Neotropical palm-inflorescence feeding moths (Lepidoptera ...

*Corresponding author 

Received: 25 February 2013 

Accepted: 29 March 2013 

The Journal 

of Research 

on the Lepidoptera 

tHe lepIDopterA reSeArCH FoUNDAtIoN, 1 MA y 2013 

Copyright: This work is licensed under the Creative Commons 

Attribution-NonCommercial-NoDerivs 3.0 Unported License. To 

view a copy of this license, visit http://creativecommons.org/ 

licenses/by-nc-nd/3.0/ or send a letter to Creative Commons, 

171 Second Street, Suite 300, San Francisco, California, 94105, 

USA. 

Volume 46: 1-21 

ISSN 0022-4324 (pr I N t) 

ISSN 2156-5457 (o N l I N e) 

Neotropical palm-inflorescence feeding moths (Lepidoptera: Batrachedridae, 

Blastobasidae, Cosmopterigidae, Gelechiidae, Pyralidae, Tineidae): a review 

of the literature and new records from Trinidad, West Indies 

MAt t H e w J.w. Co C k 1* A N D Dw A y N e H. BU r r I S 2 

1 CABI E-UK, Bakeham Lane, Egham, TW20 9TY, UK 

2 UNEP-GEF Project Mitigating the Treats of Invasive Alien Species in the Insular Caribbean, Ministry of Food Production, Land & 

Marine Affairs, Research Division, Central Experiment Station, Caroni North Bank Road, Centeno, Trinidad and Tobago; current 

address: #30 Bay View Boulevard, Bay View La-Romain, Trinidad and Tobago 

m.cock@cabi.org, mjwcock@btinternet.com, dwayne.h.b@gmail.com 

In t r o d u c t I o n 

Abstract. The insects associated with the inflorescences of palms (Arecaceae) and their roles as 

pollinators, pests and scavengers are not well known compared to foliage-feeders and trunk borers, 

especially in the Neotropical Region. A literature review of the Lepidoptera associated with palm 

inflorescences in the Neotropical Region is provided. In a preliminary survey of Lepidoptera 

associated with palm inflorescences in Trinidad (Trinidad and Tobago), more than one thousand 

moths of 12 species were reared from six of the 14 palm species sampled. Of the moths reared, three 

could not be identified beyond family, three could be identified only to genus, and the remaining 

six were identified to species. These six comprise two species that feed on flowers of Cocos nucifera 

and Roystonea oleracea (viz. Atheloca bondari, Batrachedra nuciferae) and four that are polyphagous, 

widespread detritivores (Xystrologa nigrivitta, Anatrachyntis rileyi, Erechthias minuscula, Phidotricha 

erigens). Anatrachyntis rileyi and E. minuscula are thought to be introduced moth species, while the 

others are indigenous. Biological observations are presented on the moths reared. 

Keywords: Anatrachyntis rileyi, Atheloca bondari, Batrachedra nuciferae, Erechthias minuscula, Holcocera 

sp., Neodavisia sp., Phidotricha erigens, Xystrologa nigrivitta, Arecaceae, coconut. 

The insects associated with palms (Arecaceae), 

especially Cocos nucifera (coconut), have been relatively 

well documented (Lepesme, 1947; Lever, 1979; Howard 

et al., 2001), but those associated specifically with the 

inflorescences and their roles as pollinators, pests and 

scavengers are less well known. Traditionally, palms 

were thought to be wind pollinated, but it has become 

clear in recent decades that insect pollination does occur 

and may well be the norm rather than the exception 

(Moore, 2001). The floral ecology is variable among 

species (Henderson, 1986). Some have hermaphrodite 

flowers, others are monoecious (containing flowers 

of both sexes) in the same inflorescence, and others 

are dioecious (each palm bears either male or female 

inflorescences). Some maintain male flowers for several 

days, and stagger their maturation; others mature all 

the male flowers on an inflorescence at once and drop 

them all within a day. A priori, the former are likely to 

be suitable for the development of insects, whereas the 

latter are not (Moore, 2001). 

The caterpillars of Lepidoptera found in palm 

inflorescences are either herbivorous or detritivorous 

species. The plant feeding species range from those that 

feed purely on pollen in the male flowers, to those that 

feed on the structures of the male flowers, female flowers, 

developing nuts, and inflorescence branches. Obviously, 

these have different implications for the fecundity of the 

palms, and production of nuts where this is an economic 

consideration, e.g. coconuts. However, as yet very little 

is known about the ability of palms to compensate for 

insect damage to their inflorescences (Cock et al., 1987; 

Waterhouse & Norris, 1987; Moore, 2001).

2 

The native palms of Trinidad and Tobago were 

treated by Comeau et al. (2003). There are 22 native 

species in 15 genera and they are an important feature 

of the country’s landscape and biodiversity. Coconut 

is an introduced species in Trinidad of many years 

standing. It is found in cultivation throughout the 

island, usually near dwellings or former dwellings, but 

has not naturalised on any great scale. 

This paper summarises published information on 

the Lepidoptera associated with the inflorescences 

of coconut and other palms in South America, and 

reports new observations from a preliminary survey of 

several indigenous and introduced palms in Trinidad, 

West Indies. The survey was focused on Batrachedra 

nuciferae Hodges (Lepidoptera: Batrachedridae), to 

establish its status and host range in Trinidad, to help 

assess the need and options for its management, as 

reported in Cock (2013). A second component of 

the survey was to collect information on what other 

species of small Lepidoptera feed on the same host 

palms as B. nuciferae, so that early stages and damage 

by B. nuciferae could be distinguished from those of 

other Lepidoptera. Here we present observations on 

the biology and host range of the Lepidoptera species 

associated with selected palms in Trinidad. 

Me t h o d s 

A literature survey was made of the known 

palm-inflorescence feeding Lepidoptera in the 

Neotropical Region. This was based on key reference 

works (Lepesme, 1947; Howard et al., 2001) and the 

references therein, and by searching CAB Abstracts 

to establish which species have been recorded from 

palm inflorescences in the area. Once the names of 

Lepidoptera species were found, further searches were 

made, including consulting the original descriptions 

and selected taxonomic and general works that might 

deal with these species, as well as internet searches, 

and relevant references compiled from these sources. 

The collection of the Natural History Museum, 

London (BMNH) was examined and biological and 

distribution data recorded for relevant species. 

The field and laboratory methods and palms 

sampled are described in Cock (2013), and only 

outlined here. The fourteen species of palms that 

were sampled are presented in Table 1. Whole 

inflorescences and/or dropped male flowers were 

taken for each sample (Table 2). They were inspected 

visually in the laboratory, and caterpillars and cocoons 

on them were documented. The samples were then 

set up in emergence boxes for whole inflorescences 

or in plastic rearing containers for subsamples to rear 

out adult moths, which were recorded daily. 

J. Res.Lepid. 

Because the field work was carried out during a 

short period in October 2011, it was not possible to 

sample all palm inflorescences at the same stage of 

development, and this will have affected the species 

and numbers of moths obtained. The condition of 

each sample is listed in Table 2 and notes on each 

sample are provided. 

Samples of coconut flowers were obtained from 

five localities. Coconut is monoecious, and because 

it flowers continuously (Child, 1974), there was little 

difficulty taking samples with pollen-producing male 

flowers that were attracting insects. 

The next seven species of palm are considered 

indigenous (Comeau et al., 2003). Male and female 

flowers of Attalea maripa are born on separate 

inflorescences, but the floral biology and ecology of 

the genus is poorly understood and may vary with 

the age of the palm (Henderson et al., 1995). The 

male flowers of A. maripa consist of little more than 

a bunch of pollen-bearing stamens, so they are very 

different in structure from all the other palms that 

we sampled. Although A. maripa was common at 

Bush Bush Island in Nariva Swamp, most palms were 

too tall to sample, and only an unopened spathe 

and an old dead male inflorescence from a shorter 

palm could be sampled. The unopened spathe was 

unblemished, with no signs of insect damage. 

Bactris major is monoecious; the male flowers 

release pollen for about 24 hours and then are 

dropped immediately (Essig, 1971; Henderson, 

1986). It seems likely that this short period of pollenproducing 

male flowers would make it very difficult 

for a pollen-feeding caterpillar to establish itself and 

survive. No trace of Lepidoptera feeding was found on 

the inflorescences sampled, and no moths were reared. 

Caterpillars, particularly of detritivorous species, 

theoretically could develop on the fallen flowers, but 

a 40ml sample of newly fallen flowers collected at the 

Botanic Gardens showed no signs of Lepidoptera 

feeding (visual inspection and dissection of ten 

flowers), and none were reared. Bactris simplicifrons 

has small inflorescences, each having only a few male 

and female flowers. We did not find any inflorescences 

with male flowers, but two inflorescences that had 

recently dropped their male flowers showed no trace 

of Lepidoptera or other insect damage. 

All the male flowers had dropped, and only green 

developing nuts remained on the inflorescence sample 

of Euterpe precatoria. Examination of the branches 

from which the male flowers had dropped showed no 

signs of Lepidoptera damage, and specifically no sign 

of male flowers attached to the inflorescence by silk, 

or of cocoons. One of the hundreds of developing 

nuts was black rather than green and it contained

46: 1-21, 2013 

Table 1. The palms (Arecaceae) of which inflorescences were sampled, Trinidad, October 2011. Classification follows 

Dransfield et al. (2008) as presented by Trebrown Nurseries (2011); common names and their distributions are based on 

Comeau et al. (2003). 

Subfamily, Tribe (subtribe) Species 

Arecoideae, Areceae 

(Ptychospermatinae) 

Arecoideae, Areceae 

(Ptychospermatinae) 

Arecoideae, Cocoseae 

(Attaleinae) 


(Attaleinae) 


(Attaleinae) 


(Bactridinae) 

a small Lepidoptera caterpillar. It was not reared 

successfully and no moths were obtained from the 

emergence box. 

Individual Mauritia flexuosa palms are dioecious; 

they bear inflorescences of either male or female 

flowers, not both (Ervig, 1993). We found one male 

palm at Aripo Savannah that was short enough 

to sample. It had several young, yellow-green 

inflorescences whose flowers had not yet opened, 

and several old dead, dark brown inflorescences; we 

collected one of each. The young inflorescence showed 

no sign of Lepidoptera damage when inspected, 

although moths were obtained by emergence box. 

Some feeding damage and very small caterpillars were 

found under the bracts of the old inflorescence, and 

adult moths were obtained by emergence box. 

Roystonea oleracea is monoecious. An unusual 

feature of the genus is that from before the spathe 

Common name in 

Trinidad 

Distribution in Trinidad 

Adonidia merrillii Manila palm Introduced ornamental; widespread and 

common 

Ptychosperma macarthurii Hurricane palm Introduced ornamental; widespread and 

common 

Cocos nucifera Coconut Introduced; cultivated in all except highest 

parts 

Attalea maripa Cocorite Widespread except highest parts 

Syagrus romanzoffiana Queen palm Introduced ornamental 

Bactris major Roseau, picmoc, 

black roseau 

Bactris simplicifrons Yuyu Local, scattered 


(Bactridinae) 


(Euterpeae) 

Arecoideae, Roystoneae 

Calamoideae Calameae 

(Calaminae) 

Roystonea oleracea Palmiste, royal palm, 

cabbage palm 

Calamoideae, Lepidocaryeae 

(Mauritiinae) 

Widespread but absent from Northern 

Range and Long Stretch 

Euterpe oleracea Manac Widespread at low elevations 

Possibly introduced; S & E coasts, 

widespread as cultivated 

Calamus sp. Rattan palm Introduced ornamental 

Mauritia flexuosa Moriche Local especially Nariva Swamp, Long 

Stretch 

Coryphoideae, Sabaleae Sabal mauritiiformis Carat South, widespread 

Coryphoideae, Trachycarpeae 

(Livistoninae) 

Licuala spinosa Spiny licuala palm Introduced ornamental 

Coryphoideae, Trachycarpeae 

(unplaced) 

Pritchardia pacifica Fiji fan palm Introduced ornamental 

opens, it is packed with millions of very small, fluffy, 

branched hairs (Henderson et al., 1995). We were able 

to sample one inflorescence from a relatively short palm 

at Kernahan. In this inflorescence, the male flowers had 

recently dropped and the female flowers were small. 

