The family Phylliidae (often misspelled Phyllidae) contains the
extant
Extant is the opposite of the word extinct. It may refer to:
* Extant hereditary titles
* Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English
* Extant taxon, a taxon which is not extinct, ...
true leaf insects or walking leaves, which include some of the most remarkably
camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d leaf
mimics (mimesis) in the entire animal kingdom. They occur from
South Asia
South Asia is the southern subregion of Asia, which is defined in both geographical and ethno-cultural terms. The region consists of the countries of Afghanistan, Bangladesh, Bhutan, India, Maldives, Nepal, Pakistan, and Sri Lanka.;;;;; ...
through
Southeast Asia
Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
to
Australia. Earlier sources treat Phylliidae as a much larger taxon, containing genera in what are presently considered to be several different families.
Characteristics
Leaf insects are
camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d, taking on the appearance of leaves. They do this so accurately that predators often are not able to distinguish them from real leaves. In some species, the edge of the leaf insect's body has the appearance of bite marks. To further confuse predators, when the leaf insect walks, it rocks back and forth, mimicking a real leaf being blown by the wind.

The scholar
Antonio Pigafetta
Antonio Pigafetta (; – c. 1531) was an Venetian scholar and explorer. He joined the expedition to the Spice Islands led by explorer Ferdinand Magellan under the flag of the emperor Charles V and after Magellan's death in the Philippine Islands, ...
probably was the first Western person to document the creature, though it was known to people in the tropics for a long time. Sailing with
Ferdinand Magellan
Ferdinand Magellan ( or ; pt, Fernão de Magalhães, ; es, link=no, Fernando de Magallanes, ; 4 February 1480 – 27 April 1521) was a Portuguese explorer. He is best known for having planned and led the 1519 Spanish expedition to the Eas ...
's
circumnavigation
Circumnavigation is the complete navigation around an entire island, continent, or astronomical body (e.g. a planet or moon). This article focuses on the circumnavigation of Earth.
The first recorded circumnavigation of the Earth was the Magel ...
al expedition, he studied and chronicled the
fauna
Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as ''biota''. Zoo ...
on the
island
An island or isle is a piece of subcontinental land completely surrounded by water. Very small islands such as emergent land features on atolls can be called islets, skerries, cays or keys. An island in a river or a lake island may be ...
of
Cimbonbon
Balabac Island is the southernmost island of the Palawan province, and therefore the westernmost undisputed island in the Philippines, only about north from Sabah, Malaysia, across the Balabac Strait.
Administratively, the island forms the ...
as the fleet hauled ashore for repairs. During this time he documented the ''Phyllium'' species with the following passage:
Tribes, genera and species
The subfamily Phylliinae has been divided into two
tribes
The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
since 2003. This classification is not confirmed by more recent
molecular genetics
Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
investigations. In addition to the fossil
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
''Eophyllium'', the subfamily distinguishes thirteen recent genera, eight of which have been described since 2017.
Within the genus ''Phyllium'', the subgenus ''
Pulchriphyllium'', set up by
Achille Griffini in 1898, has been distinguished from ''Phyllium'' since 1898.
With ''
Comptaphyllium'' and ''
Walaphyllium'' two more subgenera became in 2019 and 2020.
As of 2021, all three subgenera are considered separate genera.
Since 2021, in addition to
morphological, molecular genetic studies have also increasingly been included in clarification of the
phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of Phylliidae. Their results show the general relationship between the genera, but when comparing female and male representatives, they do not yet provide a clear phylogenetic picture of the recent genera.
Cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
s of the Phylliidae species determined on the basis of molecular genetics analysis and morphological investigations according to Cumming and Le Tirant (2022):
The ''Phasmida Species File (V. 5.0)'' lists the following genera in two
tribes
The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...
:
Phylliini
Auth.
* ''
Chitoniscus'' (
Pacific
The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
)
** ''
Chitoniscus feejeeanus''
** ''
Chitoniscus lobipes''
** ''
Chitoniscus lobiventris'' -
type species
In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
(as ''Phyllium lobiventre'' )
* ''
Comptaphyllium'' (
Australasia
Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecolo ...
