Rear declining species in captivity
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Overall effectiveness category Evidence not assessed
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Number of studies: 22
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Supporting evidence from individual studies
A replicated, paired, controlled study in 1964 in a fen in Cambridgeshire, UK (Duffey 1968) reported that semi-wild large copper Lycaena dispar batavus caterpillars reared at high density on small great water dock Rumex hydrolapathum plants had lower survival than caterpillars reared at lower density or on larger plants. Results were not tested for statistical significance. On small plants, the survival of large copper caterpillars reared in groups of 12/plant was 35% (67/192 survived), compared to 65% (31/48 survived) for caterpillars in groups of three/plant. However, on large plants the survival of caterpillars in groups of 12/plant was 81% (39/48 survived) compared to 75% (9/12 survived) for caterpillars in groups of three/plant. The author reported that small plants with 12 caterpillars/plant were abandoned after all the leaves had been eaten, before the caterpillars were fully grown. In a fen with a semi-wild large copper colony, four batches of 10 great water dock plants were selected. In each batch, eight plants were 50 cm tall (3 leaves/plant) and two were >100 cm high (9–20 leaves/plant). In May 1964, three or 12 large copper caterpillars were placed onto each plant, and the plants were covered with a 6-mm plastic mesh cage to exclude birds and mammals. In July 1964, all surviving caterpillars and pupae were counted.
Study and other actions testedA controlled study in 1968 at a research station in Cambridgeshire, UK (Duffey 1977) reported that large copper butterflies Lycaena dispar batava laid more eggs in a cage kept in a greenhouse than in a cage kept outside, and eggs in the greenhouse had a higher hatching success. Results were not tested for statistical significance. The number of eggs laid in a cage kept in a greenhouse (498 eggs) was higher than the number laid in a cage kept outside (126 eggs). In addition, the proportion of eggs which hatched was higher in the greenhouse (91%) than outside (40%). In summer 1968, two cages (5.40 × 1.65 × 1.80 m) were constructed from 1 × 1 cm mesh. One was kept in a greenhouse and the other was placed outside. Each cage contained 15 female and 23 male large coppers, and 20 potted great water dock Rumex hydrolapathum plants. From 17 May–14 August 1968, the mean maximum temperature in the greenhouse cage (23.8°C) was higher than in the outside cage (16.8°C), but the mean minimum temperature was similar (greenhouse: 8.7°C; outside: 8.6°C). Eggs were counted daily.
Study and other actions testedA controlled study in 1979–1980 in a laboratory in the UK (Thomas & Wardlaw 1990) found that large blue Maculinea arion caterpillars reared in ant Myrmica spp. nests without a queen present were more likely to survive than caterpillars reared in nests with a queen. The survival of caterpillars in nests without queen ants (10 out of 26 caterpillars) was higher than in nests with queen ants present (6 out of 39 caterpillars). The authors reported that caterpillars in Myrmica scabrinodis nests had lower survival than caterpillars in nests of the other species (data not presented). In 1979 and 1980, a total of 65 Myrmica ant colonies were established, containing 20–1,137 workers/colony depending on nest design (see paper for details). In each of 26 nests, 1–6 queen ants (depending on colony size) were present, and the other 39 nests did not contain queens. Most nests were Myrmica sabuleti, but four colonies were established with each of Myrmica rubra, Myrmica ruginodis and Myrmica scabrinodis. After >1 week, one newly moulted caterpillar was introduced to each nest. Caterpillar survival was monitored for >2 weeks.
Study and other actions testedA controlled study (years not given) in a laboratory in the UK or France (location not clear) (Elmes et al. 1991) found that the survival of mountain Alcon blue Maculinea rebeli caterpillars reared in captivity differed between ant Myrmica spp. species, but not between colonies with or without queens present. Mountain Alcon blue caterpillars reared with Myrmica schencki (10 out of 99 survived to pupation) had higher survival rates than caterpillars reared with Myrmica sabuleti (4/78 survived), Myrmica scabrinodis (2/43 survived), Myrmica rubra (1/112 survived), Myrmica ruginodis (4/71 survived) or Myrmica sulcinodis (1/24 survived). The survival of caterpillars reared for three weeks in colonies with queens was 31–89%, compared to 43–78% without queens (statistical significance not assessed, see paper for details on each ant species). Over five years, >800 mountain Alcon blue caterpillars were introduced to 120 ant colonies kept in small plastic “Brian” nests (no further details provided). Colonies were collected from France and England, fed fruit flies Drosophila spp. and sucrose, and kept at a constant temperature which was adjusted weekly to mimic natural temperatures. Caterpillars were collected on their food plant, and placed into the foraging areas of the ants after emergence. Caterpillar survival was monitored for between three weeks and 10 months (to pupation) in the ant nests.
