Hispines (chrysomelidae, cassidinae) of la selva biological station, costa rica.

Survey work from 1992-2001 identified 139 species of hispines at the lowland part of La Selva Biological Station, Costa Rica. The tribe Cephaloleiini was the most speciose with 58 species (41.7%) followed by the Chalepini with 55 (39.5%). The fauna is most closely related to that in South America but with some genera which are more speciose in the Nearctic Region. Plant associations are known for 88 (63.3%) of the species but many of these are merely collecting records, not host plant associations. The first plant associations are reported for Alurnus ornatus, Alurnus salvini, and Acentroptera nevermanni.

Introduction

Hispines comprise half of the subfamily Cassidinae (sensu lato) in the family Chrysomelidae within the order Coleoptera (Staines 2002b). Until recently, most authors treated the group as a separate subfamily but recent work has shown that there is no biological or morphological reason to retain sub-familial status (Staines 2002b). The combined subfamily consists of 6000 species placed in 42 tribes (Staines 2002b). See Staines (2002b) for a detailed history of the classification of the two groups.

The combination of the Hispinae with the Cassidinae (s. str.) has created difficulty in having a handy term to use for these beetles. Several have been proposed but they PageBreakare cumbersome. Until an easily used term is coined for this group, I continue to use “hispines” in the traditional sense of the genera and species in the former subfamily Hispinae (see Seeno and Wilcox 1982 for a list of genera).

The adult hispine head is opisthognathous, prominent, visible from above, at least to behind the eyes. The frons is prominent, exposed or rarely retracted. The antennae are not retractable and are closely inserted between the eyes. The pronotum is narrower than the elytra; it is more or less quadrangular or trapezoidal, with definite anterior angles which may have a small tubercle. The scutellum is always visible. The elytra lack lateral expansions or have reduced and discontinuous expansions. The margins are usually denticulate or with spines.Larvae are either leaf-miners or free living. They have eight pairs of abdominal spiracles which are well developed and dorsally placed; with the eighth abdominal segment terminal, and with free hind margin (Staines 2002b, 2006).

Ecologically, New World hispines fall into three feeding groups: external feeders; sheath, appressed or rolled-leaf feeders; and leaf-miners. In the Old World, some species have been reported as stem borers in herbaceous or semi-ligneous plants, but this has not been reported from the New World. The biology of few species has been studied; most are not associated with a host plant or plant family.

Methods

Study area

La Selva Biological Station (10°26'N, 83°59'W) is located in the Atlantic tropical lowlands of Costa Rica and is adjacent to Braulio Carrillo National Park. It is about 100 km from San José. The station comprises 1600 hectares. Habitat is a mosaic of primary forest, early secondary pasture, young secondary forest, abandoned plantations, and selectively logged primary forest. The elevation varies from 35 to 137 m. The station is near the confluence of Rio Puerto Viejo and Rio Sarapiquí. It is owned and operated by the Organization for Tropical Studies (McDade and Hartshorn 1994).

Rainfall varies from 152.0 mm (March) to 480.7 mm (July) with a total 4 m per year. The dry season is short and not severe (Sanford et al. 1994).

There are 1744 plant species documented from La Selva. The most speciose families are Pteridophyta, Orchidaceae, Araceae, Rubiaceae, Melastomataceae, Fabaceae, and Piperace (Hartshorn and Himmel 1994).

The Arthropods of La Selva (ALAS) project was started in 1991 (http://viceroy.eeb.uconn.edu/ALAS/ALAS.html). An existing building on the station was remodeled as an entomology laboratory and four technicians were trained in the National Biodiversity Institute (INBio) six-month parataxonomist course. From 1992 until 2000 the project was funded by separate grants from the U.S. National Science Foundation (Biotic Surveys and Inventories Program). From 2001–2006 the focus of the project shifted to a transect survey from La Selva Biological Station to the summit of Volcan Barva. This paper deals only with the results at the La Selva Biological Station.PageBreak

Daily operations of ALAS were conducted by the parataxonmists under the direction of the principal investigators John T. Longino, Evergreen State College, and Robert K. Colwell, University of Connecticut. Over the course of the project there were over 100 collaborating taxonomists.

Survey aethods

The ALAS survey consisted of both structured and directed sampling. Structured sampling consisted of black-lights, Malaise and flight intercept traps, and canopy fogging (see Furth et al. 2003 for summary).

Passive black-lights were utilized from 1993 to 1999 at twelve sites, six on the ground and six in the canopy. Malaise traps were used at sixteen sites from 1993 to 2000. Specimens were collected directly into ethanol and the traps emptied every two weeks. Flight intercept traps were place at sixteen sites and samples were collected every two weeks.

Canopy fogging was conducted in 1993–1994, 1996, and 2000. Sixteen trees were fogged: six trees of the most common species at La Selva, six trees of an intermediate abundant species, and trees of six different species. The tree selected had large crowns with little crown overlap and with good climbing access.

