Volume 10 Number 10 20 December 2022

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

ISSN 2643-4776 (print) / ISSN 2643-4806 (online)

Establishment of the invasive Cactus Moth, Cactoblastis cactorum

(Berg) (Lepidoptera: Pyralidae) in Pakistan: a potential threat to

cultivated, ornamental and wild Opuntia spp. (Cactaceae).

‘Muhammad Ather Rafi, 7Harry Pavulaan, “>Muhammad Islam, ‘Muhammad Ashfaq,

4Haseeb Kamran, °Walija Fayaz, “Gul Naz Parveen, ’Riffat Sultana, ‘Ahmad Zia, >Waqar

Ahmed, ’Qudrat Ullah, !°Muhammad Qasim, !'Falak Naz, §Nazeer Ahmed, !*Muhammad

Tariq Khan, "Muhammad Saeed, “Jalal Hayat Khan.

'National Insect Museum, National Agricultural Research Center, Islamabad, Pakistan; ’The International Lepidoptera

Survey, Leesburg, Virginia, USA; *Rangeland Research Institute, National Agricultural Research Centre, Islamabad,

Pakistan; *Department of Zoology, University of Education, D.G. Khan, Punjab, Pakistan; °>Department of

Entomology, Faculty of Crop Protection Sciences, The University of Agriculture, Peshawar, Pakistan; ‘Department of

Botany, Women University Swabi, Swabi, Khyber Pakhtunkhwa, Pakistan; ‘Department of Zoology, University of

Sindh, Jamshoro, Pakistan; "Department of Entomology, University of Swabi, Khyber Pakhtunkhwa, Pakistan;

’Department of Zoology, University of Peshawar, Peshawar, Pakistan; ‘Department of Zoology, Kohsar University

Murree, Punjab, Pakistan; ''Coordination & Monitoring Division, Pakistan Agricultural Research Council, Islamabad,

Pakistan; '*Department of Plant Protection, Karachi, Pakistan.

ABSTRACT. Subsequent to the significant accomplishment of biological control of Opuntia weeds in

Australia, the larvae of the cactus moth, Cactoblastis cactorum (native to parts of South America), were released in

many countries for the biological control of native Opuntia species (Simmonds & Bennett, 1966). Inauspiciously,

larvae were also released in the Caribbean, where the moth spread naturally and by the human support all over the

region (Garcia-Turudi et al, 1971). Its enhanced dissemination rate and the biological potential for invasiveness,

suggests that the cactus moth is likely to become an invasive pest of Opuntia in the Southeast United States, Mexico

and southwestern America. Its damage is restricted mainly to the plants of genus Opuntia (plants with the

characteristic of flat prickly pear pads of the former genus Platyopuntia, now considered to be the part of the genus

Opuntia). In this region, plants of this genus provide valuable resources for humans, livestock, and wildlife such as

food, medicine, and emergency fodder, while in the arid and semi-arid regions, the plants play key roles in ecosystem

processes and soil conservation. At present, the cactus moth has developed into a severe threat to the high diversity of

prickly pear cacti, all over the world for both the native and cultivated species of Opuntia IAEA, 2002).

During the year 1994, the larvae of C. cactorum were also released in Pakistan for the control of naturalized

wild Opuntia weed in the district Chakwal, Punjab. The establishment of the cactus moth was not confirmed in

Pakistan after its release in district Chakwal until December 2016 and various unconfirmed presence records of many

authors have been found (e.g., Julien & Griffiths, 1998; Zimmermann et al., 2004; Legaspi & Legaspi, 2010; EPPO,

2021). Twenty-three years after the introduction of C. cactorum larvae in Pakistan, presence of larvae was monitored

on cultivated prickly pear cactus (1.e., Opuntia ficus-indica (L.) Mill.) where O. ficus-indica was introduced in three

localities of district Chakwal for fodder purposes during 2014. Further surveys were conducted consecutively for wild

Opuntia in the four tehsils of the district until August 2020 and it was found that all plantations of wild Opuntia in

the district were severely infested with the larvae of the cactus moth. In 2019, all the three fields cultivated with

spineless species (1.e., O. ficus-indica) had completely vanished. The occurrence of C. cactorum larvae on cultivated

spineless species (O. ficus-indica) and wild Opuntia in the district of Chakwal, Punjab, Pakistan in December, 2016

1

is the first authentic record after the 1994 release of its larvae, which was not effectively studied after the release.

Additional key words: Invasive species, biological invasion, natural ecosystems, biological control,

Opuntia cactus.

INTRODUCTION

The natural ecosystems that exist today are impartially different from those that have existed

in the past and in many parts of the earth are being significantly changed owing to climate change

and enormous alterations of the land for multipurpose activities (Mooney & Cleland, 2001; Sala et

al., 2000). However, similarly drastic changes both beneficial and harmful are happening in the

population, economic activities and community structure of natural ecosystems due to the

occurrence of biological invasions (Pimentel et al., 2005; Vila et al., 2011; Blackburn et al., 2014;

Jeschke et al., 2014).

The phenomenon of biological invasion is characteristic of living organisms and has been

occurring since the origin of life (animals, plants and microbes) and long before human existence

on this planet. The incidence of biological invasion is of significant scientific attention and

recurrently of socioeconomic importance (Mitchell & Gopal, 1991). Such biological invaders are

known as “alien species” or invasive taxa occurring outside of their natural range and have potential

to disperse. They are non-native, non-indigenous, introduced, foreign or exotic. An alien species

which has become established outside its native range, in natural or semi natural ecosystems or

habitats, which can threaten native species and biological diversity, is known as an “Invasive Alien

Species” (IAS). Impacts of alien or alien invasive species on native fauna and flora can be found

in all terrestrial and freshwater environs (Parker et al., 1999; Rahel, 2000).

Historical movement of biological organisms throughout the taxonomic groups, substantial

travelling of humans and massive trade of goods and materials, helped the movement of species,

worldwide; however, others are being traded among continents or various nations (Bryan, 1996;

USBC, 1998). Recently, the introductory rate and the threat of IAS have increased immensely

because of human population growth and human activities, which alter the environment at an

alarming rate (Pimentel et al., 2000; Mooney & Cleland, 2001). The definite statistics of individuals

and species which have transported and are being transported across bio-geographical boundaries

daily are probably enormous. Unfortunately, this event has augmented species wealth in numerous

places where new species are introduced (Mooney & Cleland, 2001). However, only an

insignificant fraction of introduced species become established in a new environment, and among

those usually about 1% become pests (Williamson, 1996).

Biological invasions

The world’s most destructive 100 IAS include microorganisms, macro fungi, plants,

invertebrates, amphibians, fishes, birds, reptiles, and mammals (Lowe et al., 2000). Rapid increase

in their ranges is homogenizing the world’s flora and fauna (Mooney & Hobbs, 2000) and such

biological invasions may be observed as a form of biological pollution and major elements of global

changes and one of the major causes of severe species declines that may be followed by extinction

(Drake et al., 1999). They cause major economic losses in agriculture, forestry, many other sectors

2

of the world economy and also negatively impact on ecological integrity in countries around the

world, either by depressing growth of populations of more valuable species, or by direct impacts

on human and animals. Invasive alien species of many weeds, arthropods, and animals are the

greatest direct threats to biodiversity (Elton, 1958; Mooney & Drake, 1986; Mack & D’ Antonio,

1998; Pimentel et al., 2000; Vila et al., 2011; Blackburn et al., 2014).

In certain regions of the world enormous land and water bodies are completely dominated

by invasive alien species of plants such as Centaurea solstitialis (the yellow star thistle) which is

native to the Mediterranean Basin region, now found invasive in the rangelands of California;

Bromus tectorum (cheatgrass) is native to Europe, southwestern Asia and northern Africa, and now

invasive in the mountain ranges of the western USA; and Eichornia crassipes (water hyacinth) is

an aquatic plant native to the Amazon basin, now invasive in numerous tropical lakes and

waterways (White, 1985; Mitchell & Gopal, 1991). Besides that, invasive alien pathogens threaten

humans and animals by causing emerging infectious diseases such as highly pathogenic avian

influenza and swine influenza have the ability to mutate and threaten animals and humans (Hunter,

2009).

Many alien species have been introduced for beneficial purposes, and their beneficial

impacts cannot be ignored, such as corn (Zea mays L.); rice (Oryza sativa L.); wheat (Triticum

spp.); plantation forest; domestic chicken (Gallus spp.); cattle (Bos taurus); and many others that

provide more than 98% of the world’s food (Pimentel et al., 2001), with a value of more than US$

5 trillion per year (USBC, 1998). Similarly, the European honey bee, Apis melliferais, native to

Africa, Europe and western Asia, vigorously introduced in various countries by humans during the

17" century (Winston et al., 1981; Sammataro & Avitabile, 1998), is a highly valued species all

over the world for its production of honey and is a keystone pollinator of many economically

important crops and wild plants (Klein et al., 2007; Staveley et al., 2014; Pashte & Said, 2015).

However, many alien species are used for landscape restoration, sport, pets, food processing, and

biological control agents against pests (Mack & D’ Antonio, 1998; Pimentel et al., 2000).

In many cases biological control has been severely prejudiced by abiotic factors, such as

climatic factors (Olfert et al., 2016) which play a significant role in the growth and existence of

flora and fauna; for example, insect pest species which are minor at their origin may become

invasive in their new environment. There are numerous alien species that have been introduced as

biological control agents for specific purposes, now have unwanted non-target effects; for example,

in weed biological control the musk thistle weevil, Rhinocyllus conicus (Coleoptera:

Curculionidae), introduced for the biological control of thistle weeds in Canada, 1968; in USA:

Montana and Virginia during 1969 and in California, 1971 (Zwolfer & Harris, 1984), is now

attacking the seed-heads of native thistles over large areas in the West and Central United States

(Turner, 1985; Turner et al., 1987; Guretzky & Louda, 1997; Louda et al., 1997; Strong, 1997).

The deliberate introduction of Opuntia stricta in Australia and Africa, attained the status of weeds

(Dodd, 1940; Greathead, 1971; Foxcroft et al., 2004; Novoa et al., 2016).

The harmful effects of introduced biological control agents on organisms other than the target

pest have, rightly, been critiqued and the safety of biological control as a practice has been

questioned (Howarth, 1991; Miller & Aplet, 1993; Simberloff & Stiling, 1996; Thomas & Willis,

3

1998). Great care with strict guidelines is now required for the introduction of herbivores, mainly

for plant-feeding insects (Howarth, 1991). Although much has been learned by the impacts of

Cactoblastis cactorum in Australia, while plenty of evidences predicted the worst, but there are

also warnings that preclude a repeat of the Australian experience.

Introduction of Opuntia spp. in Australia

In Australia no native prickly pear species were present, where prickly pear cactus (Opuntia

stricta Haw.) was deliberately introduced in the year 1788 to establish a cochineal industry (Dodd,

1940). Cochineal is a valued source of dye i.e., red pigment carminic acid, mostly extracted from

the scale insect species, Dactylopius coccus Costa; however, D. coccus did not survive there and,

by the 1920s, Opuntia stricta invaded about 25 million hectares of land that was then useless for

agriculture production (Dodd, 1940).

Importance of carminic acid dye

This dye was primarily used in food colouring, cosmetics, drugs, fabrics and many other

products (Canamares et al., 2006; Chavez Moreno et al., 2009), and also used to colour the typical

red coats of the British soldiers in the past (Dodd, 1940). Eleven Dactylopius species (cochineal

insects), in the monogenetic scale insect family Dactylopiidae, commonly known as cochineal scale

insects, are a parasite of certain species of prickly pear cactus (Dapson, 2007), flourishing on

Opuntia plants are mostly known as a natural basis of crimson dye. Fresh cochineal is a red liquid

within the body of Dactylopius spp. from which crimson or carmine dye is extracted. These insects

have been used for commercial purposes since the 16" century in Central and South America, in

Mexico and in Spain (Chavez-Moreno et al., 2009; De Lotto, 1974; Pérez-Guerra & Kosztarab,

1992). Out of eleven Dactylopius species, D. coccus has been the main species which has been

cultivated and used commercially because of the quality and quantity of its pigments (Pifia, 1977);

other species were also used (de Humboldt, 1811; Anderson, 1981). Dactylopius opuntiaeisis is

also reared in many countries for the production of carmine dye. The economic importance of it is

shown through its introduction to South Africa, Australia, India, Sri Lanka, etc. to improve the dye

industry (Kumar et al., 2018). However, all the eleven species in the family Dactylopiidae are

exclusively pests of cactus species (Caryophyllales: Cactaceae). Currently, there is ongoing

research on the antioxidant and antimicrobial properties of carminic acid, to understand its potential

applications in immunology, wastewater treatment and solar cells (Gonzalez et al., 2009; El

Moselhy et al., 2011).

Other importance of Dactylopius spp.

Some Dactylopius species have been used for biological control against invasive cacti

(Volchansky et al., 1999; De Felice, 2004); Dactylopius spp. can become invasive to Opuntia

species where they are non-native (Van Dam & May, 2012).

Biological control of Opuntia weed (O. stricta Haw.) in Australia

The larvae of the cactus moth, Cactoblastis cactorum (Berg), (Lepidoptera: Pyralidae) was

selected to control the invasive weed, Opuntia stricta in Australia in 1926. The cactus moth is

4

native to the South American countries of Peru, Bolivia, Paraguay, Uruguay, Argentina and

southern Brazil. Larvae of C. cactorum were transported from Argentina to Australia, and

deliberately introduced as a biological control agent. The moth successfully controlled the exotic

Opuntia species (Dodd 1940; Mann, 1969; McFadyen, 1985; Zimmermann et al., 2000 a, b).