)

** ''
Comptaphyllium caudatum'' - type species (as ''Phyllium caudatum'' )
** ''
Comptaphyllium regina''
** ''
Comptaphyllium riedeli''
* ''
Cryptophyllium'' (
SE Asia
Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
).
Selected species:
** ''
Cryptophyllium athanysus''
** ''
Cryptophyllium celebicum
''Cryptophyllium celebicum''Haan W de (1842) In Temminck d. ''Verhandelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen'' 3: 111. is type species of leaf insect in the new (2021) genus ''Cryptophyllium'',Cumming R ...
'' - type species (as ''Phyllium celebicum'' )
** ''
Cryptophyllium westwoodii
''Cryptophyllium westwoodii''Cumming RT, Bank S, Bresseel J, Constant J, Le Tirant S, Dong Z, Soner G, Bradler S. (2021) ZooKeys 1018: 14link on www.researchgate.net/ref> is a species of leaf insect in the family Phylliidae. It is distributed f ...
''
* ''
Microphyllium'' (Northern
Philippine Islands
The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no),
* bik, Republika kan Filipinas
* ceb, Republika sa Pilipinas
* cbk, República de Filipinas
* hil, Republ ...
)
** ''
Microphyllium haskelli''
** ''
Microphyllium spinithorax'' - type species

* ''
Phyllium
''Phyllium''Illiger (1798) ''Verzeichnis der Käfer Preussens'' 499. is the largest and most widespread genus of leaf insects in the family Phylliidae (Phasmatodea). They can be found in Sundaland, Philippine Islands, Wallacea, and Australasi ...
'' (
Sundaland
Sundaland (also called Sundaica or the Sundaic region) is a biogeographical region of South-eastern Asia corresponding to a larger landmass that was exposed throughout the last 2.6 million years during periods when sea levels were lower. It ...
, Philippine Islands,
Wallacea
Wallacea is a biogeographical designation for a group of mainly Indonesian islands separated by deep-water straits from the Asian and Australian continental shelves. Wallacea includes Sulawesi, the largest island in the group, as well as ...
, Australasia).
Selected species:
** ''
Phyllium bilobatum
''Phyllium bilobatum'' is a species of leaf insect in the family Phylliidae. It is found in the Philippines and Malaysia. This species was first described in 1843 by the English zoologist George Robert Gray, who gave it the name ''Phyllium bil ...
''
** ''
Phyllium hausleithneri
''Phyllium hausleithneri'' or Hausleithner's stick insect is a species of phasmid or leaf insect of the genus '' Phyllium''. It is found in peninsular Malaysia, and Sri Lanka
Sri Lanka (, ; si, ශ්රී ලංකා, Śrī La ...
''
** ''
Phyllium jacobsoni''
** ''
Phyllium letiranti
''Phyllium letiranti'' is a species of phasmid or walking leaf of the genus '' Phyllium''. It is only found on the island of Peleng in Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceani ...
''
** ''
Phyllium siccifolium
''Phyllium''Illiger (1798) ''Verzeichnis der Käfer Preussens'' 499. is the largest and most widespread genus of leaf insects in the family Phylliidae (Phasmatodea). They can be found in Sundaland, Philippine Islands, Wallacea, and Australas ...
'' - type species (as ''Gryllus siccifolius'' )
* ''
Pseudomicrophyllium'' (Northern Philippine Islands)
** ''
Pseudomicrophyllium geryon
''Pseudomicrophyllium geryon'' is a species of phasmid or leaf insect. It is found in the Philippines, Java (Indonesia), and Sri Lanka
Sri Lanka (, ; si, ශ්රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இ� ...
''
** ''
Pseudomicrophyllium pusillulum'' - type species (as ''Pseudomicrophyllium faulkneri'' )
* ''
Pulchriphyllium'' (
Seychelles
Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
,
India
India, officially the Republic of India ( Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the ...
, Western
Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
, continental
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an ...
)
Selected species:
** ''
Pulchriphyllium bioculatum
''Pulchriphyllium bioculatum'', Seychelles leaf insect, Javanese leaf insect, or Gray's leaf insect, is a leaf insect of the family Phylliidae native to tropical Asia as well as Madagascar, Mauritius and the Seychelles. It was first described by ...