Directed collecting for chrysomelids used beating, sweeping, visual observation, known host plant observation, and use of a mid-canopy shaker net.

Results and discussion

Species richness at La Selva

As of the end on 2001, a total of 139 hispine species have been collected at La Selva Biological Station (see Table 1).

Table 1.

Hispines known from La Selva Biological Station and their plant associations (A=adult plant feeding; L=larval host plant; U=unspecified).

Tribe	Genus/Species	Plant association	Plant family	Observation	References
Alurnini	Alurnus ornatus Baly	Chamaedorea sp.	Arecaceae	A	New observation
Alurnini	Alurnus salvini Baly	Chamaedorea sp.	Arecaceae	A	New observation
Arescini	Chelobasis bicolor Gray	Heliconia sp., Heliconia latispatha Benth., Heliconia tortuosa Griggs, Heliconia cathaeta R. R. Smith	Heliconiaceae	L	Maulik 1937;<br/> Strong 1977a, 1983;<br/> Meskins et al. 2008
		Musa sp.	Musaceae	L
		Calathea latifolia Klotzsch	Marantaceae	L
Arescini	Chelobasis perplexa Baly	Heliconia imbricata (Kuntze) Baker, Heliconia latispatha, Heliconia pogonantha Cuford.,	Heliconiaceae	L	Maulik 1932;<br/> Strong and Wang 1977;<br/> McKenna and Farrell 2005; Meskins et al. 2008
		Heliconia irrasa R. R. Smith, Heliconia mariae Hook.Calathea insignis Hort. & Bull.	Marantaceae	L
Cephaloleiini	Aslamidium impurum (Boheman)	Calathea ovata Lindl., Calathea virginalis Linden, Calathea insignis, Calathea micans (Mathieru) Koern.	Marantaceae	A	Bondar 1940; Spaeth 1938;
		Heliconia sp.	Heliconiaceae	A	Windsor et al. 1992
Cephaloleiini	Cephaloleia aequilata Uhmann	Unknown
Cephaloleiini	Cephaloleia atriceps Pic	Unknown
Cephaloleiini	Cephaloleia bella Baly	Heliconia imbricata	Heliconiaceae	A	Staines 1996
Cephaloleiini	Cephaloleia belti Baly	Calathea insignis, Calathea latifolia Klotzsch, Calathea lutea (Aubl.) GFW Mey., Ischnosiphon pruinosus Peterson	Marantaceae	A	Uhmann 1930; Maulik 1932; Strong 1977b, 1982a; Meskins et al. 2008; Descampe et al. 2008; García-Robledo et al. 2010
		Heliconia imbricata, Heliconia latispatha, Heliconia pogonantha, Heliconia mariae, Heliconia tortuosa, Heliconia catheta, Heliconia irrasa, Heliconia vaginalis Benth., Heliconia wagneriana Peterson	Heliconiaceae	A, L
Cephaloleiini	Cephaloleia championi Baly	Heliconia sp.	Heliconiaceae	A	Staines 1996
Cephaloleiini	Cephaloleia congener Baly	Heliconia latispatha, Heliconia tortuosa	Heliconiaceae	A	Staines 1996
Cephaloleiini	Cephaloleia consanguinea Baly	Unknown
Cephaloleiini	Cephaloleia costaricensis Uhmann	Chusquea simpliciflora Munro	Poaceae	U	Meskins et al. 2008
Cephaloleiini	Cephaloleia deficiens Uhmann	Unknown
Cephaloleiini	Cephaloleia dilaticollis Baly	Calathea insignis, Calathea lutea, Calathea inocephala (Kuntze), H. Kennedy, Ischnosiphon pruinosus	Marantaceae	A	Staines 1996; McKenna and Farrell 2005; Meskins et al. 2008; Descampe et al. 2008; García-Robledo et al. 2010
		Renealmia sp., Renealmia alpinia (Rottb.) Maas	Zingiberaceae	A, L
Cephaloleiini	Cephaloleia disjuncta Staines	Vitex copperi Stanley	Verbenaceae	A	Staines 1998
Cephaloleiini	Cephaloleia distincta Baly	Calathea sp.	Marantaceae	A	Staines 1996
		Heliconia imbricata	Heliconiaceae	A
Cephaloleiini	Cephaloleia dorsalis Baly	Costus sp., Costus pulverulentus C. Presl., Costus malortieanus Wendl., Costus larvis Ruiz. & Pav.	Costaceae	A, L	Staines 1996; McKenna and Farrell 2005; Meskins et al. 2008; García-Robledo and Horvitz 2009; García-Robledo et al. 2010