Within a few years after its release in Australia, millions of hectares of lands infested by prickly

pear (O. stricta) were given back to agriculture and livestock farming. This was a remarkable

achievement that stimulated fruitful developments in the field of biological control of weeds mainly

by one species of the insect. After the successful attempt in Australia, the larvae of the cactus moth

were released in several other countries for the biological control of invasive Opuntia weed. (Dodd,

1940; McFadyen, 1985; Briano et al., 2012).

The cactus moth Cactoblastis cactorum (Berg)

The cactus moth, Cactoblastis cactorum

(Lepidoptera: Pyralidae) (Fig. 1) is one of the five

species in the genus Cactoblastis, namely C.

cactorum, C. bucyrus, C. mundelli, C. doddi and

C. ronnai. All of these are native to the southern

part of South America, from southern Peru to

Bolivia, Paraguay, Uruguay, Argentina and

southern Brazil (Heinrich, 1939). Except for C.

cactorum, the rest of the species have very

restricted host ranges with limited distributions.

C. cactorum has a wide host range within the Fig. 1. Cactoblastis cactorum (Cactus Moth).

Cactus genus Opuntia, and its larvae have been [Image from Wiki Commons]

reported to feed on many species of this genus in

the native countries, but its spread is controlled by native pathogens and parasites. The female

cactus moth lays eggs, one on top of the other to form spine-like “egg sticks“ that comprise, on

average, 60—100 eggs and each female usually produces about 300 eggs (Dodd, 1940; Pettey, 1948).

The newly hatched larvae jointly make a small hole in the cactus cladode and enter the cladodes

through the hole (Hoffmann & Zimmermann, 1989). The larvae feed gregariously inside the

cladode on mucilage and repeatedly facilitate secondary pathogenic infections and ultimately, death

of the plant (Heinrich, 1939; Dodd, 1940; Mann, 1969; Starmer et al., 1988; Zimmermann et al.,

2000, 2004). Larvae remain inside the cladode for about two months in summer and about four

months in winter, then exit the damaged cladode for pupation in the soil or in the leaf-litter (Dodd,

1940; Pettey, 1948). However, full grown larvae also exit from the hole during the hot season and

can be seen in a large group, in a group of 2-4, or singly (Rafi, personal observation). In Australia

and South Africa, C. cactorum occurs in temperate latitudes, where it has two or seldom three

generations per year (Pettey, 1948; Robertson, 1985). In the warmer tropical climate of the

Caribbean and Florida there may be more generations each year (Zimmermann et al., 2004).

Release of Cactoblast cactorum in several other countries to control Opuntia weed

e In 1933, larvae were released to control O. ficus-indica in South Africa, where O. ficus indica

was prominent weed species and the moth became established. In South Africa the moth played

5

only a minor role in controlling invasive cacti O. ficus-indica (Pettey, 1948; Hoffmann et al., 1998;

Julien & Griffiths, 1998).

e During the same year larvae were sent from Australia to New Caledonia (northeast of Australia),

where it became established (Zimmermann et al., 2004).

e In 1950, cactus moth larvae were transported from Australia and released in Hawaii (Fullaway,

1954) where it became established.

e Also in 1950, cactus moth larvae were released in Mauritius, having been transported from South

Africa to control Opuntia weeds, i.e. O. ficus-indica, O. tuna and O. monacantha, where it became

established (Zimmermann et al., 2004).

e In 1957, cactus moth larvae were sent to the Caribbean island of Nevis from South Africa to

control native Opuntia. The biological control was successful and its population established

successfully (Julien & Griffiths, 1998).

e In 1960, after the successful biological control on Nevis Island, larvae of C. cactorum were

released in the neighboring Caribbean islands (Antigua & Montserrat) from the population of Nevis

Island (Julien & Griffiths 1998).

e In 1966, cactus moth larvae were sent from Antigua to Kenya, but its establishment was not

confirmed until 2000 (Zimmermann et al., 2000 a, b), with its establishment reconfirmed in 2010

(Legaspi & Legaspi, 2010).

e In 1970, cactus moth larvae were sent from Nevis and Antigua to the Cayman Islands in the

Western Caribbean Sea, with its establishment confirmed (Zimmermann et al., 2004).

e In 1971, cactus moth larvae were transported from Nevis and Antigua to Saint Helena Island,

with its establishment confirmed (Zimmermann et al., 2004).

e In 1973, cactus moth larvae were sent from Saint Helena to Ascension Island to control the

native species of the Opuntia, with its establishment confirmed (Zimmermann et al., 2004).

e In 1994, cactus moth larvae were released from Australia to Pakistan to control wild Opuntia

weed, where this has not been effectively studied; hence, its establishment not confirmed until

December 2016 (Julien & Griffiths 1998; Zimmermann et al., 2004; Legaspi & Legaspi, 2010). Its

first outbreak was observed in December 2016 on cultivated spineless species of Opuntia, 1.e. O.

ficus-indica and wild Opuntia in the district Chakwal. After personal communications it was

confirmed that cactus moth larvae were released in the district Chakwal and its vicinities in 1994.

e Cactus moth larvae were sent to Israel from South Africa (year not reported), where its

establishment is unconfirmed (Julien & Griffiths, 1998; Zimmermann et al., 2004; Legaspi &

Legaspi, 2010).

Current dependable records of C. cactorum either introduced or naturally spreading

throughout the world.

e North America: Reported throughout the Caribbean islands (Garcia-Turudi, 1971; Legaspi &

Legaspi, 2010; Habeck et al., 2016); Mexico: Most of the area in the Mexico (EPPO, 2021); USA:

Florida Keys, Hawaii, Texas, Alabama, Georgia, Mississippi, Louisiana and South Carolina

6

(CABI, 2022). The cactus moth extended its range in the United States subsequent to introduction

with movement westward along the Gulf Coast and northward along the Atlantic Coast (Dickel,

1991; Jezorek et al., 2012).

e South America: Native to Peru, Bolivia, Paraguay, Uruguay, Argentina and southern Brazil

(Heinrich, 1939; Invasive Species Specialist Group (SSG), 2011).

e South Africa: Saint Helena and Ascension Island (Invasive Species Specialist Group (ISSG),

2011).

e East Africa: Kenya (Legaspi & Legaspi, 2010); Tanzania and Mauritius (EPPO, 2021)

e Oceania: Australia: New South Wales, Queensland (EPPO, 2021); New Caledonia (Invasive

Species Specialist Group (ISSG), 2011).

e Asia: India: southern India (Legaspi & Legaspi, 2010; EPPO, 2021).

e Asia: Pakistan: Present (presence records are confirmed) from district Chakwal, Punjab

Province (this report Rafi, 2022).

Unluckily, in 1957, C. cactorum was purposefully introduced into the Caribbean region for

the purpose of biological control of cacti species of the genus Opuntia. In the Caribbean region the

moth spread all over the region naturally (a potential invasion route matches historical hurricane

trajectories) and by human-supported introductions (Simmonds & Bennett, 1966; Garcia-Turudi,

1971; Andraca-Goémez et al., 2015). The moth conquered the North American continent during

the last 20-30 years, threatening the most important center of biodiversity of native Opuntia

species, where hurricanes are one of the major ecological reasons for the dispersal of C. cactorum

in the region (Andraca-Gémez et al., 2015). The moth was reported from the Florida Keys in 1989

(Dickel, 1991) and has spread throughout most of the Florida peninsula, along the Atlantic Coast

to North Carolina, and the Gulf Coast to Texas (Varone, 2020). Outbreaks of C. cactorum

populations have taken place in Mexico: Quintana Roo and on Isla Mujeres in 2006 and Isla Contoy

in 2007, and then eradicated by 2009. Due to its rapid expansion in geographical range and its

status as a voracious feeder of Opuntia species, C. cactorum continues to threaten Mexico’s

Opuntia diversity and its industry (Hight & Carpenter, 2009).

As stated above, the arrival of C. cactorum in continental North America is a major concern.

It has high potential to destroy native Opuntia spp. In the Florida Keys, Opuntia spinosissima

(Martyn) Mill., and O. tricantha (Willdenow) are rare and included in the threatened list.

Other native species, such as O. cubensis Britton & Rose, O. stricta Haw.and O. humifusa (Raf.),

in addition to the exotic species, either grow wild or as ornaments are also in danger. In the

desert regions of Texas and Mexico, wild Opuntia and Cylindropuntia (Engelmann)

Kreuzinger species, which provide food and nesting sites for a variety of wildlife and contribute to

soil stability, are at risk (Chavez-Ramirez et al., 1997). In Mexico these plants are harvested on

over three million hectares, where they grow naturally (Soberon et al., 2001; Vigueras & Portillo,

2001). In addition, 250,000 hectares of Opuntia and Nopalea species are cultivated for human

and livestock food, for fuel, medicines, crimson dye and fencing etc.; with cultivation has occurred

since 9,000 BC in Mexico (Casas et al., 1997; Vigueras & Portillo 2001). In Mexico these plants

are valued at over 80 million dollars annually (Soberon et al., 2001). Mexico, with 850 cacti

7

species, has the highest diversity of cacti, with many of them endemic (L6pez Collado et al, 2013;

Pérez-De la, 2020).

Control methods

Several tactics have been evaluated for cactus moth control, including control strategies

suggested to minimize the risk and consequences of invasion by the cactus moth; for example, hand

removal of infested cladodes and egg sticks (Zimmermann et al., 2004). Substantial research has

been conducted on the potential use of sterile insect technique “SIT” (Hight et al., 2005),

pheromones for the monitoring for the presence and occurrence of the moth (Heath et al., 2006)

and insecticidal applications (Bloem et al., 2005). Still, none of these control tactics prevented the

spreading of the C. cactorum. Painted apple moths (Teia anartoides) were successfully eradicated

at Auckland, New Zealand, using the release of partially sterilized males in combination with aerial

applications of Bacillus thuringiensis var. kurstaki (Btk) and an intensive trapping program, using

virgin female moths (Suckling et al., 2007). The same approach was used to eradicate the outbreaks

of invasive cactus moth in Mexico (Bello-Rivera et al., 2021).

The use of exotic biological control agents has been assessed as a management tactic

(Pemberton & Cordo, 2001a, b; Stiling, 2002). For example, one parasitoid, Apanteles opuntiarum

Martinez & Berta (Hymenoptera: Braconidae), has recently been reported as a potential candidate,

because its host range is restricted to the genus Cactoblastis in its country of origin (i.e. Argentina),

where it is a gregarious larval parasitoid of C. cactorum (Martinez et al., 2012; Mengoni et al.,

2014; Varone et al., 2015). Colonies of A. opuntiarum are presently maintained in a quarantine

facility in Florida undergoing host specificity testing on North American non-target species

(Srivastava et al., 2019). However, the potential areas where this braconid wasp could become

established after its release in North America are still unknown, which is an important aspect

essential to the success of a biological control agent (Pérez-De la et al., 2020). Other reported

natural enemies are the pathogen Nucleo polyhedrosis virus (CABI, 2022), the parasitoid

Phycitiplex doddi (Hymenoptera: Ichneumonidae) (CABI, 2022) and the egg parasitoids

Trichogramma fuentesi and Trichogramma_ pretiosum (Hymenoptera: Trichogrammatidae)

(Paraiso et al., 2011).

FOOD SECURITY IN ARID AND SEMI-ARID REGIONS

The shared effects of climate change and IAS have negative impacts on food security and

availability of basic needs, such as water and energy in poorer countries (Masters & Norgrove,

2010), uneven rainfall throughout the growing season, with poor nutrient soils and high

temperatures, which limit plant growth, especially of agricultural and forage crops in the arid and

semi-arid regions (Teklehaimanot & Tritschler, 2011; Louhaichi et al., 2018). Furthermore,

increases in human population, placing the burden on natural resources and livestock production

(Seto et al., 2011), result in increased risks of desertification, decline in rangeland resources;

overgrazing, ploughing of marginal land, and soil erosion (Owen et al., 2004; Pimentel et al., 2010;

Hudson et al., 2014). Increasing cropland worldwide to meet human demands, continuously

diminishes rangelands which cannot fulfill the annual feed requirements of livestock (Nefzaoui et

al., 2014). Livestock often depend on natural grazing or low-quality crop residues (e.g. straws,

8

stubbles) and expensive feed supplements when forage has been depleted and overgrazed (Aregawi

et al., 2008; Islam et al., 2019). Consequently, goats and sheep (the ideal livestock animal and the

leading producers of milk, meat, fiber, manure & skins) are facing serious nutrient shortages in arid

and semiarid areas and livestock production in such regions facing challenges (Russell & Felker,

1987; Islam et al., 2019).

