A Monograph of Conostegia (Melastomataceae, Miconieae)

Abstract A recent molecular phylogenetic analysis identified a clade containing all species of Conostegia, but that also included species of Clidemia and Miconia nested inside. A taxonomic revision of a more broadly circumscribed Conostegia is presented here. In total, 77 species of Conostegia are recognized. One species from Ecuador, Conostegia ortizae is described as new. Twenty-nine new combinations are proposed for the species of Clidemia and Miconia that fall inside Conostegia. Two new names are proposed for the two species for which the epithet was previously occupied in Conostegia. An infrageneric classification of Conostegia is proposed recognizing three sections based on the results of the molecular phylogeny. This taxonomic revision includes ample documentation of the anatomy and morphology of most species in the genus, taxonomic descriptions, a dichotomous key, and distribution maps for all species.


Introduction
Conostegia D. Don, a genus in the tribe Miconieae (Melastomataceae), is most famous for the calyptrate calyx of the fl owers of its species. Th e group was revised by Schnell (1996) who recognized 42 species, but this thorough revision unfortunately was never published. A recent molecular phylogenetic analysis based on DNA sequences from four chloroplast and two nuclear ribosomal spacers found Conostegia not to be monophyletic but identifi ed a core Conostegia clade . Th is Conostegia clade identifi ed contains all species sampled of Conostegia, in addition to some species of Clidemia D. Don and Miconia Ruiz & Pav. most of which are narrowly endemic taxa from Costa Rica and Panama. Ancestral state reconstruction of the calyptra onto the resulting molecular phylogeny of Conostegia provided evidence for the multiple origins of this structure albeit with diff erent anatomical characteristics in each origin . Th e main objective of this revision is to provide a detailed taxonomic account guided by these recent findings. For this purpose, a broadened circumscription of Conostegia is here proposed. Th is revision also includes the proposal of an infrageneric classification, extensive documentation of the anatomy and morphology in the genus, a key for the identification of its species, descriptions, phenological graphs, diagnostic illustrations for most species and distributional maps.
Some authors have suggested that the best solution to the problem of lack of monophyly of many genera of the tribe Miconieae (Michelangeli et al. 2004;Goldenberg et al. 2008;Martin et al. 2008) is to lump all of them into a giant expanded genus Miconia, which could then be recognized by the presence of berry fruits Judd and Ionta 2013;Judd and Majure 2013;Judd 2013a, 2013b). Th ese publications recommended and have started to divide the expanded Miconia into sections. Th us, the tribe Miconieae, in their view contains a single genus with about 1800 species divided into an uncertain number of sections. Given that most sections recognized to date are as small as three taxa, it is reasonable to believe the number of sections that will have to be recognized is quite large. Th is problem also applies with the large number of genera that would have to be created to accommodate all species within Miconieae. Th e case of Conostegia is diff erent in that sampling for phylogenetic and morphological studies include most of its species, something far from happening for a group as large as the Miconieae. Using sections is problematic because taxonomic databases and herbaria usually do not include infrageneric classifi cations in their organization. Th ese authors argue that lumping all taxa into Miconia will result in taxonomic stability because most species were already included in Miconia, and that fewer taxonomic changes will have to be made than if separate genera are recognized. An aspect that is less mentioned is how many new names would have to be created to accommodate species in all genera being lumped into a broadly circumscribed Miconia. In other words, what is more unstable, to create new combinations or to create altogether new names? For example, including all species of Conostegia in Miconia would require 30 new names, whereas including its species in an expanded Conostegia requires only two new names. I roughly calculated the number of new names required to accomodate all species of the Miconieae in the genus Miconia and came up with the number 354 new names. Th is means that the epithet of those taxa will change all together. It is not clear how to measure taxonomic stability, but I argue that that amount of new names is unnecessary and will result in high nomenclatural instability. Th e last phylogeny of the Miconieae included two genes for 449 species , which represents around 25% of the tribe. Greater sampling both of molecular data as well as species would be desirable when so many nomenclatural changes are being considered. If more data results in better supported clades, these could be considered as the genera to recognize. Th is point was emphasized by Goldenberg and collegues (2008) who cautioned on making major taxonomic realignments since they would be "premature" based on their results. Some authors on the other hand are choosing the alternate route and recognizing genera within the Miconieae such as Killipia and Leandra s.s. (Posada-Herrera and Mendoza-Cifuentes 2013;Reginato 2014).