		Renealmia sp.	Zingiberaceae	A
Cephaloleiini	Cephaloleia elegantula Baly	Unknown
Cephaloleiini	Cephaloleia erichsonii Baly	Calathea gymnocarpa H. Kennedy, Calathea inocephala, Calathea leucostachys Hook., Calathea insignis, Calathea latifolia, Calathea lutea	Marantaceae	A	Staines 1996; Strong 1977a; McKenna and Farrell 2005; Meskins et al. 2008; Descampe et al. 2008
		Heliconia sp., Heliconia catheta, Heliconia latispatha, Heliconia mariae, Heliconia vaginalis, Heliconia wagneriana	Heliconiaceae	A
Cephaloleiini	Cephaloleia exigua Uhmann	Unknown
Cephaloleiini	Cephaloleia fenestrata Weise	Ischnosiphon sp., Ischnosiphon cerotus Leos., Pleiostachya pruinosa K. Schum.	Marantaceae	L	Staines 1996; Strong 1977a; Johnson 2004a
Cephaloleiini	Cephaloleia flava Uhmann	Unknown
Cephaloleiini	Cephaloleia fulvolimbata Baly	Unknown
Cephaloleiini	Cephaloleia heliconicae Uhmann	Heliconia sp.	Heliconiaceae	A	Staines 1996
		Calathea insignis	Marantaceae	S
Cephaloleiini	Cephaloleia histrionica Baly	Unknown
Cephaloleiini	Cephaloleia lata Baly	Chamaedorea tepejilote Liebm., Chamaedorea wendlandiana Hemsl.	Arecaceae	A	McKenna and Farrell 2005; Meskins et al. 2008
Cephaloleiini	Cephaloleia mauliki Uhmann	Heliconia sp.	Heliconiaceae	A	Uhmann, 1930; Maulik 1932, 1937
		Calathea insignis	Marantaceae	A
Cephaloleiini	Cephaloleia metallescens Baly	Bactris major Jacq., Chamaedorea wendlandiana	Arecaceae	U	Meskins et al. 2008
Cephaloleiini	Cephaloleia nevermanni Uhmann	Calathea insignis, Calathea macrosepala K. Schumann	Marantaceae	A	Uhmann 1930; Staines 1996
		Heliconia imbracata	Heliconiaceae	A
Cephaloleiini	Cephaloleia nigricornis (Fabricius)	Unknown
Cephaloleiini	Cephaloleia ornatrix Donckier	Heliconia sp.	Heliconiaceae	A	Strong 1977a
Cephaloleiini	Cephaloleia placida Baly	Renealmia sp., Renealmia alpinia (Rottb.) Maas	Zingiberaceae	A	Staines 1996; García-Robledo and Horvitz 2009; García-Robledo et al. 2010
Cephaloleiini	Cephaloleia puncticollis Baly	Calathea insignis	Marantaceae	L	Uhmann 1930; Seifert and Seifert 1976; Staines 1996
		Heliconia imbricata, Heliconia latispatha	Heliconiaceae	L
		Musa sp.	Musaceae	L
Cephaloleiini	Cephaloleia quadrilineata Baly	Heliconia imbricata, Heliconia latispatha	Heliconiaceae	A	Staines 1996
Cephaloleiini	Cephaloleia reventazonica Uhmann	Heliconia latispatha	Heliconiaceae	A	Staines 1996
Cephaloleiini	Cephaloleia ruficollis Baly	Unknown
Cephaloleiini	Cephaloleia sallei Baly	Heliconia sp., Heliconia irrasa, Heliconia catheta, Heliconia latispatha, Heliconia mariae, Heliconia vaginalis	Heliconiaceae	L	Strong 1977a; Staines 2004a; McKenna and Farrell 2005; Meskins et al. 2008; Descampe et al. 2008
		Renealmia strobilifera
		Calathea inocephala, Calathea latifolia, Calathea lutea,	Zingiberaceae	A
		Ishnosiphon pruinosus	Marantaceae	A
Cephaloleiini	Cephaloleia semivittata Baly	Calathea marantifolia Standley	Marantaceae	A	Staines 1996
Cephaloleiini	Cephaloleia splendida Staines	Unknown
Cephaloleiini	Cephaloleia stevensi Baly	Heliconia sp.	Heliconiaceae	A	Staines 1996; McKenna and Farrell 2005; Meskins et al. 2008
		Calathea micans, Calathea inocephala, Calathea latifolia, Ishnosiphon pruinosus	Marantaceae	A
Cephaloleiini	Cephaloleia sulciceps Baly	Unknown
