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Insect Tanatocoenosis (Death Assemblage) related to the African Tulip Tree (Spathodea campanulata) in Popayán, Colombia (South America)

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Entomological Tanatocoenosis related to the African Tulip Tree (Spathodea campanulata) in Popayán, Colombia (South America) by Jaime Andrés Ochoa García (Ecologist) The following report is a shortened version of my 2001 thesis dissertation. Spathodea campanulata was introduced in Colombia in the 1930s as an ornamental plant. Nowadays, regardless of its African origin, this tree is considered a pantropical species and in Hawaii it is seen as a pest. The local government has used this species to decorate the roads of the administrative area (Cauca) and the streets of its capital city, Popayán, in southwestern Colombia (aprox. 800km southwest of Bogotá). In 1998 the local government´s plant nursery had an estimated 2000 seedlings ready to be planted. This tree grows at different altitudes in Colombia, from sea level to at least 1800 meters above sea level. The trees in the areas that were checked were in full bloom from January to June. During the years 1997, 1998, 1999, and 2000 many (about 100,000) flowers of the African Tulip Tree (Spathodea campanulata) were collected since it was presumed that many insects were dying in its flowers. I set out to confirm the observation that many insects were dying in Spathodea campanulata´s flowers and on a weekly basis, five designated areas around the city were probed. I personally collected from the ground and opened every single flower. The idea was to get all the insects that were found dead and later identify them up to the taxon: order; some families under those orders were also identified. Tanatocoenosis means death assemblage, so after four years of painstakingly opening flowers and retrieving dead insects from the inside of those flowers, it was concluded that indeed insects are found dead (thirty thousand insects were retrieved), and the most common orders of dead insects were hymenoptera, coleoptera, diptera, and lepidoptera. This research did not aim at pinpointing the cause of those deaths, despite that, I had a laboratory run a phytochemical test and it showed that the flowers contain flavonoids and alkaloids, but I cannot state that these substances are the real cause of the deaths of the insects. The inside of the flowers feel rather sticky and some contain a gooey substance. So insects, after flying or crawling into the flowers, may die after getting stuck in the inside walls of the flowers, this was not proved; a thorough analysis in this respect is needed. What can be stated is that insects play a central role in the pollination of flowers and the most recognized pollinators are precisely insects in the orders that are dying in the African Tulip tree flowers as it was corroborrated in my study; this tree species places a threat to the local insect community, if this tree colonizes tropical forests, their local entomofauna will be at risk. I have had the chance to collect African Tulip Tree flowers in other Colombian towns and cities (Piendamó, Cali, Palmira, Armenia, Tuluá, and Medellín) as well as in other countries: Ecuador and The USA (Hawaii - Oahu-). Unfortunately, I also found dead insects of the same orders in those towns and cities, so it is a death event that is related to the flowers of this tree. There have been reports that hummingbirds have been found dead in Brazil but this could not be proved. After this reasearch was done it can be said that potential local pollinators die in big numbers in the flowers of the African Tulip Tree so it is recommended that local governments reconsider planting this species of tree in order to relieve the local insect community from the ecological pressure exerted by this alien species on the colombian environment. Addendum: Eleven years after finishing my research, I still collect African Tulip tree flowers sporadically and I have to sadly report the same observations as the ones presented 11 years ago: the same taxon of insects are still dying and more of these trees are seen decorating not only the streets of Popayán but also of more colombian areas. Jaime Andrés Ochoa G.
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Entomological Tanatocoenosis related to the African Tulip tree in Popayán, Cauca (Colombia, South America). Undergraduate Thesis. 2001.
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Associations

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Spathodea campanulata is frequently visited by nectar-feeding birds, which may be important pollinators (Corlett 2005). Rangaiah et al. (2004) found that birds were the exclusive pollinators of these flowers, based on their observations. They noted that bees also fed on the floral rewards, but they tended to feed on flowers from a single tree and therefore could not be effective pollinators given the self-incompatibility of S. campanulata. Trigona bees died in the calyx water and nectar and were consumed by visiting birds.

