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Publicado en: Phytoneuron 2012-33: 2. 2012. (Phytoneuron) Name publication detailView in Biodiversity Heritage Library
 

Datos del Proyecto Nombre (Last Modified On 12/8/2022)
Aceptación : Accepted
Nota : Subfam. Dialypetalanthoideae (formerly Ixoroideae)
Datos del Proyecto     (Last Modified On 7/8/2024)
Notas :

This tribe has long been treated under the name Condamineeae, but recent changes to the International Code of Nomenclature for plant names along with changes in the circumscription of both Rubiaceae and this tribe (Kainulainen et al., 2010) now require that the name Dialypetalantheae be used (Reveal, 2012). This change is mandated by Art. 19.5, first adopted in the Melbourne Congress in 2011, which extended the use of conserved family names to infrafamilial taxa that include the type of the conserved family name. As discussed below, Kainulainen et al. (2010) demonstrated that the morphologically enigmatic genus Dialypetalanthus does not represent a separate famliy but belongs to Rubiaceae, and wtihin this family it belongs to the tribe that was long called Condamineeae: when Dialypetalanthus is included in this group, this name has priority. (If Dialypetalanthus is excluded again at some point from this tribe's circumscription, then the tribe would again take the name Condamineeae). 

Tribe Dialypetalantheae, as circumscribed by Kainulainen et al. (2010; as Condamineeae), belongs to Subfamily Ixoroideae (which should now be called Subfamily Dialypetalanthoideae) and includes about 33 or 34 morphologically diverse genera with perhaps 150 species. Most of the genera are endemic to the Neotropics, but the tribe is also found in warm temperate Asia (Emmenopterys), tropical Southeast Asia (Mussaendopsis), warm temperate North America (Pinckneya), and the Pacific region (Dolicholobium, Mastixiodendron). Plants of Dialypetalantheae are woody, and range from monocaulous shrubs to large trees, some of them canopy emergents. Raphides are absent.Stipule form varies widely among the genera, and includes interpetiolar free stipules, calyptrate fused stipules, bilobed stipules, and stipules divided into intrapetiolar segments. The inflorescences are generally cymose and variously axillary and terminal. The flowers are mostly hermaphroditic though Dioicodendron is dioecous, and Dolicholobium is variously reported to be monoecious (Smith & Darwin 1988) or dioecious (Kainulainen et al. 2010). The hermaphroditic flowers are protandrous or in a few cases protogynous (Alseis, Chimarrhis, Warszewiczia), with this floral biology consistent at the genus level. Species of several genera have well developed calycophylls, and corolla lobe aestivation varies. he corollas are divided into free petals in Dialypetalanthus, Mastixiodendron, and Mussaendopsis, and the stamens are numerous in Dialypetalanthus. The fruits are mostly dry and capsular with numerous seeds that are variously angled to flattened and winged, but the fruits are are baccate, fleshy, and indehiscent in several genera (Bothriospora, Hippotis, Pentagonia, Pogonopus, Tammsia), and drupaceous with usually a single furrowed seed and internal air spaces in Lintersemina and Masitixiodendron. Overall as discussed by Kainulainen et al. the Tribe Dialypetalantheae is difficult to characterize morphologically.

The circumscription of Tribe Dialypetalantheae has been varied and problematic, not surprisingly given the morphological variation here, and some of its genera were classified quite differently by various authors until the analysis of Bremer & Eriksson (2009) found a clearly circumscribed group within the Subfamily Ixoroideae (i.e., Dialypetalantheae). This monophyletic Tribe Dialypetalantheae was then studied in more detail by Kainulainen et al. (2010). Before these studies, the classification of the genera now included in this tribe was based on selected morphological features so these were mostly separated in various tribes (e.g., Delprete, 1999). Andersson (1995) studied morphologically the capsular-fruited genera that had been included in tribe Cinchoneae, and he separated some of the into a newly described Tribe Calycophylleae. Rova & Andersson (1995) also found support for separation of their Tribe Hippotideae. Molecular data eventually clarified the systematics of this group, and Bremer & Eriksson showed that Calycophylleae and Condamineae form a monophyletic group in Subfam. Ixoroideae (i.e., Dialypetalanthoideae) and then Kainulainen et al. (2010; as Condamineeae) found the Hippotideae group included within Dialypetalantheae. Although the Tribe Dialypetalantheae is now rather clearly circumscribed, the delimitations and relationships of the component genera are far from understood. Kainulainen et al. (2010) sampled potential member genera of the tribe well but studied relatively few species of many of them. They found  several genera that were clearly not monophyletic, but their analysis adequately support only three taxonomic rearrangements: the resurrection of a monospecific genus, the synonymization of another monospecific genus, and the transfer of six species from one genus to another, newly separated genus. Their study found several additional genera or groups that need taxonomic review, and many of the well circumscribed genera remain to be studied in detail. Most of the genera of Tribe Dialypetalantheae have never been comprehensively reviewed, and morphological diversification within this tribe is still being discovered and detailed. Recently Mendoza-Cifuentes et al. (2020) added a genus with novel morphology for the Rubiaceae to this tribe, and presented a comparative table of principal morphological character of all the genera of the tribe.