Raising of livestock is critical for the survival and livelihood for the people in these areas

(Gusha et al., 2015). Hence, it is essential to find alternative non-conventional feed resources to

sustain food production for human and livestock and improve the feed-water problem during dry

and low-producing seasons. The utilization of forage species that adapt to the changing and poor

growing conditions contributes to ensure livestock security, in the face of an increase in both

livestock and human population (Burke et al., 2009). Under these challenging conditions, planting

of drought-tolerant and water- efficient plants, especially under the genus Opuntia, that are hardy

and well adapted as buffer food reserves, for example as a form of food assurance for the frequent

droughts and to bridge a regular annual period of shortage of food for human and improve livestock

production in various arid and semi- arid zones of the world, is also a favorable way of lessening

the demand for water (Louhaichi et al., 2018). Several studies have been confirmed that cactus

cladodes have a good nutritional value when used as supplements in small ruminant diets (Misra et

al., 2006; Aguilar-Y afiez et al., 2011; Gusha et al., 2015).

Cactus pears have the benefit of being a source of ample water for animals,

mostly during the dry season; these plants are also tolerant of poor soil situations, and yield high

biomass with acceptable palatability to animals (Russell & Felker, 1987). Cactus species have the

capability to survive prolonged drought, high temperatures, as well as wind and water loss; hence,

they are ideal for agricultural and livestock growth in areas affected by the world’s

two biggest environmental problems: desertification and climate change, as they relate to higher

temperatures and reduced soil moistness accessibility (Ben Salem & Abidi, 2009). Cacti are

generally used as survival fodder, and, given its availability in conditions not suitable for

other agriculture or better forage production, it is important to find options to make their use

more efficient, framing it within a context of sustainable animal production (Atti et al., 2009).

Cultivation of prickly pear cactus in Pakistan

In arid and semi-arid regions of Pakistan livestock production remains a critical source of

income for the rural population. Sheep and goats raised in these areas are mostly confronted with

severe malnutrition, in addition to the negative impacts of climate change (Devendra, 1998; 1999;

McDermott et al., 2010). Increasing numbers of livestock is one of the vital changes for the survival

and sustainable livelihood for the people of these areas. To examine the effects of adding cacti as

a supplement to sheep grazing on degraded rangelands of the district Chakwal, Pakistan. Spineless

cactus (Opuntia ficus-indica) was introduced to the farmers in the village of Chakwal, as a succulent

and drought-tolerant species with great potential to provide fodder reserves to fill the gap during

the periods of low feed accessibility and provision of spineless cactus. As a supplement to sheep,

this has a positive impact on live-weight gain, compared with only grazing poor rangeland

conditions, and it was found that cactus is important to develop diets that maximize the full

potential of low-nitrogen, yet water- and carbohydrate-rich as this unconventional feed source

9

(Islam et al., 2019).

After these promising results to promote cactus cultivation in the arid and semi-arid regions

of Pakistan, in 2014, the Pakistan Agriculture Research Council (PARC) (an apex scientific and

research organization at the national level in the agriculture sector) imported many cultivar/varieties

of spineless cacti from Brazil (Table 1, Fig. 2). Cladodes of all imported cultivar/varieties of

spineless cacti were initially planted at National Agricultural Research Centre (NARC), Islamabad

under the Rangeland Research Institute (RRI), NARC. The growth of all the cultivar/varieties was

found satisfactory and a plan was prepared to cultivate the spineless prickly pear 1.e., Opuntia ficus-

indica in the arid and semi-arid areas of the district Chakwal.

Table 1: List of varietal/cultivars clones of cactus imported from Brazil

Serial Scientific name Common Name Origin

No.

| 06 | Nopaleacochenillifera__|__—Jaguarear_——————s|_~S—Arcoverde

[08 [Opuntia undulata Griffiths} African elephant ear_[ _Caruaru

| 09 | Opuntia atropesRose__ | __——sF8-Forage |S Carvarn

10

; : if ~ ’

AAR 1 key

Aa Be

Fig. 2. Imported Cacti germplasm at National Agricultural Research Centre (NARC), Pakistan.

The district Chakwal is located at 32°56'17"N, 72°51'30.71"E., on the Pothohar Plateau,

Punjab, Pakistan (Fig. 3). It is situated in the north of the Punjab province, bounded by district

Khushab to its South, district Rawalpindi to its northeast, district Attock to its northwest, district

Jhelum to its East and district Mianwali to its West. District Chakwal is comprised of four tehsils

namely Kallar Kahar, Chakwal, Choa Saidan Shah and Talagang. Dominant flora of the Chakwal

districts consists of Acacia modesta, Acacia nilotica and Dalbergia sissoo. Approximately 50%

of the area is uncultivated and primarily used for grazing animals, and about 11% is natural grasses

and shrubs. District Chakwal is the largest peanut producer in Pothohar region, where the peanut

is largely grown in the tehsil of Chakwal and Talagang, and peanut stubble is the main stock to the

small ruminants in the area. Due to high demand of peanut straw in large cities, the shrubby

vegetation mainly supports the grazing of goats and sheep.

Fig. 3. District Chakwal, Punjab-Pakistan

11

MATERIALS AND METHODS

Selection of study areas

For the cultivation of spineless cactus pear, i.e. Opuntia ficus-indica, three fields were

selected in the vicinities of the villages Latifal and Beghal within the tehsil and district Chakwal

and the village Dhulli in tehsil Talagang, district Chakwal. The fields were selected by three

gradients of rainfall, high, medium, and low, and population of small ruminants in selected areas,

in consultation with local livestock extension departments of each tehsil. Selected villages

represented the larger territory of Pothohar regions. The main purpose for selection of different

sites was to identify the situations that are representative of the larger domain of rain-fed Pothohar

regions with respect of small ruminant farming.

Target villages:

e Village Latifal was selected in the high rainfall zone, located at 33°09'48”N, 72°50'28”E, and

situated in tehsil Chakwal about 41 kilometres from Chakwal city near the boundary of

Rawalpindi and Chakwal districts.

e Village Begal was selected in the medium rainfall zone, located at 33°2'57”°N, 72°39'11”E. This

village is also a part of tehsil Chakwal.

e Village Dhulli was selected in the low rainfall zone, located at 32°51'31.02”N, 72°11'28.22”E.

This village is a part of tehsil Tatagang.

Plantation of Opuntia ficus-indica in fields of selected Villages

After the 2014 monsoon, Rangeland Research Institute, NARC under Agriculture

Innovation Program (AIP) introduced spineless prickly pear species, i.e. Opuntia ficus-indica to

cultivate in the fields of the targeted villages on approximately 30 ha (Fig. 4).

a a | "2 : Se

By ies a. P| ae i

ro * 1 * 5 —

as

Fig. 4. Cultivation of spineless cactus O. ficus-indica under AIP in Chakwal for fodder purposes.

12

Survey after plantation

After planting, all fields were visited once a month. The growth of O. ficus-indica was

found to be excellent in all the targeted fields until November 2016. However, during the

December 2016 survey it was observed that a few cladodes of O. ficus-indica were found dead

with infestation of an unknown pest (Fig. 5). All infested pads were removed and burned. In

December 2017, the same infestation was again reported and the majority of the cladodes were

found to be infested. Infested cladodes were examined in the field and some lepidopterous larvae

were found inside the cladodes (Fig. 6). Larvae were brought to the laboratory of the National

Insect Museum for identification. However, in July 2019 it was found that cultivated O. ficus-

indica at all the three localities had completely vanished due to the heavy attack by cactus moth

larvae (Figs. 7 & 8).

ne i hd :

Caneel ks

Fig. 5. Infested cladode of spineless O. ficus-indica Fig. 6. Larvae inside the cladode of O. ficus-indica

in cultivated field in District Chakwal. in District Chakwal.

i | i . ea ee es “Me

Ne ee | Py tim Wa

Fig. 7. Dead cladode of O. ficus-indica in vanished Fig. 8. Completely vanished field of O. ficus-indica

field in District Chakwal. in District Chakwal.

13

RESULTS

The field collected larvae were brought to the laboratory of the National Insect Museum at

the National Agricultural Research Centre for identification. On the basis of morphological

characteristics and with the help of published literature (i.e., Mann, 1969; McFadyen, 1985;

Zimmermann et al., 2004; Habeck et al., 2016; and Folgarait et al., 2018), the field collected larvae

were confirmed to be the renowned cactus moth (C. cactorum). Adult C. cactorum look very

similar to several other native species of the subfamily Phycitinae, and identification of the adults

is often difficult (Heinrich, 1956). However, the best means of identification can be accomplished

with confirmation of the fully grown larvae.

Surveys for wild cactus in vicinities of study areas:

After confirmation that the cladodes of cultivated O. ficus-indica were infested by the

invasive moth, C. cactorum, plans were made to conduct detailed surveys for the whole district of

Chakwal. Surveys were conducted for all tehsils of district Chakwal to discover the level of

infestation of C. cactorum on wild Opuntia species which was introduced naturally/deliberated

(Figs. 8 & 9). Surveys were consecutively conducted from July 2019 to July 2021. During

consecutive surveys over two years, it was found that sporadic plantation of wild cactus throughout

the district Chakwal was in the stage of re-growing (Fig. 11) from the debris of the cactus, which

were damaged through the heavy attack of the larvae. However, larvae were found at various

localities in tehsil Talagang of district Chakwal on wild cactus. In all surveyed localities wild

cactus was severely infested by the cactus moth and threatened (Figs 7 & 10). During this study

establishment of C. cactorum has been confirmed after the introduction of the cactus moth larvae

in the district Chakwal, Pakistan during 1994.

Fig. 9. C. cactorum infested cladode of wild cactus in

District Chakwal.

14

i a=

Fig. 11. Regrowth of cladode from debris/damaged cactus in District Chakwal.

STATUS OF OPUNTIA CACTI IN INFESTED REGIONS

Cacti (Caryophyllales: Cactaceae) are a large and diverse group of succulent plants, Latin

American in origin. Historically, they are among the most popular horticultural plants having close

relationships with the local population, with different life forms such as arborescent, columnar,

globular, barreliform, articulated, cylindrical and flat cladode (Gibson & Nobel, 1986; Terrazas-

Salgado & Mauseth, 2002). Cacti have been used by native peoples as food for humans and

animals, medicines, cochineal dye (red pigments), and fencing (Vigueras & Portillo, 2001). Cacti

are a very famous component of America's deserts and inhabit a wide range of ecosystems such as

plains, deserts, sandy beaches, savannas, dry broad-leaved forests, high alpine-steppes, and

tropical rain forests. However, they mainly occur in warm and arid lands of North and South

America, excepting the high latitude boreal forest and tundra vegetation of North America and the

temperate rain forests of southern Chile (Nyffeler, 2002; Barthlott et al., 2015).

Economic importance of Cacti

Cacti have evolved physiological and morphological adaptations which allow them to grow

in habitats which are too dry for other plants (Mauseth, 2000). Despite the various challenges,

cacti are distinctive and unusual plants, which can adapt to extremely arid and hot environments

(Shetty et al., 2012). They are a well-known crop for cultivation in arid and semi-arid regions,

mostly for their expressive adaptive capability, through the prompt defense mechanisms against

abrupt changes in temperature, CO2 atmospheric concentration, soil water accessibility and

photosynthetic active radiation (Ferraz et al., 2017). Most of the cactus species have been

recognized for Crassulacean Acid Metabolism (CAM) due to the high-water use efficiency found

in fast growing species of the genus Opuntia such as O. ficus-indica, O. megacantha, and O.

amychlea, which are usually considered drought resistant species because they store considerable

15

amounts of water in their shoots (Nobel, 1994, 1995). The morphology and anatomy of the shoots

have evolved to serve this function.

Cacti are especially drought-tolerant plants (Casas et al., 1997). They are highly productive

even though they consume very little water and are extremely tolerant of dehydration when it

occurs (Sowell, 2001; El Obeidy, 2004). Throughout the world outside their native ranges many

species of columnar cacti have been widely introduced as a drought tolerant crop in arid and semi-

arid lands. For example, Australia has water limitations in many cities; hence, among drought-

tolerant plants, cacti are gaining popularity there (Shetty et al., 2012). Similarly columnar

cacti have been planted as drought-tolerant plants in United Arab Emirates (El Obeidy, 2004).

Cacti are very important to eliminate desertification; for example, cactus pears are able to grow

on land where no other crops are able to grow; they can be used to restore degraded land and, in

many countries, such as Ethiopia, it is the only crop that can be relied on when everything else

fails (Inglese et al., 2017). In Eritrea and Ethiopia, where it is cultivated on a small scale or

in natural stands; cactus pear is a needed source of food (fruits and its derivatives) for the rural

poor and fodder for their livestock (Mora, 2017). Furthermore, cacti are widely introduced for

ornamental purpose as well (Barthlott & Hunt, 1993; Mizrahi, 1999; FAO, 2001).

Prickly pear cacti preserve high moisture (88-91%), high protein and fiber contents in their

cladodes. Their nectar is an excellent source of dietary fiber, potassium, vitamin C & B, calcium,

magnesium, copper, taurine, flavonoids, polyphenols, and betalains (Pimienta, 1990; Fuentes-

Rodriguez, 1997; Kumar et al., 2018). The main components are carbohydrate-containing

polymers, which comprise a mix of mucilage and pectin (Gabriel et al., 2014). Moreover, prickly

pears are also comprised of crude protein (CP), crude fiber (CF), ether extract (EE), and nitrogen

free extract (NFE). Fiber fractions including neutral detergent fiber (NDF), acid detergent fiber

(ADB), and acid detergent lignin (ADL) have been reported (Van Soest et al., 1991). Prickly pear

possesses antioxidant, anti-lipidemic and antimicrobial properties (Gengatharan et al., 2015). The

prickly pear pulp contains 7.61% CP, 3.88% Ash, 85.75% carbohydrates, and 1.92% EE

(Rodriguez-Garcia et al., 2007; Atef et al., 2013; Hassan et al., 2019) and about 60% of the total

energy necessities of animals can be provided by prickly pear (L6pez-Garcia et al., 2001). Prickly

pear pulp (PPP) and prickly pear fruit (PPF) are rich in vitamins A and E and free from alkaloids

that are well-known anti-nutritional factors (Guevara-Figueroa et al., 2010; Yahia & Mondragon-

Jacobo, 2011; El-Mostafa et al., 2014). Antioxidant activities of different colored PPF are strongly

associated with total phenolics, betalains and ascorbic acid concentrations (Fernandez- Lopez et

al., 2010; Yahia and Mondragon-Jacobo, 2011).

