**2.1.2 Yeasts associated with digestive tract of ants and termites**

Ants, belonging like bees to the order *Hymenoptera*, have been also associated closely with variety of yeasts harbouring their nests (Carreiro et al., 1997; Rodrigues et al., 2009; Pagnocca et al., 2010). Some yeasts can pass into the infrabuccal pocket, a pouch in the ants'

Microorganisms live in the diverse habitats of the world. In course of evolution, some microbes adapted to the extreme environment prevailing in the gastrointestinal ecosystem of human and animals. Gastrointestinal tracts of mammals (Hooper & Gordon, 2001; Bauer et al., 2006; Ley et al., 2008) and insects (Dillon & Dillon, 2004; Hongoh, 2010; Grunwald et al., 2010) harbour vast bacterial communities which undoubtedly play an important role for the maturation and proper function of mucosal and systemic immune systems, nutrient metabolism and host health. In contrast, the knowledge of yeasts which naturally occur in

Insects are among the most diverse group of animals that has been found worldwide (Chapman, 2009) and they unavoidably come into contact with yeasts widespread in various habitats like soil (Botha, 2006), plants (Fonseca & Inàcio, 2006) and fresh and marine water (Nagahama, 2006). This has been confirmed by the fact that yeasts can be found on body surface as well as in the entrails of insects: beetles, bees, flies, lacewings, termites, and mosquitoes; and their larvae. Table 1 summarises the yeast species that were recently discovered in the GIT of insects and that have particularly been identified using molecular

In general, insects can be considered as either as a vector carrying yeasts on the body surface or the consumers of the yeasts (for review see Phaff & Starmer, 1987; Ganter, 2006). For instance, since yeasts regularly occur in flowers they are considered as autochthonous for this environment, and so they are closely associated with flower-visiting insects (Lachance et al., 2001; Brysch-Herzberg, 2004). In floristic nectar, ascomycetous yeasts belonging to the genera *Metschnikowia*, *Kodamaea, Wickerhamiella* have been found in higher abundance, whereas basidiomycetous yeasts (e.g. *Cryptococcus* spp., *Rhodotorula* spp., *Pseudozyma* spp.) were rarely isolated (Lachance et al., 2001; Brysch-Herzberg, 2004). Brysch-Herzberg (2004) counted an astonishing number of yeast cells (up to 16,000 cells/µl nectar) from the nectar samples of *Digitalis purpurea*. In this study, yeasts were isolated from nectar, plant materials, honey and from body of bumblebees, but unfortunately not from the GIT. In another study (Batra et al., 1973), the same yeasts (species of *Candida, Endomycopsis, Oidium, Hansenula, Rhodotorula, Saccharomyces, Schizosaccharomyces, Pichia,* and *Zygosaccharomyces*) were found in nectar and in the crops of bees, however, in the last niche yeasts were determined to be in higher, about 10 to 100 fold, concentration. Gilliam et al. (1974) summarized the data of yeasts isolated from the digestive tract of honey bees and also reported about their own

Seven species were observed in the study, but three: *Candida* (*Torulopsis*) *magnoliae*, *Candida parapsilosis*, and *Candida* (*Torulopsis*) *glabrata* were found most frequently providing evidence

Ants, belonging like bees to the order *Hymenoptera*, have been also associated closely with variety of yeasts harbouring their nests (Carreiro et al., 1997; Rodrigues et al., 2009; Pagnocca et al., 2010). Some yeasts can pass into the infrabuccal pocket, a pouch in the ants'

**2. Biodiversity of yeasts in the gastrointestinal ecosystem** 

intestine and thereby belong to the intestinal microbiota still remains deficient.

**2.1 Yeasts' diversity in the alimentary tract of insects** 

**2.1.1 Yeasts associated with flowers and gut of bees** 

investigations on yeasts in intestines of 388 adult worker honey bees.

**2.1.2 Yeasts associated with digestive tract of ants and termites** 

for their dominance in this environment.

methods.


Biodiversity of Yeasts in the Gastrointestinal

*Candida kunorum* 

*Candida lessepsii* 

*Candida lycoperdinae* 

sp. nov.

sp. nov.

sp. nov.

*Candida maxii* 

*Candida michaelii* 

*Candida pallodes* 

*Candida panamensis* 

*Candida panamericana* 

*Candida picachoensis*

*Candida pimensis*

*Candida plutei* 

sp. nov.

sp. nov.

sp. nov.