Th e recognition in this study of a broad Conostegia is based on the best sampled and only phylogenetic study of Conostegia which places all species in a clade within the Miconieae, and since most currently recognized species fall in this clade, and most can easily be recognized by the presence of the calyptra, it is deemed more useful to broaden Conostegia than to lump its species in a giant Miconia. Lastly, the species of Clidemia and Miconia that fall within Conostegia are almost all endemic to Costa Rica and Panama evidencing the strong and useful geographic component that together with morphology can be used in identifying species of this group.

Taxonomic history
Th e idea of separating a group of calyptrate species in the Melastomataceae was fi rst suggested by Bonpland (1806Bonpland ( -1816, who intended to group them into the new genus Calyptres, but never validly published the name. Subsequently, and apparently following Bonpland's idea, David Don (1823) formally described the genus Conostegia emphasizing the presence of the calyptrate calyx. Th e name derives from conus (meaning 'cone') and tectum (meaning 'roof '). Don (1823) included nine species in his concept of Conostegia all of which were known as Melastoma at the time, namely M. calyptrata Desr., M. extinctorum Bonpl., M. glabratum Sw., M. montanum Sw., M. procera Sw., M. xalapense Bonpl. and the following three species which were undescribed at the time and each followed in Don's publication: M. superbum Bonpl., M. cucullata Pav. ex D. Don, and M. holosericea Steud. Ex Triana (Schnell 1996). Don (1823) did not choose a type because this was not customary at the time, but he provided a description, diagnosis and discussion of the affi nities of Conostegia to Miconia. Schnell (1996) informally designated M. procera as the type of Conostegia. Th e combinations of Don's species to Conostegia were done by Candolle (1828) but the latter author credited the names in Conostegia to Don. Candolle (1828) provided descriptions for sixteen species of Conostegia and was one of only two monographers to provide an infrageneric classification in Conostegia. He organized the genus into two sections. Th e first section Eriostegia DC. contained only one species that had been described as Melastoma mutisii Bonpl. and was proposed as Conostegia mutisii (Bonpl.) DC. Th e globose, hispid and abrupt apex of the calyptra characterized this section. Melastoma mutisii was eventually included in the capsular fruited genus Centronia (Triana, 1872) and very recently included in Meriania (Mendoza-Cifuentes & Fernández-Alonso, 2012). Th e second section he called Euconostegia and characterized it based on the shape of the calyptra, which was abruptly acuminate and also hispid. Th e next worker to treat Conostegia was Naudin (1850) who provided detailed descriptions for 11 species and cited an additional seven names. Most of these taxa are currently recognized in Conostegia. Naudin (1850) did not provide an infrageneric classification for Conostegia. After Naudin, Triana (1872) treated 23 species of Conostegia in his monograph of the family which included detailed descriptions for seven of them. Most of those taxa are also recognized today. Lastly, before a break in the revision of the genus that would last about 100 years, Cogniaux (1891) treated 34 species of Conostegia. Later Schnell (1996) provided the only detailed classification within Conostegia in which he recognized 42 species and divided them in three subgenera (Conostegia, Lobatostigma and Ossaeiformis) and further divided subgenus Conostegia into six sections (Axilifl ora, Conostegia, Dasystegia, Notostegia, Parvistigma and Tomentostegia). Unfortunately Schnell's work was never published. Th e genus had always been thought of as a morphologically distinct group (Bonpland 1806(Bonpland -1816Don 1823;Schnell 1996) and in one of the few thorough revisions of the taxonomy of genera with terminal inflorescences, Judd and Skean (1991) concluded that Conostegia was likely monophyletic. After the work of Schnell (1996), the most comprehensive study that included species of Conostegia has been that for the Flora Mesoamericana by Almeda (2009). Th e latter study followed very closely the work of Schnell (1996).