Cactus pears are a very flexible food source; fresh cladodes/nopales of spineless varieties

are mostly used as vegetables by chopping young cladodes, while the flour of cladode, jams and

juices are used as processed food (Mora, 2017). The most widely used and economically important

species of Opuntia for food is O. ficus-indica (L.) Mill., known as Indian fig or forage palm

(Spodek et al., 2014; Ferraz et al., 2017). O. ficus-indica has been used traditionally in central

Mexico as a vegetable (nopalitos) and a fruit (tunas) (Maki et al., 2015). Many other edible species

16

are used in Mexico, including O. megacantha Salm-Dyck, O. stricta Haw., O. dillenii (Ker Gawl.)

Haw., O. schumannii Weber, O. robusta Wendl., and O. albicarpa Scheinvar (Marquez et al.,

2012; Arba et al., 2017). Mexico leads the world in prickly pear cactus food production (74%) and

is also the principal consumer. Demand for nopalitos is increasing in foreign markets such as in

the United States and Canada (Maki et al., 2015). Mexico grows nopalitos on 12,853 hectares

(SYBAP, 2020), with a food supply chain that supports many families across the country. In the

northeast region of Brazil, plantations totalling over 400,000 ha serves as a key element in

supporting livestock production in the country’s semi-arid regions. In the United States of

America, cacti are grown mainly as ornamentals in Arizona, California, Nevada, New Mexico,

and Texas (Perez- Sandi, 2001; Vigueras & Portillo, 2001).

Among prickly pear cacti, plants of Opuntia species are undoubtedly truly multipurpose

and their ability to adapt to different environments and has allowed these plants to be cultivated in

many continents such as Africa, North and South America, Asia, Australia and Europe

(Barbera, 1995). Cacti were introduced to the Indian subcontinent by the British as host plants

for cochineal scale insects (Dactylopius coccus) for the production of cochineal dye, but these

plantations steadily vanished due to pests and flooding of the areas (Singh, 2006).

Important cactus-pear producing countries are Italy, South Africa, Argentina, Chile,

Bolivia, Peru, Colombia, United States of America, Brazil, Malta, Mexico, Morocco, Algeria,

Libya, Yemen, Lebanon, Syria, Saudi Arabia, Tunisia, Egypt, Jordan, Pakistan, Israel, Greece,

Spain, Portugal, Turkey, France, Bulgaria, Albania, and Cyprus (Novoa et al., 2015; Mora, 2017;

Kumar et al., 2018). However, in a significant number of these countries, particularly in Africa,

cactus pear fruits are considered a by-product, for example the plant is mainly used for the

conservation of degraded soils (Mora, 2017). Other countries, including Morocco, Ethiopia, South

Africa, Kenya, India and Pakistan, are increasing their production and usage (Inglese et al., 2017).

The cultivation of cactus pear for use as fodder is also being adopted in sub-Saharan Africa and

South Asia. Furthermore, its medicinal properties and industrial uses are being researched and

promoted (Inglese et al., 2017). Prickly pear cacti are receiving attention throughout the world

(Hoffmann, 1995; Soberon et al., 2001; Makumbe, 2010; Louhaichi et al., 2018) because of their

huge potential with a wide range of applications; have the enormous potential to be a future food

(Shetty et al., 2012). Among cactus pears, Opuntia ficus-indica is gradually achieving the status

of recognized crop and cultivated on a commercial scale in many countries (Inglese et al., 2017;

Mondrag6n Jacobo & Méndez Gallegos, 2017).

Cacti as an invasive weed

In spite of economic gains, the cacti are among the most damaging alien invasive species

(AIS) by establishing a noteworthy tangible and imaginable vulnerability to conservation and

agricultural production (Cronk & Fuller, 1995; Bright, 1998), causing economic losses throughout

the world (Novoa et al., 2015). The tolerance of its vegetative growth, short root structure, and

the ability for CAM photosynthesis make it a harmful weed, which competes with many other

native species (Cronk & Fuller, 1995; FAO, 2001). Its management and elimination is problematic

17

due to the existence of large and small hairlike spines. Hence, before the introduction of cacti

species, appropriate knowledge of the diversity of introducing cactus species, management and its

potential utilization, especially in arid and semi-arid lands where they have become adopted are

vital. The deliberate introduction of prickly pear in Australia, South Africa, and Europe, attained

the status of weeds (CABI, 2016; Foxcroft et al., 2004; Greathead, 1971; Novoa et al., 2015; Novoa

et al., 2016; Shackleton et al., 2017).

CONCLUSIONS

The bio-control agents which are specialist consumers turn into active invaders, known as

escaped bio-control species such as Cactoblastis cactorum, which is an oligophagous species, 1.e.,

consumes plants of only the genus Opuntia (Schartel & Brooks, 2018). The areas presently

invaded by the cactus moth, have adequate levels of host plants of the genus Opuntia (Johnson &

Stiling, 1996; Schartel & Brooks, 2018). The successful establishment and spread of invasive

species depends on the presence of an appropriate host in the new environment, particularly those

with some level of host specificity such as C. cactorum (Schartel & Brooks, 2018). For instance,

the adverse effects on non-target species through the introduction of C. cactorum for biological

control of weeds were seriously apparent through the introduction of prickly pear in Australia in

1926 (Dodd, 1940), after the successful biological control of Opuntia weed. Later on, the larvae

of C. cactorum were introduced in many countries, including Pakistan to control the Opuntia weed.

The larvae of the cactus moth were released during 1994 in the district Chakwal, Pakistan, where

it was not well studied after its release and unconfirmed presence record(s) of the cactus moth from

Pakistan were reported by many authors (e. g. Julien & Griffiths 1998; Zimmermann et al., 2004;

Legaspi & Legaspi, 2010 & EPPO, 2021). On the other hand, 21 years after its introduction, the

first incidence of larvae of this moth was observed in cultivated spineless species of Opuntia ficus-

indica. Further surveys were conducted consecutively for wild Opuntia in the four tehsils of district

Chakwal until October 2021, and it was found that the whole district was severely infested with

the larvae.

The main concern is the fast rate of its dispersal in surveyed localities of the district

Chakwal, Punjab province of Pakistan. This rate of dispersal is significantly faster than the first

observations of C. cactorum in December 2016 in Chakwal. Further studies on C. cactorum

dispersal will provide some clues of its dispersal either by natural spread, anthropogenic activities,

or by any other means, particularly in the district Malakand, which is located between mountain

ranges, despite the fact that the mountain ranges may have prevented the cactus moth from

spreading even if suitable Opuntia host-species is in abundance (Hoffmann, 1989). However, it

has not spread naturally due to the large cultivation of O. ficus-indicain in its native range, 1.e.,

Argentina, Paraguay, Uruguay and southern Brazil (Arruda et al., 1999). Nevertheless, its larvae

are able to move short distances from one host plant to another, but these insignificant movements

are irrelevant in the situation of the overall spread of the moth. If suitable hosts are in abundance,

the adults rarely range far, but as food plants decrease in density the moths travel more widely

(Dodd, 1940; Pettey, 1948; Robertson, 1985). There is a record of individual females flying as

far as 24 km (15 miles) to oviposit (Dodd, 1940). In Australia, the cactus moth spread unaided,

18

from the release points, for about 16—24 km (10-15 miles) in dense O. stricta infestations in 2.5

years (Dodd, 1940).

The cactus moth has tolerated a wide climatic range in many countries including Pakistan.

In temperate latitudes of Australia and South Africa, where C. cactorum occurs, there are two or

three generations per year (Pettey, 1948; Robertson, 1985), whereas in the warm tropical climate

of the Florida and the Caribbean there might be more generations annually. Existing information

suggests that the cactus moth’s establishment and its devastating effects are likely across huge

areas throughout the world, with cactus plantations particularly in the southwest U.S. and Mexico.

Cacti plants of the genus Opuntia are commonly, increasingly grown as ‘wonder-plants’ in

various countries, i.e., North, South and East Africa, the Mediterranean countries, the Middle East,

India, China, and Pakistan. But, the cultivated prickly pears in these countries are vulnerable to

attack by C. cactorum (Zimmermann, 2000; Inglese et al., 2017). In the future there is a high risk

of economic, social, and environmental damages in the areas where Opuntia plants (cultivated,

ornamental and wild) maintain a valuable role in agriculture, and in areas sustaining unique

ecosystems and wildlife in Pakistan.

The establishment of a C. cactorum population in the district of Chakwal is alarming.

Consequently, the future prospects of large-scale cultivation of prickly pears cacti in Pakistan, for

fodder and other purposes will be seriously affected. Besides that, throughout Pakistan, countless

nurseries imported several expensive species of ornamental Opuntia, which are under threat by the

attack of cactus moth. Moreover, throughout the world several cacti spp. of the genus Opuntia,

commonly are grown as ‘wonder- plants’ in various countries, i.e., North, South and East Africa,

the Mediterranean countries, the Middle East, India, China including Pakistan. However, the

cultivated prickly pear in these countries is at present vulnerable to the attack of C. cactorum.

The potential impact of C. cactorum on economic and social environment, ecosystems and

biodiversity requires immediate action to monitor and prevent the spread of this major

environmental pest affecting both agriculture and the environment (Stohlgren et al., 2005).

Response options are limited as chemical pesticides are largely ineffective and potential biological

control organisms pose risks to native species. Although the impact was only partially assessed,

available data clearly indicate that social, environmental and economic costs of invasion are likely

to be high. An active response depends on the best available knowledge of history, biology,

identification, surveillance, and dispersal range, and control of the moth is inevitable.

Key findings of this initial study

1. Prevalence of Cactoblastis cactorum is established in district Chakwal, Punjab-Pakistan,

where it was introduced 22 years ago in 1994. Moreover, it prevails in the districts of

Malakand, Mardan, and Nowshera during 2019.

2. Cultivated spineless species of Opuntia (i.e., O ficus-indica (L.) Mill.) and naturalized wild

cactus experienced a severe attack of C. cactorum. Resultantly, both the cultivated and wild

19

Opuntia species have completely vanished in district Chakwal.

3. Highly significant economic, social, and environmental damage is foreseen if C. cactorum

spreads and establishes in other parts of Pakistan.

4. Large scale cultivation of Opuntia spp. in arid and semi-arid regions of Pakistan for fodder

and various other purposes will also be badly affected.

5. Immediate remedial action is inevitable to curb the further spread of C. cactorum.

6. In near future, C. cactorum can also pose a serious threat to the ornamental Opuntia in

several nurseries throughout Pakistan.

7. Robust surveillance programs are required for early detection of further spread of the

invasive moth on naturalized, cultivated and ornamental Opuntia species.

8. Integrated, effective response based on data exchange on an international level, situation

analysis, and rapid response programs is immediately required.

CONTRIBUTIONS

The following scientists conducted field surveys throughout District Chakwal for two years

(July 2019 to July 2021): Dr. Muhammad Ather Rafi Ex. Director; Dr. Ahmad Zia, Senior

Scientific Officer, National Insect Museum, National Agricultural Research Center, Islamabad,

Pakistan; Dr. Muhammad Islam; Dr. Falak Naz, Principal Scientific Officer; and Dr. Muhammad

Qasim, Assistant Professor. Dr. Muhammad Islam Ex. Director and Mr. Jalal Hayat Khan

Scientific officer, Rangeland Research Institute also helped in monitoring larvae of C. cactorum

in cultivated fields of Opuntia ficus-indica at three localities of District Chakwal. Mr. Haseeb

Kamran, Mr. Jalal Hayat Khan and Dr. Nazeer Ahmed, Assistant Professor helped in GIS mapping.

Dr. Muhammad Ather Rafi, Dr. Muhammad Ashfaq, Principal Scientific officer, Muhammad

Saeed, Associate Professor, Dr. Gul Naz Parveen, Associate Professor, Dr. Muhammad Tariq

Khan, Director Plant Protection and Dr. Riffat Sultana, Associate Professor helped in preparation

of the manuscript. Thanks to Crispin Guppy for final review of the manuscript. Following students

Miss Walija Fayaz, Mr. Wagar Ahmed and Mr. Qudrat Ullah keenly participated in surveys and

collection for their future research.

ACKNOWLEDGEMENTS

Highly recognized Dr. Muhammad Islam, Ex. Director, Rangeland Research Institute,

National Agricultural Research Center, Islamabad, Pakistan, for his interest and information about

C. cactorum outbreak in cultivated spineless Opuntia (.e., O. ficus-indica). Mr. Muhammad

Ashraf Laboratory Assistant, Muhammad Tahair and Majid Bilal, Field Assistant acknowledged,

National Insect Museum, National Agricultural Research Center, Islamabad, Pakistan, for the

collection of infested cladodes of the cacti and dissection of the cladodes of wild spiny prickly

pear at survey localities of the district Chakwal. Mr. Muhammad Ashraf also raised C. cactorum

larvae in the laboratory of the National Insect Museum. Special thanks to Mr. Muhammad Israr to

provide transport during surveys.