Cephaloleiini	Cephaloleia suturalis Baly	Costus malorticenus H. Wendl., Costus sp., Costus pulverulentus	Costaceae	A	Uhmann 1930; Maulik 1937; Meskins et al. 2008
Cephaloleiini	Cephaloleia tenella Baly	Unidentified	Areaceae	A	Staines 1996
Cephaloleiini	Cephaloleia trimaculata Baly	ginger lily, Renealmia sp.	Zingiberaceae	A	Uhmann 1950; McKenna and Farrell 2005; Meskins et al. 2008
		Costus pulverulentus	Costaceae	A
Cephaloleiini	Cephaloleia trivittata Baly	Calathea haamelii H. Kennedy, Calathea macrosepala	Marantaceae	A	Staines 1996
Cephaloleiini	Cephaloleia vicina Baly	Heliconia spp., Heliconia latispatha, Heliconia imbricata	Heliconiaceae	A	Strong 1977a, 1977b, 1981
		Calathea spp., Ischnospihon spp.	Marantaceae	A
Cephaloleiini	Cephaloleia sp. 1	Unknown
Cephaloleiini	Demotispa nevermanni Uhmann	Unknown
Cephaloleiini	Demotispa strandi Uhmann	Spermacoce sp.	Rubiaceae	U	Flowers and Janzen 1997; Staines 2006a
		Calathea sp.	Marantaceae	A
Cephaloleiini	Demotispa sp. 1	Unknown
Cephaloleiini	Demotispa sp. 2	Unknown
Cephaloleiini	Homalispa gracilis Baly	Unknown
Cephaloleiini	Homalispa nevernmanni Uhmann	Oenocarpus panamanus Bailey	Arecaceae	U	Meskins et al. 2008
Cephaloleiini	Homalispa sp. 1	Unknown
Cephaloleiini	Imatidium rufiventre Boheman	Inga marginata Willd.	Fabaceae	A	Gilbert et al. 2001
Cephaloleiini	Imatidium thoracicum Fabricius	Calathea insignis, Calathea ovata, Calathea virginalis, Calathea lutena	Marantaceae	A	Spaeth 1938; Bondar 1940; Windsor et al. 1992; Meskins et al. 2008
		Heliconia latispatha, Heliconia catheta, Heliconia irrasa, Heliconia wagneriana	Heliconiaceae	A
Cephaloleiini	Solenispa leptomorpha (Baly)	Unknown
Cephaloleiini	Stenispa graminicola Uhmann	Unknown
Cephaloleiini	Stenispa sallei Baly	Unknown
Cephaloleiini	Stenispa vespertina Baly	Cyperus sp.	Cyperaceae	L	Bondar 1931b
Cephaloleiini	Stilpnaspis rubiginosus (Boheman)	Unknown
Chalepini	Anisostena pilatei (Baly)	Unknown
Chalepini	Baliosus productus (Baly)	Unidentified	Bignoniaceae	L	Hespenheide and Dang 1999
Chalepini	Baliosus sp.1	Urera bogataense ?	Urticaceae	L	Hespenheide and Dang 1999
Chalepini	Baliosus sp. 2	Unknown
Chalepini	Carinispa nevermanni Uhmann	Malpighia glabra L., Bunchosia sp.	Malpighiaceae	L	Uhmann 1934; Flowers and Janzen 1997
Chalepini	Chalepus amiculus Baly	Unknown
Chalepini	Chalepus angulosus Baly	Unknown
Chalepini	Chalepus assmani Uhmann	Unknown
Chalepini	Chalepus bellulus (Chapuis)	Digitaria eriantha Steud., Oryza sp.	Poaceae	L	Maes and Staines 1991; Staines 1997; Flowers and Janzen 1997
		Unidentified	Arecaceae	U
		Phaseolus sp.	Fabaceae	U
Chalepini	Chalepus brevicornis (Baly)	Unknown
Chalepini	Chalepus consanguineus (Baly)	Lasiacis sp. Unidentified	Poaceae	L	Uhmann 1935; Hespenheide and Dang 1999
Chalepini	Chalepus digressus Baly	Lasiacis sp.	Poaceae	L	Memmott et al. 1993
Chalepini	Chalepus nigripictus Baly	Unknown
Chalepini	Chalepus pici Descarpentries & Villiers	Unknown
Chalepini	Chalepus similatus Baly	Unknown
Chalepini	Chalepus tappesi Chapuis	Unknown
Chalepini	Chalepus verticalis (Chapuis)	Phaseolus sp.	Fabaceae	U	Maes and Staines 1991
Chalepini	Chalepus sp. 1	Unknown
Chalepini	Chalepus sp. 2	Unknown
Chalepini	Charistena ruficollis (Fabricius)	Zea mays L., Paspalum conjugatum Berg	Poaceae	U	Bondar 1931a; Schlottfeldt 1944; Maulik 1937; Maes and Staines 1991
		Glycine max (L.) Merr.	Fabaceae	U
		Coffea sp.	Rubiaceae	U