Phylogeny and infrageneric classification
Th e most recent hypothesis of relationships in Conostegia based on DNA sequences from four chloroplast and two nuclear ribosomal regions, resolved the genus as paraphyletic with species of Clidemia and Miconia nested inside . Nonetheless, all species of Conostegia fall in a major clade regardless of the type of analysis conducted (concatentation vs. concordance). A summary of the relationships within Conostegia based on the hypothesis derived from the concatenated analysis is presented in Figure 1. Th ese results also show the paraphyly of Schnell's (1996) subgenus Conostegia since the species of his subgenus Lobatostigma form a clade nested inside of it. With respect to Schnell's sectional classification of subgenus Conostegia his groups are not monophyletic except sections Parvistigma and Tomentostegia. For this reason, the subgeneric classification adopted here includes only sections corresponding to the three major clades in Conostegia identifi ed by both concatentation and concordance analyses and proposed as Conostegia sections Australis, Conostegia, and Geniculatae, respectively (Fig. 1). Section Australis has calyptrate calyces that break at one side, abundant sclereids in the hypanthium, mucilage in the ovary, exserted styles, lack of filament geniculation, and all but one species sampled lack a vascular cylinder in the style. Even then, the cylinder in Conostegia ortizae has a single vascular bundle (unlike species in section Conostegia which have more). Species in section Conostegia are almost unique within the genus in that their styles are not exserted (fl owers not herkogamous). Th is short style is only shared with some populations of C. monteleagreana in section Australis and the clade composed of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata and C. xalapensis within section Geniculatae. An additional character almost unique to section Conostegia and  Schnell (1996). To the right of the phylogeny is the sectional classification proposed here. Th e phylogeny is a slightly modifi ed version of the majority rule consensus tree derived from a Bayesian analysis of DNA sequences from four chloroplast regions and two nuclear ribosomal spacers  pruned to one accession per species. not shared with any of the latter short-styled taxa is the presence of a vascular cylinder within the style. Additional characteristics of section Conostegia include the abundant sclereids in the hypanthium and presence of ovary mucilage (shared with section Australis). Based on anatomy and morphology, section Australis and section Conostegia are quite similar. Th ese two sections also share the frequent pleiostemonous condition and the lack of an evident filament geniculation. Th e two groups diff er in that species of section Conostegia do not have exserted styles, and all have vascular cylinders in the style whereas species of section Australis have exserted style and all but one species lack the vascular cylinder in the style. Lastly, many species in section Australis have an "anther shoulder" that species of section Conostegia lack. Section Geniculatae can be recognized by the lack of calyptra in most of its species. Only the following taxa have a calyptra: C. cinnamomea, C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis. Th e calyptra of these species diff ers from those of the other two sections in that they lack sclereids. Additional characteristics of section Geniculatae include the filament geniculation, the exserted style (in most of its species), small flower size, diplostemonous flowers of most of its species, and the frequently papillose seed testa. Lastly, it is noteworthy that many of the species in this section have leaves that are strongly plinerved and frequently asymmetric. Also, see the Biogeography section for remarks on the marked geographical patterns at the sectional level.

Biogeography
Of the three main clades (sections) of Conostegia, the smallest one, Australis, is noticeable for its species being primarily South American, i.e., C. apiculata, C. centronioides, C. dentata, C. extinctoria, C. lancifolia, C. ortizae and C. rubiginosa. Th is is almost the only clade to contain species endemic to South America except for C. ecuadorensis and C. foreroi of section Geniculatae. Other species of section Australis (e.g. C. lasiopoda and C. tenuifolia ) reach southern Central America and are common in that region, but none of the species in section Australis ranges beyond Nicaragua and none are present in the Caribbean. Th e only species of section Australis to reach an oceanic island is C. lasiopoda, which occurs on Cocos Island in the Pacific Ocean.