LITERATURE REFERENCES

Aguilar-Yanez, M., Hernandez-Mendo, O., Guerrero-Legarreta, I., Ramirez-Bribiesca, J.,

Aranda-Osorio, G. & Crosby-Galvan, M. 2011. Productive response of lambs fed with

fresh or dehydrated spineless cactus (Opuntia ficus-indica L.). J Prof Assoc Cactus, 13:

23-35.

20

https://www.researchgate.net/profile/Gilberto-Aranda-

Osorio/publication/271965788 Productive _response_of lambs fed with fresh or dehy

drated_spineless cactus Opuntia _ficus-

indica_L/links/566f305508ae486986b701ab/Productive-response-of-lambs-fed-with-

fresh-or-dehydrated-spineless-cactus-Opuntia-ficus-indica-L.pdf

Anderson, A. J. O. 1981. Florentine codex: general history of the things of New Spain. Book 11-

earthly things. Salt Lake City, UT, USA: University of Utah Press.

Andraca-Gomez, G., Ordano, M., Boege, K., Dominguez, C. A., Pinero, D., Pérez-Ishiwara,

R., Pérez-Camacho, J., Canizares, M. & Fornoni, J. 2015. A potential invasion route of

Cactoblastis cactorum within the Caribbean region matches historical hurricane

trajectories. Biological Invasions,17: 1397-1406.

Arba, M., Falisse, A., Choukr, R. & Sindic, M. 2017. Biology, flowering and fruiting of the

cactus Opuntia spp.: A review and some observations on three varieties in Morocco. Braz.

Arch. Biol. Technol., 60: 1-11.

https://www.scielo.br/j/babt/a/hR Y X4rKvOgChy Ghdhj8bbMK/?lang=en&format=html

Aregawi, T., Melaku, S. & Nigatu, L. 2008. Management and utilization of browse species as

livestock feed in semi-arid district of North Ethiopia. Livestock Res Rural Dev. 20(6): 86.

Arruda, G. P., de Arruda, F. G. P. & Warumby, J. F. 1999. Introducci6n y utilizacion de las

cactaceas Nopalea cochenillifera (L.) y Opuntia ficus-indica en Brasil.Proceedings of the

VIII Congreso Nacional y VI Internacional sobre Conocimiento y Aprovechamiento de

Nopal (ed. by J.R.A. Rivera and J.A.R. Agiiero), pp. 101-102. University of San Luis

Potosi, Mexico.

Atef, A. M., Abou-Zaid, N., Ibrahim, I., Mostafa, T. R. & Nadir, A. 2013. Quality evaluation

of sheets, jam and juice from prickly pear and melon blends. Life Sci. J., 10(2): 200-208.

https://doi.org/10.7537/marslsj100213.31

Atti, N., Mahouachi, M., Zouaghi, F. & Rouissi, H. 2009. Incorporation of cactus (Opuntia

ficus-indica f. inermis) in young goat’s diets: 1. Effects on intake, digestion, growth and

carcass composition. Livestock Res. for Rural Devel. 21(12):1-13.

http://www. lrrd.org/Irrd21/12/atti21217.htm

Barbera, G. 1995. History, economic and agro-ecological importance, pp. 1-11 Jn G. Barbera, P.

Inglese, and E. Pimienta-Barrios (eds.). Agro-ecology, cultivation and uses of cactus pear.

FAO Plant Prod. & Protec. Paper No. 132. Rome, Italy.

Barthlott, W. & Hunt D. R. 1993. Cactaceae: In: Kubitzki, K. (Ed.): The families and genera of

flowering plants, Berlin, Heidelberg, New York: Springer. 2: 161-197.

Barthlott, W. K., Burstedde, J. L., Geffert, P. L., Ibisch, P. L., IbischKorotkova, N, Miebach,

A., Rafiqpoor, M. D., Stein, A. & Mutke, J. 2015. Biogeography and biodiversity of

cacti. Schumannia: 205 pp.

Bello-Rivera, A., Pereira, R., Enkerlin, W., Bloem, S., Bloem, K., Hight, S. D., Carpenter, J.,

Zimmermann, E., Sanchez-Anguiano, H. G., Zetina-Rodriguez, R. & Trujillo-

Arriaga, F. J. 2021. Successful area-wide programme that eradicated outbreaks of the

invasive cactus moth in Mexico, pp. 561-580, Jn: J. Hendrichs, R. Pereira and M. J. B.

Vreysen (eds.), Area-wide integrated pest management. Development and field

application. CRC Press, Boca Raton, FL, USA.

Ben Salem, H. and Abidi, S. 2009. Recent advances on the potential use of Opuntia spp. in

livestock feeding. Acta Hortic 811: 317-326.

https://doi.org/10.17660/ActaHortic.2009.811.43.

21

Blackburn, T. M., Essl, F., Evans, T., Hulme, P. E., Jeschke, J. M., Kithn, I., Kumschick, S.,

Markova, Z., Mrugala, A., Nentwig, W., Perel, J., PySek, P., & Rabitsch, W. 2014. A

unified classification of alien species based on the magnitude of their environmental

impacts. {PLoS} biology, 12(5):1—11. https://do1.org/10.1371/journal.pbio. 1001850.

Bloem, S., Mizel, R. F. HI, Bloem, K. A., Hight, S. D. & Carpenter, J. E. 2005. Laboratory

evaluation of insecticides for control of the invasive Cactoblastis cactorum (Lepidoptera:

Pyralidae). Fla. Entomol. 88(4): 395—400.

https://journals.flvc.org/flaent/article/view/75462/73120

Briano, J., Varone. L., Logarzo, G., & Villa, C. 2012. Extended geographical distribution and

host range of the cactus moth, Cactoblastis cactorum (Lepidoptera: Pyralidae), in

Argentina. Fla. Entomol. 95(1): 233-237.

https://journals.flvc.org/flaent/article/view/78889/76270

Bright, C. 1998. Life out of bounds. The World Watch Environmental Alert Series, (New York),

NY: Norton, 182: 89-92.

Bryan, R. T. 1996. Alien species and emerging infectious diseases: past lessons and future

applications. Jn: Sandlund, G. T., Schel, P. J., Viken, A. (Eds.), Proceedings of the

Norway/UN Conference on Alien Species, 1-5 July. Norwegian Institute for Nature

Research, Trondheim, Norway, pp. 74-80.

Burke, M. B., Lobell, D. B. & Guarino, L. 2009. Shifts in African crop climates by 2050, and

the implications for crop improvement and genetic resources conservation. Glob. Environ.

Change: 317-325.

https://assets.publishing.service.gov.uk/media/57a08b57e5274a27b2000abd/burke_et_al

GEC_2009.pdf

CABLI. 2022. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

https://www.cabi.org/isc/datasheet/10680#todistribution (accessed on January 21, 2022).

CABI. 2016. Opuntia stricta. In: Invasive species compendium. CAB International, Wallingford.

www.cabi.org/isc (accessed on September 20, 2016).

Canamares, M. V., Garcia Ramos, J. V., Domingo, C. & Sanchez Cortes, S. 2006. Surface-

enhanced Raman scattering study of the anthraquinone red pigment carminic acid. Vib.

Spectrosc., 40: 161- 167.

Casas, A., Pickersgill, B., Caballero J. & Valient-Banuet, A. 1997. Ethnobotany and

domestication in Xoconochtli, Stenocereus stellatus (Cactaceae), in the Tehuacan Valley

and La Mixteca Baja, Mexico. Economic Botany, 51(3): 279-292.

Chavez-Moreno, C. K., Tecante, A., & Casas, A. 2009. The Opuntia (Cactaceae) and

Dactylopius (Hemiptera: Dactylopiidae) in Mexico: a historical perspective of use,

interaction and distribution. Biodiversity and Conservation, 18: 3337-3355.

Chavez-Ramirez F., Wang, X. G., Jones, K., Hewitt, D., & Felker, P. 1997. Ecological

characterization of Opuntia clones in South Texas: Implications for wildlife herbivory and

frugivory. Journal of the Professional Association for Cactus Development, 2: 9-19.

Cronk, Q. C. B. & J. C. Fuller. 1995. Plant invasions: the threat to natural ecosystems. Chapman

and Hall, London. 241 pp.

Dapson, R.W. 2007. The history, chemistry and modes of action of carmine and related

dyes. Biotechnic and Histochemistry, 82:4, 173- 187, doi: 10.1080/10520290701704188.

De Felice, M.S. 2004. Prickly pear cactus, Oputia spp. A spine-tingling tale. Weed

Technol., 18: 869-877.

de Humboldt, A. 1811. Political essay on the kingdom of New Spain, Vol. 1. London, England.

289pp.

22

https://archive.org/details/b29350451_ 0001/page/n10/mode/1lup?ref=ol&view=theater

De Lotto, G. 1974. On the status and identity of the cochineal insects (Homoptera: Coccoidea:

Dactylopiidae). Journal of the Entomological Society of Southern Africa, 37: 167-193.

Devendra, C. 1998. Improvement of small ruminant production systems in rainfed agro-

ecological zones of Asia. Ann. Arid Zone 37(3): 215-232.

Devendra, C. 1999. Goats: challenges for increased productivity and improved livelihoods.

Outlook Agric. 28(4): 215-226. https://doi.org/10.1177/003072709902800404

Dickel, T.S. 1991. Cactoblastis cactorum in Florida (Lepidoptera: Pyralidae: Phycitinae).

Trop. Lepid. 2(2): 117-118. 86685 (archive.org)

Dodd, A. P. 1940. The biological campaign against prickly pear. Commonwealth Prickly Pear

Board, Brisbane, Australia: 1-177 pp.

Drake, J. A., Mooney, H. A., di-Castri, F., Groves, R., Kruge, F., Rejmanek, M. & Ensernik,

M. 1999, Predicting invasions: biological invaders sweep in. Science, 285: 1834-1836.

http://www.columbia.edu/itc/cerc/danoffburg/invasion_bio/invbio_readings.html.

El Moselhy, M. M., Sengupta, A. K. & Smith, R. 2011. Carminic acid modifid anion exchanger

for the removal and preconcentration of Mo(VI) from wastewater. J. Hazard.

Mater., 185: 442-446.

El-Mostafa, K., El Kharrassi, Y., Badreddine, A., Andreoletti, P., Vamecq, J., El Kebbaj, M.

S., Latruffe, N., Lizard, G., Nasser, B. & Cherkaoui Malki, M. 2014. Nopal Cactus

(Opuntia ficus-indica) as a Source of Bioactive Compounds for Nutrition, Health and

Disease. Molecules, 19: 14879-14901 https://www.mdpi.com/1420-3049/19/9/14879

El Obeidy, A. A. 2004. Introducing and growing some fruiting columnar cacti in a new arid

environment. J. Fruit Omam, Plant Res., 12: 127-136.

http://www.inhort.pl/files/journal_pdf/journal_2004/full2004-16.pdf

Elton, C. S. 1958. The ecology of invasions by animals and plants. London: Methuen. Jn:

Classics in physical geography revisited. Prog. In Phys. Geog. 31(6) (2007): 659-666.

EPPO. 2021. EPPO Global database. Jn: EPPO Global database. Paris, France.

http://gd.eppo.int/

FAO. 2001. Cactus (Opuntia spp.) as forage. FAO Plant production and Protection Paper 169.

FAO International Technical Cooperation Network on Cactus Pear. Food and Agriculture

organization of the United Nations, Rome. www.fao.org

Fernandez-Lopez, J. A., Almela, L., Obon, J. M. & Castellar, R. 2010. Determination of

antioxidant constituents in Cactus Pear fruits. Plant Food Hum. Nutr., 65(3): 253-259.

https://www.researchgate.net/profile/Luis-Almela-

2/publication/46111496 Determination of Antioxidant Constituents in Cactus Pear F

ruits/links/544684b50cf2fl 4fb80f4dd2/Determination-of-Antioxidant-Constituents-in-

Cactus-Pear-Fruits.pdf

Ferraz, R. L. S., Costa, P. S., Neto, J. D., Anjos, F. A., Barreto, N. P., Soares, L. S.,

Magalhaes, I. D & Barbosa, M. A. 2017. Opuntia ficus-indica (L.) Mill. (Cactaceae) in

Climate Change Scenarios and Its Potential for Wastewater. Bioremediation in:

Semi-Arid Regions: A Systematic Review and Meta-Analysis. Journal of Experimental

Agriculture International 18(3): 1-11

Folgarait, P. J., Montenegro, G. A., Plowes, R. M. & Gilbert, L. 2018. A study of Cactoblastis

cactorum (Lepidoptera: Pyralidae) in its native range: further insights into life cycle, larval

identification, developmental parameters, natural enemies, and damage to the host plant

Opuntia ficus-indica (Caryophyllales: Cactaceae). Florida Ent. 101(4): 559-572.

https://doi.org/10.1653/024.101.0416

23

Foxcroft, L. C, Rouget, M., Richardson, D. M., & MacFayden, S. 2004. Reconstructing 50

years of Opuntia stricta invasion in the Kruger National Park, South Africa: environmental

determinants and propagule pressure. Divers. and Distrib. 10(5-6): 427-437.

https://onlinelibrary.wiley.com/doi/full/10.1111/j.1366-9516.2004.00117.x

Fullaway, D. T. 1954. Biological control of cactus in Hawaii. Journal of Economic Entomology,

47: 696-700.