One of the morphologically oddest genera of Rubiaceae is Dialypetalanthus, which was long included in its own family, Dialypetalanthaceae, and of unclear taxonomic affinity. It was variously included in Rosales, Myrtales, Cornales, and a narrowed circumscription of Gentianales that excluded Rubiaceae, but now clearly is placed in Rubiaceae by molecular (Kainulainen et al., 2010) and floral ontogeny (Vrijdaghs et al., 2022). Dialypetalanthus is anomalous in Rubiaceae in its pubescence with some stellate or dendritic trichomes, petals that are secondarily free and imbricated with usually (though not always) one external and apparently one internal, fully developed flowers with 16-25 stamens that are fused at their bases into a ring and held erect in a tube-like arrangement, quite large poricidal anthers, and the absence of a nectary disk on top of the ovary. For more details on this genus see its genus page. 

As noted above, the Dialypetalantheae genera are unusual in their wide morphological variation in morphological features that often characterize a tribe of Rubiaceae. The key below treats only the capsular-fruited Neotropical genera, which have been the most widely confused of this group as to their circumscriptions and diagnostic features. Because of the morphological variation in this group, most of the genera are separated by a combination of vegetative, flowering, and fruiting characters. The key below summarizes these characters; a key that uses only flowers or only fruits is, by necessity, a key to individual species rather than an outline of generic separations. 

Author: C.M. Taylor The content of this web page was last revised on 8 July 2024.
Taylor web page: http://www.mobot.org/MOBOT/Research/curators/taylor.shtml

Distribución : Humid and seasonal woody vegetation mainly in the Neotropics, also in the tropical Pacific Islands and warm temperate eastern Asia and southeastern North America.
Referencias :
Taxa Included Here :

Genera of Dialypetalantheae, sensu Kainulainen et al. (2010) and Mendoza-Cifuentes et al. (2020):
Alseis Schott
Bathysa C. Presl
Bothriospora Hook.f.
Calycophyllum DC.
Capirona Spruce
Chimarrhis Jacq.
Condaminea DC.
Dialypetalanthus Kuhlm.
Dioicodendron Steyerm.
Dolichodelphys K. Schum. & K. Krause
Dolicholobium A. Gray, Southeast Asia & Pacific Region
Elaeagia Wedd.
Emmenopterys Oliv.
Ferdinandusa Pohl
Hippotis Ruiz & Pav.
Holtonia Standl.
Lintersemina H. Mend., Á. Celis & M. Gonzalez
Macbrideina Standl.
Macrocnemum P. Browne
Mastixiodendron Melch., Pacific Islands
Mussaendopsis Baill., Southeast Asia
Parachimarrhis Ducke
Pentagonia Benth.
Picardaea Urb.
Pinckneya Michx.
Pogonopus Klotzsch
Rustia Klotzsch
Schizocalyx Wedd.
Semaphyllanthe L. Andersson = Calycophyllum DC.
Simira Aubl.
Sommera Schltdl.
Tammsia H. Karst.
Tresanthera Urb. = Rustia
Warszewiczia Klotzsch
Wittmackanthus Kuntze


 

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Key to Capsular-Fruited Genera of Dialypetalantheae in the Neotropics
(Note: some genera are diagnosed by a combination of flower and fruit characters)

1. Inflorescences axillary.

   2. Corolla lobes thinly imbricated in bud; capsules septicidal; seeds suborbicular, 0.3--1 mm long..... Chimarrhis Jacq.

   2'. Corolla lobes convolute in bud; capsules loculidal through the apical portion; seeds cylindrical to navicular, 22--26 mm long..... Lintersemina H. Mend., Á Celis, & M. Gonzalez

1'. Inflorescences terminal and sometimes also in axils of uppermost leaves.