Section Conostegia is mostly restricted to Central America and the Caribbean. Roughly speaking there are three distinct areas where endemic species of this section occur. Th e area with the most endemics is southern Central America (Costa Rica and Panama). Th e mountains in these two countries include the volcanic ranges in Costa Rica and the Talamanca mountains that start in Costa Rica and end in Panama, harboring endemics such as C. bigibbosa, C. chiriquensis, C. fragrantissima, C. macrantha, C. micrantha, C. muriculata, C. oerstediana, C. pittieri, C. rhodopetala, and C. vulcanicola. Some of these species reach their northernmost distribution on volcanoes of Nicaragua and some also reach the lowlands of these three countries. Th e second area of endemism for species of section Conostegia is in northern Central America, both in the mountains of southern Mexico as well as in some lower-elevation and drier valleys. Some of these northernmost endemics include C. arborea, C. caelestis, and C. jaliscana. Lastly the third area of endemism, which could potentially be divided into two, are the Caribbean islands of Hispaniola and Cuba (where the endemic C. lindenii grows), and the island of Jamaica where three endemics occur (C. balbisiana, C. procera, and C. pyxidata).
Section Geniculatae stands out biogeographically because most of its species (80%) are endemic to the southern Central American countries of Costa Rica and Panama. Th e rest of the species occur in northern Central America with endemics of that area including C. fulvostellata, C. oligocephala, and C. plumosa. Other species of section Geniculatae (e.g. C. speciosa, C. subcrustulata, and C. xalapensis) are more-or-less widespread in Central America reaching South America. Just like in section Australis, only one species of section Geniculatae (i.e., C. ombrophila) reaches an oceanic island, i.e. Cocos Island in the Pacifi c.
Ongoing work in the tribe Miconieae to obtain a dated phylogeny will provide a time calibrated hypothesis that will enable a thorough biogeographical analysis of Conostegia.

Natural history Herbivory
Many insects especially in the orders Coleoptera, Homoptera, Hymenoptera, and Lepidoptera have been documented interacting with species of Conostegia. Herbivory mostly of the leaves of species of Conostegia by larvae of Lepidoptera has been well documented in Costa Rica. Th e data base of lepidopteran herbivores of Janzen and Hallwachs (2013) from northern Costa Rica includes around 4420 records of 134 Lepidoptera species in 84 genera of 27 families reared from species of Conostegia. Conostegia xalapensis stands out for the large number of species feeding on it. Other lepidopteran species can induce galls such as as species in the family Momphidae (Fig. 2). Hymenopterans have been documented parasitizing larvae in some of these galls, and some have been named after Conostegia. Chrysonotomyia conostegiae Hansson parasitizes Momphidae larvae in galls of Conostegia xalapensis as well as larvae of gall midges (Cecidomyiidae) (Hansson 2004). Another hymenopteran herbivore also named after Conostegia is Allorhogas conostegiae Marsh and Shaw (Chavarría et al. 2009). Th is is one of the few phytophagous braconid wasps that has ever been reported (Chavarría et al. 2009). It was reared from the fruits of C. xalapensis. During this study, beetles of the family Curculionidae were frequently encountered in flowers of Conostegia species. As a result of pickling flowers in the field, several of these beetles remained at the bottom of the pickle jars of diff erent species (Fig. 3).
Th e beetles appear to be of the genus Phylothrox sp. (Curculioninae: Acalyptini) (Franz 2006). Other curculionid beetles that have been reared from Conostegia include one species of the genus Anthonomus from C. oerstediana (Chacón-Madrigal et al. 2012). In another family of the beetles, the Chrysomelidae, two species of the genus Margaridisia have been found, one in C. schlimii and one in C. xalapensis (Flowers and Janzen 1997). From these studies it is clear that much is to be learned about Conostegia species and their herbivores.