Gabriel, A., Victor, N. & James Du Preez, C. 2014. Cactus pear biomass, a potential

lignocellulose raw material for Single Cell Protein production (SCP): A Review. Int. J.

Curr. Microbiol. App. Sci., 3(7): 171-197.

https://www.researchgate.net/profile/Gabriel-Akanni-

2/publication/344775248 Cactus pear biomass a potential lignocellulose raw materia

| for Single Cell Protein production SCP A Review/links/S5f8flal4a6fdccfd7b71cf53

/Cactus-pear-biomass-a-potential-lignocellulose-raw-material-for-Single-Cell-Protein-

production-SCP-A-Review.pdf

Garcia-Turudi, J. C., Martorell, L. F.,. Medina Guad, S. 1971. Geographical distribution and

host plant list of the cactus moth, Cactoblastis cactorum (Berg), in Puerto Rico and the

United States Virgin Islands. J. Agric. Univ. Puerto Rico, 55: 130-134.

Gengatharan, A., Dykes, G. A. & Choo, W. S. 2015. Betalains: Natural plant pigments with

potential application in functional foods. LWT-Food Sci. Technol., 64(2): 645-649.

Gibson, A. C. & Nobel, P. S. 1986. The cactus primer. Harvard University Press, Cambridge: 85

Pp.

Gonzalez, E. A., Garcia, E.M. & Nazareno, M.A. 2009. Free radical scavening capacity

and antioxidant activity of cochineal (Dactylopius coccus C.) extracts. Food Chem.,

119: 358-362.

Greathead, D. J. 1971. A review of biological control in the Ethiopian region. Tech. Comm. No.

5, Commonwealth Inst. of Biol. Contr., Commonwealth Agric. Bur., Slough: 1-162.

Guevara-Figueroa, T.; Nez-Islas, H. J.; Reyes-Escogido, M. L.; Mortensen, A. G.; Bente, B.;

Laursen, B. B.; Lin, L.; De Leo ‘n-Rodri guez, A.; Fomsgaard, I. S. & De La Rosa, A.

B. 2010. Proximate composition, phenolic acids, and flavonoids characterization of

commercial and wild nopal (Opuntia spp.). J. Food Compos. Anal., 23: 525-532.

Guretzky, J. A. & Louda, S. M. 1997. Evidence for natural biological control: insects decrease

survival and growth of a native thistle. Ecological Applications, 7: 1330-1340.

https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1007&context=bioscilouda

Gusha, J., Halimani, T., Katsande, S. & Zvinorova, P. 2015. The effect of Opuntia ficus indica

and forage legumes based diets on goat productivity in smallholder sector in Zimbabwe.

Small Ruminant Res. 125: 21-25.

Habeck, D. H., Bennett, F. D. & Miller, C. 2016. Cactus Moth, Cactoblastis cactorum (Berg)

(Insecta: Lepidoptera: Pyralidae). Department of Entomology and Nematology, UF/IFAS

Extension, EENY-056. http://edis.ifas.ufledu

Hassan, F. A., Mobaraz,S. M., Basyony, M. M., Mahrose, K. M & El-Medany, S. A. 2019.

Effect of using prickly pear and its by-products as alternative feed resources on

performance of growing rabbit. Egyptian Journal of Rabbit Science, 29(1): 99-124.

https://ejrs.journals.ekb.eg/article_45677_2259f8647ec9b79393c8975f04bafab5.pdf

Heath, R. R., Teal, P. E. A., Epsky, N. D., Dueben, B. D., Hight, S. D., Bloem, S., Carpenter,

J. E., Weissling, T. J., Kendra, P. E., Cibrian-Tovar, J. & Bloem, K. A. 2006.

Pheromone-based attractant for males of Cactobalstis cactorum (Lepidoptera: Pyralidae).

Envir. Entom., 35(6): 1469-1476. https://academic.oup.com/ee/article/35/6/1469/387276

24

Heinrich, C. 1939. The cactus-feeding Phycitinae: a contribution toward a revision of the

American pyralidoid moths of the family Phycitidae. Proc. U.S. National Museum, 86: 331-

413, pls. 23-51. https://repository.si1.edu/bitstream/handle/10088/16296/1/USNMP-

86 3053 _1939.pdf

Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. National Museum Bulletin

No. 207: 581 pp.

Hight, S. D. & Carpenter, J. E. 2009. Flight phenology of male Cactoblastis cactorum

(Lepidoptera: Pyralidae) at different latitudes in the Southeastern United States. Fla.

Entomol., 92(2): 208-216. https://doi.org/10.1653/024.092.0203

Hight, S. D., Carpenter, J. E., Bloem, S. & Bloem, K. A. 2005. Developing a Sterile Insect

Release Program for Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae): Effective

over flooding ratios and release-recapture field studies. Environ. Entomol., 34(4): 850-856

https://academic.oup.com/ee/article/34/4/850/785338

Hoffmann, J. H. & Zimmermann, H. G. 1989. Ovipositional and feeding habits in cactophagous

pyralids: prediction for biological control of cactus weeds. Proceedings of the VII

Symposium on Biological Control of Weeds (ed. by E.S. Delfosse), MAF, Rome: 395-—

399.

Hoffmann, J. H., Moran V. C. & Zeller D. A. 1998. Evaluation of Cactoblastis cactorum

(Lepidoptera: Phycitidae) as a biological control agent of Opuntia stricta (Cactaceae) in

the Kruger National Park, South Africa. Biological Control, 12: 20-24.

Hoffmann, W. 1995. Ethnobotany, pp. 12-19. In: G. Barbera, P. Inglese, & E. Pimienta-Barrios

(eds.). Agro-ecology, cultivation and uses of cactus pear. FAO Plant Produc. and Protec.

Paper No.132. Rome, Italy.

Howarth, F. G. 1991. Environmental impacts of classical biological control. Annual Review of

Entomology, 36: 485-509. http://www.ask-force.org/web/Ecology/Howarth-

Environmental-Impacts-Classical-1991.pdf

Hudson, L. N., Newbold, T., Contu, S. and Purvis, A. 2014. The PREDICTS database: a global

database of how local terrestrial biodiversity responds to human impacts. Ecol. Evol. 4(24):

4701- 4735. https://onlinelibrary.wiley.com/doi/full/10.1002/ece3.1303

Hunter, K. 2009. Report on Agriculture Research Needs of Invasive Species. National

Agricultural Research, Extension, Education and Economics Advisory Board. U.S. Dept.

of Agric. 4 pp.

International Atomic Energy Agency (IAEA) 2002. Mitigating the Threat of Cactoblastis

cactorum to International Agriculture and Ecological Systems and Biodiversity. Report

and Recommendations of a Consultants’ Group Meeting organized by the Technical Co-

operation Department of the IAEA and the Joint FAO/IAEA Division of Nuclear

Applications in Food and Agriculture, Vienna, Austria, July 2002.

Inglese, P., Mondragon, C. & Nefzaoui, A. 2017. Crop ecology, cultivation and uses of cactus

pear. The Food and Agric. Org. of the United Nations (FAO) and the Intern. Center for

Agric. Res. in the Dry Areas (ICARDA) Rome. 225 pp.

Invasive Species Specialist Group (ISSG), 2011. Global Invasive Species Database (GISD). In:

Global Invasive Species Database (GISD), Auckland, New Zealand: University of

Auckland. http://www.issg.org/database

Islam, M., Razzaq, A., Hassan, S., Qamar, I. A., Rischkowsky, B., Ibrahim, M. N. M. &

Louhaichi, M. 2019. Assessment of different supplemental feeding strategies including

cactus (Opuntia ficus-indica) to enhance sheep productivity in Chakwal, Pakistan. In: C.

Saenz et al., (eds). CAM Crops for a Hotter and Drier World. Acta Hortic. 1247: 117-122.

25

Jeschke, J. M., Bacher, S., Blackburn, T. M., Dick, J. T. A., Essl, F., Evans, T., Gaertner, M.,

Hulme, P. E., Kiihn, I., Mrugal, A., Pergl, J., PySek, P., Rabitsch, W., Ricciardi, A.,

David, M., Richardson, D. M., Sendek, A., Vila, M., Winter, M. & Kumschick. S.

2014. Defining the impact of Non-Native species. Cons. Biol., 28(5): 1188-1194.

https://doi.org/10.1111/cobi.12299

Jezorek, H., Baker, A. J. & Stiling, P. 2012. Effects of Cactoblastis cactorum on the survival

and growth of North American Opuntia. Biol. Invasions, 14: 2355-2367.

Johnson, D. M. and Stiling, P. D. 1996. Host specificity of Cactoblastis cactorum (Lepidoptera:

Pyralidae), an exotic Opuntia-feeding moth, in Florida. Environ. Entom. 25(4): 743-748.

https://doi.org/10.1093/ee/25.4.743

Julien, M. H. & Griffiths, M. W. 1998. Biological control of weeds. A world catalogue of agents

and their target weeds, 4" Edition. CABI Publishing, Wallingford: 1-233.

Klein, A. M., Vaissiere, B. E., Cane, J. H., Steffan-Dewenter, I., Cunnigham, S. A., Kremen,

C. & Tscharntke, T. 2007. Importance of pollinators in changing landscapes for world

crops. Proceedings of the Royal Society B, London: Biological Sciences, 274: 303-313.

https://royalsocietypublishing.org/doi/10.1098/rspb.2006.3721

Kumar, K., Singh, D. & Singh, R. S. 2018. Cactus Pear: Cultivation and Uses. ICAR-Central

Institute for Arid Horticulture, Bikaner, Rajasthan, India. Tech. Bull. No. 73: 38 pp.

https://www.researchgate.net/profile/Kamlesh-Kumar-

9/publication/330104598 Cactus Pear Cultivation _and_uses/links/Sc2dbca4a6fdccfc707

922f2/Cactus-Pear-Cultivation-and-uses.pdf

Legaspi, Jr., B.C. & Legaspi, J. C. 2010. Field-Level validation of a CLIMEX Model for

Cactoblastis cactorum (Lepidoptera: Pyralidae) using estimated larval growth rates. Quant.

Ecol.: 395-377. https://academic.oup.com/ee/article/39/2/368/478699

Lopez Collado, C. J., Malpica Vazquez, A., Lopez Collado, J., Garcia Pérez, E. & Sol

Sanchez, A. 2013. Crecimiento de Opuntia ficus-indica (L.) Mill. en la zona central de

Veracruz. Rev. Mex. Cienc. Agric., 4(5): 1005-1014.

https://www.scielo.org.mx/scielo.php?pid=S2007-09342013000900012&script=sci_arttext

Lopez-Garcia, J. J., Fuentes-Rodriguez, J. M. & Rodriguez, R. A. 2001. Production and use

of Opuntia as forage in Northern Mexico. Jn: Cactus (Opuntia spp.) as forage. Edited by

Mondragoén-Jacobo, C. and Perez Gonzalez, S. FAO Plant Production and Protection

Paper 169: 29-36.

https://www.google.com/books/edition/Cactus_Opuntia_Spp_as_Forage/e_ bloAXLdKgC?

hl=en& gbpv=1&dg=Cactus+(Opuntia+spp.)+as+forage.&printsec=frontcover

Louda, S. M., Kendall, D., Connor, J. & Simberloff, D. 1997. Ecological effects of an insect

mairouueed for tlie biological control of weeds. Science, 277: 1088-1090.

https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1000&context=biosciecology

Louhaichi, M., Kumar, S., Tiwari, S., Islam, M., Hassan, S., Yadav, O. P., Dayal, D., Moyo,

H. P., Dev, R. & Sarker, A. 2018. Adoption and Utilization of Cactus Pear in South Asia-

Smallholder Farmers’ Perceptions. Sustainability 10(10), 3625: 1-15.

doi:10.3390/su10103625

Lowe, S., Browne, M., Boudjelas, S., De Poorter, M. 2000. 100 of the Worlds Worst Invasive

Alien Species. A Selection from the Global Invasive Species Database; Invasive Species

Specialist Group: Auckland, New Zealand: 12pp.

Mack, M. C. & D’ Antonio, C. M. 1998. Impacts of biological invasions on disturbance regimes.

Trends in Ecol. Evol., 13(5): 195-198. https://doi.org/10.1016/S0169-5347(97)01286-X

26

Maki, G., Pena, C. B., Garcia, R., Arévalo, M. L., Calderon, G. & Anaya, S. 2015. Physical

and chemical characteristics of nopal vegetable (Opuntia ficus-indica) for export and

national consumption. Agrociencia, 49(1): 31-51.

https://www.scielo.org.mx/scielo.php?pid=S 1405-

31952015000100003 &script=sci_arttext

Makumbe, M. T. 2010. A review of the distribution, use and potential of cactus pear (Opuntia

ficus- indica (I.) mill.) as ruminant feed in Zimbabwe. Jn Improved Utilization of Cactus

Pear for Food, Feed, Soil and Water Conservation and Other Products in Africa:

Proceedings of International Workshop Held in Mekelle, Ethiopia, 19-21 October 2009.

Tigray Agricultural Research Institute: Mekelle, Ethiopia: 14-19.