Ayensu (1974) reported that he had observed several bat visits to these flowers over several years in Ghana. He observed Micropteropus pusillus inserting their heads into the cup-like flowers to lap large quantities of rather dilute nectar. On several occasions, rather than enter the corolla tube, the bats tore the basal part of the flower cup, thereby obtaining an easy flow of nectar, which they lapped up. (Ayensu 1974)

Bito (2007) studied moth communities colonizing S. campanulata invading secondary rain forest vegetation in Melanesia. They found 54 species of folivorous Lepidoptera on S. campanulata. Most were generalists, feeding on more than just a single native plant family. However, the three most abundant species, representing 83% of all individuals (Acherontia lachesis, Hyblaea puera complex, and Psilogramma menephron) were relatively host-specific, feeding mainly on a single native family (native hosts: Rubiaceae, Verbenaceae, and Loganiaceae, respectively). Most of the 23 species analysed in detail had a wide geographic distribution, including 13 species wih distributions spanning the entire 1000 km study transect. Despite its phylogenetic isolation from the native vegetation, Spathodea campanulata was rapidly colonized (on a scale of decades) by folivorous Lepidoptera communities with a resulting species richness and dominance structure indistinguishable from the assemblages feeding on native hosts. Although most species were generalists, the highest population densities were reached by relatively specialized species, similar to the communities on native hosts. The species turnover across distances from 10 to 1000 km was relatively low as most of the species had wide geographic ranges.

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Brief Summary

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The African Tuliptree (Spathodea campanulata) is native to tropical forests in a broad area of sub-Saharan Africa. However, it is now grown far more widely than this throughout the world's tropics and has been reported to be an invasive colonizer in the South Pacific (e.g., Tahiti and Rarotonga, Meyer 2004), Puerto Rico, and Brazil. (Bittencourt et al. 2003 and references therein). In Tahiti, for example, S. campanulata often dominates lowland mesic forests as well as native upland wet forests up to 1,200 m in elevation (Meyer 2004). It was first reported from the Pacific Islands (Hawaii) prior to the 20th century (Whistler 1995). These trees grow very rapidly (measured trees in Puerto Rico increased their trunk diameter by as much as 2 inches per year), but require nearly full sun (Little and Wadsworth 1964). Under good conditions, they may begin flowering as young as 3 or 4 years of age, with trees grown in the open flowering when they are about 5 m tall (Orwa et al. 2009).

Spathodea campanulata has large bright red flowers with the edges of the corolla lobes colored yellow ("corolla" is a collective term for all the petals in a flower). The flowering heads appear in circular masses with packed buds. The buds on the outer portion of the head open together, surrounding the inner buds. (Ayensu 1974)

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Distribution

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In its introduced range in the eastern Pacific, Spathodea campanulata grows in mesic-wet forests from 10 to 1200 m (Meyer 2004).

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Ecology

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Spathodea campanulata has been a very successful invader in many regions to which it has been introduced. For example, together with Piper aduncum, it is the most successful woody invader of secondary forests in the northern New Guinea lowlands and adjacent Bismarck Archipelago. The tree invades early stages of rain forest succession developing in abandoned gardens from swidden (slash-and-burn) agriculture or after natural disturbance, such as the opening of large forest gaps from treefalls and landslides, but it does not penetrate into closed primary forests. (Bito 2007) Whistler (1995) writes that it escapes and becomes naturalized in disturbed lowland areas such as pastures and shrublands in Fiji, Samoa, Hawaii, Guam, and perhaps elsewhere in the Pacific Islands. However, Meyer (2004) reported that this species is indeed able to penetrate into apparently pristine undisturbed forests in the Society Islands in the eastern Pacific.