Chalepini	Euprionota aterrima Guérin-Méneville	Unknown
Chalepini	Glyphuroplata nigella Weise	Valota sp., Eriochloa gracilis (Fourn.) Hitchc.	Poaceae	L	Riley 1985; Hespenheide and Dang 1999
		Mimosa sp.	Fabaceae	U
Chalepini	Heptispa limbata (Baly)	Cassia grandis L., Cassia fruitcosa Mill., Inga sp., Machaerium sp.	Fabaceae	L	Uhmann 1934, 1937; Memmott et al. 1994; Hespenheide and Dang 1999
		Serjania sp.	Sapindaceae	U
		Olyra latifolia	Poaceae	U
Chalepini	Heterispa vinula (Erichson)	Triumfetta josefina Polak,	Tilaceae	L	Uhmann 1934, 1937; Maulik 1937; Hespenheide and Dang 1999; Casari and Teizeira 2004
		Apeiba membranacea Spruce ex. Benth.
		Guazuma ulmifolia L.	Sterculiaceae	L
		Sida sp. Sida rhombifolia L., Sida carpinifolia K. Schum.	Malvaceae	L
		Infigofera sp.	Fabaceae	L
Chalepini	Octhsipa bimaculata Uhmann	Stigmaphyllum lindenianum A. Juss.	Malphigiaceae	L	Hespenheide and Dang 1999
Chalepini	Octhispa decepta (Baly)	Stigmaphyllum lindenianum	Malphigiaceae	L	Hespenheide and Dang 1999
Chalepini	Octhispa elegantula Baly	Serjania sp., Paullinia sp.	Sapindaceae	L	Uhmann 1937; Hespenheide and Dang 1999
		Pithecoctenium echinatum K. Schum.	Bignoniaceae	U
Chalepini	Octhispa elevata (Baly)	Paullinia sp.	Sapindaceae	L	Uhmann 1934; Maulik 1937; Hespenheide and Dang 1999
		Pithecoctenium echinatum	Bignoniaceae	U
Chalepini	Octhispa haematopyga Baly	Colubrina spinosa Don. Sm.	Rhamnaceae	L	Hespenheide and Dang 1999
Chalepini	Octhispa nevermanni Uhmann	Ochroma lagopus Rowlee	Bombaceae	L	Hespenheide and Dang 1999
Chalepini	Oxychalepus alienus (Baly)	Centrosema macrocarpum Benth., Cassia fruticosa	Fabaceae	L	Flowers and Janzen 1997; Hespenheide and Dang 1999
Chalepini	Oxychalepus posticatus (Baly)	Cassia oxyphylla Kunth., Cassia hayesiana Standl., Cassia fruticosa	Fabaceae	L	Uhmann 1937; Memmott et al. 1994; Hespenheide and Dang 1999
Chalepini	Oxyroplata nr. bellicosa Uhmann	Bamosteroa argentea Spreng.	Malphighiaceae	L	Uhmann 1937
Chalepini	Pentispa explanata (Chapuis)	Pithecoctenium sp.	Bignoniaceae	L	Uhmann 1934
Chalepini	Pentispa fairmairei (Chapuis)	Chusquea sp.	Poaceae	U	Uhmann 1937; Maulik 1937; Morris et al. 2004
		Calea urticaefolia (P. Mill.) DC, Calea axillaries DC., Vernonia mollis H.B.K., Verbesina sp., Eupatorium populifolium Hook. & Arn., Clibadium sp., Lepidaploa tortuosa (L.) H. Rob.	Asteraceae	L
		Elephantopus spicatus Aubl., Malpighia glabra	Malpighiaceae	U
		Serjania sp.	Sapindaceae	U
Chalepini	Pentispa sp. 1	Unknown
Chalepini	Pentispa sp. 2	Unknown
Chalepini	Platocthispa championi (Baly)	Piper sp.	Piperaceae	L	Hespenheide and Dang 1999
Chalepini	Platocthispa emorsitans (Baly)	Calathea sp., Calathea insignis,	Marantaceae	U	Staines 2004a; Meskins et al. 2008
		Calathea latifolia
		Costus sp.	Costaceae	U
		Heliconia catheta, Heliconia irrasa, Heliconia latispatha	Heliconiaceae	U
Chalepini	Platocthispa sp. 1	Ochroma lagopus	Bombaceae	L	Hespenheide and Dang 1999
Chalepini	Probaenia armigera (Baly)	Piptocarpha chontalensis Baker in Mart.	Asteraceae	L	Hespenheide and Dang 1999
Chalepini	Probaenia pici Uhmann	Mikania guaco Humb. & Bonpl.	Asteraceae	L	Hespenheide and Dang 1999
Chalepini	Probaenia sp. 1	Arrabidaea chica (Humb. & Bonpl.) Verl.	Bignoniaceae	L	Hespenheide and Dang 1999
Chalepini	Sumitrosis amica (Baly)	Heliconica sp.	Heliconiaceae	L	Hespenheide and Dang 1999