Mann, J. 1969. Cactus-Feeding Insects and Mites. Smithsonian Institution Press, Washington

DC. USNM Bull. 256: 158 pp. https://doi.org/10.5479/si1.03629236.256.1

Marquez, S. R., Torcuato, C., Almaguer, G., Colinas, M. T. & Gardez, A. K. 2012. The

productive system of prickly pear cactus (Opuntia albicarpa and O. megacantha) in

Axapusco, State of Mexico. Problematic and alternatives. Rev. Chapingo Ser. Hortic.

2012, 18: 81-93.

Masters, G., & Norgrove, L. 2010. Climate change and invasive alien species. CABI Working

Paper 1: 30 pp. https://www.cabi.org/cabi-publications/climate-change-and-invasive-

alien-species/

Mauseth, J. D. 2000. Theoretical aspects of surface-to-volume ratios and water-storage

capacities of succulent — shoots. Am. J. Botany, 88: 1107-1115.

https://bsapubs.onlinelibrary.wiley.com/doi/epdt/10.2307/2656647

McDermott, J. J., Staal, S. J.. Freeman, H. A., Herrero, M., & Van de Steeg, J. A. 2010.

Sustaining intensification of smallholder livestock systems in the tropics. Livest. Sci. 130

(1-3): 95-109 https://doi.org/10.1016/j.livsci.2010.02.014

McFadyen, R. E. 1985. Larval characteristics of Cactoblastis spp. (Lepidoptera: Pyralidae) and

the selection of species for biological control of prickly pears (Opuntia spp.). Bull. of

Entomol. Res., 75: 159-168.

Mengoni, C., Varone, L., Logarzo, G. A., Guala, M., Rodriguero, M. S., Hight, S. D. &

Carpenter, J. E. 2014. Geographical range and laboratory studies on Apanteles

opuntiarum (Hymenoptera: Braconidae) in Argentina, a candidate for biological control of

Cactoblastis cactorum (Lepidoptera: Pyralidae) in North America. Fla. Entomol., 97:

1458-1468. https://doi.org/10.1653/024.097.0421

Miller, M. & Aplet, G. 1993. Biological control: a little knowledge is a dangerous thing. Rutgers

Law Review 45: 285-334.

Misra, A., Mishra, A., Tripathi, M., Chaturvedi, O., Vaithiyanathan, S., Prasad, R. &

Jakhmola, R. 2006. Intake, digestion and microbial protein synthesis in sheep on hay

supplemented with prickly pear cactus (Opuntia ficus-indica (L.) Mill., with or without

groundnut meal. Small Ruminant Res. 63(1):125-134.

Mitchell, D. S., and Gopal, B. 1991. Invasion of tropical freshwaters by alien aquatic plants. In:

Ramakrishnan, P.S. (editor), Ecology of Biological Invasion in Tropics. International

Scientific Publications, India: 139-154

Mizrahi, M. 1999. Beating the Odds: New Exotic Desert-Grown Fruits. Examines a koubo

orchard in Israel's inhospitable Western Negev Desert.

Mondragoén Jacobo, C. & Méndez Gallegos, S. J. 2017. Nopalitos or vegetable cactus,

production and utilization. Pages 93-103 In: Inglese, P., Mondragon, C. & Nefzaoui, A.

(eds.) 2017. Crop ecology, cultivation and uses of cactus pear. The Food and Agric. Org.

of the United Nations (FAQ) and the Intern. Center for Agric. Res. in the Dry Areas

27

(ICARDA), Rome: 225 pp

Mooney, H. A, & Drake, J. A. 1986. Ecology of Biological Invasion of North America and

Hawaii. Springer-Verlag. New York. Ecological Studies 58: 321 pp.

Mooney, H. A. & Cleland, E. E. 2001. The evolutionary impact of invasive species. PNAS

98(10): 5446-5451. www.pnas.org/cgi/doi/10.1073/pnas.09 1093398.

Mooney, H. A. & Hobbs, R. J. (eds.). 2000. Invasive Species in a Changing World. Island Press,

Washington, D.C.: 384 pp.

Mora, M. 2017. Marketing and communication constraints and strategies. Pages 195-202, In:

eds: Inglese, P., Mondragon, C. & Nefzaoui, A. (eds.). Crop ecology, cultivation and uses

of cactus pear. The Food and Agric. Org. of the United Nations (FAQ) and the Intern.

Center for Agric. Res. in the Dry Areas (ICARDA) Rome. 225 pp

Nefzaoui, A., Louhaichi, M. & Ben Salem, H. 2014. Cactus as a tool to mitigate drought and to

combat desertification. J. Arid Land Stud., 24: 121-124.

Nobel, P.S. 1994. Remarkable agaves and cacti. Oxford Univ. Press, New York: 180 pp.

Nobel, P. S. 1995. Environmental biology. Pages 36-48 Jn: G. Barbera, Inglese, P. & Pimenta

Barrious, E. (eds.), Agro-ecology, cultivation and uses of cactus pear. FAO Plant

Production and Protection Paper No. 132. Rome: 216 pp.

Novoa, A., Kaplan, H., Wilson, J.R. & Richardson, D. M. 2016. Resolving a prickly situation:

involving stakeholders in invasive cactus management in South Africa. Environ Manag.

57: 998-1008.

Novoa, A., Le Roux, J. J., Robertson, M. P., Wilson J. R. U. & Richardson, D. M. 2015.

Introduced and invasive cactus species: a global review. AoB Plants, 7: plu0O78: 14

pp. https://doi.org/10.1093/aobpla/plu078

Nyffeler, R. 2002. Phylogenetic relationships in the cactus family (Cactaceae) based on evidence

from trnK/matK and trnL-trnF sequences. Amer. Jour. Bot., 89: 312-326.

https://bsapubs.onlinelibrary.wiley.com/doi/full/10.3732/ajb.89.2.3 12

Olfert, O., Haye, T., Weiss, R., Kriticos, D. & Kuhlmann, U. 2016. Modelling the potential

impact of climate change on future spatial and temporal patterns of biological control

agents: Peristenus digoneutis (Hymenoptera: Braconidae) as a case study. Can.

Entomol.,148: 579-594.

Owen, E., Smith, T., Steele, M., Anderson, S., Duncan, A. & Herrero, M. 2004. Responding

to the livestock revolution: the role of globalisation and implications for poverty

alleviation. British Society of Animal Science, Pub. 33. Nottingham University Press,

Nottingham. 361 pp.

Paraiso, O., Hight, S. D., Kairo, M. T. K. & Bloem, S. 2011. Egg parasitoids attacking

Cactoblastis cactorum (Lepidoptera: Pyralidae) in North Florida. Florida Entomologist,

94(1): 81-90. View of Egg Parasitoids Attacking <I>Cactoblastis cactorum</I>

(Lepidoptera: Pyralidae) in North Florida (flvc.org)

Parker, M. H., Simberloff, I. M., Lonsdale, D., Goodell, W. M., Wonham, K., Kareiva, M.,

Williamson, P. M., von Holle, B., Moyle, P. B., Byers, J. E. & Goldwasser, L. 1999.

Impact: toward a framework for understanding the ecological effects of invaders.

Biological Invasions, 1: 3-19.

Pashte, V. V. & Said, P. P. 2015. Honey bees: Beneficial Robbers. Inter. J. Agr. Sci.and Res.

(IJASR), 5(5): 343-352.

Pemberton, R. W. & Cordo, H. A. 2001a. Potential and risks of biological control of

Cactoblastis cactorum (Lepidoptera: Pyralidae) in North America. Fla. Entomol.,

84(4): 513-526. https://journals.flvc.org/flaent/issue/view/3566

28

Pemberton, R. W., Cordo, H. A., 2001b. Nosema (Microsporida: Nosematidae) species as

potential biological control agents of Cactoblastis cactorum (Lepidoptera: Pyralidae):

surveys for the Microsporidia in Argentina and South Africa. Florida Entomologist, 84(4):

527-530. https://journals.flvc.org/flaentarticle/view/74999/72657

Pérez-De la O, N. B., Espinosa-Zaragoza, S., L6pez-Martinez, V., Hight, S. D. & Varone, L.

2020. Ecological Niche Modeling to Calculate Ideal Sites to Introduce a Natural Enemy:

The case of Apanteles opuntiarum (Hymenoptera: Braconidae) to Control Cactoblastis

cactorum (Lepidoptera: Pyralidae) in North America. Insects, 11(7): 454.

doi:10.3390/insects 11070454

Perez-Guerra, G. & Kosztarab, M. 1992. Biosystematics of the family Dactylopiidae

(Homoptera: Coccinea) with emphasis on the life cycle of Dactylopius coccus Costa.

Bulletin of the Virginia Agricultural Experiment Station, 92: 1-90.

https://vtechworks.lib.vt.edu/bitstream/handle/10919/29024/LD5655.V856_1991.P464.p

dfsequence=1

Perez-Sandi, C. M. 2001. Addressing the threat of Cactoblastis cactorum (Lepidoptera:

Pyralidae), to Opuntia in Mexico. Florida Ent. 84: 499-502.

https://journals.flvc.org/flaent/article/view/74994/72652

Pettey, F.W. 1948. The biological control of prickly pear in South Africa. Science Bulletin,

Department of Agriculture of the Union of South Africa, 271: 1-163.

Pimentel, D., Lach, L. Zuniga R. & Morrison, D. 2000. Environmental and economic costs

associated with non-indigenous species in the United States. BioScience, 50(1): 53-65.

http://www.rexano.org/Federal_ Canada/PimentelEtalOOCostExotics.pdf

Pimentel, D., McNair, S., Janecka, J., Wightman, J., Simmonds, C., O’Connell, C., Wong,

E., Russel, L., Zern, J., Aquino, T. & Tsomondo, T. 2001. Economic and environmental

threats of alien plant, animal, and microbe invasions. Agriculture, Ecosystems and

Environment, 84: 1-20.

https://www.forestpests.org/pdf/Economic %20and% 20environmental%20threats %200f%

20alien%20plant,%20animal,%20and%20microbe%20invasions.pdf

Pimentel, D., Whitecraft, M., Scott, Z. R., Zhao, L., Satkiewicz, P., Scott, T. J., Phillips, J.,

Szimak, D., Singh, G. & Gonzalez, D. O. 2010. Will limited land, water, and energy

control human population numbers in the future? Hum Ecol. 38(5): 599-611.

Pimentel, D., Zuniga, R. & Morrison, D. 2005. Update on the environmental and economic

costs associated with alien-invasive species in the United States. Ecological Economics,

52: 273-288. https://www.sci-hub.ru/10.1016/j.ecolecon.2004. 10.002

Pimienta, E. 1990. El nopal tunero. Universidad de Guadalajara, Guadalajara, Jalisco, Mexico:

246 pp.

Pina, L. I. 1977. La grana 0 cochinilla del nopal. Monografias LANFI, No. 1. Publicaciones de

los Lab. Nales. de Fomento Ind. de México: 55 pp.

Rahel, F. J. 2000. Homogenization of fish faunas across the United States. Science, 288(5467):

854-856. se180000854p (uwyo.edu)

Robertson, H. G. 1985. The ecology of Cactoblastis cactorum (Berg) (Lepidoptera: Phycitidae)

in relation to its effectiveness as a biological control agent of prickly pear and jointed cactus

in South Africa. PhD Thesis, Rhodes University, Grahamstown, South Africa.

Rodriguez-Garcia, M. E., De Lira, C., Hernandez-Becerra, E., Cornejo Villegad, M. A.,

Palacios- Fonseca, A. J., Rojas-Molina, I., Reynoso, R., Quintero, L. C., Del Real, A.,

Zepeda, A. & Munoz-Torres, C. 2007. Physicochemical characterization of Nopal pads

(Opuntia ficus indica) and dry vacuum nopal powder as a function of the maturation. Plant

Food Hum. Nutr., 62: 107-112.

29

Russell, C. E. & Felker, P. 1987. The prickly-pears (Opuntia spp., Cactaceae): a source of human

and animal food in semiarid regions. Econ Bot. 41(3): 433-445.

https://www.researchgate.net/profile/Peter-Felker/publication/225175919 The Prickly-

pears Opuntia spp Cactaceae_A Source _of Human and Animal Food _ in Semiarid

Regions/links/00b4952f3b06006089000000/The-Prickly-pears-Opuntia-spp-Cactaceae-

A-Source-of-Human-and-Animal-Food-in-Semiarid-Regions.pdf

Sala, O. E., Chapin, F. S., Armesto, J. J., Berlow, E., Bloomfield, J., Dirzo, R., Hubersanwald,

E., Huenneke, L. F., Jackson, R. B., Kinzig, A., Leemans, R., Lodge, D. M., Mooney,

H. A., Oesterheld, M., Poff, N. L., Sykes, M. T., Walker, B. H., Walker, M. & Wall,

D. 2000. Global biodiversity scenarios for the year 2100. Science, 287: 1770-1774.

Sammataro, D. & Avitabile, A. 1998. The Beekeeper’s Handbook, 3” edition. Ithaca, New

York, USA: Comstock Publishing Associates: 190 pp.