*Candida labiduridarum* 

Ecosystem with Emphasis on Its Importance for the Host 281

**Species Host organism References**  *Candida kruisii* sap-feeding beetles (*Nitidulidae*) (Suh et al., 2006b)

sp. nov. sap-feeding beetle (*Nitidulidae*) (Suh et al., 2004b)

earwig (*Labiduridae*); cricket

sap-feeding beetle (*Nitidulidae*);

(*Passalidae*); scarab beetle

sp. nov. darkling beetle (*Tenebrionidae*) (Suh et al., 2004b)

(*Corydalidae*); green lacewings

*Candida nodaensis* mosquitoes (*Culicidae*) (Gusmão et al., 2010) *Candida oregonensis* bark beetles (*Scolytinae*) (Rivera et al., 2009)

sp. nov*.* sap-feeding beetles (*Nitidulidae*) (Suh et al., 2006b)

beetle (*Tenebrionidae*)

sp. nov*.* green lacewings (*Chrysopidae*) (Suh et al., 2004a)

*Candida piceae* bark beetles (*Scolytinae*) (Rivera et al., 2009)

sp. nov. green lacewings (*Chrysopidae*) (Suh et al., 2004a)

sp. nov. rove beetle (*Staphylinidae*) (Suh & Blackwell,

*Candida pseudorhagii* click beetle (*Elateridae*) (Suh et al., 2008) *Candida quercitrusa* dobsonfly (*Corydalidae*) (Nguyen et al., 2007)

sap-feeding beetles (*Nitidulidae*);

pleasing fungus beetle (*Erotylidae*); rove beetle (*Staphylinidae*); darkling

handsome fungus beetle

*Candida maltosa* click beetles (*Elateridae*); bess beetles

*Candida membranifaciens* cranefly (*Tipulidae*); dobsonfly

(*Scarabaeidae*)

(*Chrysopidae*)

(*Gryllidae*); owlflies (*Ascalaphidae*) (Suh et al., 2008)

unidentified beetle (Suh et al., 2005b)

bark beetles (*Scolytinae*) (Rivera et al., 2009)

scarab beetle (*Scarabaeidae*) (Suh et al., 2006b)

(*Endomychidae*) (Suh et al., 2005b)

darkling beetle (*Tenebrionidae*) (Suh et al., 2006b)

leaf beetles (*Chrysomelidae*) (Nguyen et al., 2006)

cotton bollworm (*Noctuidae*) (Molnar et al., 2008)

(Suh et al., 2008)

(Suh et al., 2005b; Nguyen et al., 2007)

(Suh et al., 2004b)

(Nguyen et al., 2006)

2005)


pleasing fungus beetles (*Erotylidae*); darkling beetles (*Tenebrionidae*)

longhorned beetle (*Cerambycidae*);

(*Labiduridae*); cricket (*Gryllidae*)

darkling beetles (*Tenebrionidae*); minute tree-fungus beetles (*Ciidae*);

fungus weevil (*Anthribidae*)

sp. nov. leaf beetles (*Chrysomelidae*) (Nguyen et al., 2006)

sp. nov. dobsonflies and fishflies (*Corydalidae*) (Nguyen et al., 2007)

sp. nov. dobsonflies (*Corydalidae*) (Nguyen et al., 2007)

pleasing fungus beetles (*Erotylidae*);

sp. nov. click beetle (*Elateridae*) (Suh & Blackwell,

handsome fungus beetle

handsome fungus beetle

*Candida ernobii* bark beetles (*Scolytinae*) (Rivera et al., 2009)

scarab beetle (*Scarabaeidae*)

handsome fungus beetle (*Endomychidae*); elephant beetle

sp. nov. sap-feeding beetles (*Nitidulidae*) (Suh et al., 2006b)

sp. nov. click beetles (*Elateridae*) (Suh et al., 2008)

*Candida intermedia* mosquitoes (*Culicidae*) (Ricci et al., 2011a)

pleasing fungus beetles (*Erotylidae*);

(*Scarabaeidae*)

tooth-necked fungus beetles

(*Derodontidae*)

(*Endomychidae*)