Many of the male flowers and the fluffy matrix were 

caught up in the inflorescence and the spathe, which 

remained in place below the inflorescence. More than 

600 moths were reared from this one inflorescence. 

It seems likely that the fluffy matrix contained in 

inflorescences of R. oleracea provides a food source that 

can support many caterpillars of some of these moths, 

although we did not attempt to establish this. 

Sabal mauritiiformis is unusual amongst the 

indigenous palms sampled in that the small flowers 

are bisexual (Henderson et al., 1995), and hence they 

are not dropped once pollen production is complete. 

Two palms were examined at Kernahan; one had an 

3

4 

Date Palm species Location GPS Details 

8 Oct Adonidia merrillii* Curepe, CABI N10°39.159 W61°24.065 1 inflorescence 

12 Oct Attalea maripa Nariva Swamp, Bush 

Bush Is. (North) 

inflorescence that included open male flowers and 

this was sampled. Small caterpillars of a Batrachedra 

sp. were found. 

The following six palms are introduced, ornamental 

species. Adonidia merrillii is monoecious, but the 

opening of the male flowers is staggered, so that only a 

few are open at a time. No sign of Lepidoptera feeding 

could be found in the inflorescence of the palms 

sampled, and none were obtained by emergence box. 

J. Res.Lepid. 

Table 2. Collections of palm inflorescences made in Trinidad, 8-18 October 2011. All sample sites were less than 50m asl. 

N10°23.726 W61°02.381 1 unopened male inflorescence, 1 old 

male inflorescence 

18 Oct Bactris major Botanic Gardens N10 40°426 W61°30.913 Fallen male flowers 

12 Oct Bactris major Nariva Swamp, Bush 


N10°23.730 W61°02.375 1 old inflorescence 

14 Oct Bactris simplicifrons Nariva Swamp, Bush 

Bush Is. (South) 

N10 23°390 W61°02.711 2 old inflorescence 

18 Oct Calamus sp.* Botanic Gardens N10 40°460 W61°30.888 2 old inflorescence 

12 Oct Cocos nucifera Nariva Swamp, Bush 


1 inflorescence 

10 Oct Cocos nucifera Centeno Fallen male flowers 

8 Oct Cocos nucifera Curepe, CABI (palm 

NL1) 

N10°39.182 W61°24.069 1 inflorescence 


TRT08) 



TRT09) 

1 inflorescence 

12 Oct Cocos nucifera Nariva Swamp, 

Kernahan** 

N10 21°490 W61°00.856 1 inflorescence 

10 Oct Cocos nucifera Waller Field, Gafoor’s 

Plantation 


14 Oct Euterpe precatoria Aripo Savannah N10 35°476 W61°11.338 1 old inflorescence 

18 Oct Licuala spinosa* Botanic Gardens N10 40°424 W61°30.913 1 old inflorescence 

14 Oct Mauritia flexuosa Aripo Savannah, KP N10 36°352 W61°12.384 1 young inflorescence, 1 old 

Quarry section 

inflorescence 

18 Oct Pritchardia pacifica* Botanic Gardens N10 40°427 W61°30.984 2 inflorescence (pooled) 

18 Oct Ptychosperma macarthurii* Botanic Gardens N10 40°423 W61°30.939 1 inflorescence 

8 Oct Ptychosperma macarthurii* Curepe, CABI N10°39.170 W61°24.086 several bits of inflorescence 

18 Oct Roystonea oleracea Botanic Gardens N10 40°428 W61°30.924 Fallen male flowers 

12 Oct Roystonea oleracea Nariva Swamp, 

Kernahan** 


12 Oct Sabal mauritiformis Nariva Swamp, 

Kernahan** 


18 Oct Syagrus romanzoffiana* Botanic Gardens N10 40°422 W61°30.933 1 inflorescence; fallen male flowers 

*Introduced ornamental species. 

**Also spelt Kernaham. 

Our sample of Calamus sp. in the Royal Botanic 

Gardens was two old, dry male inflorescences, 

comprising a densely packed mass of dry dead 

flowers. Examination showed some webbing and 

caterpillar frass in most parts of the inflorescence, and 

caterpillars with a dark head and dark purplish body 

were associated with this. Two cocoons were found 

but nothing emerged from these or in the emergence 

box, so we cannot confirm whether this species is a

46: 1-21, 2013 

suitable food plant for any of the Lepidoptera found 

in our survey, but certainly it is host to at least one 

species of Lepidoptera. 

Licuala spinosa has bisexual flowers, and although 

each f lower produces pollen only brief ly, the 

inflorescence produces mature flowers over a period 

of about a month (Barfod et al., 2003). We were able 

to obtain only an old inflorescence, from which most 

of the flowers were long gone, and on which some nuts 

were beginning to develop. The inflorescence showed 

no sign of Lepidoptera feeding, webbing or cocoons, 

but probably it was too old to expect to see this. 

Pritchardia pacifica is monoecious. In addition to 

two inflorescences, we collected male flowers that 

had dropped and caught amongst the branches of 

an older inflorescence; there was light webbing and 

frass amongst these dropped male flowers and moths 

were obtained by emergence box. 

Ptychosperma macarthurii is monoecious, and the 

male flowers are not synchronised. Thus we were 

able to find small numbers of scattered male flowers 

on the inflorescences that we sampled, and some 

flowers that dropped and got caught up between 

the inflorescence branches in one sample. There 

was no trace of webbing or frass on either the fresh 

or dropped male flowers, and no Lepidoptera were 

obtained by emergence box. 

Syagrus romanzoffiana is monoecious but the one 

that we sampled had already dropped most of the 

male flowers. The inflorescence seemed completely 

healthy except that the ends of many apical 

secondary branches were trapped in the narrow 

apex of the spathe, together with many male flowers 

that had started to become mouldy. This situation 

seemed ideal for attack by Lepidoptera yet there was 

no trace of caterpillar damage. A small collection of 

dropped male flowers was also made from the base 

of the palm and set up separately. No Lepidoptera 

emerged from either sample. 

Obtaining samples was a significant challenge, 

due to limited availability of flowers at the selected 

sites in the week of the survey, and those palms which 

were flowering were often too tall to sample safely. 

The samples obtained were not necessarily at the best 

stage to look for flower feeding caterpillars, especially 

those that feed on pollen, such as Batrachedra spp. 

Nevertheless, because Lepidoptera feeding leaves 

recognisable traces, such as webbing, frass and cocoons, 

old inflorescences where the dead male flowers were 

still present provided useful information. 

Examination of inflorescences in the laboratory 

gave fairly reliable insight into the species present 

and their damage, but overlooked eggs or very young 

individuals, and risked overlooking very low density 

populations. As a measure of population density, 

taking a sample at a single time point as we did, 

unavoidably makes no allowance for eggs that would 

have been laid in the inflorescence after collection, 

nor for moths that had already emerged before 

the sample was taken. By carefully examining the 

material before setting it up in the emergence box, 

we have a qualitative control for the latter aspect, but 

we could not control for the former. 

Nevertheless, there was wide variation in the 

numbers of moths obtained from the emergence 

boxes, and large numbers of moths were obtained 

in several cases, so that this method does provide a 

time-efficient and cost-effective way to recognise the 

relative density of populations of Lepidoptera in the 

different inflorescences at the time of sampling. Any 

more rigorous approach would inevitably be much 

more expensive in time and money, so we believe this 

is a practical and appropriate approach. 

re s u lt s: lI t e r at u r e s u r v e y 

Until recently, there has been little work on the 

Lepidoptera that breed in palm inflorescences in the 

Neotropical Region, and almost none of a general 

nature. Hence, the early work of Bondar (1940a, 

1940b) in Bahia, Brazil, is still an important source 

of information on Lepidoptera that develop in the 

inflorescences of coconut (Cocos nucifera) and other 

palms. Bondar deals with Batrachedra nuciferae (as 

B. perobtusa Meyrick, a misidentification), and two 

more damaging species: Atheloca bondari Heinrich (as 

Hyalospila ptychis Dyar, a misidentification or synonym, 

see below) and Cadra cautella (Walker) (as Ephestia 

cautella). In addition, there is limited information 

on a small number of other moths recorded from 

coconut, mostly noted by Lepesme (1947) in his 

standard reference Les Insectes des Palmiers. The 

summary by Moore (2001) in the recent equivalent 

title Insects on Palms (Howard et al., 2001) adds very 

little for the Neotropical Region to what Bondar and 

Lepesme reported. Species not previously recorded 

from palm inflorescences were reared in this survey, 

and the available literature on these species is also 

summarised here. An overview of the literature survey 

is provided as Table 3. 

Atheloca sp(p). (Pyralidae, Phycitinae) 

Atheloca bondari was described by Heinrich (1956) 

in his revision of the American Phycitinae, based on 

Bondar’s specimens in the United States National 

Museum. There is a very similar species, A. subrufella 

(Hulst) (=Hyalospila ptychis), found in Florida (USA), 

5

6 

Cuba and the Virgin Is., which differs primarily in 

the absence of ‘a strong, dorsal, yellow hair tuft 

from the base of the male hind tibia’ found only in 

A. bondari (Heinrich, 1956). Heinrich illustrates 

the male and female genitalia of A. subrufella, 

which show ‘no essential differences’ from those of 

A. bondari. Pictures of adult moths of A. subrufella 

can be seen at MPGNA (2012). Schotman (1989) 

reports A. subrufella from French Guiana and St. 

Lucia. Although the two species were considered 

distinct in Shaffer’s (1995) treatment in the checklist 

of Neotropical Lepidoptera and though they do 

not seem to have been formally synonymised 

since (e.g. Beccaloni et al., 2003), they are likely 

to be synonyms (M.A. Solis, pers. comm., 2012). 

J. Res.Lepid. 

Table 3. An overview of the Lepidoptera associated with palm inflorescences in the Neotropical Region, based on this 

literature review. 

Species Classification Host plants Feeding on Distribution Key references 

Anatrachyntis rileyi 

(Walsingham) 

Atheloca bondari 

(Heinrich) 

Atheloca subrufella 

(Hulst) 

Batrachedra nuciferae 

Hodges 

Cadra cautella 

(Walker) 

Erechthias minuscula 

(Walsingham) 

Holcocera ochrobathra 

(Meyrick) 

Phidotricha erigens 

(Ragonot) 

Xystrologa nigrivitta 

(Walsingham) 

Cosmopterigidae Many plant hosts Detritus Old World 

origin, southern 

USA, Caribbean 

Pyralidae, 

Phycitinae 

Pyralidae, 

Phycitinae 

Cocos nucifera, 

Syagrus spp., Attalea 

spp. 

Cocos nucifera, Sabal 

palmetto, Serenoa 

repens 

Batrachedridae Cocos nucifera 

(unconfirmed: 

Syagrus spp., Attalea 

spp.) 