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Morphology

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The African Tuliptree (Spathodea campanulata) produces tubular tulip-like orange-red to scarlet flowers around 9 to 13 cm long and 7 to 8 cm across in erect clusters, mainly at the top of the tree's crown. Each flower has four pale yellow stamens 5 to 6 cm long with dark brown anthers. The flowers are subtended by a spathe-like calyx is covered with velvety rusty-brown hairs. The fruits are large erect green to dark brown pods around 13 to 25 cm long, 4 cm wide, and 2 cm thick, which point upwards at the ends of the branches. The oppositely arranged pinnate leaves are 30 to 60 cm long with usually 11 to 17 (sometimes 5 to 19) opposite elliptic 8 to 15 cm long leaflets on short stalks. The terminal inflorescenses (flower clusters) are around 10 to 25 cm long with the flowers clustered at the top. The fruit is an oblong capsule, 16 to 24 X 3.5 to 6 cm, which splits open to release numerous winged seeds, 2 to 2.5 cm long (including the membranous wings). The tree may reach 15 to 25 m in height and 30 to 45 cm in trunk diameter, with a dense irregular crown of large spreading branches that are evergreen or nearly deciduous. (Little and Wadsworth 1964; Whistler 1995)

The inflorescence of Spathodea campanulata includes around 40 to 50 flowers with acropetal maturation (i.e., the terminal flower is usually the first to mature, while the others tend to mature starting from the bottom of the stem). The showy orange ornithophilous (bird-pollinated) flowers have a flower fairly typical of the family Bignoniaceae, with a zygomorphic (bilaterally symmetrical) corolla, four didynamous stamens (i.e., 4 stamens in 2 pairs of unequal length), and a bicarpellate pistil with a 7.0 to 7.5 cm long style terminating in an exserted, bilamellate, touch-sensitive stigma. The ovary contains around 1000 ovules. (Bittencourt et al. 2003)

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Reproduction

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The African Tuliptree is an obligate outcrosser and requires pollinators for pollen flow between conspecific trees (Rangaiah et al. (2004). Bittencourt et al. (2003) carried out controlled pollinations of Spathodea campanulata in Brazil. Cross-pollination yielded 55% fruit set, whereas self-pollination resulted in no fruit set. Through careful embryological and histological investigations, Bittencourt et al. showed that the majority of ovules in selfed pistils were penetrated and fertilized within 48 hours, so the factors causing self-incompatibility and failure to set fruit are late-acting. According to Rangaiah et al. (2004), the natural fruit set of S. campanulata is very low, but is at least somewhat compensated for by a large seed crop.

Spathodea campanulata has winged seeds that are wind dispersed (Meyer 2004). The fruit is a capsule and dehisces naturally when mature, releasing small, light and winged seeds into the air. Seeds are dispersed efficiently by wind during the dry season. (Rangaiah et al. (2004)

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Uses

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Spathodea campanulata is used by humans mainly as a striking ornamental. Unopened flower buds contain water (foul smelling and tasting) which squirts out when the buds are squeezed or pricked and children play with them like water pistols. (Little and Wadsworth 1964)

According to Orwa et al. (2009), the seeds are edible and are used in many parts of Africa, but the hard central portion of the fruit is poisonous and used to kill animals.

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Derivation of specific name

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campanulata: campanulate, referring to the shape of the corolla.
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Hyde, M.A., Wursten, B.T. and Ballings, P. (2002-2014). Spathodea campanulata P. Beauv. Flora of Zimbabwe website. Accessed 28 August 2014 at http://www.zimbabweflora.co.zw/speciesdata/species.php?species_id=164250
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Description

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Tree, to 20 m. Leaves 1-pinnate, imparipinnate, opposite, large (to 50 cm long); leaflets ovate to elliptic. Flowers orange to red. Calyx spathaceous, densely hairy. Corolla lobes ± undulate. Fruit a capsule, 15 - 30 cm, fusiform, held erect. Seeds winged.
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Hyde, M.A., Wursten, B.T. and Ballings, P. (2002-2014). Spathodea campanulata P. Beauv. Flora of Zimbabwe website. Accessed 28 August 2014 at http://www.zimbabweflora.co.zw/speciesdata/species.php?species_id=164250
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Frequency