*Candida fermentati* sap-feeding beetle (*Nitidulidae*);

pleasing fungus beetles (*Erotylidae*);

scarab beetle (*Scarabaeidae*) (Suh et al., 2008)

minute tree-fungus beetle (*Ciidae*) (Suh et al., 2004b)

(*Endomychidae*) (Suh et al., 2005b)

fishfly (*Corydalidae*) (Nguyen et al., 2007) mosquitoes (*Culicidae*) (Gusmão et al., 2010)

scarab beetle (*Scarabaeidae*) (Suh et al., 2004b)

(Suh et al., 2004b)

(Nguyen et al., 2007; Suh et al., 2008)

(Suh et al., 2004b)

(Suh & Blackwell,

(Suh et al., 2004b)

(Suh & Blackwell,

(Suh et al., 2008)

2005)

2004)

2004)

**Species Host organism References** 

*Candida* cf *neerlandica* owlfllies (*Ascalaphidae*); earwig

*Candida bribrorum* 

*Candida buenavistaensis* 

*Candida chickasaworum* 

*Candida choctaworum* 

*Candida chrysomelidarum* 

*Candida corydali* 

*Candida derodonti* 

*Candida dosseyi* 

*Candida elateridarum* 

*Candida emberorum* 

*Candida frijolesensis* 

*Candida gatunensis* 

*Candida gigantensis* 

*Candida guaymorum* 

*Candida endomychidarum* 

sp. nov.

sp. nov.

sp. nov.

sp. nov.

sp. nov.

sp. nov.

sp. nov.

sp. nov.

sp. nov.


Biodiversity of Yeasts in the Gastrointestinal

*Geotrichum carabidarum* 

*Geotrichum cucujoidarum*

*Geotrichum histeridarum* 

*Kodamaea laetipori* 

*Metschnikowia andauensis* 

*Metschnikowia chrysoperlae* 

*Metschnikowia corniflorae* 

*Metschnikowia noctiluminum* 

sp. nov.

sp. nov.

sp. nov.

sp. nov.

Ecosystem with Emphasis on Its Importance for the Host 283

ground beetle (*Carabidae*); geometrid larva (*Geometridae*); pleasing fungus

minute tree-fungus beetle (*Ciidae*); cucujoid beetle (*Cucujoidae*); hining

clown beetles (*Histeridae*); pleasing fungus beetle (*Erotylidae*); geometrid larvae (*Geometridae*); tiger moths

fungus beetle (*Scaphidiinae*)

*Hanseniaspora uvarum* crambid snout moths (*Crambidae*) (Molnar et al., 2008)

*Hanseniaspora vineae* dobsonflies and fishflies (*Corydalidae*) (Nguyen et al., 2007) *Issatchenkia orientalis* scarab beetle (*Scarabaeidae*) (Rao et al., 2007)

> darkling beetles (*Tenebrionidae*); scarab beetles (*Scarabaeidae*)

*Kuraishia capsulata* bark beetles (*Scolytinae*) (Rivera et al., 2009) *Kuraishia cf. molischiana* bark beetles (*Scolytinae*) (Rivera et al., 2009) *Lachancea fermentati* dobsonflies and fishflies (*Corydalidae*) (Nguyen et al., 2007) *Lachancea thermotolerans* dobsonflies and fishflies (*Corydalidae*) (Nguyen et al., 2007) *Lodderomyces elongisporus* bark and ambrosia beetle (*Scolytinae*) (Suh et al., 2008)

sp. nov. cotton bollworm (*Noctuidae*) (Molnar & Prillinger,

sp. nov. soldier beetles (*Cantharidae*) (Nguyen et al., 2006)

cotton bollworms (*Noctuidae*)

sp. nov. green lacewings (*Chrysopidae*) (Nguyen et al., 2006) *Metschnikowia pulcherrima* dobsonfly (*Corydalidae*) (Nguyen et al., 2007)

sp. nov. green lacewings (*Chrysopidae*) (Suh et al., 2004a)

beetle (*Nitidulidae*); pleasing fungus

leaf beetle (*Chrysomelidae*) (Molnar et al., 2008)

mosquitoes (*Culicidae*) (Ricci et al., 2011a)

mosquitoes (*Culicidae*) (Gusmão et al., 2010)

dobsonfly (*Corydalidae*) (Nguyen et al., 2007)

green lacewings (*Chrysopidae*) (Woolfolk & Inglis,

(Suh & Blackwell,

(Suh & Blackwell,

(Suh & Blackwell,

(Suh & Blackwell,

(Suh & Blackwell, 2005; Nguyen et al.,

2006)