Pyralidae, 

Phycitinae 

Tineidae, 

Erechthiinae 

Blastobasidae, 

Holcocerinae 

Pyralidae: 

Epipaschiinae 

Tineidae, 

?Meesiinae 

Usually on stored 

products 

Male and female 

flowers 


flowers 

Pollen of 

male flowers 

(unconfirmed: 

female flowers) 


flowers 

Brazil, Trinidad 

(this survey) 

Florida, 

Caribbean 

(unconfirmed: 

French Guiana) 

Brazil, 

Venezuela, 

Trinidad, 

probably 

widespread in 

South America 

Many plant hosts Detritus Old World 

origin, southern 

USA, Caribbean 

Walsingham, 1882; 

Busck, 1917; Heinrich, 

1921; Hodges, 1978; 

Zimmerman, 1978 

Bondar, 1940a, 1940b; 

Heinrich, 1956. As A. 

subrufella: Santana, 2008; 

Santana et al., 2009, 2010, 

2011 

Heinrich, 1956; Kimball, 

1965; Habeck & Nickersen, 

1982; Schotman, 1989; 

Bento et al., 2006 

Bondar, 1940a, 1940b; 

Hodges, 1966; Arnal et al., 

1998; Sanchéz Soto, 2004; 

Sánchez-Soto & Nakano, 

2002, 2004a, 2004b, 2008; 

Cock 2013 

Cosmopolitan Bondar, 1940a, 1940b 

Walsingham, 1897; Swezey, 

1909; Lepesme, 1947; 

Clarke, 1971; Zimmerman, 

1978; Clarke, 1986 

Cocos nucifera Detritus Guyana, Florida Meyrick 1921; Bodkin 

1922; Adamski, 2002b; 

Heppner, 2003 

Polyphagous Reproductive 

parts and leaves of 

diverse plants 

Several plant hosts Bracket fungus, 

Roystonea oleracea 

inflorescence (this 

survey); probably 

diverse materials 

Neotropical Solis, 1993, 2011; Diniz & 

Morais, 2002 

Caribbean Davis et al., 2012 

Nevertheless, we maintain both names here in line 

with formal taxonomy and pending a critical study 

of the question. 

Bondar (1940a, 1940b) found that the caterpillars 

of A. bondari feed on both male and female flowers 

of coconut, as well as on the flowers of the native 

palms of the genera Syagrus (as Cocos) and Attalea. 

He illustrated the damage to male coconut flowers, 

which includes holes eaten through the sides of the 

male flowers; damage to the female flowers causes 

abortion or early nut fall. 

Cabbage palm (Sabal palmetto) and saw palmetto 

(Serenoa repens) (Arecaceae) have been reported 

as food plants of A. subrufella in Florida (Kimball, 

1965), and Habeck & Nickersen (1982) subsequently

46: 1-21, 2013 

described the biology on coconut thus: ‘Larvae feed 

on newly emerged inflorescences and destroy flower 

buds and young developing coconuts. Coconuts up 

to 1.5 inches in diameter may be completely hollowed 

out. Larvae spin silk over the inflorescences … and 

incorporate frass and plant material into the webs. 

Pupation occurs amid the plant material under the 

webbing.’ Bento et al. (2006) describe the biology 

and mating behaviour in more detail. S.W.J. de 

Santana studied the life history and bioecology of A. 

subrufella in Pernambuco State, Brazil for her PhD 

thesis (Santana, 2008), and published accounts of 

the rearing method (Santana et al., 2011), thermal 

biology (Santana et al., 2010) and interaction with 

coconut mite, Aceria guerreronis Keifer (Santana et al., 

2009). Santana (2008) and Bento et al. (2006) both 

treat Atheloca bondari and the reports on its biology as 

though it is a synonym of A. subrufella. 

Cadra cautella (Walker) (Pyralidae, Phycitinae) 

Cadra cautella is a well-known stored products 

pest found throughout tropical and temperate areas 

(Heinrich, 1956; Mound, 1989). Bondar (1940a, 

1940b) reports that the caterpillars feed on male 

and female flowers of ‘Cocos spp.’ (i.e. C. nucifera and 

Syagrus spp.) and Attalea spp. It should be noted that 

Cadra includes several similar species, with similar 

biology as stored product pests, and dissection of 

the genitalia is usually needed to confirm their 

identification (e.g. Goater, 1986; Mound, 1989); 

accordingly this identification needs species level 

confirmation. For the identification of the larvae, 

see Mound (1989) and Solis (2011). 

Phidotricha erigens (Ragonot) (Pyralidae: 

Epipaschiinae) 

Phidotricha erigens, described from Puerto Rico 

(Ragonot, 1888), is found throughout the Caribbean 

and adjoining mainland, south to Brazil (Solis, 

1993, 2011). In the past P. erigens has been treated 

as a synonym of Pococera atramentalis Lederer (1863). 

This would have been because Phidotricha erigens was 

identified as Pococera atramentalis in the collection 

of the Natural History Museum, London (BMNH). 

However, although the BMNH series of Pococera 

atramentalis are Phidotricha erigens (or were until 

curated), the holotype of Pococera atramentalis (also in 

the BMNH) is a different species (Solis, 1993). 

Phidotricha erigens is known to be polyphagous on 

the leaves and reproductive parts of a wide variety 

of plants including Anacardiaceae, Calophyllaceae, 

Cucurbitaceae, Fabaceae, Oxalidaceae, Malvaceae, 

Rutaceae, Passifloraceae, Phytolaccaceae, Poaceae, 

Rosaceae, Zingiberaceae (Solis, 1993, 2011), 

Burseraceae, Celastraceae, Erythroxylaceae, Fabaceae, 

Vochysiaceae (Diniz & Morais, 2002), Asparagaceae 

(Velázquez et al., 2010), and Caryocaraceae (Carregaro, 

2007), but not hitherto from Arecaceae. The 

possibility that these records represent several cryptic 

species with more specialised food plant preferences 

has not been investigated. 

This species has previously been reported from 

Trinidad, based on specimens reared by F.W. Urich 

as Pococera atramentalis (Kaye & Lamont, 1927). Some 

of F.W. Urich’s specimens are in the United States 

National Museum, Washington; they were reared 

in November 1922 from Albizia saman (Fabaceae). 

It also has been reared from flowers of Tephrosia 

sp. (Fabaceae), in October 1954 at St. Augustine, 

Trinidad, by F.D. Bennett (specimen in University of 

the West Indies Zoology Museum). The species occurs 

also in Tobago (M.J.W. Cock unpublished). 

Batrachedra spp. (Batrachedridae) 

The genus Batrachedra is usually placed in the family 

Batrachedridae of the Gelechioidea (Hodges, 1999; 

Brown et al., 2004; Kaila et al., 2011; van Nieukerken 

et al., 2011), but has also been placed in the subfamily 

Batrachedrinae of the Coleophoridae (Becker, 1984; 

Kaila, 2004). The family Batrachedridae is considered 

paraphyletic (Kaila, 2004; Kaila et al., 2011), so further 

changes may follow. 

Hodges (1966) revised the American Batrachedra 

spp. and recognised three groups of species within 

the genus. One of these groups includes three species 

feeding on palm inflorescences, one feeding on the 

base of the pineapple fruiting body, and six of unknown 

biology. Positive identification of these species is only 

possible based on dissection and examination of the 

genitalia, those of the female being more diagnostic 

than those of the male. Batrachedra nuciferae is 

discussed below; B. mathesoni Busck occurs in Florida 

and the caterpillars feed on coconut flowers; and B. 

decoctor Hodges also occurs in Florida where the palm 

Serenoa repens is a food plant. Other Batrachedra spp. 

of this group have been described from Puerto Rico, 

Jamaica, St Lucia and Central America, etc. Batrachedra 

arenosella (Walker) is the name applied to a Batrachedra 

species in South-East Asia and Australasia which also 

has been reported to feed on coconut inflorescence, 

causing insignificant damage (Corbett & Gater, 

1924; Kalshoven & van den Laan, 1981). However, 

most probably this name is incorrectly applied, as B. 

arenosella was described from New Zealand, where it is 

known as a scale predator (Moore, 2001). 

7

8 

At least two species of Batrachedra feed as caterpillars 

in the inflorescences of palms in Trinidad (Cock, 

2013), B. nuciferae on coconut and Roystonea oleracea, 

and an unidentified species on Sabal mauritiformis. 

It seems likely that other species of this group of 

American Batrachedra spp. will be found to feed on 

palms, probably showing some specialisation as to 

species or genera that are acceptable as food plants. 

Batrachedra nuciferae Hodges (Batrachedridae) 

Batrachedra nuciferae was first recognised by Bondar 

(1940a, 1940b) who described its biology in Bahia 

State, Brazil (as B. perobtusa). It was subsequently 

described as a new taxon, based on Bondar’s material 

reared from male coconut flowers, in Hodges’ (1966) 

revision of the American Batrachedra. The species 

description was based only on material reared 

from coconut and Hodges did not refer to material 

from the other palms which Bondar (1940a, 1940b) 

records as food plants. Moore (2001) summarises 

Bondar’s observations under the name Ifeda perobtusa, 

overlooking the name change in Hodges’ (1966) 

revision. In 1998, B. nuciferae was recorded from 

Venezuela as a new pest of coconut (Arnal et al., 1998). 

In 2006, B. nuciferae was correctly reported from 

Trinidad as a new pest of coconut (MALMR, 2006, 

2008), although the adult moth shown in MALMR 

(2008) is Anatrachyntis rileyi (Walsingham) (reported 

below as reared from palm inflorescences). 

Bondar (1940a, 1940b) reported that the caterpillars 

of B. nuciferae rest in the male flowers of coconut where 

they feed on pollen, and they are also common in the 

flowers of several other palms: Syagrus coronata (=Cocos 

coronata), S. vagans (=C. vagans), S. schizophylla (=C. 

schizophylla), Attalea funifera and A. piassabossu. He 

considered that the damage to male flowers reduced 

the probability of fertilization of female flowers and 

hence could adversely affect nut production, but 

presented no evidence for this conclusion. He gives 

brief descriptions of the caterpillar and pupa, states 

that the cocoon is formed on a solid substrate or 

amongst the fallen male flowers in the leaf axil below, 

and that the life cycle takes 15-18 days. 

Since 1940, there was almost no published work on 

the coconut moth, until the work of S. Sanchéz-Soto 

in São Paulo State, Brazil, this century. The moth was 

the subject of his research thesis (Sanchéz Soto, 2004) 

and publications on the distribution (Sánchez-Soto & 

Nakano, 2002, 2004a), morphology (Sánchez-Soto & 

Nakano, 2004b), and biology (Sánchez-Soto & Nakano, 

2008). The egg, caterpillar (including chaetotaxy), 

pupa and adult are illustrated in both Sánchez Soto 

(2004) and Sánchez-Soto & Nakano (2004b). 

J. Res.Lepid. 

Arnal et al. (1998) reported the presence of the 

moth in several parts of Venezuela. Carneiro et al. 

(2004) stated that in the Município de Parnaíba, 

Piauí, north-east Brazil, the caterpillars eat both 

male and female f lowers. Observations from 

Trinidad were reported by Cock (2013), who found 

no evidence that female flowers were damaged. 

His observations showed that the inflorescence of 

Roystonea oleracea is also used as a food source, but 

that no B. nuciferae were obtained from inflorescences 

sampled from 12 other indigenous and introduced 

palms, including Attalea maripa. 

Holcocera ochrobathra (Meyrick) (Blastobasidae, 

Holcocerinae) 

Although some recent works treat Blastobasidae 

as a subfamily of Coleophoridae (Hodges, 1999), it is 

retained as a family here in line with van Nieukerken 

et al. (2011). Species of Blastobasidae are usually 

considered to be scavengers or detritivores on a variety 

of substrates, but some are herbivorous (Adamski & 

Brown, 1989). The North American species have been 

revised and arranged in two subfamilies and several 

genera (Adamski & Brown, 1989), but the South 

American species are still poorly known (Adamski, 

2002b), apart from those of Costa Rica (Adamski, 

2002a, 2013). There are many undescribed species 

(Adamski & Brown, 1989; Adamski, 2002b). 

Meyrick (1921) described H. ochrobathra from 

Guyana, in the genus Blastobasis, based on specimens 

reared from coconut flowers by L.D. Cleare Jr. in 

1920, and stated that the type was in the ‘Brit. Mus.’ 

(BMNH). There are four such specimens in the 

BMNH, of which a male is designated lectotype and 

has been dissected and illustrated by Clarke (1963), 

and a female has been designated paralectotype. 