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Common
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Hyde, M.A., Wursten, B.T. and Ballings, P. (2002-2014). Spathodea campanulata P. Beauv. Flora of Zimbabwe website. Accessed 28 August 2014 at http://www.zimbabweflora.co.zw/speciesdata/species.php?species_id=164250
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Worldwide distribution

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Tropical Africa
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Hyde, M.A., Wursten, B.T. and Ballings, P. (2002-2014). Spathodea campanulata P. Beauv. Flora of Zimbabwe website. Accessed 28 August 2014 at http://www.zimbabweflora.co.zw/speciesdata/species.php?species_id=164250
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Spathodea campanulata

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Spathodea is a genus in the plant family Bignoniaceae. The single species it contains, Spathodea campanulata, is commonly known as the African tulip tree.[1] The tree grows between 7–25 m (23–82 ft) tall and is native to tropical dry forests of Africa. It has been nominated as among 100 of the "World's Worst" invaders.[2]

This tree is planted extensively as an ornamental tree throughout the tropics and is much appreciated for its very showy reddish-orange or crimson (rarely yellow), campanulate flowers. The generic name comes from the Ancient Greek words σπαθη (spathe) and οιδα (oida),[3] referring to the spathe-like calyx.[4] It was identified by Europeans in 1787 on the Gold Coast of Africa.[5]

Description

Nandi flame tree in Kerio Valley

The flower bud is ampule-shaped and contains water. People sometimes play with these buds because of their ability to squirt water.[6] The sap sometimes stains yellow on fingers and clothes. The open flowers are cup-shaped and hold rain and dew, making them attractive to many species of birds.

Flower anatomy

The African tulip tree flower produces large flamboyant reddish-orange flowers that have approximately five petals and are 8-15 cm long. The flowers are bisexual and zygomorphic. These are displayed in a terminal corymb-like raceme inflorescence. Its pedicel is approximately 6 cm long. This flower also has a yellow margin and throat. The pistil can be found at center of four stamens that is inserted on the corolla tube. This flower has a slender ovary that is superior and is two celled. The seeds of this tree are flat, thin, and broadly winged.

Species associations

In Neotropical gardens and parks, their nectar is popular with many hummingbirds, such as the black-throated mango (Anthracothorax nigricollis), the black jacobin (Florisuga fusca), or the gilded hummingbird (Hylocharis chrysura).[7] The wood of the tree is soft and is used for nesting by many hole-building birds such as barbets. [8] Unfortunately the flowers have a natural defence killing bees, and it is thought various other species who harvest its pollen.

Geographic distribution

It has become an invasive species in many tropical areas, such as Hawaii, Queensland (Australia), Fiji, Papua New Guinea, South Africa[10] and the wet and intermediate zones of Sri Lanka.[11]

Spathodea campanulata is a declared class 3 pest species in Queensland, Australia, under the Land Protection (Pest and Stock Route Management) Act 2002.[12] It is known to be toxic to Australian native stingless bees, such as Lipotriches (Austronomia) flavoviridis.[13]

Pests and diseases

In Uganda, two lepidopteran species, two termite species, and one bark beetle attack S. campanulata. In Puerto Rico nine insect species in the orders Hemiptera, Hymenoptera, Lepidoptera, and Thysanoptera have been reported as feeding on various parts of S. campanulata. The species is quite susceptible to butt and heart rot; wood of the tree rots quickly when in contact with the ground.

Use

Its wood is too weak for construction use but ideal enough for boxes and toothpicks. Fijian children sometimes use its flower buds as water pistols.[14]