2006)

2006)

2005)

2007)

2005)

2005)

2004)

(Molnar & Prillinger,

**Species Host organism References** 

beetle (*Erotylidae*)

(*Arctiidae*)

*Kodamaea ohmeri* dobsonfly (*Corydalidae*); sap-feeding

*Metschnikowia fructicola* crambid snout moths (*Crambidae*);

beetle (*Erotylidae*)

*Cryptococcus zeae* crambid snout moths (*Crambidae*);


(*Endomychidae*); cerambycid larvae

endomychid larva (*Endomychidae*);

bark-gnawing beetle (*Trogossitidae*);

elephant beetle (*Scarabaeidae*); pygmy

pleasing fungus beetles (*Erotylidae*);

(*Passalidae*); ichneumon wasps (*Ichneumonidae*); dobsonfly and fishfly (*Corydalidae*); roach (*Blattidae*)

sp. nov. pleasing fungus beetle (*Erotylidae*) (Suh et al., 2004b)

sp. nov. pleasing fungus beetle (*Erotylidae*) (Suh et al., 2004b) *Clavispora lustansiae* blister beetle (*Meloidae*) (Rao et al., 2007)

beetles (*Chrysomelidae*)

cotton bollworm (*Noctuidae*); leaf

*Cryptococcus oeirensis* leaf beetle (*Chrysomelidae*) (Molnar et al., 2008) *Cryptococcus victoriae* crambid snout moth (*Crambidae*) (Molnar et al., 2008)

Iphiclus beetle (*Erotylidae*)

sp. nov. sap-feeding beetles (*Nitidulidae*) (Suh et al., 2006b)

sp. nov. darkling beetle (*Tenebrionidae*) (Suh et al., 2004b)

*Candida tenuis* minute tree-fungus beetle (*Ciidae*) (Suh et al., 2005b)

sp. nov. pleasing fungus beetle (*Erotylidae*) (Suh et al., 2004b)

cotton bollworm (*Noctuidae*) (Molnar et al., 2008)

bess beetle (*Passalidae*) (Suh et al., 2005b)

grasshopper (*Tetrigidae*) (Suh et al., 2008)

scarab beetle (*Scarabaeidae*) (Suh et al., 2006b)

leaf beetle (*Chrysomelidae*) (Molnar et al., 2008)

leaf beetles (*Chrysomelidae*) (Molnar et al., 2008)

green lacewings (*Chrysopidae*) (Woolfolk & Inglis,

green lacewings (*Chrysopidae*) (Woolfolk & Inglis,

(Suh et al., 2005b)

(Suh & Blackwell,

(Nguyen et al., 2007; Suh et al., 2008)

(Molnar et al., 2008)

2004)

2004)

2004)

**Species Host organism References** 

(*Cerambycidae*)

*Candida sake* crambid snout moths (*Crambidae*);

*Candida tropicalis* owlfly (*Ascalaphidae*); bess beetles

*Candida xestobii* crambid snout moths (*Crambidae*);

*Cryptococcus flavescens* crambid snout moth (*Crambidae*);

*Cryptococcus luteolus* crambid snout moths (*Crambidae*);

*Candida sinolaborantium* handsome fungus beetle

*Candida smithsonii* 

sp. nov.

sp. nov.

sp. nov.

sp. nov.