Adamski (2002b) reported four further paratypes 

in the US National Museum. In the British Guiana 

Department of Science and Agriculture Annual 

Report for 1920, Bodkin (1922, as abstracted in 

Review of Applied Entomology) reported “In one district 

the blossoms of the palms were found to be infested 

by the larvae of a small moth, Blastobasis ochrobathra, 

Meyr.” Although this species did not appear 

in the Lepidoptera of North America checklist 

(Hodges et al., 1983), it is recorded from Florida 

(Heppner, 2003). In his synopsis of the Neotropical 

Blastobasidae, Adamski (2002b) transferred 

ochrobathra to Holcocera, but noted that it is known 

only from the type locality. Heppner (2003) placed 

this species in Blastobasis rather than Holcocera, but 

we follow Adamski (2002b) here. We note that 

there is a Barbados specimen in the BMNH reared

46: 1-21, 2013 

from castor oil seeds by R.W.E. Tucker, December 

1937, although its identity has not been confirmed 

by dissection. 

There is a similar species, H. grenadensis 

(Walsingham, 1891), described from Grenada. 

The female lectotype and four paralectotypes from 

Grenada and Dominica are in the BMNH (Adamski, 

2002b), together with specimens from Barbados 

and the Bahamas, which have not been dissected 

to confirm their identity. Adamski (1998) treated 

this species and transferred it to the genus Holcocera. 

Some authorities still (or again) place it in Blastobasis 

(e.g. Lee & Brown, 2009a), but we follow Adamski’s 

(2002b) treatment here. On external appearance, 

H. grenadensis differs from H. ochrobathra primarily in 

that the forewings are irregularly streaked with pale 

brown scales (Adamski, 2002b), but both species are 

variable. The two species can also be differentiated 

by characters of the genitalia (Adamski, 2002b). The 

male and female genitalia have been figured (Clarke, 

1963; Adamski, 1998, 2002b). Given the general 

similarities of species in this subfamily, even between 

genera, support from DNA barcoding (Hajibabaei, 

et al., 2007; Janzen et al., 2009; Adamski et al., 2010) 

might simplify identifications in future. 

Anatrachyntis rileyi (Walsingham) (Cosmopterigidae) 

Anatrachyntis (Meyrick, 1915a) is a genus of more 

than 50 species of small moths, almost exclusively from 

the Old World. The species of known biology seem 

to be scavengers and several have been associated 

with palm inflorescences. Anatrachyntis simplex 

(Walsingham) was described from Africa, but is now 

found in many parts of the tropics, and recognised as 

a polyphagous scavenger on various crops including 

cotton and coconut (Lepesme, 1947). Other species 

of this genus have been recognised as feeding on 

coconut flowers in the Old World, including, A. 

paroditis (Meyrick) in South-east Asia (Corbett, 1922), 

the Pacific (Lever, 1938), and the Seychelles (Vesey- 

Fitzgerald ,1941) etc., and A. dactyliota (Meyrick) is 

recorded in Malaysia (Meyrick, 1931). 

Although A. rileyi was described from Georgia, 

USA (Walsingham, 1882), and is known from the 

USA and several Caribbean Islands, it is likely to be 

of Old World origin, perhaps from Africa (Meyrick, 

1915a, p. 326; Zimmerman, 1978) and is widespread 

from southern Asia through the Pacific. At different 

times it has been placed in the genera: Batrachedra 

(Walsingham, 1882), Sathrobrota (Hodges, 1962), and 

Pyroderces (Hodges, 1978). At about the same time 

that Hodges (1978) placed rileyi in Pyroderces in his 

treatment of the family in the standard reference The 

Moths of America North of Mexico, Zimmerman (1978) 

transferred rileyi to Anatrachyntis. The latter has 

become accepted by European authors (e.g. Koster 

& Sinev, 2003; Heckford & Sterling, 2004), while 

Pyroderces is still commonly used in North America 

(e.g. Lee & Brown, 2009b). 

There is a similar species, A. badia (Hodges) in the 

USA, which was described in 1962 and has a similar 

range of food materials but with little documented 

overlap of actual food plant species (Hodges, 1962); 

the two are separated by markings on the hind leg tibia 

(Hodges, 1978). Where the two species occur together 

they could easily be confused, and this would have 

been the case with publications from the first half of 

the 20 th century (Zimmerman, 1978). The caterpillars 

of P. rileyi have been described by Busck (1917) and 

Heinrich (1921) and those of P. badia by Adamski et 

al. (2006), who could find no diagnostic characters to 

separate caterpillars of the two species. 

The caterpillars are detritivores reared from 

a wide variety of plant materials including cotton 

bolls (Walsingham, 1882; Busck, 1917; Heinrich, 

1921), as well as flowers, beans and pods of Ricinus 

(castor oil; Euphorbiaceae), flowers of Hyptis sp. 

(Lamiaceae), Colocasia esculenta (dasheen; Araceae), 

maize (corn husks and tassels, stored corn), many 

kinds of old leguminous pods, aloe, coffee beans, 

coffee cherries, eggplant, banana, dead Panicum 

torridum, pineapple (dried parts, fruits, stored seeds), 

Rochea (Crassulaceae), Samanea saman (Fabaceae- 

Mimosoideae), Sapindus oahuensis (Sapindaceae), 

and tamarind (Hodges, 1962, 1978; Zimmerman, 

1978; Garraway et al., 2007). Although A. badia has 

been reported from ‘blossoms of coconut’ in Florida 

(Hodges, 1962), here A. rileyi is reported from coconut 

inflorescence for the first time. 

Erechthias minuscula (Walsingham) (Tineidae, 

Erechthiinae) 

Erechthias minuscula was described from Jamaica, 

the Virgin Islands and Grenada (Walsingham, 1897), 

and it is known from various Caribbean Islands 

(Clarke, 1971, 1986; specimens in the BMNH), 

North America (MPGNA, 2012), Africa, southern 

Asia and the Pacific (Lepesme, 1947; Clarke, 1971; 

Zimmerman, 1978; Clarke, 1986). Meyrick (1915b, p. 

367) considered that it is probably Oriental in origin, 

and therefore introduced in Africa, the Americas and 

Pacific. It is predominantly a detritus feeder found 

associated with dead or decaying tissue of a wide 

range of plants from many families (Swezey, 1909, 

1910; Lepesme, 1947; Clarke, 1971; Zimmerman, 1978; 

Plumbley & Rees, 1983; Clarke, 1986; material in 

9

10 

BMNH), including coconut (Lepesme, 1947; material 

in BMNH from Fiji and the Solomon Islands). It is 

also recorded as a predator or scavenger of scale 

insects, especially Pseudaulacaspis pentagona (Targioni- 

Tozzetti), Lepidosaphes pinnaeformis (Bouché), Icerya 

purchasi Maskell, Aspidoproctus bouvieri Vayssière, and 

Orthezia insignis Browne (Swezey, 1909; Lepesme, 

1947). In contrast to all other reports, Harris (1935) 

stated that ‘there is no doubt as to its ability to feed 

on living coconut tissues’ in Tanzania, but this has 

not been confirmed. 

A related species, E. flavistriata (Walsingham) 

found from South-East Asia and the Pacific, has been 

recorded from coconut flowers (Meyrick, 1928), but is 

primarily associated with leaf sheaths, dead leaves and 

fibrous parts from a variety of plants, but in particular 

sugar cane (Zimmermann, 1978). 

Xystrologa nigrivitta (Walsingham) (Tineidae, 

?Meesiinae) 

The following is based on a recent paper on the 

West Indian species of Xystrologa by Davis et al. (2012). 

This Neotropical genus comprises six described and 

several undescribed species. Two species occur in 

the West Indies: X. grenadella (Walsingham) and X. 

nigrivatta. The former has been reared from branches 

of Sabal causiarum (Arecaceae) in Bermuda, from bark 

mulch used as a potting media and roots of orchids 

in nurseries in Florida, from damaged areas on the 

trunks of bonsai Ficus trees in Florida, on the roots of 

pineapple in Puerto Rico, and pupae have been found 

under bark of an unidentified tree in Dominica. As 

an introduced species in Germany, it was recently 

reared from caterpillars ‘found in dead wet wood of 

Robinia, on which are arranged Tillandsia and other 

Bromeliaceae, and on palm (Washingtonia sp.)’ in a 

large greenhouse. 

Xystrologa nigrovitta has been reared from an 

unidentified bracket fungus in Dominica, but has not 

previously been associated with a palm inflorescence. 

Thus, the available records point to members of this 

genus being opportunistic detritivores. Xystrologa 

nigrovitta is known from several West Indian Islands, 

including Trinidad, and it probably occurs throughout 

the West Indies. It was reared from the inflorescence 

of Roystonea oleracea in the survey reported below. 

Other species 

Lepesme (1947) includes a record of Tirathaba 

complexa (Butler) (=Harpagoneura complexa) (Pyralidae, 

Galleriinae) from Brazil. This is one of several species 

names included under the common name coconut 

J. Res.Lepid. 

spike moth, but it is unclear whether this includes 

several similar species or one variable species under 

the general name T. rufivena (Walker) (Waterhouse 

& Norris, 1987). This species or group of species 

from South-East Asia and the Pacific is considered a 

pest of coconut inflorescence, although the impact 

on yield is questionable (Corbett, 1931; Taylor, 

1930; Cock et al., 1987; Waterhouse & Norris, 1987). 

However, the record from Brazil is likely to be an 

error or misidentification, as there have been no 

subsequent observations of this relatively conspicuous 

inflorescence feeder from South America. 

Additional species recorded below, for which 

there is no published information include a species 

of Gelechiidae reared from Attalea maripa and two 

species of Cosmopterigidae reared from A. maripa 

and Mauritia flexuosa. 

re s u lt s: FI e l d s u r v e y a n d l a b w o r k In 

trInIdad 

More than one thousand moths of 12 species were 

reared from six of the 14 palm species (Table 4). 

Moths emerged from the samples for up to two months 

after collection of the inflorescence sample (Fig. 1). 

Atheloca bondari Heinrich (Pyralidae, Phycitinae) 

(Fig. 2a) 

Adults of A. bondari were identified as an Atheloca 

sp. from the photographs on MPGNA (2012), and 

to species from Heinrich (1956). The Trinidad 

specimens have the strong hair tuft at the base of the 

hind leg tibia, which is the distinguishing character 

for A. bondari, and absent in A. subrufella (Heinrich, 

1956). Accordingly, this species is treated as A. bondari, 

although it is recognised that this may prove to be a 

synonym of A. subrufella (see literature review). 

This species was reared from Roystonea oleracea 

at Kernahan, Nariva Swamp. The caterpillars were 

characterised as having a dark brown head and 

pronotum, purple-brown body with paler dorsal 

and lateral lines (Fig. 4), but these preliminary 

observations need confirmation based on systematic 

rearing of documented individual caterpillars. 

Caterpillars were observed to make webbing amongst 

flowers, tie together dead flowers, and feed amongst 

the fluffy padding of R. oleracea inflorescences and 

on male flowers. Caterpillars are larger than those 

of Batrachedra nuciferae and produce correspondingly 

larger frass. 

Similar caterpillars were found on coconut at 

Curepe, but none were individually reared successfully

46: 1-21, 2013 

Table 4. Summary of moths reared from palm inflorescences. No moths were reared from Adonidia merrillii (Curepe, CABI), 

Bactris major (Bush Bush Island, Botanic Gardens), Bactris simplicifrons (Bush Bush Island), Calamus sp. (Botanic Gardens), 

Euterpe precatoria (Aripo Savannah), Licuala spinosa (Botanic Gardens), Ptychosperma macarthurii (Curepe, CABI; Botanic 

Gardens), or Syagrus romanzoffiana (Botanic Gardens). In addition, a single specimen of Phidotricha erigens was reared from 

the young male inflorescence of Mauritia flexuosa. 

Palm species Location 

Atheloca bondari 

Neodavisia sp. 