Chalepini	Sumitrosis fryi (Baly)	Eupatorium populifolium	Asteraceae	L	Uhmann 1937
Chalepini	Sumitrosis instabilis (Baly)	Unknown
Chalepini	Sumitrosis pallescens (Baly)	Chamaecrista fasciculata (Michx.) Greene, Chamaecrista nictitans (L.) Moench.	Caesalpiniaceae	U	Cavey 1994
Chalepini	Sumitrosis terminatus (Baly)	Unidentified	Fabaceae	L	Hespenheide and Dang 1999
Chalepini	Uroplata fusca Chapuis	Pithecactenium echinatum, Arrabidaea mollisima Bureau & K. Schm.	Bignoniaceae	L	Uhmann 1934, 1937; Memmott et al. 1994
		Malpighia glabra	Malpighiaceae	U
Chalepini	Uroplata sculptilis Chapuis	Clibadium aspersum DC, Synedrella nodiflora Gaertn.	Asteraceae	L	Uhmann 1934, 1937; Hespenheide and Dang 1999
		Inga edalis Mart.	Fabaceae	L
		Gouania adenophora Pilg.	Rhamnaceae	L
Chalepini	Uroplata sp. 1	Unknown
Chalepini	Uroplata sp. 2	Unknown
Chalepini	Xenochalepus amplipennis (Baly)	Unidentified	Fabaceae	L	Hespenheide and Dang 1999
Chalepini	Xenochalepus erythroderus (Chapuis)	Coussapoa nymphaeifolia Standl., Coussapoa villosa Poepp. & Endl., Cecropia insignis Liebm, Pourouma bicolor (Standl.) C.C. Berg & E.C. van Heusden	Cecropiaceae	L	Hespenheide and Dang 1999
Chalepini	Xenochalepus rufithorax (Baly)	Unknown
Prosopodontini	Prosopodonta distincta (Baly)	Unknown
Prosopodontini	Prosopodonta dorsata (Baly)	Costus sp.	Costaceae	U	Uhmann 1930; Meskins et al. 2008
		Chamaedorea wendlandiana, Cryosophila warscewiczii Bartl., Oenocarpus panamanus Bailey	Arecaceae	L
Sceloenoplini	Acentroptera strandi Uhmann	Pentaclethra macroloba Kuntze	Fabaceae	U	New observation
Sceloenoplini	Ocnosispa humerosa Staines	Conceveiba pleiostemona Donn. Smith	Euphorbiaceae	A	Staines 2002a
Sceloenoplini	Pseudispa fulvolimbata (Baly)	Unknown
Sceloenoplini	Sceloenopla antennata (Baly)	Unknown
Sceloenoplini	Sceloenopla bicolorata Staines	Sterculia recordiana papyracea E. Taylor	Sterculiaceae	A	Staines 2002a
Sceloenoplini	Sceloenopla bidentata Staines	Unknown
Sceloenoplini	Sceloenopla erudita (Baly)	Anthurium sp.	Araceae	L	Uhmann 1944; Hespenheide and Dang 1999
		Cupania sp.	Sapindaceae	L
Sceloenoplini	Sceloenopla godmani (Baly)	Clusia flava Planch. & Triana	Clusiaceae	L	Hespenheide and Dang 1999
Sceloenoplini	Sceloenopla gracilenta (Baly)	Unknown
Sceloenoplini	Sceloenopla lampyridiformis Staines	Unidentified	Viscaceae	L	Staines 2002a
Sceloenoplini	Sceloenopla longula (Baly)	Unidentified	Araceae	L	Hespenheide and Dang 1999
Sceloenoplini	Sceloenopla lutena Staines	Virola koschnyi Warb.	Myristicaceae	A	Staines 2002a
Sceloenoplini	Sceloenopla minuta Staines	Unknown
Sceloenoplini	Sceloenopla multistriata Uhmann	Virola koschnyi	Myristicaceae	U	Staines 2002a; Maes 2004
		Phoradendron sp.	Loranthaceae	U
		Persea americana P. Mill.	Lauraceae	U
Sceloenoplini	Sceloenopla nevermanni Uhmann	Anthurium sp.	Araceae	L	Uhmann 1944; Hespenheide and Dang 1999
		Cupania sp.	Sapindaceae	L
Sceloenoplini	Sceloenopla nigropicta Staines	Virola koschnyi	Myristicaceae	A	Staines 2002a
Sceloenoplini	Sceloenopla obscurovittata (Baly)	Philodendron radiatum radiatum Schott, Monstera tenuis K. Koch	Araceae	L	Hespenheide and Dang 1999
Sceloenoplini	Sceloenopla proxima (Baly)	Unknown
Sceloenoplini	Sceloenopla scherzeri (Baly)	Davilla nitida (Vahl) Kubitzki	Dilleniaceae	L	Bondar 1937; Hespenheide and Dang 1999
		Persea gratissima Gaertn.	Lauraceae	L
Sceloenoplini	Sceloenopla subparallela (Baly)	Unknown

