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''Retrophyllum'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Podocarpaceae Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber Pre ...
.Christopher N. Page. 1990. "Podocarpaceae" pages 332-346. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and Genera of Vascular Plants'' volume I. Springer-Verlag: Berlin;Heidelberg, Germany. It contains five generally recognized extant species with a
disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
in the Southern Hemisphere, found in
Papuasia Papuasia is a Level 2 botanical region defined in the World Geographical Scheme for Recording Plant Distributions (WGSRPD). It lies in the southwestern Pacific Ocean, in the Melanesia ecoregion of Oceania and Tropical Asia. It comprises the ...
and also in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
.Kew World Checklist of Selected Plant Families
/ref>James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. . ''Retrophyllum'' are evergreen trees typically occurring in tropical
rainforests Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainforest ...
and
cloud forests A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud ...
.


Etymology

The name ''Retrophyllum'' is derived from the Latin ''retro'', meaning "backward" or "reversed", and the Greek ''phyllos'', meaning "leaf". The name refers to the unique phyllotaxis where the adaxial surfaces of the leaves face up on one side of the shoot and down on the other.


Description

''Retrophyllum'' are
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s. They range in size from dwarfed to very large, reaching heights in excess of 40 and potentially 60 meters.
Resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
canals are found in both leaves and the seed cones. The bark is usually smooth at first, becoming fissured or flaking with age. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are generally flat with a decurrent base and a spreading blade, but leading and cone-bearing shoots may also have small appressed scale-like leaves. The base
phyllotaxis In botany, phyllotaxis () or phyllotaxy is the arrangement of leaf, leaves on a plant stem. Phyllotactic spirals form a distinctive class of patterns in nature. Leaf arrangement The basic leaf#Arrangement on the stem, arrangements of leaves ...
or leaf arrangement is spiral though the leaves usually form subopposite and nearly decussate pairs. The leaves of a lateral shoot are further twisted at their petioles to form two pectinate rows in a horizontal plane around the shoot. The leaf petioles in ''Retrophyllum'' are uniquely twisted on the lateral shoots in opposite directions on each side of a shoot orienting the leaf blades with the adaxial or ventral surface upwards on one side of the shoot and the abaxial or dorsal surface upwards on the opposite side of the shoot. The leaf blade varies in shape from lanceolate to narrowly ovate. The leaves have conspicuous midribs and are amphistomatic with stomata present on both sides. ''Retrophyllum'' are
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
with male pollen cones and female seed cones on separate individual trees. The male pollen cones may be axillary or terminal and solitary or grouped. They have glabrous peduncles. A pollen cone consists of many spirally arranged microsporophylls each with two pollen sacs producing bisaccate
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
. The female
seed cones A conifer cone (in formal botanical usage: strobilus, plural strobili) is a seed-bearing organ on gymnosperm plants. It is usually woody, ovoid to globular, including scales and bracts arranged around a central axis, especially in conifers an ...
develop from axillary buds. They are often solitary but may also be paired. The cones consist of several basal sterile cone scales and a single apical fertile scale. The basal scales are fused. The fertile scale has one
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
producing
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
. The single seed of the cone is covered by a modified ovuliferous scale known as the epimatium. The epimatium becomes fleshy and drupe-like at maturity. It varies in shape from elliptic to ovoid or pyriform and may be red, violet or purplish brown in color.


Distribution

''Retrophyllum'' has a naturally disjunct distribution covering the
Maluku Islands The Maluku Islands (; Indonesian: ''Kepulauan Maluku'') or the Moluccas () are an archipelago in the east of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located ...
of
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 ...
,
New Guinea New Guinea (; Hiri Motu Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea). It is a simplified version of ...
, New Britain and New Ireland in the Asia-Pacific region,
Fiji Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists ...
,
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
and
Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (french: link=no, République de Vanuatu; bi, Ripablik blong Vanuatu), is an island country located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of no ...
in the Pacific and parts of
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
,
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
,
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...
,
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
and
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
in South America. The species ''
Retrophyllum minus ''Retrophyllum minus'', the ''bois bouchon'', is a species of conifer in the family Podocarpaceae. It is endemic to New Caledonia. It is a sparsely branched small tree with a flared trunk which tapers towards the top and has rough bark. It pr ...
'' occurs in riparian and lacustrine habitats on
ultramafic Ultramafic rocks (also referred to as ultrabasic rocks, although the terms are not wholly equivalent) are igneous and meta-igneous rocks with a very low silica content (less than 45%), generally >18% MgO, high FeO, low potassium, and are composed ...
soils in New Caledonia. The other species usually grow in tropical lowland or montane rainforests or cloud forests.


Fossil record

''Retrophyllum'' shoots have a distinctive morphology in which the leaves are in subopposite pairs, and twisted in such a way that the abaxial surface of one leaf is up, and in the other it is down. This feature, added to a distinctive epidermal morphology means that well-preserved specimens can be easily identified in the fossil record. The fossil record shows that ''Retrophyllum'' was present in the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
.Pole, M.S., 1997. Miocene conifers from the Manuherikia Group, New Zealand. Journal of the Royal Society of New Zealand, 27: 355-370. It is now extinct in those places.


Taxonomy

In a 2009 book on conifers, ''Retrophyllum piresii'' was treated as
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...
with ''Retrophyllum rospigliosii''. The author stated "One specimen from low elevations in Brazil was separated as a distinct species, but it falls well within the range of variation of the species as a whole and is geographically close to some locations in Peru."


Species

There are five generally recognized species.


References


External links


Retrophyllum
At

At
The Gymnosperm Database
{{Taxonbar, from=Q138257 Podocarpaceae Podocarpaceae genera Dioecious plants