Floral biology
Floral longevity in the Melastomataceae has not been thoroughly studied. Stratton (1989) documented floral longevity in Monteverde, Costa Rica, and included nine species of Melastomataceae. He found the mean floral longevity for the family to be 1.24 days. Th is study included three species of Conostegia. Th is short fl oral life span is expected in fl owers of the family because most are buzz pollinated (Renner 1989). Plants that are buzz pollinated tend to be homogamous (off er pollen reward at the same time they receive it) because the lack of rewards such as nectar inhibit them from being dichogamous. Dichogamous plants such as protandrous ones that reward nectar can change the position of the sexual organs through time and off er a reward in both male and female phases. Because melastomes are generally not dichogamous, their fl owers need not live as long as those that change sexual expression.
Floral scent has been noted on several specimen labels of several species of Conostegia. To obtain some descriptive statistics on scent production within Conostegia, I compiled information from both the Instituto Nacional de Biodiversidad, Costa Rica (INB) and Th e New York Botanical Garden herbarium for all Melastomataceae. In total, about 52,000 specimens were databased which had label information. Of this total, 389 specimens had some report of fragrance after searching with key words "olor",  "odor", "scent", "aroma", "fragr", and "cheiro". Some 60 of the specimens with some report of scent are in the Conostegia. Schnell (1996) noted that some species such as C. montana and C. oerstediana can have fragrant or odorless populations. Indeed more thorough observations are needed to assess the systematic significance and distribution of floral scent in Conostegia. Also, for taxa with scented flowers, it remains to be determined where the scent is being emitted. I have perceived strong pleasant scent emission in fl owers of C. cuatrecasii, C. macrantha and C. oerstediana.

Reproductive biology
Within Conostegia, at least two species are known to be self-compatible, C. montana (Tanner, 1982) and C. oerstediana (Schnell, 1996) (Table 1). Another species (C. macrantha) is reported to be self-incompatible (Renner 1989). Th e main way in which species within Conostegia are thought to avoid self pollination is through herkogamy (Renner 1989). Herkogamy is the spatial separation of sexual parts within fl owers and is usually manifested within Conostegia by its species having exserted styles. Th is type of herkogamy falls within the "homomorphic" type of flower in the classification of Webb and Lloyd (1986). Within the homomorphic type they belong to the "ordered" type in which there is usually a single contact between the pollinator with the stigma and stamens. Lastly, within the "ordered" type, flowers with exserted styles fall in the "approach" category in which the stigma is placed forward in the pollinator's path. Two species, C. fragrantissima and C. pittieri are unusual in that they appear always to have their style bent opposite the stamens. A third species C. schlimii also can have its style bent away from the stamens. Although this position of the style is uncommon in Conostegia, it is present in several other groups of Melastomataceae (e.g. Kriebel 2015). Th e evolution of the giant stigma appears to correspond to a diff erent kind of flower altogether. Taxa with giant stigma tend to have large flowers with spreading petals and with the large stigma probably functioning as a large bullseye for the bee to touch when approaching the flower. Th is kind of flower fits those termed "dish-blossoms" in the classification of Webb and Lloyd (1986), characterized by having many stamens and a large central stigmatic region. Th ey were considered as having "homomorphic, ordered type" flowers like the others but which may be of the "unordered" type in which the pollinator may contact the stigma and anthers several times and in no particular order. It should be noted that there is a second hypothesis (besides avoiding self pollination and promoting outcrossing) for the evolution of herkogamy (Webb and Lloyd 1986). Th is second hypothesis proposes that herkogamy evolved as a way to avoid interference between the function of the stamens and that of the style (Webb and Lloyd 1986;Fetscher 2001;Barrett 2002).