Quantitative inventory by non-specialists using standard sampling techniques can capture about half of the fauna. Individual methods are needed to sample the rest of the community. Sweeping, beating, and host plant sampling are the best methods. Fogging, Berlese funnels, and Malaise traps capture a few species usually not otherwise collected but are not sufficient in themselves to indicate the actual fauna.

Major lineages

The most recent classification of hispines is by Würmli (1975) and Staines (2002b). There are 24 extant tribes of hispines, of which six have been found at La Selva (see Table 1).

Over 40% of the 139 hispine species and 25% of the genera are in the tribe Cephaloleiini. The Cephaloleiini is a New World tribe of 16 genera and 382 species (Staines 2002b). Over 200 species are in the genus Chevrolat (Uhmann 1957, Staines 1996).

PageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakAt La Selva is the most speciose genus with 44 species from La Selva. The biology of various species has been studied by Strong (1977a, b, 1982a, 1983), Seifert and Seifert (1976), Strong and Wang (1977), Auerbach and Strong (1981), and Morrison and Strong (1981). Since the only identification aid available to these workers was Baly (1885), which covered less than half of the species known from Central America, some of the published names are not associated with the correct species. However the published information does give valuable data on the general biology and ecology of species. Staines (2004a) attempted to associate the biological data with the correct species. Additional biological work and host plant associations have been done by Johnson (2004a, b), Johnson and Horvitz (2005), McKenna and Farrell (2005), Descampe et al. (2008), Meskins et al. (2008), García-Robledo and Horvitz (2009, in press), and García-Robledo et al. (2010).

eggs are flat, with a thin chorion; hence they are subject to desiccation. Eggs are laid on host surfaces. Oviposition sites vary among beetle species and host plant. The most common oviposition sites are leaf surfaces, petioles of immature leaves or inflorescence bracts. Eggs hatch in 10 to 20 days. Larvae begin feeding immediately upon the part of the plant on which the egg was laid. larvae have a water penny-like appearance. They are flat and well adapted to moving between the wet surfaces of Zingiberales leaves, stems, and flowers. Larvae grow very slowly and go through up to eight molts depending on the size of the species and the part of the plant fed on. During their development, larvae of leaf and stem-feeding species utilize several leaves or even leaves on adjacent plants. Inflorescence-feeding larvae are restricted to a single inflorescence. Larvae of species feed on the plant by dragging their mandibles across the plant surface while they crawl forward. This results in an irregularly shaped feeding scar and a trail of frass. Pupation occurs above ground, usually on the stalk of the host plant and lasts about 20 days. Adult are found in the same habitat as larvae and cause similar feeding damage. Several different species as well as other genera may utilize the same leaf, so larval associations require rearing (Strong 1977a, b, 1982a, 1983; Strong and Wang 1977; Auerbach and Strong 1981; and Morrison and Strong 1981).

Seven other genera of Cephaloleiini containing 14 species are known from La Selva. Most of these species are poorly known and not associated with their host plant.

The tribe Arescini consists of four genera and 17 species from the Neotropics (Staines 2002b). One genus and two species are known from Mesoamerica. None of the genera have been revised and little work has been done on the biology. Gray and Baly are found at La Selva. Strong (1977a, 1983) reported the larval host plants of as Benth. and Griggs (Heliconiaceae). Strong (1983) reported on the biology of this species indicating that eggs are laid on wet, tender tissue of the host plant and hatch in about 20 days. Larvae begin feeding in rolled leaves immediately after hatching. Development is slow, requiring at least eight months until pupation. Larvae require more than one leaf-roll to complete development and move from maturing leaf-rolls to more tender ones at night. If they are between leaf-rolls at daylight, they hide between the petiole PageBreakand stalk until nightfall. Adults are polymorphic (in color and size) and long-lived; in mark-recapture studies adults were found 18 months after marking.

is known to feed on Hort. & Bull. (Marantaceae) and (Kuntze) Baker in Costa Rica (Maulik 1932). Strong and Wang (1977) and Auerbach and Strong (1981) reported as a larval host plant. The biology of this species is similar to that of .

The tribe Alurnini consists of six genera and 29 species (Staines 2002b) and contains some of the largest chrysomelids (25–45 mm). The tribe was revised by Fischer (1935) and I am in the process of revising it. Published life histories record various genera and species feeding on palms (Arecaceae) (Fischer 1935, Villacis Santos 1968, Macedo et al. 1994). Both Mesoamerican species, Baly and Baly, have been collected at La Selva. is the more commonly collected species.

The New World tribe Prosopodontini contains the genus Baly with 26 species found from Nicaragua to Ecuador (Staines 2002b). The genus is in need of revision.

Two species, (Baly) and (Baly), have been collected at La Selva. McCoy (1984, 1985) reported (as ) adults and larvae feeding in accumulated leaf debris on the top of leaves in Costa Rica and Ecuador. All other species of have been reported as leaf-miners on various Arecaceae (Jolivet and Hawkeswood 1995). The photograph in McCoy (1984) is a adult however the larval photograph does not resemble the known larvae (Maulik 1931). All other species of are associated with Arecaceae and I have only found on unfurled palm fronds, never on .

