults of Orthaltica syzygium and Orthaltica terminalia utilise holes previously made by larger herbivores, in the leaves of their respective hosts, as shelter. They then create feeding trenches which radiate from the leaf hole which is their shelter (Fig. 1). Beetles could be seen inside holes during most of the day when they were not feeding. They came out of their leaf-hole shelters to feed, forming irregular trenches radiating from the hole. When threatened by means of a finger or a stick, beetles in holes immediately shifted their position to the reverse side of laminae, thus making themselves invisible to the enemy on the opposite side. It was observed that whenever an ant appeared on one side of the lamina, the occupant of a leaf-hole shelter also shifted its position to the reverse side.

Leaf beetles of the subfamily Eumolpinae are the most common primary hole makers on Syzygium and Terminalia species. At Kudremukha National Park it was observed that a Basilepta sp. (Eumolpinae) leaf beetle chewed holes in the laminae of Syzygium caryophyllatum (Fig. 3). Orthaltica syzygium released on the leaf readily occupied the leaf-hole as a shelter (Fig. 4). By the second day feeding trenches were observed radiating from the leaf-hole shelter (Fig. 5).

The triangular artificial holes, made in the laminae of Syzygium travancoricum with a punch, were accepted as shelters (Fig. 6). It was observed that the shape and size of the holes were not exactly what the beetles required. Holes were then resized by partitioning—walls were constructed using faecal pellets (Fig. 7). A single occupant was observed in a hole in most cases. However, when high population densities occurred, several beetles could be seen in a single hole that was large enough to accommodate them (Figs 1, 8).

In the case of the triangular artificial leaf-holes, beetles had a distinct preference for the narrow vertex of the triangle, above its wider base (Fig. 7). This may be because their size allowed them to easily fit inside the narrow apical angle. On 20th May it was found that, of the 316 triangular holes examined on 53 leaves after a month, 235 were occupied. Similarly on 20th August, after a period of four months, of the 319 triangular holes on 58 leaves, 240 were occupied.

Even when suitable leaf-holes were available on leaf laminae, non-shelter forming species never occupied such holes (Fig. 2). Orthaltica syzygium confined on leaves without holes fed normally on the adaxial surface of the laminae.

Leaf-hole shelters provide a roosting site that offers a certain degree of camouflage as well as protection. In the field it was observed that on sensing the presence of an enemy on one side of the leaf the occupant of a leaf-hole shelter could easily shift to the other side, making itself invisible to the intruder. It may also be presumed that larger predators cannot pass through the hole in pursuit of the occupant. The leaf-hole, as well as the surrounding area with feeding trenches, turn dark brown and provide a dark background from which the beetle cannot be easily differentiated by a potential predator. On the other hand, the leaf-hole shelter can also provide sufficient cues for a specialist enemy that only needs to learn to locate the leaf-hole shelter to get the inmate. This is the case where insect collectors are concerned—it is easy for them to locate leaf-hole shelter and collect the beetles utilising them.

Leaf-hole shelters were observed in two of the five Orthaltica species found in southern India. This is certainly a behavioural novelty of evolutionary significance. It can be seen as an example of Lorenz’s (1937) discovery that a behaviour pattern can be treated as an anatomical organ, which Dawkins (1982) further synthesized and developed into the hypothesis of “the extended phenotype”. It is likely that this extracorporeal adaptation evolved in two lineages of Orthaltica in response to interactions with other leaf feeding beetles. Further investigation into the prevalence and diversity of herbivorous beetles on the host plants of Orthaltica, and their interactions with each other, might provide the answer.

Adult chrysomelids, unlike their larvae, never carry their faeces in the form of defensive shields. However, females of many species are known to defecate on their eggs after oviposition. Adults are also known to defecate when seized (Müller and Hilker 2004). The use of excreta for the construction of defensive structures or retreats was, until now, not known for adult leaf beetles.

Acknowledgements

PKD’s work on leaf beetles is supported by the Kerala State Council for Science, Technology and Environment, Trivandrum as well as the Indian Council of Agricultural Research through the Network Project on Insect Biosystematics. We are indebted to A. K. Pradeep (University of Calicut, Kerala, India) for identification of the host plants. Eumolpinae specimens were identified by A. Moseyko (Zoological Institute, St. Petersburg, Russia). A. K. Tishechkin (Santa Barbara Museum of Natural History), and N. Woodley (Systematic Entomology Laboratory, USDA, ARS) kindly reviewed a preliminary version of this manuscript and provided valuable suggestions. USDA is an equal opportunity provider and employer.