Attalea maripa (old male) Bush Bush Is. (North) 26 11 21 58 

Batrachedra nuciferae 

Cocos nucifera 

(combined) 

Curepe, CABI 40 10 15 1 66 

Cocos nucifera Centeno 20 1 1 22 

Cocos nucifera 

(combined) 

Waller Field 121 116 7 244 

Cocos nucifera Bush Bush Is. (North) 1 1 

Cocos nucifera Kernahan 1 26 27 

Mauritia flexuosa (old 

male) 

Aripo Savannah 52 52 

Mauritia flexuosa (young 

male) 

Aripo Savannah 8 1 9 

Pritchardia pacifica* Botanic Gardens 16 3 19 

Roystonea oleracea Kernahan 10 57 16 41 339 135 598 

Roystonea oleracea Botanic Gardens 2 2 4 

Sabal mauritiformis Kernahan 12 12 

TOTAL 11 26 265 12 34 192 353 135 11 21 52 1,112 

* Introduced ornamental species. 

to confirm the identification. One adult was reared 

from a caterpillar isolated from coconut at Kernahan; 

amongst the terminal male flowers of one secondary 

inflorescence branch, it had tied three dead male 

flowers to the branch with silk. Since A. bondari is 

reported to attack coconut inflorescences (Bondar, 

1940a, 1940b; Heinrich, 1956) it seems safe to anticipate 

that this species does attack coconut more widely in 

Trinidad, although not as frequently as some of the 

other species considered here, such as B. nuciferae. 

Furthermore, as noted by Bondar (1940a, 1940b), the 

caterpillar is more damaging than that of B. nuciferae, 

causing direct damage to male and female flowers. 

Cocoons are formed completely enclosed in 

loose debris, but were not distinguished from those 

of Anatrachyntis rileyi. The adult moths are typical 

Batrachedra sp. 

Holcocera sp(p). 

Anatrachyntis rileyi 

Erechthias minuscula 

Xystrologa nigrivitta 

phycitine moths (Fig. 2a), and no other members of 

this subfamily were obtained in our survey. 

Unidentified Gelechiidae 

Unidentified Cosmopterigidae 1 

Phidotricha erigens (Ragonot) (Pyralidae, 

Epipaschiinae) (Fig. 2b) 

Unidentified Cosmopterigidae 2 

Total 

11 

This species (Fig. 2b) was reared just once from 

old, dead male inflorescence of Mauritia flexuosa. 

Given the range of alternative hosts reported, it 

is probably an occasional facultative herbivore or 

detritivore in palm inflorescences. 

Neodavisia sp. (Pyralidae, Pyralinae) (Fig. 2c) 

This appears to be an undescribed species, 

probably in the genus Neodavisia (Fig. 2c); it will be

12 

J. Res.Lepid. 

Figure 1. Number of moths emerging daily, counting from the day of sample collection. Results are pooled for all collections. 

Each colour line is related to a different axis scale. The single specimen of Phidotricha erigens is not shown; it emerged 27 

days after the inflorescence was collected. 

described in a forthcoming paper dealing with this 

and two similar, perhaps synonymous, genera from 

the Americas (M.A. Solis, pers. comm., 2012). It was 

reared from the dead male inflorescence of Attalea 

maripa and associated with dark caterpillars similar 

to those of Atheloca bondari and Xystrologa nigrivitta 

feeding amongst the stamens, and cocoons spun 

up amongst and completely covered by the stamens. 

Because the inflorescence was old and dead, most 

likely the caterpillars feed on dead plant material. 

Batrachedra nuciferae Hodges (Batrachedridae) 

(Figs. 2d, 3b) 

Cock (2013) reports observations on B. nuciferae 

from this survey, and illustrates the early stages. 

Caterpillars were found on coconut and Roystonea 

oleracea, but not on other palms; they were considered 

to be primarily restricted to the male flowers in which 

they are pollen feeders. 

Batrachedra sp. unidentified (Batrachedridae) (Fig. 2e) 

This species was reared from carat palm, Sabal 

mauritiiformis, at Kernahan, Nariva Swamp. It 

resembles B. nuciferae, but is significantly smaller (Fig. 

2d). Examination of parts of the inflorescence that 

included open flowers revealed webbing and frass, 

forming a protective tunnel or tube on the stem (Fig. 

5). At one end this tunnel covered a shallow groove 

in the inflorescence branch, and the webbing here 

incorporated yellow debris from the branch in a more

46: 1-21, 2013 

Figure 2. Pinned moths reared from palm inflorescences, Trinidad, Oct 2011; not to scale, the wingspan for each is given; 

a, Atheloca bondari, male, 16.6mm; b, Phidotricha erigens, 9.7mm; c, Neodavisia sp., 10.5mm; d, Batrachedra nuciferae, 

10mm; e, Batrachedra sp. indet. reared from Sabal mauritiiformis, 7mm; f, Anatrachyntis rileyi, 9mm; g, Erechthias minuscula 

female, 9.7mm; h, Holcocera sp., 13.7mm; i, Xystrologa nigrivitta female, 8.0mm; j, unidentified Cosmopterigidae sp. 2 (from 

Mauritia flexuosa), 5.2mm.; k, unidentified Gelechiidae (from Attalea maripa), 8.9mm; l, unidentified Cosmopterigidae sp. 1 

(from Attalea maripa), 5.5mm 

or less continuous cover or shelter. The remainder 

of the tunnel was more transparent, with scattered 

frass incorporated. Each tunnel sheltered a small 

pale caterpillar, resembling those of B. nuciferae, 

but smaller and with the head and pronotum pale 

brown. These caterpillars bore into the base of 

individual flowers and move from flower to flower, 

mostly in their tunnels. In due course some of these 

caterpillars made cocoons in the style of B. nuciferae, 

but smaller. The adult is similar to that of B. nuciferae 

13 

(fig. 2d) but smaller. Three adults were reared from 

these caterpillars and a further nine by emergence 

box over two weeks. 

Holcocera sp. (Meyrick) (Blastobasidae, 

Holcocerinae) (Fig. 2h) 

A Holcocera sp. was reared in this survey from 

inflorescences of three palm species: coconut, 

Roystonea oleracea and dead male inflorescence of

14 

Mauritia flexuosa. Specimens were identified as an 

unknown Holcocera sp. by D. Adamski, United States 

Department of Agriculture. The first author examined 

the male genitalia of a specimen from each palm and 

considered them to represent just one species, closely 

related to, but apparently distinct from, H. ocrobathra, 

the species described from coconut flowers in Guyana, 

J. Res.Lepid. 

Figure 3. Dead moths (except for photo a, which was alive) in more or less normal resting positions, reared from palm 

inflorescences, Trinidad, Oct 2011; not to scale, see legend to Figure 2 for wingspans; scale squares = 2mm. a, living Anatrachyntis 

rileyi; b, Batrachedra nuciferae; c, Xystrologa nigrivitta; d, Erechthias minuscula mating pair; e, unidentified Gelechiidae (from 

Attalea maripa); f, unidentified Cosmopterigidae sp. 1 (from Attalea maripa); g, unidentified Cosmopterigidae sp. 2 (from 

Mauritia flexuosa). Figure a (from MALMR 2008), photo by Research Division Central Experimental Station, Centeno, Trinidad 

and Tobago. 

and H. grenadensis, described from Grenada and 

Dominica (see literature review). 

Caterpillars of this species were found on and 

reared from amongst the fluffy matrix and dead 

male flowers of R. oleracea at Kernahan, but we did 

not establish which part of this served as their food 

source. They were characterised as similar to those

46: 1-21, 2013 

of Atheloca bondari (Fig. 4) but darker. Holcocera spp. 

generally feed on detritus and dead plant material, 

and this is probably the role of this species in palm 

inflorescences. The cocoons were not distinguished 

from those of A. bondari. The adults (Fig. 2h) are 

easily distinguished from the other species obtained 

in this survey by their wing shape and colour. 

Anatrachyntis rileyi (Walsingham) (Cosmopterigidae) 

(Figs. 2f, 3a) 

This species was initially identified by comparison 

with the type and the BMNH series. All specimens 

were checked against the diagnostic features in 

Hodges (1978, p. 47), to ensure that no A. badia 

were present. A subsample of pinned moths was 

examined by D.R. Davis, Smithsonian Institution, who 

confirmed their identity. The adult of this moth (Fig. 

3a) was illustrated incorrectly as Batrachedra nuciferae 

in MALMR (2008). It is attractively coloured and 

marked in tones of orange and brown (Figs. 2f, 3a) 

and easily distinguished amongst the moths reared 

from the survey. 

In this survey, A. rileyi was reared from the 

inflorescences of five palm species (Table 4). The 

caterpillar was characterised as having the head 

chestnut brown, pronotum and anal plate brown, 

body pink-brown (Fig. 6), but these preliminary 

observations need confirmation based on systematic 

rearing of documented individual caterpillars, an 

exercise beyond our resources on this occasion. 

The cocoon was similar to that of Atheloca bondari, 

but smaller. The exact food and feeding style was 

not established, but the record from dead male 

inflorescence of M. flexuosa, and the fact that adults 

emerged later from the emergence boxes than did 

those of the pollen-feeding B. nuciferae, supports the 

expectation that it feeds on dead plant material, as 

has been previously documented. 

Erechthias minuscula (Walsingham) (Tineidae, 

Erechthiinae) (Figs. 2g, 3d) 

This species was initially identified by comparison 

with the BMNH series. A subsample was examined 

by D.R. Davis, Smithsonian Institute, who confirmed 

this identification. It was obtained from Kernahan 

by emergence box from Roystonea oleracea in large 

numbers and once from coconut (Table 4). Amongst 

the material reared, it was distinctive due to the 

reflexed wing tips (Fig. 3d). 

Early stages have not been unequivocally associated, 

but caterpillars found on coconut at Kernahan (Fig. 7) 

are likely to be this species. This caterpillar webbed 

Figure 4. Caterpillar of Atheloca bondari amongst dropped 

male flowers and fluffy matrix of inflorescence of palmiste, 

Roystonea oleracea, Kernahan. 

Figure 5. Caterpillars and signs of Batrachedra sp. indet. 

on carat palm, Sabal mauritiiformis, Kernahan. Note the 

webbing and frass to the right. 

15

16 

Figure 6. Caterpillar of Anatrachyntis rileyi amongst silk 

webbing on dropped male flowers of palmiste, Roystonea 

oleracea, Kernahan. 

Figure 7. Provisionally associated early stages of Erechthias 

minuscula on coconut, Kernahan. a, dead male flowers 

attached to inflorescence with silk webbing; b, caterpillar, 

dorsolateral view. 

J. Res.Lepid. 

together dead male flowers attaching them to the 

inflorescence branch, sometimes in a pendulous 

chain (Fig. 7a). One had attached a dead male flower 

to the base of a female flower, and although the 

female flower was marked, it did not appear to have 

been significantly damaged on this occasion. These 

caterpillars were characterised as having a chestnut 

brown head, dark brown pronotum with a narrow pale 

dorsal line, body dull pale brown with darker dorsal, 

dorsolateral and lateral longitudinal lines; body with 

erect setae with the base dark. 

Xystrologa nigrivitta (Walsingham) (Tineidae, 

?Meesiinae) (Figs. 2i, 3c) 

This species was reared commonly from the 

inflorescence of Roystonea oleracea from Kernahan 

(Table 4). Emergence did not start until a month after 

the inflorescence was collected (Fig. 1). No observations 

were made on the early stages, but given the known 

biology of the genus (Davis et al., 2012) they are likely 

to be detritivores in inflorescences of R. oleracea. 

Unidentified Cosmopterigidae sp. 1 and Gelechiidae 

sp. from Attalea maripa (Figs. 2k, 2l, 3e, 3f) 

Two more species were reared from the old dead 

male inflorescence of Attalea maripa, where they were 

probably feeding on dead plant material. The old male 

inflorescence, in which the flower parts, including 

the long pollen-bearing stamens, were completely 

dead and dry, showing a great deal of old feeding 

damage by Lepidoptera, based on the amount of frass 

and webbing that was observed (Fig. 8). The flower 

remains contained many small 3mm long, plain white 

Lepidoptera cocoons, which were associated with an 

unidentified Cosmopterigidae species (sp. 1, Figs. 