'' (
** ''
Pulchriphyllium giganteum'' (
** ''
Pulchriphyllium pulchrifolium'' - type species (as ''Phyllium pulchrifolium'' )
* ''
Rakaphyllium'' (
New Guinea
New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...
and
Ayu Islands
The Ayu Islands (Kepulauan Ayu), are a small archipelago south of the Asia Islands and north of Raja Ampat Islands in Indonesia. This group is formed by two coral atolls. The highest point is 106 m.Pub164, 2004 Sailing Directions (Enroute): New Gu ...
)
** ''
Rakaphyllium exsectum''
** ''
Rakaphyllium schultzei'' – type species (as ''Pulchriphyllium schultzei'' )
* ''
Trolicaphyllium'' (Pacific)

** ''
Trolicaphyllium brachysoma'' - type species (as ''Phyllium brachysoma'' )
** ''
Trolicaphyllium erosus''
** ''
Trolicaphyllium sarrameaense''
* ''
Vaabonbonphyllium'' (New Guinea and
Solomon Islands
Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania, to the east of Papua New Guinea and north-west of Vanuatu. It has a land area of , and a population of approx. 700,000. Its ca ...
)
** ''
Vaabonbonphyllium groesseri'' ( – type species (as ''Phyllium groesseri'' )
** ''
Vaabonbonphyllium rafidahae''
* ''
Walaphyllium'' (Australasia)
** ''
Walaphyllium lelantos''
** ''
Walaphyllium monteithi''
** ''
Walaphyllium zomproi'' - type species (as ''Phyllium zomproi'' )
Nanophylliini

Auth.
* ''
Acentetaphyllium'' (New Guinea)
** ''
Acentetaphyllium brevipenne'' – type species (as ''Phyllium brevipennis'' )
** ''
Acentetaphyllium larssoni''
** ''
Acentetaphyllium miyashitai''
** ''
Acentetaphyllium stellae''
* ''
Nanophyllium'' (Southern Indonesia, New Guinea, NE Australia)
** ''
Nanophyllium adisi''
** ''
Nanophyllium asekiense''
** ''
Nanophyllium australianum''
** ''
Nanophyllium chitoniscoides ''
** ''
Nanophyllium daphne''
** ''
Nanophyllium frondosum''
** ''
Nanophyllium hasenpuschi''
** ''
Nanophyllium keyicum''
** ''
Nanophyllium pygmaeum'' – type species
** ''
Nanophyllium rentzi''
** ''
Nanophyllium suzukii''
Captivity
Several species have gained in popularity as
pet
A pet, or companion animal, is an animal kept primarily for a person's company or entertainment rather than as a working animal, livestock, or a laboratory animal. Popular pets are often considered to have attractive appearances, intelligence ...
s including ''
Cryptophyllium celebicum
''Cryptophyllium celebicum''Haan W de (1842) In Temminck d. ''Verhandelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen'' 3: 111. is type species of leaf insect in the new (2021) genus ''Cryptophyllium'',Cumming R ...
'', ''
Cryptophyllium westwoodii
''Cryptophyllium westwoodii''Cumming RT, Bank S, Bresseel J, Constant J, Le Tirant S, Dong Z, Soner G, Bradler S. (2021) ZooKeys 1018: 14link on www.researchgate.net/ref> is a species of leaf insect in the family Phylliidae. It is distributed f ...
'', ''
Phyllium jacobsoni'', ''
Phyllium ericoriai
''Phyllium ericoriai'' is a species of leaf insects
The family Phylliidae (often misspelled Phyllidae) contains the extant true leaf insects or walking leaves, which include some of the most remarkably camouflaged leaf mimics (mimesis) in t ...
'', ''
Phyllium siccifolium
''Phyllium''Illiger (1798) ''Verzeichnis der Käfer Preussens'' 499. is the largest and most widespread genus of leaf insects in the family Phylliidae (Phasmatodea). They can be found in Sundaland, Philippine Islands, Wallacea, and Australas ...