References
Basu CR, Sengupta T (1978) The Indian species of Livolia (Coleoptera: Chrysomelidae: Alticinae). Oriental Insects 12: 481-487. doi: 10.1080/00305316.1978.10432533
Basu CR, Sengupta T (1985) A new species of Orthaltica (Coleoptera: Chrysomelidae: Alticinae) from India. Bulletin of the Zoological Survey of India 7: 45-48.
Chaboo CS, Grobbelaar E, Larsen A (2007) Fecal ecology in leaf beetles: novel records in the African arrow-poison beetles, Diamphidia Gerstaecker and Polyclada Chevrolat (Chrysomelidae: Galerucinae). Coleopterists Bulletin 61: 297-309. doi: 10.1649/0010-065X(2007)61[297:FEILBN]2.0.CO;2
Chaboo CS, Brown CG, Funk DG (2008) Fecal case architecture in the gibbosus species group of Neochlamisus Karren (Coleoptera: Chrysomelidae: Cryptocephalinae: Chlamisini). Zoological Journal of the Linnean Society 152: 315-351. doi: 10.1111/j.1096-3642.2007.00343.x
Dawkins R (1982) The Extended Phenotype. Oxford University Press, New York, 313 pp.
Gilbert AJ, Andrews FG, Staines CL (2001) An unusual host record for Imatidium and new distributional records for I. rufiventre Boheman and I. thoracicum Fabricius (Coleoptera, Chrysomelidae, Hispinae: Cephaloleiini). Pan-Pacific Entomologist 77: 278-280.
Hansell MH (2005) Animal architecture. Oxford University Press, New York, 322 pp. doi: 10.1093/acprof:oso/9780198507529.001.0001
Hansell MH (2007) Built by animals. Oxford University Press, New York, 256 pp.
Konstantinov AS (1998) Revision of the Palearctic Species of Aphthona Chevrolat and Cladistic Classification of the Aphthonini (Coleoptera: Chrysomelidae: Alticinae). Memoirs on Entomology International, Associated Publishers, Florida, 429 pp.
Lorenz K (1937) Uber die Bildung des Instinktbegriffes. Die Naturwissenschaften 25: 289-300. doi: 10.1007/BF01492648
Maulik S (1919) The fauna of British India including Ceylon and Burma. Coleoptera: Chrysomelidae (Hispinae and Cassidinae). Taylor and Francis, London, 439 pp.
Medvedev LN (1998) New Alticinae and Galerucinae (Coleoptera, Chrysomelidae) from South Asia. Acta Zoologica Hungarica 45: 297-301.
Müller C, Hilker M (2004) Ecological relevance of fecal matter in Chrysomelidae. In: Jolivet P, Santiago-Blay JA, Schmitt M (Eds) New Developments in the Biology of Chrysomelidae. SPB Academic Publishing bv, The Hague: 693-705.
Prathapan KD, Chaboo CS, Karthikeyan K (2009) Natural History and Leaf Shelter Construction of the Asian Rice Leptispa Beetle Leptispa pygmaea Baly (Coleoptera: Chrysomelidae: Cassidinae: Leptispini). Zoological Studies 48: 625-631.
Prathapan KD, Chaboo CS (2011) Biology of Blepharida-group flea beetles with first notes on natural history of Podontia congregata Baly, 1865 an endemic flea beetle from southern India (Coleoptera, Chrysomelidae, Galerucinae, Alticini). ZooKeys 157: 95-130. doi: 10.3897/zookeys.157.1472
Reid CAM (1990) Micrantipha Blackburn, a synonym of Orthaltica Crotch (Col., Chrysomelidae). Entomologist’s monthly Magazine 126: 205-206.
Scherer G (1971) Das Genus Livolia und seine umstrittene Stellung im System. Entomologische Arbeiten aus dem Museum G. Frey 22: 1-37.
Seeno TN, Wilcox JA (1982) Leaf Beetle Genera (Coleoptera: Chrysomelidae). Entomography 1: 1-221.
Vencl FV, Morton TC, Mumma RO, Schultz JC (1999) Shield defense of a larval tortoise beetle. Journal of Chemical Ecology 25: 549-566. doi: 10.1023/A:1020905920952
Voronova HB, Zaitsev YM (1982) The genus Leptispa– pests of bamboo in Vietnam. In: Medvedev LN (Ed) Animal world of Vietnam. Nauka Press, Moscow, 115-124. [in Russian]
Weiss RS (2006) Defecation behavior and ecology of insects. Annual Review of Entomology 51: 635-661. doi: 10.1146/annurev.ento.49.061802.123212