All species within section Conostegia and a few within section Geniculatae have consistently short styles. Flowers that lack a distance between the anthers and the stigma may be called non-herkogamous flowers. Recently the term plesiogamous was proposed for this fl ower type (Nesom 2012). Th is term derives from the Greek, plesios, meaning near, alluding to the proximity of the stamens and stigma (Nesom 2012). Th e evolution of plesiogamy has occurred at least twice within Conostegia  and is known to have evolved in other clades of Melastomataceae (e.g., Miconia section Hartigia).
Little has been published on the pollination of Conostegia. Renner (1989) did not report any bee observations for Conostegia in her review of melastome reproductive biology. However, because of their poricidal yellow anthers they are all thought to be buzz pollinated, and some observations of buzz pollination are available (Schnell Table 1. 1996). Th roughout the present study, photographs of potential pollinators were gathered in the following Costa Rican localities (province in parenthesis): Escazú (San José), San Miguel Arriba de Grecia (Alajuela), La Selva Biological Station (Heredia), and the Osa Peninsula (Puntarenas). In the larger flowered taxa which tend to occur at higher elevations, bees of the genus Bombus as well as an unidentified genus were observed buzzing the flowers of C. pittieri and C. oerstediana. At lower elevations, bees of the genus Melipona were observed buzzing the flowers of C. cinnamomea, C. schlimii, and C. subcrustulata. Also at lower elevations, a species of Auglochlora (family Halictidae) was observed buzzing the flowers of C. bracteata, C. consimilis, C. grayumii, and C. setosa. Photographs taken in Ecuador by Darin Penneys of a flowering tree of C. ortizae also show a halictid bee buzzing the flowers. In the lowlands of Costa Rica, species in the orchid bee genus Euglossa have been observed buzzing the flowers of C. bracteata and C. subcrustulata (Fig. 4, Table 1). During this study it was observed that in small flowered species with exserted styles such as C. cinnamomea and C. grayumii, the bees cover the stigma with their abdomen before buzzing the flower (Fig. 4). Th ese taxa tend to have their flowers facing downward which appears to force the bees into this position. On the other hand short styled species such as C. bracteata and C. subcrustulata have their flowers positioned horizontally and the bees grasp the stamen but do not cover the stigma. In large flowered species such as C. oerstediana, the flower tends to be either upright or horizontally positioned and the stigma tends to bend downwards potentially being landed on or touched by the abdomen of large bees. Pollen thieves of the genus Trigona were also encountered both eating the anthers as well as sticking their tongues into the anther pores of C. oerstediana (Fig. 4). Pollen robbing by Trigona bees has been well documented in the Melastomataceae (Renner 1983). In summary, the poricidal anther dehiscence, and the observation of only bees visiting fl owers of species in all three of the major clades of Conostegia suggests that most if not all of the species are buzz pollinated by bees.

Dispersers
Seeds of species of Conostegia are mostly dispersed by birds (Schnell 1996). Seed dispersal in Conostegia volcanalis by the rodents Peromyscus aztecus and Reithrodontomys fulvescens has also been reported (Vázquez et al. 2004). Two cases of fruit dispersal by bats have also been documented, one by Hylonycteris underwoodi dispersing C. xalapensis (Castro-Luna and Sosa, 2009) and one by an unidentified species of bat dispersing C. oerstediana (Harvey, 2000). Bats are thought not to be important dispersal agents of melastomes (Schnell 1996).