2l, 3f) obtained by emergence box. An unidentified 

species of Gelechiidae (Figs. 2k, 3e) was obtained by 

emergence box only. The relative contribution of 

the different species reared to the damage observed 

is not clear. Although no other distinctive remains 

in terms of cocoons were found when examining the 

inflorescence, it is possible that additional species 

may have completed development and emerged and 

dispersed before our sample was taken. 

In addition to these small species, a long, tough 

silk-lined gallery or tunnel was found running 

through the dry mass of stamens, more than 30cm 

long and in places 1cm in diameter (Fig. 9). A cast 

head capsule, 2.4mm wide x 2.7mm high, was found in 

this gallery. No other sign of the caterpillar or its pupa 

were found, but we assume it was the maker of this 

tunnel. The purpose of the tunnel and life style of the

46: 1-21, 2013 

caterpillar that made it is open to speculation. The 

tunnel would probably provide protection from small 

vertebrate predators that are likely to be attracted to 

the large number of caterpillars in the inflorescence. 

It would enable the inhabitant to move around safely 

over long distances within the inflorescence; could 

the builder be a predator itself? 

Unidentified Cosmopterigidae sp. 2 from Mauritia 

flexuosa (Figs. 2j, 3g) 

Some feeding damage and very small caterpillars 

were found under the bracts of the old male 

inflorescence of Mauritia flexuosa. More than 50 

moths of a very small unidentified Cosmopterigidae 

species (sp. 2, Figs. 2j, 3g) were subsequently reared 

by emergence box from the old male inflorescence, 

but not from the young inflorescence sampled from 

the same palm. We suppose that caterpillars of this 

species feed as detritivores on dead plant tissue in the 

old inflorescence. 

dIscussIon a n d c o n c l u s I o n s 

In total, more than one thousand moths of 12 

species were reared from six of the 14 palm species 

sampled (Table 4). All the previously reported genera 

of palm-feeding Lepidoptera were obtained apart 

from Cadra. Although C. cautella has been recorded 

from Trinidad (Kaye & Lamont, 1927), this species 

was not found in the survey. Given the observations 

of Bondar (1940a, 1940b), it seems likely that further 

surveys will show that one or more Cadra spp. develop 

in palm inflorescences in Trinidad. 

Of the moths reared, three could not be identified 

beyond family, three could only be identified to genus, 

and the remaining six were identified to species. These 

six comprise two species associated with coconut 

(Atheloca bondari, Batrachedra nuciferae) and four that 

are polyphagous and widespread (Xystrologa nigrivitta, 

Anatrachyntis rileyi, Erechthias minuscula, Phidotricha 

erigens). Of these, P. erigens has long been known 

from Trinidad (Kaye & Lamont, 1927), B. nuciferae 

was reported recently (MALMR, 2006, 2008), and 

X. nigrivitta was reported from Trinidad only after 

the survey was completed (Davis et al., 2012). The 

remaining species (Atheloca bondari, Anatrachyntis 

rileyi, E. minuscula and the partially identified species) 

have not previously been reported from the island. 

This supports the suggestion of Cock (2003) that a 

large number of species from the families of smaller 

moths remain to be identified from Trinidad, and that 

surveys of specialist niches will rapidly generate new 

information on these. 

Figure 8. Views of old male inflorescence of Attalea 

maripa, from Bush Bush Island showing Lepidoptera frass, 

and stamen remains held together with silk webbing. 

17 

Anatrachyntis rileyi and E. minuscula are considered 

to be of Old World origin and so must have been 

introduced with trade, probably long ago. Batrachedra 

nuciferae was suspected to be an introduced species 

spreading in South America (MALMR, 2006, 2008), 

but Cock (2013) suggests it is more likely to be an 

indigenous species that has been overlooked. The 

remaining species, including those only identified to 

genus and family are likely to be indigenous species, 

not previously reported. 

Some of the moths appear to be specialists 

associated with particular palm species, while 

others are generalists. Those thought to feed on 

living tissue (Atheloca bondari, Batrachedra spp.) 

are relatively specialised on two species. Those 

thought to be detritivores appear to be a mixture 

of specialists (X. nigrivitta on Roystonea oleracea,

18 

unidentified Gelechiidae species and unidentified 

Cosmopterigidae sp. 1 on Attalea maripa, unidentified 

Cosmopterigidae sp. 2 on Mauritia flexuosa) and 

generalists (Holcocera sp., Anatrachyntis rileyi, and 

E. minuscula, each on several palms). However, X. 

nigrivitta has been reared from a bracket fungus 

(Davis et al., 2012), so it is not a specialist as this 

survey suggested, and this also may prove to be 

the case when more is known about the three 

unidentified species. 

There were three different patterns for the delay 

of emergence of moths from the collection date 

for the samples (Fig. 1). Atheloca bondari, the two 

Batrachedra spp. and unidentified Cosmopterigidae 

sp. 2 (from old M. flexuosa) all emerged within the 

first month; the unidentified Gelechiidae species, 

unidentified Cosmopterigidae sp. 2 and Neodavisia 

sp. (all from dead Attalea maripa) emerged at a fairly 

steady, low rate over two months; and Holcocera sp., 

X. nigrivitta, Anatrachyntis rileyi, and E. minuscula 

emerged primarily in the second month after the 

emergence boxes were set up. We suggest these 

patterns represent two different life styles. The first 

group, those species feeding on fresh plant material, 

rapidly completed their development and emerged 

within a month, whereas the last group, developing 

as detritivores, completed their development more 

slowly, either because they started later or because 

the poorer food quality dictated slower development, 

and emerged after the plant-feeding species. The 

middle group also are detritivores, reared from dead 

inflorescences of Attalea maripa and M. flexuosa, but 

because the sample of inflorescence was already 

dead, the moths had completed development and 

started to emerge immediately after collection. 

ac k n o w l e d g e M e n t s 

J. Res.Lepid. 

Figure 9. Half of the ‘tunnel’ made by an unidentified Lepidoptera species found amongst the old male inflorescence of cocorite, 

Attalea maripa, Bush Bush Island. Scale squares = 4mm. 

The first author was responsible for the design of this work, 

literature review and writing; both authors (and collaborators) 

carried out the field work and initial laboratory work; the second 

author was responsible for the bulk of the laboratory work, including 

sorting, recording and preparation of the reared moths. 

We thank the following for support and help with aspects of 

this study: Ms Velda Ferguson-Dewsbury, Mr Farzan Hosein, Ms 

Petal Ram and staff of the Forestry Department and Royal Botanic 

Gardens, Ministry of Food Production, Land & Marine Affairs 

of Trinidad & Tobago; Dr Elizabeth S Johnson, Ms Suzy Wood, 

CABI; Dr Yasmin Comeau and Mr Winston Johnson, National 

Herbarium; Mr Imran Gaffoor of Waller Field, Trinidad and 

Tobago; Dr M Alma Solis, Dr John W Brown and Dr David Adamski, 

United States Department of Agriculture, Systematic Entomology 

Laboratory; Dr Don R. Davis, Smithsonian Institution. 

Live and dead insect material was exported under permit from 

the Plant Protection Organisation of Trinidad and Tobago. Live 

material was imported to the UK and held in CABI’s UK quarantine 

facility for emergence under licence from the Department for 

Environment, Food and Rural Affairs. The field visits to Nariva 

Swamp and Aripo Savannah were made under permit from the 

Forestry Department, Trinidad and Tobago. This study was 

undertaken on behalf of the Ministry of Food Production, Land & 

Marine Affairs of Trinidad and Tobago, in support of their role in 

a United Nations Environment Programme – Global Environment 

Facility funded project: “Mitigating the Treats of Invasive Alien Species 

in the Insular Caribbean” Project Number GFL/ 2328–2713-4A86 

(PMS: GF/1030-09-03), which provided the financial support. 

CABI’s support in Trinidad and the preparation of this paper was 

funded by the CABI Development Fund, with contributions from 

the Australian Centre for International Agricultural Research, the 

UK’s Department for International Development and the Swiss 

Agency for Development and Cooperation. 

lI t e r at u r e c I t e d 

ADA M S k I, D. 1998. On the identity of Holcocera guilandinae 

(Busck 1900) (Lepidoptera: Gelechioidea: Coleophoridae: 

Blastobasinae). Proceedings of the Entomological Society of 

Washington 100: 731–741.

46: 1-21, 2013 

ADA M S k I. D. 2002a. Holcocerini of Costa Rica (Lepidoptera: 

Gelechioidea: Coleophoridae: Blastobasinae). Memoirs of the 

Entomological Society of Washington 24: 147 pp. 

AD A M S k I, D. 2002b. A synopsis of described Neotropical Blastobasinae 

(Lepidoptera–Gelechioidea–Coleophoridae). Thomas Say 

Publications in Entomology: Monographs. Entomological 

Society of America, Lanham, Maryland, 150 pp. 

ADA M S k I, D. 2013. Review of the Blastobasinae of Costa Rica 

(Lepidoptera: Gelechioidea: Blastobasidae). Zootaxa 3618(1): 

1–223. 

AD A M S k I, D. & r.l. Br o w N . 1989. Morphology and systematics of 

North American Blastobasidae (Lepidoptera: Gelechioidea). 

Mississippi Agricultural and Forestry Experiment Station 

Technical Bulletin 165: 1–70. 

AD A M S k I, D., J.w. Br o w N & w.H. wH I t e. 2006. Description of the 

immature stages of Pyroderces badia (Hodges) (Lepidoptera: 

Cosmopterigidae), with a new host record from Louisiana. 

Proceedings of the Entomological Society of Washington 108: 

341–346. 

AD A M S k I, D., r.S. Co p e l A N D, S.e. MI l l e r, p.D.N. He B e rt, k. DA r r o w & 

Q. lU k e. 2010. A review of African Blastobasinae (Lepidoptera: 

Gelechioidea: Coleophoridae), with new taxa reared from 

native fruits in Kenya. Smithsonian Contributions to Zoology 

630, vi + 68 pp. 

Ar N A l, e.; J. Cl Av I J o, e. So t o & F. rA M o S . 1998. Batrachedra nuciferae 

Hodges, 1966 (Lepidoptera: Momphidae) a new pest of coconut 

in Venezuela. Boletin de Entomologia Venezolana, Serie 

Monografias 13: 69–71. 

BAr F o D, A.S., t. BU r H o lt & F. Bo r C H S e N I U S . 2003. Contrasting 

pollination modes in three species of Licuala (Arecaceae: 

Coryphoideae). Telopea 10: 207–223. 

BeC C A l o N I, G.w., M.J. SC o B l e , G.S. ro B I N S o N & B. pI t k I N (e D S.) 

2003. The Global Lepidoptera Names Index (LepIndex). 

http://www.nhm.ac.uk/entomology/lepindex (last accessed 

1 November 2011). 

Be C k e r, v.o. 1984. 24. Coleophoridae, pp. 42–43. In: Heppner, 

J.B. (ed.), Atlas of Neotropical Lepidoptera. Checklist: Part 

1 Micropterigoidea – Immoidea. Dr W Junk Publishers, The 

Hague, Netherlands. 

Be N t o, J.M.S., D.e. NAvA, M.C.M. CH A G A S & A.H. Co S tA. 2006. Biology 

and mating behavior of the coconut moth Atheloca subrufella 

(Lep.: Phycitidae). Florida Entomologist 89: 199–203. 

Bo D k I N, G.e. 1922. Government economic biologist. Report of 

the Department of Science & Agriculture, British Guiana 1920: 

51–58. [Review of Applied Entomology 1922 10: 561]. 

Bo N D A r, G. 1940a. Insetos nocivos e moléstias do coqueiro (Cocos 

nucifera) no Brasil. Tipographía Naval, Brasil, 160 pp. 

Bo N D A r, G. 1940b. Notas entomológicas da Bahia. Revista de 

Entomologia, Rio de Janeiro 11: 199–214. 

Bro w N , J.w., D. AD A M S k I, r.w. Ho D G e S, & S.M. BA H r II. 2004. 