*Candida stri* 

*Candida taliae* 

*Candida temnochilae* 

*Candida terraborum*

*Candida tetrigidarum* 

*Candida wounanorum* 

*Candida yuchorum* 

*Candida tritomae* 


Biodiversity of Yeasts in the Gastrointestinal

Ecosystem with Emphasis on Its Importance for the Host 285

oral cavity (Hansen & Klotz, 2005). Based on physiological characteristics using a microbial identification system BIOLOG, Mankowski & Morrell (2004) identified 19 species from 155 yeast isolates collected from nest, surrounding soil and frass as well as from exoskeleton and infrabuccal pockets of carpenter ants. From 17 isolates found in the infrabuccal pockets, ten were identified as *Debaryomyces polymorphus* and other species (*Pichia guilliermondii*, *Candida ergatensis*, *Candida edax*, *Bulleromyces* spp. and *Cryptococcus laurentii*) occurred only once or twice. Besides of soil samples, *D. polymorphus* was the most often isolated yeast from the all analysed materials. Further social insects, such as termites may harbour high yeasts' numbers (107-108 cells/ml) in their gut (Schäfer et al., 1996). Schäfer et al. (1996) cultured 35 yeast isolates from the intestinal contents of termite species, *Zootermopsis nevadensis* and *Neotermes castaneus*, but the authors reported presence of only 15 yeast strains, as their enzymatic activity were significant to the study. These phenotypes were related to the genera *Candida*, *Sporothrix*, *Pichia* and *Debaryomyces*. In another study, *Debaryomyces hansenii*  and *Sporothrix albicans* as well as species of *Trichosporon* and *Rhodosporidium* could also be found in the hindgut of the termites from families: *Mastotermitidae*, *Hodotermitidae*,

In the gut of some maize' pests (*Diabrotica virgifera*, *Helicoverpa armigera* and *Ostrina nubialalis*), Molnar et al. (2008) isolated 97 yeast strains; furthermore they detected yeasts as well as other fungi of the genera: *Acremonium*, *Aspergillus*, *Cladosporium* and *Fusarium* by means of cloning and denaturing gradient gel electrophoresis (DGGE). The occurence of clones was given in percents. All methods reveald that *Metschnikowia* spp., closely related to *Metschnikowia pulcherrima*, *Cryptococcus* spp. (*Cr. luteolus, Cr. zeae* and *Cr. flavescens*) as well as *Candida* spp., bearing close similarity to *C. xestobii* or *C. sake,* and *Pseudozyma* spp. were the most frequently identified yeasts. *Pichia guiliermondii* and *Rhodotorula* species were less common. Some of occassionaly found yeasts e.g. *Aureobasidium pullulans, Candida quercitrusa, Hanseniaspora uvarum, Sprobolomyces coprosmae, Tilletiopsis washingtonensis* were detected however only via culturing. There are some publications reporting presence of the yeasts in the gut of mosquitos (*Diptera*: *Culicidae*), which are known to be vectors of many diseases in humans. Gusmão et al. (2007; 2010) identified *Pichia caribbica, Pichia guilliermondii, Pichia* (syn. *Kodamaea*) *ohmeri, Candida fermentati* and *Candida nodaensis* in the diverticulum of *Aedes aegypti*. Ricci et al. (2011a) investigated yeasts in the gut of *Anopheles stephensi* using molecular and cultivation-dependent methods. Forty six clones that expressed fragments of the 18S rRNA gene retrieved from the gut samples of 6 adults were sequenced. Eleven clones were identified as *Wickerhamomyces anomalus*, known also as *Pichia anomala*, while others could be assigned either to genus *Candida* or *Pichia* or to unidentified fungus. Moreover, 100 colonies were cultured from 10 insect speciemens, classified based on their morphology and identified as *Candida intermedia*, *Hanseniaspora uvarum* and *W. anomalus* (77%, 15% and 8% respectively) by sequencing analysis of 18S and 26S rRNA genes and ITS fragments. *W. anomalus* was detectable using both approaches. Furthermore, Ricci et al. (2011a; 2011b) observed the presence of *W. anomalus* in the midgut of different mosquitos species *Anopheles stephensi, Anopheles gambiae, Aedes albopictus* and *Aedes aegypti*  of both sexes as well as on larvae, pupae and gonads, thereby supposed close relationship

*Kalotermitidae* and *Rhinotermitidae* and roaches (Prillinger et al., 1996).

**2.1.3 Yeasts associated with the gut of some pests** 

between this yeast species and mosquitos.