Study and other actions testedA replicated, controlled study (years not given) in a laboratory in Spain (Thomas et al. 1993) found that mountain Alcon blue Maculinea rebeli caterpillars reared in ant Myrmica rubra nests at low density had higher survival rates than caterpillars reared at higher densities. The survival of mountain Alcon blue caterpillars reared at low density (five caterpillars/nest: 18 of 20 caterpillars survived) was higher than for caterpillars reared at higher densities (10 caterpillars/nest: 29/40 survived; 25 caterpillars/nest: 22/100 survived). At higher densities, more caterpillars survived in ant nests founded from a colony containing winged females (10: 19/20 survived; 25: 17/50 survived) than from a colony without winged females (10: 10/20 survived: 25: 5/50 survived), but surviving individuals were lighter in colonies with winged females (22–27 mg) than without them (37–50 mg). Caterpillars reared at low density weighed 39–46 mg. Two Myrmica rubra nests in the Pyrenees were excavated and used to establish 12 colonies, each containing 50 workers and 10 ant larvae. Colonies were kept in “Brian” nests with abundant food (no further details provided). When excavated, one nest contained a large number of winged females while the other contained none. In August, on the evening of their final moult, 160 mountain Alcon blue caterpillars were collected from the same site, and introduced to the ant colonies at three densities: five, 10 and 25 caterpillars/colony. From October–March, nests were overwintered in a cool room, after which caterpillars grew for another eight weeks until pupation. The survival and weights of caterpillars were recorded before pupation.
Study and other actions testedA study in 1994 in Cape Town, South Africa (Heath & Brinkman 1995a) reported that a female Dickson’s copper Oxychaeta dicksoni butterfly laid eggs in captivity in the presence of black cocktail ants Crematogaster perengueyi collected from the site where she emerged, but not in the presence of ants from 10 km away. The female laid ~100 eggs in captivity over 10 days when she was placed with ants that had been collected from the same site as her, but when these ants were swapped for others collected from a site 10 km away she stopped laying. When these ants were replaced with others from the original site, she laid another 20 eggs. Thirteen out of 20 eggs kept in captivity hatched 18–20 days after being laid but none of the larvae survived to adulthood. A freshly emerged female Dickson’s copper was collected from the wild after she was observed mating. She was placed in a small container (dimensions not provided), covered with a nylon net, containing Phylica stems and two black cocktail ants from either the same site or a site 10km away. Ten of the resulting larvae were placed in a container with a small ants’ nest taken from the same site as the female butterfly and a potted Phylica plant. The other three larvae were placed in a petri dish with two black cocktail ants and some dry twigs.
Study and other actions testedA study in 1995 in a captive setting in the Western Cape, South Africa (Heath & Brinkman 1995b) reported that two wild-caught, gravid female scarce mountain copper butterflies Argyrocupha malagrida brought into captivity laid eggs, but none hatched. Two gravid females of the scarce Table Mountain copper subspecies Argyrocupha malagrida maryae and Argyrocupha malagrida paarlensis, collected from the wild, laid eggs in captivity (approximately 12 each). Time from capture to laying is not reported. Eight months after laying no eggs had hatched but dissection revealed the larvae inside were still alive. In January 1995, one Argyrocupha malagrida maryae was caught near Bredasdorp and one Argyrocupha malagrida paarlensis on the Perdeberg mountain. They were each placed in their own container covered with netting and containing pieces of buchu plant Agathosma sp. And three common pugnacious ants Anoplolepis custodiens from where the butterflies were caught. Six weeks after laying, the eggs were exposed to direct sunlight for short periods daily, but this was stopped after 3–4 days. Eggs were misted with water once every two months. An egg was dissected at three, six and eight months after hatching.