Catalog of the type specimens of Gelechioidea (Lepidoptera) 

in the collection of the National Museum of Natural History, 

Smithsonian Institution, Washington, DC. Zootaxa 510: 

1–160. 

BU S C k, A. 1917. The pink bollworm, Pectinophora gossypiella. Journal 

of Agricultural Research 9: 343–370, plates 7–12. 

CA r N e I r o, J.S., p.H.S. D A SI lvA, C.C.p. No G U e I r A & H.U. D e So U S A. 

2004. Morfologia externa, biologia, danos e sugestões de 

controle da traça-dos-frutos-docoqueiro Batrachedra nuciferae. 

Comunicado Técnico, Ministério da Agricultura Pecuária e 

Abasticimento, (Teresina, PI, Brazil) 159: 4 pp. 

CAr r e G A r o, J.B. 2007. Insetos herbívoras im botões florais de 

Caryocar brasiliense Camb (Caryocaraceae): comparação entre 

duas áreas de cerrado de Brasilia, D.F. MSc thesis, Universidade 

de Brasilia, v + 35 pp. 

CHI l D , r. 1974. Coconuts. 2nd ed. Longmans, Green & Co., 

London, xvi + 335pp. 

19 

ClA r k e, J.F.G. 1963. Catalogue of the type specimens of 

Microlepidoptera in the British Museum (Natural History) 

described by Edward Meyrick. Vol. 4: Phalonidae, Carposinidae, 

Chlidanotidae, Oecophoridae, Blastobasidae, Momphidae, 

Epermeniidae, Strepsimanidae, Physoptilidae. The Trustees 

of the British Museum, London, 521 pp. 

Cl A r k e, J.F.G. 1971. The Lepidoptera of Rapa Island. Smithsonian 

Contributions to Zoology 56: 282 pp. 

ClA r k e, J.F.G. 1986. Pyralidae and Microlepidoptera of the 

Marquesas Archipelago. Smithsonian Contributions to Zoology 

416: 485 pp. 

CoC k , M.J.w. 2003. On the number of species of moths 

(Lepidoptera) in Trinidad and Tobago. Living World, Journal 

of the Trinidad and Tobago Field Naturalists’ Club 2003: 

49–58. 

CoC k , M.J.w. IN p r e S S. Batrachedra nuciferae, an inflorescencefeeding 

moth associated with coconut, Cocos nucifera, and 

palmiste, Roystonea oleracea, in Trinidad, West Indies. Journal 

of Insect Science. 

Co C k, M.J.w., C.N. GA l l e G o & H.C.J. Go D F r A y. 1987. Biological 

control of Tirathaba rufivena in the Philippines, pp. 147–157. 

In: Ferrar, P. & D.-H. Stechmann (eds.), Biological control in 

the South Pacific. Report on an International Workshop held 

at Government Experimental Farm, Vaini, Kingdom of Tonga, 

17-25 October 1985. Ministry of Agriculture, Fisheries & Forests 

of Tonga, Nuku’alofa, Tonga. 

Co M e A U , p.l., y.S. Co M e A U & w. Jo H N S o N. 2003. The palm book 

of Trinidad and Tobago including the Lesser Antilles. The 

International Palm Society, 108 pp. 

Cor B e t t, G.H. 1922. Moths from coconut spikes. Malayan 

Agricultural Journal 10: 184–185. 

Co r B e t t, G.H. 1931. The control and economic importance of 

the greater spike moth (Tirathaba rufivena Walk.). Department 

of Agriculture of the Straits Settlements and Federated Malay 

States, Scientific Series 8: 1–14. 

Co r B e t t, G.H. & B.A.r. GAt e r. 1924. Batrachedra arenosella. Walk., 

(Cosmopterygidae) in relation to the nut-fall of coconuts. 

Malayan Agricultural Journal 12: 115–122. 

DAv I S, D.r., M.M. DAv I S & C. MA N N I o N. 2012. Neotropical Tineidae 

IX: A review of the West Indian Xystrologa and biology of 

Xystrologa grenadella (Wlsm.), an invasive pest of cultivated 

greenhouse plants in southern Florida, USA and Germany 

(Lepidoptera: Tineoidea). Proceedings of the Entomological 

Society of Washington 114: 439–455. 

DINIz, I.r. & H.C. Mo r A I S. 2002. Local pattern of host plant 

utilization by lepidopteran larvae in the cerrado vegetation. 

Entomotropica 17: 115–119. 

Dr A N S F I e l D, J., N.w. UH l, C.B., ASMUSSeN, w.J. BA k e r, M.M. HA r l e y 

& C.e. le w I S. 2008. Genera Palmarum - The evolution and 

classification of palms. Kew Publishing, Kew, UK, 744 pp. 

er v I G, F. 1993. Notes on the phenology and pollination of the 

dioecious palms Mauritia flexuosa (Calamoideae) and Aphandra 

natalla (Phytelephantoideae) in Ecuador, pp. 7–12. In: 

Barthlott W., C. Naumann, C. Schmidt-Loske & K. Schuchmann 

(eds.), Animal-plant interactions in tropical environments. 

Zoologisches Forschungsinstitut und Museum Alexander 

Koenig, Bonn, Germany. 

eSSIG, F.B. 1971. Observations on pollination in Bactris. Principes 

15: 20–24. 

GA r r AwA y, e., D.C. He N r y & B.r. Cr I t C H l e y. 2007. The biology, 

ecology and economic importance of the pink scavenger 

caterpillar Pyroderces rileyi (Lepidoptera: Cosmopterigidae) on 

banana in Jamaica. Tropical Science 47: 231–243. 

GoA t e r , B. 1986. British pyralid moths. A guide to their 

identification. Harley Books, Colchester, UK, 175 pp. 

HA B e C k, D.H. & J.C. NI C k e r S o N. 1982. Atheloca subrufella (Hulst), 

a pest of coconuts (Lepidoptera: Pyralidae: Phycitinae).

20 

Entomology Circular No. 241. Florida Department of Agriculture 

and Consumer Services, Division of Plant Industry, 2 pp. 

HA J I B A B A e I, M., G.A.C. SI N G e r, p.D.N. He B e rt & D.A. HI C k e y. 2007. 

DNA barcoding: how it complements taxonomy, molecular 

genetics and population genetics. Trends in Genetics 23: 

167–172. 

HA r r I S, w.v. 1935. Report of the assistant entomologist, 1934. 

Report. Department of Agriculture, Tanganyika 1935: 84–89. 

He C k F o r D, r.J. & p.H. St e r l I N G. 2004. Anatrachyntis badia (Hodges, 

1962) (Lepidoptera: Cosmopterigidae) an adventive species 

new to the British Isles, possibly the second record from Spain 

and a larval description. Entomologist’s Gazette 55: 81–89. 

HeINrICH, C. 1921. Some Lepidoptera likely to be confused with the 

pink bollworm. Journal of Agricultural Research 20: 807–836, 

plates 93–109. 

HeINrICH, C. 1956. American moths of the family Phycitinae. 

United States National Museum Bulletin 207: viii + 581 pp. 

He N D e r S o N, A. 1986. A review of pollination studies in the Palmae. 

Botanical Review 52: 221–259. 

He N D e r S o N, A., G. GA l e A N o & r. Be r N A l. 1995. Field guide to the 

palms of the Americas. Princeton University Press, Princeton, 

New Jersey, USA, ix + 353 pp. + 64 plates. 

He p p N e r, J.B. 2003. Lepidoptera of Florida. Part 1 Introduction and 

catalog. Arthropods of Florida and Neighboring Land Areas 

Volume 17. Florida Department of Agriculture & Consumer 

Services, Gainesville, Florida, USA, x + 670pp. 

Ho D G e S, r.w. 1962. A revision of the Cosmopterigidae of America 

north of Mexico, with a definition of the Momphidae and 

Walshiidae (Lepidoptera: Gelechioidea). Entomologica 

Americana 42 (n.s.): 166 pp, 199 figs. 

HoD G e S , r.w. 1966. Review of the New World species of 

Batrachedra with description of three new genera (Lepidoptera: 

Gelechioidea). Transactions of the American Entomological 

Society 92: 585–651. 

Ho D G e S, r.w. 1978. Moths of America North of Mexico, Fascicle 6.1: 

Walshiidae and Cosmopterigidae. E.W. Classey Ltd. and Wedge 

Entomological Research Foundation, London, 166 pp. 

HoD G e S, r.w. 1999. 9. The Gelechioidea, pp. 131–158. In: 

Kristensen, N.P. (ed.), Lepidoptera, moths and butterflies. 

Volume 1: Evolution, systematics, and biogeography. WaIter 

de Gruyter, Berlin, Germany. 

Ho D G e S, r.w. et a l. (e D S.) 1983. Checklist of the Lepidoptera of 

North America north of Mexico. E. W. Classey, London, xxiv 

+ 284 pp. 

Ho wA r D, F.w.; D. Mo o r e, r.M. GIBlIN-DAv I S & r.G. AB A D. 2001. 

Insects on palms. CABI Publishing, Wallingford, Oxon, UK, 

400 pp. 

JA N z e N, D.H., w. HA l lwA C H S, p. Bl A N D I N, J.M. BU r N S, J.-M. CA D I o U, I. 

CH A C o N, t. DA p k e y, A.r. De A N S, M.e. ep S t e I N, B. eS p I N o z A, J.G. 

Fr A N C l e M o N t, w.A. HA B e r, M. HA J I B A B A e I,; J.p.w. HA l l, p.D.N. 

He B e rt, I.D. GA U l D, D.J. HA r v e y, A. HA U S M A N N, I.J. kItCHING, 

D. lA F o N tA I N e, J.-F. lA N D r y, C. le M A I r e, J.y. MI l l e r, J.S. MI l l e r, 

l. MI l l e r, S.e. MI l l e r, J. Mo N t e r o, e. MU N r o e, S.r. Gr e e N, S. 

rAt N A S I N G H A M, J.e. rAw l I N S , r.k. ro B B I N S, J.J. ro D r I G U e z, r. 

ro U G e r I e, M.J. SH A r k e y, M.A. SM I t H , M.A. So l I S, J.B. SU l l I vA N, 

p. tH I A U C o U r t , D.B. wA H l , S.J. we l l e r, J.B. wH I t F I e l D, k.r. 

wIl l M o t t, D.M. wo o D , N.e. wo o D l e y & J.J. wI l S o N . 2009. 

Integration of DNA barcoding into an ongoing inventory of 

complex tropical biodiversity. Molecular Ecology Resources 

9 (Suppl. 1): 1–26. 

kAI l A, l. 2004. Phylogeny of the superfamily Gelechioidea 

(Lepidoptera: Ditrysia): an exemplar approach. Cladistics 

20: 303–340. 

kAI l A, l., M. MU t A N e N & t. Ny M A N . 2011. Phylogeny of the 

mega-diverse Gelechioidea (Lepidoptera): Adaptations 

and determinants of success. Molecular Phylogenetics and 

Evolution 61: 801–809. 