Common names

  • Afrikaans: fakkelboom, Afrika-vlamboom
  • Bangla: rudrapalash (রুদ্রপলাশ)
  • English: African tulip tree, fountain tree, Nandi flame, Nile flame, squirt tree, tulip tree, Uganda flame, Flame Tree of Thika
  • Fijian: pasiu, pisipisi[14]
  • French: immortel étranger
  • Hindi: rugtoora (रुग्तूरा)
  • Kannada: Neerukayi mara, Jeerkolavi mara (ನೀರುಕಾಯಿ ಮರ, ಜೀರ್ಕೊೞವಿ ಮರ, ಜೀರ್ಕೊಳವಿ ಮರ)
  • Luganda: kifabakazi
  • Luhya: muzurio
  • Malay: panchut-panchut
  • Malayalam: African Poomaram (ആഫ്രിക്കൻ പൂമരം)
  • Sinhala: kudaella gaha, kudulu
  • Spanish: amapola, espatodea, mampolo, tulipán africano, in Puerto Rico meaíto, in Guatemala llama del bosque.
  • Swahili: kibobakasi, kifabakazi
  • Telugu: Neeti Budda (నీటి బుడ్డ), Gonuganta (గోనుగంట)
  • Tamil: pasadi (பாசடி)
  • Trade name: Nandi flame

Gallery

Footnotes

  1. ^ USDA, NRCS (n.d.). "Spathodea campanulata". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 24 November 2015.
  2. ^ "GISD".
  3. ^ Gledhill, D. (2008). The Names of Plants (4 ed.). Cambridge University Press. p. 357. ISBN 978-0-521-86645-3.
  4. ^ Quattrocchi, Umberto (2000). CRC World Dictionary of Plant Names. Vol. 4 R-Z. Taylor & Francis US. p. 2526. ISBN 978-0-8493-2678-3.
  5. ^ African Tulip Tropical Tree
  6. ^ Hargreaves, Dorothy; Hargreaves, Bob (1964). Tropical Trees of Hawaii. Kailua, Hawaii: Hargreaves. p. 2. ISBN 9780910690027.
  7. ^ Baza Mendonça & dos Anjos (2005)
  8. ^ de QUEIROZ, A. C. M., VENTURIERI, G., VENTURIERI, G., & de OLIVEIRA JUNIOR, M. C. M. (2012). Effect of toxicity of nectar and pollen of african tulip tree (Spathodea campanulata) on Melipona fasciculata and M. seminigra (Apidae, Meliponini). In Embrapa Amazônia Oriental-Resumo em anais de congresso (ALICE). In: ENCONTRO SOBRE ABELHAS, 10., 2012, Ribeirão Preto. Anais... Ribeirão Preto: FUNPEC, 2012
  9. ^ "ফুটেছে রুদ্রপলাশ | প্রথম আলো".
  10. ^ "African tulip tree – Invasive Species South Africa". Retrieved 2022-02-17.
  11. ^ Invasive Species Compendium and Lalith Gunasekera, Invasive Plants: A guide to the identification of the most invasive plants of Sri Lanka, Colombo 2009, pp. 70–71.
  12. ^ Land Protection (Pest and Stock Route) Regulation 2003 (Qld) - Schedule 2
  13. ^ O'Flaherty, Antonia (2021-02-17). "Brisbane beekeeper creates editable map to track African tulip trees killing native stingless bees". ABC News. Australian Broadcasting Corporation. Retrieved 2021-02-18.
  14. ^ a b Keppel, Gunnar; Ghazanfar, Shahina A. (2011). Trees of Fiji: A Guide to 100 Rainforest Trees (third, revised ed.). Secretariat of the Pacific Community & Deutsche Gesellschaft für Technische Zusammenarbeit. pp. 58–9.

References

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Spathodea campanulata: Brief Summary

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Spathodea is a genus in the plant family Bignoniaceae. The single species it contains, Spathodea campanulata, is commonly known as the African tulip tree. The tree grows between 7–25 m (23–82 ft) tall and is native to tropical dry forests of Africa. It has been nominated as among 100 of the "World's Worst" invaders.

This tree is planted extensively as an ornamental tree throughout the tropics and is much appreciated for its very showy reddish-orange or crimson (rarely yellow), campanulate flowers. The generic name comes from the Ancient Greek words σπαθη (spathe) and οιδα (oida), referring to the spathe-like calyx. It was identified by Europeans in 1787 on the Gold Coast of Africa.

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