Study and other actions testedA study in 1995–1996 in a captive setting in the Western Cape, South Africa (Edge & Pringle 1996) reported that 27% of wild-caught Brenton blue butterfly Orachrysops niobe eggs developed into adults. Of 15 eggs collected from the wild and kept in captivity, four developed into adult butterflies. However, captive reared adults were smaller (18–20 mm wingspan) than average wild-reared adults (26–30 mm). Some caterpillars died from a fungal infection (numbers not provided). In November 1995, researchers observed Brenton blue butterflies ovipositing at the Brenton Blue Butterfly Reserve near Knysna and collected 15 freshly-laid eggs to be reared in captivity. Eggs were placed in small air-tight plastic containers and provided with a sprig of Indigofera porrecta every 3–4 days throughout development. Eggs, all caterpillar instars, pupae and adults were measured. Survival data is only provided for adults.
Study and other actions testedA study in 1996–1997 in a captive setting in the Western Cape, South Africa (Claassens & Heath 1997) reported that wild-caught final instar Cape Peninsula butterfly Thestor yildizae caterpillars placed in an artificial pugnacious ant Anoplolepsis custodiens nest successfully pupated and eclosed, but captive-hatched first instar Cape Peninsula and Riley’s skolly butterfly Thestor rileyi caterpillars placed next to an artificial ant nest did not survive to pupation. Three wild-caught final instar Cape Peninsula butterfly caterpillars were placed inside an artificial ant nest, were fed by ants with oral fluids, and survived to adulthood. However, first instar caterpillars of Cape Peninsula and Riley’s skolly butterflies (numbers not provided), which hatched from eggs laid in captivity and were placed next to the artificial ant nest, were not observed being fed by ants, and died before pupation. Final instar Cape Peninsula caterpillars were collected from the wild and three were placed inside an artificial pugnacious ant nest attached to a glass arena. Adult female Cape Peninsula and Riley’s skolly butterflies were also collected in the wild and put in containers to oviposit (numbers not given). Some of the hatching first instar caterpillars were placed in the arena next to the nest. In both experiments ant feeding behaviour, and caterpillar development times and survival were observed.
Study and other actions testedA review in 1998 (Wardlaw et al. 1998) reported that four species of large blue butterfly Maculinea spp. were bred in captivity using ant Myrmica spp. colonies, with varying success. Alcon blue Maculinea alcon and mountain alcon blue Maculinea rebeli caterpillars were successfully reared in captive ant colonies on different occasions over 20 years (data not presented). Large blue Maculinea arion caterpillars were sometimes reared successfully using two methods, but 13 other nest designs failed (data not presented). Scarce large blue Maculinea teleius caterpillars were reared for up to eight months, using common red ant Myrmica rubra and Myrmica scabrinodis in two nest designs. Wild butterfly eggs were collected on flowering stems of food plants (gentian Gentiana spp., wild thyme Thymus spp., oregano Origanum spp., great burnet Sanguisorba officinalis). Ant colonies (see paper for six species) were collected by excavating nests. Caterpillars were introduced to ant colonies after they dropped from the flower heads. A variety of rearing methods were used, with different nest box designs (see paper for details).
Study and other actions testedA site comparison study in 1997–1998 in a laboratory in the UK (Nicholls & Pullin 2000) found that wild- and captive-laid large copper Lycaena dispar batavus eggs and caterpillars had similar survival in captivity. Both the survival to overwintering of wild-laid eggs (19 of 20 caterpillars), and the overwinter survival of these caterpillars (4 of 19 caterpillars), were statistically similar to the survival to overwintering (15 of 20 caterpillars) and overwinter survival (3 of 15 caterpillars) of captive-laid eggs. In September 1997, twenty wild-laid eggs were collected from a lowland bog in the Netherlands, and 20 captive-laid eggs were obtained from a 25-year-old glasshouse-reared colony at Woodwalton Fen. Eggs were reared to overwintering under controlled conditions (10 hours light, 14 hours dark, 20°C) in a laboratory. Immediately before overwintering, caterpillars were transferred to great water dock Rumex hydrolapathum pot plants and maintained in an overwinter environment (10 hours light, 14 hours dark, 5°C) for 20 weeks. Emergence was stimulated by increasing light by 15 minutes, and increasing temperature by 2°C, every two days for eight days, and survival was recorded.