   3. Flowers bisexual and protogynous..... Warszewiczia Klotzsch

   3'. Flowers bisexual and protandrous, or unisexual on dioecious plants.

        4. Scandent or clambering plants; stipules interpetiolar and rounded; plants dioecious with unisexual flowers.....Dioicodendron Steyerm.

       4'. Erect shrubs and trees; stipules rounded to acute, variously interpetiolar, intrapetiolar, or calyptrate; plants with bisexual flowers.

           5. Stipules fused at base into a tube, with their upper parts splitting into two intrapetiolar segments or two bilobed interpetiolar segments; capsules loculicidal.

               6. Trichomes of mixed form, with some simple and some stellate or dendritic; corolla divided to base into secondarily free lobes or petals; stamens 16--25..... Dialypetalanthus Kuhlm.

               6'. Trichomes all simple; corolla fused in basal portion into a tube; stamens 5.

                    7. Leaves without domatia or with domatia present only along costa; corollas with the lobes half or more as long as the tube and widely spreading at anthesis; capsules dehiscing in a short apical slit through the beak portion..... Elaeagia Wedd.

                    7'. Leaves with regularly developed domatia along costa and usually also in axils of tertiary veins; corollas with the lobes a third as long as the tube or shorter and only shortly spreading at anthesis; capsules dehiscing from top through main body of the fruit into two valves..... Holtonia Standl.

               5'. Stipules free or shortly fused at the base with their upper parts interpetiolar and unlobed, or fused completely into a calyptrate cap that splits along one side or irregularly; capsules loculicidal or septicidal.

                      8. Capsules septicidal. 9. Capsules subglobose; seeds angled to discoid, sometimes with thin margins but not evidently winged..... Bathysa C. Presl

                            9'. Capsules ellipsoid to obovoid or subglobose; seeds fusiform to ellipsoid and flattened, with marginal wing.

                                 10. Calycophylls absent; corolla lobes in bud convolute and overlapping for 1/3 or more of their width, at anthesis becoming emarginate to bilobed at tip..... Ferdinandusa Pohl

                                 10'. Calycophylls absent or present; corolla lobes in bud imbricated or thinly convolute and overlapping for at most 1/10th of their width, at anthesis acute to obtuse at tip.

                                       11. Stipules interpetiolar and free, or fused completely into a calyptrate cap; corolla white to cream, internally pubescent in upper part of tube; anthers basifixed; calyx limb persistent on capsule..... Calycophyllum DC. (including Semaphyllanthe L. Andersson)

                                       11'. Stipules interpetiolar and free or shortly fused at base; corolla pink, lilac, or purple, internally glabrous throughout or pubescent in lower part of tube; anthers dorsifixed; calyx limb deciduous from capsule..... Wittmackanthus Kuntze

                      8'. Capsules loculicidal. 12. Capsules dehiscent along side margins, with valves remaining fused at top; corolla lobes reduplicate-valvate in bud..... Macrocnemum P. Browne

                         12'. Capsules dehiscent from apex into two valves or only through apical disk portion; corolla lobes imbricated, convolute, valvate, or perhaps open in bud.

                              13. Corolla red to purple, with lobes valvate in bud; Greater Antilles..... Picardaea Urb.

                              13'. Corolla white, cream, or green and sometimes marked with dull purple, with lobes imbricated, convolute, or perhaps open in bud; Mexico, Central America, and South America.

                                      14. Stipules interpetiolar; corolla lobes imbricated or open in bud; capsule subglobose; seeds flattened and elliptic, with well developed wing.

                                            15. Internal tissues of stems and fruits not oxidizing purple; capsules 0.5--1 cm in diameter; trees of inundated forest along rivers in the Amazon basin..... Parachimarrhis Ducke

                                             15'. Internal tissues of stems and fruits oxidixing purple; at least some capsules 1.1--9 cm in diameter; trees and shrubs of various habitats from Mexico to southern Brazil..... Simira Aubl.

                                      14' Stipules interpetiolar or fused into a calyptrate cap; corolla lobes convolute in bud; capsule subglobose to ellipsoid or obovoid; seeds angled to or flattened and fusiform, unwinged or with irregularly developed narrow wings.

                                              16. Stipules interpetiolar..... Macbrideina Standl.

                                              16'. Stipules fused into a calyptrate cap..... Schizocalyx Wedd.

 
 
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