Table 1. Yeasts detected in the alimentary tract of insects

*Pichia americana* bark beetles (*Scolytinae*) (Rivera et al., 2009) *Pichia canadensis* bark beetles (*Scolytinae*) (Rivera et al., 2009) *Pichia caribbica* pleasing fungus beetle (*Erotylinae*) (Rao et al., 2007)

*Pichia glucozyma* bark beetles (Scolytinae) (Rivera et al., 2009)

crambid snout moths (*Crambidae*);

*Pichia mexicana* bark beetles (*Scolytinae*) (Rivera et al., 2009)

*Pichia scolyti* bark beetles (*Scolytinae*) (Rivera et al., 2009)

*Pseudozyma flocculosa* cotton bollworms (*Noctuidae*) (Molnar et al., 2008) *Pseudozyma prolifica* cotton bollworms (*Noctuidae*) (Molnar et al., 2008) *Rhodotorula aurantiaca* leaf beetle (*Chrysomelidae*) (Molnar et al., 2008)

*Saccharomyces cariocanus* dobsonflies (*Corydalidae*) (Nguyen et al., 2007) *Saccharomyces cerevisiae* dobsonflies and fishflies (*Corydalidae*) (Nguyen et al., 2007) *Saccharomyces fermentans* fishfly (*Corydalidae*) (Nguyen et al., 2007) *Saprochete gigas* dobsonfly and fishfly (*Corydalidae*) (Nguyen et al., 2007) *Sporobolomyces coprosmae* leaf beetles (*Chrysomelidae*) (Molnar et al., 2008) *Tilletiopsis washingtonensis* leaf beetle (*Chrysomelidae*) (Molnar et al., 2008) *Torulaspora delbrueckii* fishfly (*Corydalidae*) (Nguyen et al., 2007)

bess beetles (*Passalidae*); scarab

darkling beetles (*Tenebrionidae*);

*anomalus* (*Pichia anomala*) mosquitoes (*Culicidae*) (Ricci et al., 2011a)

beetles (*Scarabaeidae*)

handsome fungus beetle

mosquitoes (*Culicidae*) (Gusmão et al., 2007)

owlfly (*Ascalaphidae*) (Nguyen et al., 2007) bark beetle (*Scolytinae*) (Rivera et al., 2009)

leaf beetle (*Chrysomelidae*) (Molnar et al., 2008) mosquitoes (*Culicidae*) (Gusmão et al., 2010)

(*Endomychidae*) (Suh et al., 2005b)

leaf beetles (*Chrysomelidae*) (Molnar et al., 2008)

leaf beetle (*Chrysomelidae*) (Molnar et al., 2008)

lower termite (*Mastotermitidae*) (Molnar et al., 2004)

bess beetles (*Passalidae*) (Gujjari et al., 2010)

(Fuentefria et al.,

2008)

**Species Host organism References** 

*Pichia guilliermondii* fishfly and dobsonfly (*Corydalidae*);

*Pseudozyma apsidi* cotton bollworms (*Noctuidae*);

*Rhodotorula glutinis* crambid snout moths (*Crambidae*);

Table 1. Yeasts detected in the alimentary tract of insects

*Pichia nakazawae var.* 

*Trichosporon insectorum* 

*Trichosporon xylopini* 

*Wickerhamomyces* 

sp. nov.

sp. nov.

sp. nov.

*Trichosporon mycotoxinivorans* 

*akitaensis* 

oral cavity (Hansen & Klotz, 2005). Based on physiological characteristics using a microbial identification system BIOLOG, Mankowski & Morrell (2004) identified 19 species from 155 yeast isolates collected from nest, surrounding soil and frass as well as from exoskeleton and infrabuccal pockets of carpenter ants. From 17 isolates found in the infrabuccal pockets, ten were identified as *Debaryomyces polymorphus* and other species (*Pichia guilliermondii*, *Candida ergatensis*, *Candida edax*, *Bulleromyces* spp. and *Cryptococcus laurentii*) occurred only once or twice. Besides of soil samples, *D. polymorphus* was the most often isolated yeast from the all analysed materials. Further social insects, such as termites may harbour high yeasts' numbers (107-108 cells/ml) in their gut (Schäfer et al., 1996). Schäfer et al. (1996) cultured 35 yeast isolates from the intestinal contents of termite species, *Zootermopsis nevadensis* and *Neotermes castaneus*, but the authors reported presence of only 15 yeast strains, as their enzymatic activity were significant to the study. These phenotypes were related to the genera *Candida*, *Sporothrix*, *Pichia* and *Debaryomyces*. In another study, *Debaryomyces hansenii*  and *Sporothrix albicans* as well as species of *Trichosporon* and *Rhodosporidium* could also be found in the hindgut of the termites from families: *Mastotermitidae*, *Hodotermitidae*, *Kalotermitidae* and *Rhinotermitidae* and roaches (Prillinger et al., 1996).