Study and other actions testedA study (year not specified) in two captive-rearing facilities in Warwickshire and Oxfordshire, UK (Lewis & Thomas 2001) reported that large white Pieris brassicae were successfully reared in captivity for >25 years, but found some morphological changes occurred. A population of large white were bred in captivity for >25 years (100–150 generations). However, long-term captive-bred butterflies were heavier (1.9 g) than butterflies in their third generation in captivity (1.8 g), and had smaller, shorter and broader wings (see paper for details). Captive-bred females laid more eggs (340 eggs/female) than females new to captivity (30 eggs/female). Caterpillars in a long-term captive population, originally caught in southern England >25 years ago, were reared on a synthetic diet at 23–25°C. Adults were kept in cages (45 × 80 × 48 cm) with 150–200 adults/cage. Wild egg batches were collected in Glamorgan, UK, and reared through two generations in captivity in the same conditions. Data were collected on the number of eggs laid in the first 16 days after emergence by 15 females kept with 15 males of each group, and the weight and wing size of freshly emerged adults (number not specified).
Study and other actions testedA study in 1999 in a captive setting in the KwaZulu-Natal province, South Africa (Lu & Samways 2001) reported that 93% of wild-caught Karkloof blue butterfly Orachrysops ariadne eggs hatched and survived for up to three months in the presence of their host plant Indigofera woodii Bolus var. laxa, but none of the caterpillars survived to adulthood. Of 30 Karkloof blue butterfly eggs collected from the wild, 28 successfully hatched. Of the 18 resulting caterpillars that were kept in the laboratory, all were recorded feeding on the host plant and survived for up to three months, moulting two or three times, before dying from an unidentified fungal infection. In May–June 1999, 30 eggs were collected from three grassland reserves in the KwaZulu-Natal province and taken into captivity. Upon hatching each caterpillar was placed in a small container with fresh Indigofera shoots that were replaced every 1–2 days. Eighteen caterpillars were monitored until death for feeding behaviour, moulting occurrence and presence of fungal patches. Ten were returned to the field but not monitored.
Study and other actions testedA study in 2003 in a laboratory in the Western Cape Province, South Africa, (Edge & van Hamburg 2003) reported that Brenton blue butterfly Orachrysops niobe caterpillars reared on stems of their larval host plant Indigofera erecta with no ants became dwarf adults, but one of two larvae reared on a whole Indigofera erecta plant, with a Camponotus baynei ant colony, became a full-sized adult. Brenton blue caterpillars that were reared on cut stems of Indigofera erecta emerged as dwarf adults (average 10–13 mm forewings, number of individuals not provided). Of the two larvae that were reared on a full Indigofera erecta plant with an ant colony, one emerged as a full-sized adult (17 mm) and one died before pupation. No statistics were conducted. For the cut stem treatment, Indigofera erecta stems containing eggs found at the Brenton Blue Butterfly Reserve were cut and placed in containers. After hatching, fresh stems were provided every few days throughout larval development. Measurements of surviving caterpillars were taken at each instar and during and after pupation. For the whole plant and ant colony treatment, two caterpillars were each placed in connecting containers with an Indigofera erecta plant each. These containers were linked to another compartment containing an ant nest collected from the wild. The surviving caterpillar was measured as an adult.
Study and other actions testedA study (years not given) in a captive setting in the Western Cape, South Africa (Edge et al 2008) reported that wild-collected Brenton blue butterfly Orachrysops niobe eggs hatched and caterpillars kept with only Phyllanthus incurvatus leaves died before completing the first instar of development but all caterpillars also given Indigofera erecta leaves developed normally through to the fourth instar. Caterpillars in captivity were seen feeding on the leaves of Indigofera erecta but not Phyllanthus incurvatus. No data on number of eggs collected or number of caterpillars in each treatment are provided. Cuttings were taken from Phyllanthus incurvatus plants in the wild where Brenton blue butterflies had laid eggs, and the cutting and eggs were placed in air-tight plastic containers. Some containers were left with only their original Phyllanthus incurvatus cutting, whereas an Indigofera erecta cutting was placed in others once the eggs had hatched. Feeding behaviour and survival to the fourth instar was recorded.