The tribe Sceloenoplini contains five genera and 299 species, with 154 species in the genus Chevrolat (Staines 2002b). They are leaf-miners in a variety of plant families. This tribe is represented at La Selva by four genera and 20 species (see Table 1). There are 17 species of known from La Selva. The biology is unknown for all species.

The tribe Chalepini consists of 55 genera and nearly 1000 species in the New World (Staines 2002b). Very few genera have been revised. All species studied are leaf-miners and appear to prefer dicots (Jolivet and Hawkeswood 1995). This tribe is represented at La Selva by 18 genera and 55 species (see Table 1). is the most speciose genus with 14 species.

Habitat specificity

Hispines can be found in most non-aquatic habitats at La Selva. There are 46 species which feed on rolled leaves and inflorences of Zingiberales. This one feeding guild accounts for 33% of the hispine species known from La Selva.

Most hispines species seem to be restricted to understory to mid-canopy level plants. Work on hispines has shown many species to be monophagous or narrowly oligophagous. These species are found mostly in relation to their host plants. Other hispines are broadly oligophagous or polyphagous and can be found in many habitats. PageBreakA continuing problem in inventory work is determining if the specimen collected was actually on its host plant or was a transient. Much of the earlier literature on host associations does not specify whether the insect was feeding as an adult, was breeding on the plant, or merely resting on it.

Relatively few species have only been collected from canopy fogging but these have been almost always undescribed species. Some of these species may actually be breeding in epiphytes rather than the fogged tree. (Olivier) and Guérin-Méneville have been associated with bromeliads (Bromeliaceae) in South America (Lowman et al. 1996; Mantovani et al. 2005).

Biogeography

Most of the La Selva hispine fauna is closely related to South American species. Some species have distributions throughout the Neotropics such as (Boheman), (Fabricius), and Fabricius. However, the genera Weise and Uhmann are most speciose in the Nearctic and the La Selva specimens are part of the southern extension of the genera (Staines 2002b). No La Selva hispines are exotic.

Many species appear to be Central American Atlantic lowland wet forest endemics but with congeners in South America. Staines, Staines, Staines, Staines, and Staines appear to fall into this category.

Specimen identification

Of the 139 hispine species known from La Selva, 125 (89.9%) are described species with published names, one is a morphospecies which is known to be new, and 14 (11.2%) are morphospecies in groups whose taxonomy is too poorly known to determine whether they are new or not.

La Selva hispine species can be identified using the key to the genera in Staines (2002b). All genera and species of La Selva hispines are in the “hispines of La Selva” web site (http://viceroy.eeb.uconn.edu/ALAS/ALAS.html). This site includes a summary of hispines, species lists, keys to species, references to revisions and other taxonomic publications, and individual species accounts with images and natural history data.

Suggestions for future Research

What do hispines eat? A little more than half (63.3%) of La Selva hispines have any host plant association. Many of these have only been noted as being collected on a plant rather than actually feeding on it (listed as adult on Table 1). Since hispines arePageBreak intimately tied to their host plant, determining the food plant will give a much better picture of their distribution and abundance. Additional leaf-miner rearing work such as that of Hespenheide and Dang (1999) is needed to make larval host plant associations. Johnson (2004a, b), Johnson and Horvitz (2005), García-Robledo and Horvitz (2009, in press), and García-Robledo et al. (2010) worked on the biology and ecology of several species at La Selva.

What is the biology and ecology of hispines? Very little work has been done on the biology and ecology of La Selva hispines. Kirkendall (1984) studied the mating behavior of the North American (Thunberg). Eberhard (1994) mentioned a hispine in his study of insect and spider courtship behavior. Staines and Staines (2001) and Flowers and Hanson (2003) suggested chrysomelids as potential indicator species assemblages for natural area monitoring. Farrell and Erwin (1988) showed that chrysomelids are a good indicator of local species richness. None of these ideas have been applied to hispines at La Selva.

What are the hispine host plant interactions? Strauss (1988) demonstrated that chrysomelids are a useful group for studing these interactions. Some work by Strong and his students (Strong 1977a, 1977b, 1981, 1982a, 1982b, Strong and Wang 1977), Horvitz and Schemske (2002), García-Robledo and Horvitz (2009, in press), and García-Robledo et al. (2010) have added to our knowledge of this but much remains to be done.

How do pathogens, predators, and parasitoids influence hispine populations? Hispines are parasitized by various wasps and flies (Cox 1994) and mites (Santiago-Blay and Fain 1994). They also have a few recorded predators (Cox 1996) and pathogens (Balazuc 1988, Hazarika and Puzari 1990). Memmott and Godfray (1993), Memmott et al. (1993), and Lewis et al. (2002) developed food and parasitism webs for some hispine species. A great deal of work needs to be done on how these organisms interact and what effect they have on hispine populations and distribution.

How do hispine populations and distributions change over time? Staines (2004b) studied the changes in chrysomelid populations over time on Plummers Island, Maryland. With the baseline inventory data and local knowledge at La Selva, a similar project could be started.