'', ''
Phyllium letiranti
''Phyllium letiranti'' is a species of phasmid or walking leaf of the genus '' Phyllium''. It is only found on the island of Peleng in Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceani ...
'', ''
Phyllium monteithi
''Phyllium monteithi'' is a species of phasmatodea in the family Phylliidae (leaf insect). ''P. monteithi'' is found in tropical Queensland in Australia. The type specimen was collected from Mount Lewis, near Julatten. Reproduction occurs throug ...
'', ''
Phyllium philippinicum '', ''
Phyllium rubrum
''Phyllium''Illiger (1798) ''Verzeichnis der Käfer Preussens'' 499. is the largest and most widespread genus of leaf insects in the family Phylliidae (Phasmatodea). They can be found in Sundaland, Philippine Islands, Wallacea, and Australas ...
'', ''
Phyllium tobeloense
''Phyllium''Illiger (1798) ''Verzeichnis der Käfer Preussens'' 499. is the largest and most widespread genus of leaf insects in the family Phylliidae (Phasmatodea). They can be found in Sundaland, Philippine Islands, Wallacea, and Australas ...
'', ''
Pulchriphyllium bioculatum
''Pulchriphyllium bioculatum'', Seychelles leaf insect, Javanese leaf insect, or Gray's leaf insect, is a leaf insect of the family Phylliidae native to tropical Asia as well as Madagascar, Mauritius and the Seychelles. It was first described by ...
'' and ''
Pulchriphyllium giganteum'' .
Extinct species
A 47-million-year-old fossil of ''
Eophyllium messelensis'', a prehistoric ancestor of Phylliidae, displays many of the same characteristics of modern leaf insects, indicating that this family has changed little over time.
References
[ Cumming, R. T.; Bank, S.; Bresseel, J.; Constant, J; Le Tirant, S.; Dong, Z.; Sonet, G. & Bradler, S.: ]
Cryptophyllium, the hidden leaf insects – descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae)
'ZooKeys 1018: 1–179 (2021)
[Cumming, R. T.; Le Tirant, S. & Büscher, T. H. (2021) ''Resolving a century-old case of generic mistaken identity: polyphyly of Chitoniscus sensu lato resolved with the description of the endemic New Caledonia ''Trolicaphyllium'' gen. nov. (Phasmatodea, Phylliidae)'', ZooKeys 1055: 1–41 (2021), ]
[Cumming, R. T.; Teemsma, S. N. & Valero, P. (2018) ]
Description of Phyllium (Phyllium) conlei, new species, and a first look at the Phylliidae (Phasmatodea) of the Lesser Sunda Islands, Indonesia
'. Insecta mundi. Center for Systematic Entomology, Inc., Gainesville, FL USA. 2018
[Cumming, R. T.; Le Tirant, S. & Hennemann F. H. (2019) ]
Review of the Phyllium Illiger, 1798 of Wallacea, with description of a new subspecies from Morotai Island (Phasmatodea: Phylliidae: Phylliinae)
', Faunitaxys, 7(4), Saint-Etienne, 2019: 1 – 25.
[Cumming, R. T.; Thurman, J. H.; Youngdale, S. & Le Tirant, S. (2020) ]
Walaphyllium subgen. nov., the dancing leaf insects from Australia and Papua New Guinea with description of a new species (Phasmatodea, Phylliidae)
' ZooKeys 939:5 (2020)
[Bank, S.; Cumming, R. T.; Li, Y.; Henze, K.; Le Tirant, S. & Bradler, S. (2021]
A tree of leaves: Phylogeny and historical biogeography of the leaf insects (Phasmatodea: Phylliidae)
/ref>
[Cumming, R. T. & Le Tirant, S. (2022) ''Three new genera and one new species of leaf insect from Melanesia (Phasmatodea, Phylliidae)'', ZooKeys 1110: 151–200 (2022), doi:10.3897/zookeys.1110.80808]
External links
*
Phasmid Study Group: Phylliidae
{{Taxonbar, from1=Q282198, from2=Q15240868
Phasmatodea families
Taxa named by Ludwig Redtenbacher
pl:Liściec