Study and other actions testedA controlled study in 2003–2006 in two captive-breeding facilities in Washington, USA (Schultz et al. 2009) found that captive-reared Puget blue butterflies Icaricia icarioides blackmorei were smaller than wild-born individuals, and caterpillars kept in refrigerators overwinter had lower survival than other treatments. There was no significant difference between the survival to adulthood of eggs collected from the wild (17/200 eggs) and eggs laid in captivity (39/548 eggs), or of caterpillars kept in environmental chambers (49/514 caterpillars) or outdoor enclosures (49/450) overwinter. However, all 308 caterpillars kept in refrigerators overwinter died. Captive-reared butterflies were smaller than wild-caught butterflies, but adult size was similar between all captive treatments (see paper for details). In June 2003, forty-eight female butterflies were collected from the wild and 39 laid 1,879 eggs in captivity. Overwinter, surviving caterpillars were kept in one of three treatments: a refrigerator, an environmental chamber with light, humidity and temperature approximating optimal rearing conditions, or an outdoor enclosure experiencing ambient conditions (see paper for housing details). In 2004, surviving caterpillars were reared on netted sickle-keeled lupine Lupinus albicaulis. In spring 2005, sixty female butterflies were collected from a second site and 51 laid 548 eggs in captivity. In addition, lupine leaves with 200 wild-laid eggs were collected and reared in captivity. All caterpillars overwintered in outdoor enclosures. In 2004–2006, captive-reared and wild-caught adults were weighed and measured.
Study and other actions testedA replicated, controlled study (years not given) in a laboratory in Belgium (Turlure et al 2010) found that cranberry fritillary Boloria aquilonaris caterpillars reared at 20 °C had higher survival to pupation than those reared at 25 °C, but there was no effect on survival of the amount of sphagnum moss Sphagnum spp. that caterpillars were reared with. Ninety percent of 68 cranberry fritillary caterpillars kept at 20 °C survived to become pupae, compared to 65% of 68 kept at 25 °C. However, the number of pieces of sphagnum moss that caterpillars were kept with did not significantly affect survival (1 piece = 70–94% survival, 6 pieces = 60–85%). One-hundred-and-thirty-six first, second and third instar wild-collected caterpillars were kept individually in petri dishes with small cranberry Vaccinium oxycoccus leaves. All caterpillars were kept at 10 °C at night, but half were kept at 20 °C during the day, and half at 25 °C. Within each temperature regime, half were kept with one piece of sphagnum moss and half with six pieces. This was primarily done to manipulate humidity, since the enclosures with one piece of sphagnum moss were less humid than those with six pieces. Every two days the petri dishes were cleaned and small cranberry leaves and sphagnum moss pieces were replaced. Survival to pupation was recorded.
Study and other actions testedA study in 2004–2007 in six captive-breeding sites in Cumbria, UK (Porter & Ellis 2011) reported that a captive population of marsh fritillary Euphydryas aurinia increased in size over two years. Results were not tested for statistical significance. Two years after 250 caterpillars were taken into captivity, the captive population was estimated at 50,000 caterpillars. In September 2004, the only two caterpillar webs (containing 155 individuals) remaining locally were taken into captivity. In addition, 95 caterpillars from 19 populations (five from each location) in west Scotland were collected. Caterpillars were checked for infection with the parasitoid Cotesia melitaearum. Caterpillars were kept at six separate locations, and reared in natural conditions using large netted cages and pot-grown devil’s-bit scabious Succisa pratensis, supplemented with garden varieties of honeysuckle Lonicera spp., snowberry Symphoricarpos albus and wild honeysuckle Lonicera periclymenum. The number of caterpillars in the captive population was estimated in spring 2007.
Study and other actions testedA replicated, site comparison study in 2014 in a laboratory in Poland (Witek et al. 2016) found that scarce large blue Maculinea teleius caterpillars reared by ants Myrmica scabrinodis from sites where the butterfly occurs survived longer than caterpillars raised by ants from sites where the butterfly does not occur, but all caterpillars ultimately died. The survival of scarce large blue caterpillars raised in ant colonies collected from sites where scarce large blue occurs was higher than in colonies collected from sites where scarce large blue does not occur (data presented as model results). However, no caterpillars survived >35 days. In August 2014, ten ant colonies were collected from each of four wet meadows, 110–470 km apart: two where scarce large blue and other ant parasites occurred and two where they did not. Each colony (50 old and 50 young workers with 15 ant larvae) was placed in a plastic box (20 × 12 × 7 cm) containing a patch of wet plaster covered by a flowerpot saucer with an entrance notch. Great burnet Sanguisorba officinalis stems were collected from one site, and placed in water with the flowerheads bagged in eight bunches of 25 stems. Bunches were shaken each morning to collect fourth instar caterpillars, and one caterpillar was placed in each ant colony. Fifteen ant larvae were added to each colony each week as food. The survival of caterpillars was checked every 1–2 days until all caterpillars had died.