Schartel, T. E. & Brooks, C. P. 2018. Biotic constraints on Cactoblastis cactorum (Berg) host

use in the southern US and their implications for future spread. Food Webs, 26: 1-9.

https://doi.org/10.1016/).fooweb.2018.e00083

Seto, K. C., Fragkias, M., Giineralp, B. & Reilly, M. K. 2011. A meta-analysis of global urban

land expansion. PloS one, 6(8): e23777.

https://journals.plos.org/plosone/article/file?id=10.1371/journal.pone.0023777&type=pri

ntable

Shackleton, R. T., Witt, A. B. R., Piroris, F. M. & Wilgen, B. W. V. 2017. Distribution and

socio—ecological impacts of the invasive alien cactus Opuntia stricta in eastern Africa.

Biol. Invas. 19(8): 2427-2441.

https://link.springer.com/article/10.1007/s10530-017-1453-x

Shetty, A. A., Rana, M. K. & Preetham, S. P. 2012. Cactus: a medicinal food. Jour. Food Sci.,

Technol. 49(5): 530-536. doi: 10.1007/s13197-011-0462-5

Simberloff, D. & Stiling, P. 1996. How risky is biological control? Ecology, 77: 1965-1974.

Simmonds, F. J. & Bennett, F. D. 1966. Biological control of Opuntia spp. by

Cactoblastis cactorum in the Leeward Islands (West Indies). Entomophaga, 11: 183-189.

Singh, G. 2006. An overview of cactus pear research and development in India. Acta Hortic.,

728: 43-50. https://www.actahort.org/books/728/728 4.htm

Soberon, J., Golubov, J., & Sarukhan, J. 2001. The importance of Opuntia in México and

routes of invasion and impact of Cactoblastis cactorum (Lepidoptera: Pyralidae). Florida

Entomol. 84(4): 486-492.

https://kuscholarworks.ku.edu/bitstream/handle/1808/16761/SoberonJ FE 84%284%29486.

pdf?sequence=1 &isAllowed=y

Sowell, J. 2001. Desert ecology. An introduction to life in the arid Southwest. University of

Utah Press, Salt Lake City, Utah: 192 pp.

Spodek, M., Ben-Dov, Y., Protasov, A., Carvalho, C. J. & Mendel, Z. 2014. First record of

Dactylopius opuntiae (Cockerell) (Hemiptera: Coccoidea: Dactylopiidae) from Israel.

Phytoparasitica 42: 377-379. doi:10.1007/s12600-013-0373-2

Srivastava, M., Srivastava, P., Karan, R., Jeyaprakash, A., Whilby, L., Rohrig, E., Howe, A.

C., Hight, S. D. & Varone, L. 2019. Molecular detection method developed to track the

koinobiont larval parasitoid Apanteles opuntiarum (Hymenoptera: Braconidae) imported

from Argentina to control Cactoblastis cactorum (Lepidoptera: Pyralidae). Fla.

Entomol.,102(2): 329-335. https://doi.org/10.1653/024.102.0207

Starmer, W. T., Aberdeen, V. & Lachance, M. A. 1988. The yeast community associated with

decaying Opuntia stricta (Haworth) in Florida with regard to the moth, Cactoblastis

cactorum (Berg). Florida Sci., 51: 7-11.

30

Staveley, J. P., Law, S. A, Fairbrother, A. & Menzie, C. A. 2014. A causal analysis of observed

declines in managed Honey Bees (Apis mellifera). Human and Ecological Risk

Assessment. Taylor & Francis, 20(2): 566-591. doi:10.1080/10807039.2013.831263

Stiling, P. 2002. Potential non-target effects of a biological control agent, prickly pear moth,

Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae), in North America, and possible

management actions. Biol. Invasions, 4: 273-281.

Stohlgren, T. J., Tyler, L., Gregg, W., Muir, R. & Garrett, L. J. 2005. Preliminary assessment

of the potential impacts and risks of the invasive cactus moth, Cactoblastis cactorum Berg,

in the U.S. and Mexico. Final Report to the International Atomic Energy Agency, IAEA:

32 pp.

Strong, D. R. 1997. Fear no weevil? Science, 277: 1058-1059.

Suckling, D. M., Barrington, A. M., Chhagan, A., Stephens, A. E. A., Burnip, G. M., Charles,

J.G. & Wee, S. L. 2007. Eradication of the Australian painted apple moth Teia anartoides

in New Zealand: trapping, inherited sterility, and male competitiveness. Pages 603-615

in: Vreysen, M. J. B., Robinson, A. S. & Hendrichs J. (eds.), Area-Wide Control of Insect

Pests. From research to field implementation. Springer, Dordrecht, the Netherlands: 789

pp. https://www.1iaea.org/sites/default/files/area-wide-control-insect-pests-book.pdf

SYBAP (Statistical Year Book of Agri. Prod.). 2020. https://nube.siap.gob.mx/cierreagricola/

(accessed on 10 April 2020).

Teklehaimanot, H. S. & Tritschler, J. P. 2011. Evaluation of Spineless Cactus (Opuntia ficus-

indicus) as an Alternative Feed and Water Source for Animals during Dry Season in

Eritrea. Jn: Sustainable Agricultural Development. Behnassi, M., Shahid, S. & DSilva, J.,

(eds.) Springer, Dordrecht, the Netherlands: 245-252.

Terrazas-Salgado, T. & Mauseth, J.D. 2002. Shoot anatomy and morphology. Jn: Nobel, P. S.

(ed.), Cacti, Biology and Uses. Univ. of California Press, Los Angeles: 125-141.

Thomas, M. B. & Willis, A. J. 1998. Biological control - risky but necessary? Trends in Ecology

and Evolution, 13: 325-329.

Turner, C. E. 1985. Conflicting interests in biological control of weeds. Proceedings of VI

International Symposium on Biological Control of Weeds, E. S. Delfosse (ed.), Agriculture

Canada, Vancouver, Canada: 203-225.

Turner, C. E., Pemberton, R. W. & Rosenthal, S. S. 1987. Host utilization of native Cirsium

thistles (Asteraceae) by the introduced weevil Rhinocyllus conicus (Coleoptera:

Curculionidae) in California. Environmental Ent.,16: 111-115.

https://www.researchgate.net/profile/Robert-Pemberton-

2/publication/233568006_ Host Utilization _of Native Cirsium Thistles Asteraceae_by

the_Introduced_Weevil_ Rhinocyllus_conicus Coleoptera Curculionidae_in_ California/l

inks/5f9ec4f6a6fdccfd7b909eb2/Host-Utilization-of-Native-Cirsium-Thistles-Asteraceae-

by-the-Introduced-Weevil-Rhinocyllus-conicus-Coleoptera-Curculionidae-in-

California.pdf

USBC, 1998. Statistical Abstract of the United States. US Bureau of the Census, US

Government Printing Office, Washington, DC. (accessed January 1, 2021).

https://www.census.gov/library/publications /1998/compendia/statab/1 18ed.html

Van Dam, A. & May, B. 2012. A new species of Dactylopius Costa (Dactylopius gracilipilus

sp. nov.) (Hemiptera: Coccoidea: Dactylopiidae) from the Chihuahuan Desert, Texas,

U.S.A. Zootaxa, 3573(1): 33-39.

https://www.biotaxa.org/Zootaxa/article/view/zootaxa.3573.1.3

31

Van Soest, P. J., Robertson, J. B. & Lewis, B. A. 1991. Methods for dietary fiber, neutral

detergent fiber and non-starch polysaccharides in relation to animal nutrition. J. Dairy Sc1.,

74: 3583-3597.

Varone, L., Gonalons, C. M., FaltIhauser, A. C., Guala, M. E., Wolaver, D., Srivastava,

M. & Hight, S. D. 2020. Effect of rearing Cactoblastis cactorumon on an artificial diet

on the behavior of Apanteles opuntiarum. Appl. Entomol., 144: 278-286.

Varone, L., Logarzo, G., Martinez, J. J., Navarro, F., Carpenter, J. E. & Hight, S. D. 2015.

Field host range of Apanteles opuntiarum (Hymenoptera: Braconidae) in Argentina, a

potential biocontrol agent of Cactoblastis cactorum (Lepidoptera: Pyralidae) in North

America. Fla. Entomol., 98(2): 803-806. https://doi.org/10.1653/024.098.0265

Vigueras, G. & Portillo, L. 2001. Uses of Opuntia species and the potential impact of

Cactoblastis cactorum (Lepidoptera: Pyralidae) in Mexico. Fla. Entomol., 84(4): 493-

498. https://journals.flvc.org/flaent/article/view/74993/72651

Vila, M., Espinar, J. L., Hejda, M., Hulme, P. E., Jarosik, V., Maron, J. L., Pergl, J.,

Schaffner, U., Sun, Y., & PySek, P. 2011. Ecological impacts of invasive alien plants: a

meta-analysis of their effects on species, communities and ecosystems. Ecology Letters,

14(7): 702-708.

Volchansky, C. R., Hoffmann, J. H. & Zimmermann, H. G. 1999. Host-plant affinities of two

biotypes of Dactylopius opuntiae (Homoptera: Dactylopiidae) enhanced prospects for

biological control of Opuntia stricta (Cactaceae) in South Africa. J. Appl. Ecol., 36: 85-

Od:

https://besjournals.onlinelibrary.wiley.com/doi/full/10.1046/}.1365-2664.1999.00381.x

White, J. 1985. Studies in Plant Demography. Academic Press, London, 393 pp.

Williamson, M. H. 1996. Biological Invasions. Chapman and Hall, London: 268 pp.

Winston, M., Dropkin, J. & Taylor. O. 1981. Demography and life history characteristics of

two honey bee races (Apis mellifera). Oecologia, 48: 407-413.

Yahia, E. M. & Mondragon-Jacobo, C. 2011. Nutritional components and anti-oxidant capacity

of ten cultivars and lines of cactus pear fruit (Opuntia spp.). Food Res. Int., 44: 2311-

2318.

https://www.sciencedirect.com/science/article/abs/pii/S096399691100144X?via%3Dihub

Zimmermann, H., Bloem, S. & Klein, H. 2004. Biology, history, threat, surveillance and control

of the cactus moth, Cactoblastis cactorum. FAO. IAEA: 40 PP.

https://www-pub.iaea.org/mtcd/publications/pdf/faobsc_web.pdf

Zimmermann, H. G., Moran V. C & Hoffmann, J. H. 2000a. The renowned cactus moth,

Cactoblastis cactorum: its natural history and threat to native Opuntia floras in Mexico and

the United States of America. Diver. and Distrib., 6(5): 259-269.

https://onlinelibrary.wiley.com/doi/full/10.1046/j.1472-4642.2000.00088.x

Zimmermann, H. G.; Pérez, M.; Goluvob, J.; Soberén, J.; & Sarukhan, J. 2000b.

Cactoblastis cactorum, una nueva plaga de muy alto riesgo para las opuntias de México.

Biodiversitas, 33: 1-14.

http://www.conabio.gob.mx/institucion/conabio_espanol/doctos/biodiver33.pdf

Zwolfer, H. & Harris, P. 1984. Biology and host specificity of Rhynocyllus conicus (Froel.)

(Col., Curculionidae), a successful agent for the biocontrol of thistle Carduusnutans L.

Zeitschriftfiir Angewandte Entomologie, 97: 36-62.

32

The Taxonomic Report

is a platinum open access peer-reviewed publication of

The International Lepidoptera Survey (TILS)

The International Lepidoptera Survey is registered as a non-profit Limited Liability Company (LLC) in the state

of Virginia, U.S.A. The Taxonomic Report (TTR), ISSN 2643-4776 (print) / ISSN 2643-4806 (online) is

published for the purpose of providing a public and permanent scientific record. Articles are peer-reviewed but

not necessarily through the anonymous editor-mediated review process. Typically, the authors are encouraged to

solicit reviews of their manuscripts from knowledgeable lepidopterists before submission. TTR appears in digital,

open-access form, is disseminated as a hardcopy to select institutional repositories, and is available as printed copy

upon request at the discretion of authors and/or the editor. Printing and postage charges may apply. An initial run

of 25 copies is printed on paper to meet ICZN Code recommendation 8B. All published TTR articles are freely

available at the archival TTR website (http:/Aepsurvey.carolinanature.com/report.html) and via the following

digital repositories:

Internet Archive (https://archive.org/)

Biodiversity Heritage Library (https://www.biodiversitylibrary.org)

Zobodat (https://www.zobodat.at/)

Zenodo (https://zenodo.org)

TILS Purpose

TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific discovery,

determination, and documentation, without which there can be no preservation.

TILS Motto

“As a world community, we cannot protect that which we do not know”

Articles for publication are sought

Manuscripts may deal with any area of research on Lepidoptera, including faunal surveys, conservation topics,

life histories and foodplant records, matters of nomenclature, descriptions of new taxa, methods, etc. Taxonomic

papers are particularly welcome. There are no publication charges for authors. Before submitting a manuscript,

email TTR editor, Harry Pavulaan, 606 Hunton Place NE, Leesburg, VA, 20176, USA at

intlepsurvey @ gmail.com (cc: to harrypav @ hotmail.com if you do not receive a reply within one week) to initiate

discussion on how to best handle your material for publication, and to discuss peer review options.

Visit The International Lepidoptera Survey on the World Wide Web at:

http:/Aepsurvey.carolinanature.com

&

Join the discussion at our list serve on Groups.io at:

https://groups.io/g/TILS

You can subscribe by sending an email to: TILS+subscribe @ groups.io

&

Join The International Lepidoptera Survey on Facebook at:

https://www.facebook.com/groups/1072292259768446

33