J. Res.Lepid. 

kAl S H o v e N , l.G.e. & p.A. vA N D e N lA A N. 1981. The pests of crops 

in Indonesia. P.T. Ichtiar Baru-Van Hoeve, Jakarta, Indonesia, 

701 pp. 

kA y e, w.J. & N. lA M o N t . 1927. A catalogue of the Trinidad 

Lepidoptera Heterocera (moths). Memoirs of the Department 

of Agriculture, Trinidad and Tobago 3: 144 pp. 

kI M B A l l , C.p. 1965. Lepidoptera of Florida. Arthropods of Florida 

and neighbouring land areas. Volume 1. State of Florida 

Department of Agriculture, Gainesville, Florida, 363 pp. 

koS t e r , S. & S. SI N e v . 2003. Momphidae, Batrachedridae, 

Stathmopodidae, Agonoxenidae, Cosmopterigidae, 

Chrysopeleiidae. Microlepidoptera of Europe (5). Apollo 

Books, Stenstrup, Denmark, 387 pp. 

le D e r e r, J. 1863. Beitrag zur Kenntnis der Pyralidinen. Wiener 

Entomologische Monatschrift 7: 243–280, 331–378, 379–426, 

427–502, plates 2–18. 

le e, S. & r.l. Br o w N . 2009a. Revised checklist of Coleophoridae 

in Neotropical Region. Updated date: 10 February 2009. 

http://mississippientomologicalmuseum.org.msstate.edu/ 

Researchtaxpages/Lepidoptera/Coleophoridae/Checklists/ 

Neotropical-Coleophoridae.html 

le e, S. & r.l. Br o w N . 2009b. Revised checklist of Cosmopterigidae 

in Neotropical Region. Updated date: 10 February 2009. 

http://mississippientomologicalmuseum.org.msstate.edu/ 

Researchtaxpages/Lepidoptera/Cosmopterigidae/Checklists/ 

Neotropical-Cosmopterigida.html 

le p e S M e p. 1947. Les insectes des palmiers. Paul Lechevalier, Paris, 

France, 909 pp. 

lev e r, r.J.A.w. 1938. Three moths from coconut flowers. 

Agricultural Journal of Fiji 9: 23. 

le v e r, r.J.A.w. 1979. Pests of the Coconut Palm. FAO, Rome, 

Italy, xiii + 190 pp. 

MAlMr (MINIStry o F AG r I C U lt U r e, lA N D S A N D MA r I N e re S o U r C e S) 

2006. 1.1.8 Coconut moth (Batrachedra nuciferae), pp. 86–87. 

In: Ministry of Agriculture, Land and Marine Resources. 

Annual Report 2006. Ministry of Agriculture, Land and Marine 

Resources, Centeno, Trinidad & Tobago. 

MAlMr (MINIStry o F AG r I C U lt U r e, lA N D S A N D MA r I N e re S o U r C e S) 

2008. “Coconut Moth” Batrachedra nuciferae - A New Pest 

of Coconuts. Coconut Moth Fact Sheet # 1. Ministry of 

Agriculture, Lands and Marine Resources – Entomology, 

Centeno, Trinidad & Tobago, 2 pp. 

Me y r I C k, e. 1915a. Exotic Microlepidoptera. Vol. 1. Part 11. The 

author, Marlborough, UK, pp. 321–352. 

Me y r I C k, e. 1915b. Exotic Microlepidoptera. Vol. 1. Part 12. The 


Me y r I C k, e. 1921. Exotic Microlepidoptera. Vol. 2. Part 15. The 


Me y r I C k, e. 1928. The micro-Lepidoptera of the “St. George” 

expedition. Transactions of the Entomological Society of 

London 76: 489–521. 

Me y r I C k, e. 1931. Exotic Microlepidoptera. Vol. 4, Part 2. The 


Mo o r e, D. 2001. Insects of palm flowers and fruits, pp. 233–266. 

In: Howard, F.W.; Moore, D., Giblin-Davis, R.M. & Abad, R.G., 

Insects on palms. CABI Publishing, Wallingford, Oxon, UK. 

MoU N D , l. (e D.) 1989. Common insect pests of stored food 

products. Economic Series No. 15. (7th Edn). British Museum 

(Natural History), London, UK, pp. 68. 

MpGNA (Mo t H pH o t o G r A p H e r S Gr o U p o F No r t H AM e r I C A ) 

2012. Digital guide to moth identification. http:// 

mothphotographersgroup.msstate.edu/MainMenu.shtml 

plU M B l e y, r.A. & D.p. re e S. 1983. An infestation by Araecerus 

fasciculatus (Degeer) (Coleoptera: Anthribidae) and Decadarchis 

minuscula (Walsingham) (Lepidoptera: Tineidae) on stored 

fresh yam tubers in South-East Nigeria. Journal of Stored 

Products Research 19: 93–95.

46: 1-21, 2013 

rAG o N o t , e.-l. 1888. Untitled. [M. E.-L. Ragonot donne les 

diagnoses de cinq espèces nouvelles de Microlépidoptères de 

Porto-Rico.] Bulletin des Séances et Bulletin Bibliographique de 

la Société entomologique du France, Series 6, 8: cxxxviii–cxl. 

Sá N C H e z So t o, S. 2004. Morphologia, biologia e fototropisma de 

Batrachedra nuciferae Hodges (Lepidoptera: Coleophoridae). 

Master’s Thesis, Universidade de São Paulo, Brazil, xi + 40 pp. 

Sá N C H e z-So t o, S. & o. NA k A N o. 2002. Ocorrência de Batrachedra 

nuciferae Hodges (Lepidoptera: Coleophoridae) no Estado de 

São Paulo. Neotropical Entomology 31: 657–658. 

Sá N C H e z-So t o, S. & o. NA k A N o. 2004a. Presença de Batrachedra 

nuciferae Hodges na cultura da coqueira no estado de São Paulo. 

Revista de Agricultura 79: 155–157. 

Sá N C H e z-So t o, S. & o. NA k A N o. 2004b. Estudo morfológico de 

Batrachedra nuciferae Hodges (Lepidotera: Coleophoridae). 

Neotropical Entomology 33: 173–178. 

Sá N C H e z So t o, S. & o. NA k A N o. 2008. Biología de Batrachedra nuciferae 

Hodges (Lepidoptera: Coleophoridae), plaga del cocotero en 

América del Sur. Boletín del Museo de Entomología de la 

Universidad del Valle 9: 22–26. 

SA N tA N A, S.w.J. 2008. Criação e bioecologia de Atheloca subrufella 

(Hulst) (Lepidoptera: Phycitidae). Tese de doutorado. 

Universidade Federal Rural de Pernambuco, Recife, 60 pp. 

SA N tA N A, S.w.J., r. BA r r o S, J.B. to r r e S & M.G.C. Go N D I M Jr. 2010. 

Exigências Térmicas da Praga do Coqueiro Atheloca subrufella 

(Hulst). (Lepidoptera: Phycitidae). Neotropical Entomology 

39: 181–186. 

SA N tA N A, S.w.J., J.B. to r r e S, M.G.C. Go N D I M Jr. & r. BA r r o S. 2009. 

Infestation of coconut fruits by Aceria guerreronis enhances the 

pest status of the coconut moth Atheloca subrufella. Annals of 

Applied Biology 155: 277–284. 

SA N tA N A, S.w.J.; r. BA r r o S, J.B. to r r e S & M.G.C. Go N D I M Jr. 2011. 

Técnica de criação e aspectos biológicos de Atheloca subrufella 

(Hulst) (Lepidoptera: Phycitidae) em frutos de coqueiro. 

Neotropical Entomology 40: 14–19. 

SHAFFer, J.C. 1995. Phycitinae, pp. 93–105. In: Heppner, J.B. (ed.), 

Atlas of Neotropical Lepidoptera. Checklist: Part 2 Hyblaeoidea 

– Pyralopidea – Tortricoidea. Association for Tropical 

Lepidoptera and Scientific Publishers, Gainesville, Florida. 

SC H o t M A N, C.y.l. 1989. Plant pests of quarantine importance to 

the Caribbean. RLAC-PROVEG, No. 21, Food and Agriculture 

Organisation of the United Nations, 80 pp. 

So l I S, M.A. 1993. A phylogenetic analysis and reclassification of the genera 

of the Pococera complex (Lepidoptera: Pyralidae: Epipaschiinae). 

Journal of the New York Entomological Society 101: 1–83. 

So l I S, M.A. 2011. Key to selected Pyraloidea (Lepidoptera) larvae 

intercepted at U. S. ports of entry: revision of Pyraloidea in 

“keys to some frequently intercepted lepidopterous larvae” 

by Weisman 1986. http://www.ars.usda.gov/Services/docs. 

htm?docid=21551. 58 pp. 

21 

Sw e z e y, o.H. 1909. The Hawaiian sugar cane bud moth (Ereunetis 

flavistriata) with an account of some allied species and natural 

enemies. Hawaiian Sugar Planters’ Association Experimental 

Station, Entomological Bulletin 6: 1–40, plates 1–4. 

Sw e z e y, o.H. 1910. The feeding habits of Hawaiian Lepidoptera. 

Proceedings of the Hawaiian Entomological Society 2: 

131–143. 

tA y l o r, t.H.C. 1930. Early nutfall from coconut palms in Fiji with 

special reference to insects attacking the flowers. Bulletin of 

the Department of Agriculture, Fiji 17: 1–42. 

tr e B r o w N NU r S e r I e S. 2011. Classification of Palm Trees [Arecaceae]. 

http://www.trebrown.com/palms_arecaceae_order.php (Last 

accessed 19 Dec 2012). 

vA N NI e U k e r k e N, e.J., l. kA I l A, I.J. kItCHING, N.p. kr I S t e N S e N, D.C. 

le e S, J. MI N e t, C. MI t t e r, M. MU tA N e N, J.C. re G I e r, t.J. SI M o N S e N , 

N. wA H l B e r G, S.-H. ye N, r. zA H I r I, D. AD A M S k I, J. BA I x e r A S, D. 

BA rt S C H, B.A. BeNGtSSoN, J.w. Br o w N , S.r. BU C H e l I, D.r. DAv I S, 

J. De pr I N S, w. De pr I N S, M.e. ep S t e I N, p. Ge N t I l I-po o l e, C. GI e l I S, 

p. Hä t t e N S C H w I l e r, A. HA U S M A N N, J.D. Ho l l o w A y, A. kA l l I e S, o. 

kA r S H o lt, A.y. kAwA H A r A, S. ko S t e r, M.v. ko z l o v, J.D. lA F o N tA I N e, 

G. lA M A S, J.-F. lA N D r y, S. le e, M. NU S S, k.-t. pA r k, C. pe N z, J. 

ro tA, A. SC H I N t l M e I S t e r, B.C. SC H M I D t , J.-C. So H N, M.A. So l I S, 

G.M. tA r M A N N, A.D. wA r r e N, S. we l l e r, r.v. yA k o v l e v, v.v. 

zo l o t U H I N & zw I C k , A. 2011. Order Lepidoptera Linnaeus, 

1758, pp. 212–221. In: Zhang, Z.-Q. (ed.), Animal biodiversity: 

An outline of higher-level classification and survey of taxonomic 

richness. Zootaxa 3148: 1–237. 

ve l á z Q U e z, J., J.l. GA r C í A, y. ro M e r o & M. Me D I N A. 2010. Phidotricha 

erigens Ragonot, 1888 (Lepidoptera: Pyralidae), causando daño 

en Agave cocui Trelease, en el estado Falcón, Venezuela. Algunos 

aspectos de su biología y su control natural. Entomotropica 

25: 117–124. 

ve S e y-FI t z G e r A l D, D. 1941. Some insects of economic importance in 

Seychelles. Bulletin of Entomological Research 32: 153–160. 

wA l S I N G H A M, lo r D. 1882. Notes on Tineidae of North America. 

Transactions of the American Entomological Society 10: 

165–204. 

wAl S I N G H A M, lo r D . 1897. Revision of the West-Indian Micro- 

Lepidoptera with descriptions of new species. Proceedings of 

the Zoological Society of London 1897: 54–183. 

wA l S I N G H A M, lo r D. 1891. On the Micro-Lepidoptera of the West 

Indies. Proceedings of the Zoological Society of London 1891: 

491–549, plate XLI. 

wA t e r H o U S e , D.F. & k.r. No r r I S. 1987. 26. Tirathaba rufivena 

(Walker), pp. 211–218. In: Waterhouse, D.F. and K.R. Norris, 

Biological Control: Pacific Prospects. Inkata Press, Melbourne, 

Australia. 

zI M M e r M A N , e. C. 1978. Microlepidoptera. Part 2, Gelechioidea. 

Insects of Hawaii 9(2), pp. 883–1903, figs 608–1355. University 

of Hawaii Press, Honolulu, Hawaii.

Neotropical palm-inflorescence feeding moths (Lepidoptera ...

Create successful ePaper yourself

Delete template?

Save as template?