Study and other actions testedA study in 2008–2016 in a zoo in Essex, UK (Gardiner et al 2017) reported that Fisher’s estuarine moth Gortyna borelii lunata eggs collected from the field successfully survived to adulthood, bred successfully in captivity and the resulting captive population continued for at least eight generations. Data on survival and breeding success were not provided. In 2008, ten batches of Fisher’s estuarine moth eggs were collected from a natural population and placed in 2 m3 stainless steel mesh cages. In April–May, post-hatching, caterpillars were placed in cages containing one or two potted hog’s fennel Peucedanum officinale plants, with 1–2 caterpillars/cage. In September–October, as adults, they were placed in cages containing a potted hog’s fennel plant and coarse grass to mate and lay eggs (numbers/cage not provided). In March, some eggs were placed in pots in a cold frame until they hatched and used to continue the captive breeding programme and some were translocated to six sites in Essex as part of a reintroduction plan (proportions of eggs kept and reintroduced not provided). Every three years after 2008, eggs were collected from the original donor site to supplement the captive-bred population (numbers of eggs not provided).
Study and other actions testedA replicated, randomized, paired, controlled study in 2014–2016 in a greenhouse in Iowa, USA (Pocius et al. 2017) reported that monarch butterflies Danaus plexippus were successfully reared in captivity for 12 generations, and found that caterpillar survival differed between milkweed Asclepias species. A population of monarchs was bred in captivity for 12 generations. However, more caterpillars fed on butterfly milkweed Asclepias tuberosa (75%) or poke milkweed Asclepias exaltata (72%) survived to adulthood than caterpillars fed on tall green milkweed Asclepias hirtella (31%) or prairie milkweed Asclepias sullivantii (36%). In May–June 2014, a total of 253 wild monarch eggs and young caterpillars were collected. Caterpillars were fed on common milkweed Asclepias syriaca in the summer, and a tropical milkweed Asclepias curassavica in the autumn and winter. Adults were tested for parasites Ophryocystis elektroscirrha before being allowed to mate. In the 13th generation, individual, newly hatched caterpillars were placed on an 8-week-old milkweed plant grown from seed. Thirty-six blocks, each containing one plant of nine milkweed species (butterfly, poke, tall green, prairie, common, swamp Asclepias incarnata, showy Asclepias speciosa, whorled Asclepias verticillata and honeyvine Cynanchum leave milkweed), were placed in a pop-up cage (57 × 37 × 55 cm) and netting in a greenhouse. From day 12, cages were checked daily, and pupae were moved to a laboratory until emergence.
Study and other actions testedA replicated, randomized, controlled study in 2014–2015 on a farm in Galilee, Israel (Berman et al. 2018) found that captive spring webworm Ocnogyna loewii caterpillars fed vegetation from grazed paddocks had a similar growth rate to caterpillars fed vegetation from ungrazed paddocks. Over five days, the growth rate of caterpillars fed on vegetation from cattle-grazed paddocks (0.12 mg/mg/day) was similar to caterpillars fed vegetation from ungrazed pastures (0.11 mg/mg/day). Sixty wild, fourth instar caterpillars were collected and weighed, and placed in individual plastic containers (12 cm diameter, 8 cm height) with a perforated lid. Caterpillars were randomly divided into six groups, and fed daily with fresh plants from one of six paddocks (three grazed, three ungrazed). Caterpillars were re-weighed after five days, and their growth rate calculated.
Study and other actions tested
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This Action forms part of the Action Synopsis:
Butterfly and Moth ConservationRelated Actions
| Actions | Effectiveness | Studies | Category | |
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Release captive-bred individuals to the wild Action Link |
Evidence not assessed | 13 |
Synopsis Link |
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Rear declining species in captivity Action Link |
Evidence not assessed | 22 |
Synopsis Link |
Butterfly and Moth Conservation - Published 2023
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