Of the known bird-dispersed species, Harvey (2000) documented sixteen species of birds feeding on C. oerstediana and one species feeding on C. xalapensis in Monteverde, Costa Rica. Loiselle and Blake (1990) found seeds of C. subcrustulata to be the most common in fecal samples of birds in second-growth forest at La Selva, Costa Rica. Loiselle and Blake (1990) also found that red-capped manakins, white collared manakins and scarlet rumped tanagers to have a special preference for the fruits of C. subcrustulata. To attract birds dispersers, species of Conostegia have purple berries that contrast with the foliage (Schnell 1996). Some species such as C. monteleagreana and C. rhodopetala have red to pink inflorescence branches that may provide further contrast with the color of the berries. Th e display of the berries in taxa with terminal inflorescences contrasts with that of taxa with axillary inflorescences. Th ese two different types of fruit display result in diff erent guilds of bird dispersers (Schnell 1996). Schnell (1996) noted that the presence of berries on individuals of Conostegia is infrequent both in herbarium specimens and in the field. He suggested that the reason for this pattern might have to do with optimizing limited resources through selective fruit abortion, as has been documented in many families of flowering plants (Stephenson 1981). Th en, the few berries that are actually produced are quickly dispersed. Schnell (1996) recognized two fruiting patterns in Conostegia. In the first one, plants produce berries directly after flowering. Th is behavior results in the presence of flowers and fruits on the same inflorescence/infructescence. Species with this fruiting type include C. lasiopoda, C. macrantha, C. montana, C. rhodopetala, C. setosa, and C. subcrustulata. Th e second type of fruiting behavior is that in which fruiting occurs several months after flowering. Examples of species with this second type of fruiting behavior are C. pittieri and C. rufescens.
Lastly, Schnell (1996) hypothesized that there are three possible advantages gained from bird dispersal: (1) displacement of seeds over a range of distances; (2) improved habitat selection; and (3) enhanced seed germination after passage of seeds through the gut. With respect to the first hypothesis, he found evidence that dispersal distances are modest. Th is is because retention of seeds by birds of species with small berries and small seeds tends to be short (Levey 1987). As for the second hypothesis of reaching suitable sites, Schnell (1996) could not find support, but noted that little is known about the behavior of birds after fruit ingestion. Lastly, Schnell (1996) found some evidence to suggest that germination is not enhanced by passage of seeds through the birds gut as demonstrated by Ellison et al. (1993).

Materials and methods
In total about 2000 sheets were studied for this revision from the following herbaria: CAS, CR, INB, NY, PMA, and USCG. In addition, digital images were studied from the following herbaria: BM, COL, F, GH, MO, P, and US. About 665 specimens were collected during the study of the group on several trips to Costa Rica, and one trip each to both Guatemala and Panama. In order to produce maps, many specimens were georeferenced and additional ones identifi ed by experts (mainly Charles Schnell and Frank Almeda) added to improve the distribution maps. Th e maps and phenology diagrams were made using the R package monographaR (Reginato 2016).

Leaf and floral anatomy
Leaves and floral buds were fixed in 70% ethanol in the field and at maximum two weeks later were brought into the lab and fixed in formalin-acetic acid-ethanol (FAA; 3.7% formaldehyde; 5% glacial acetic acid; 50% ethanol), vacuum-infiltrated overnight, and then stored in 70% ethanol. For light microscopy, fixed material was dehydrated through an alcohol-toluene series in a Leica TP-1020 automatic tissue processor, and embedded in Paraplast X-tra (Fisher Healthcare, Houston, Texas, USA). Th e samples were sectioned at 10 μm with an AO Spencer 820 rotary microtome (GMI Inc. Minnesota, USA). Sections were stained with Johansen's safranin (Johansen 1940) (2% w/v in 50% ethanol) and 0.5% Astra Blue in 2% tartaric acid w/v in distilled water (Maácz and Vágás 1961;Kraus et al. 1998) and mounted in Permount (Fisher Scientific, Pittsburgh, Pennsylvania, USA). Sections were viewed and digitally photographed with a Zeiss Axioplan compound microscope equipped with a Nikon DXM1200C digital camera with ACT-1 software.

Scanning electron microscopy
To thoroughly document trichomes, floral parts, and seeds, Scanning Electron Microscopy (SEM) was used. For the study of floral parts, flowers collected in the field were brought to the lab and transferred to acetone via an ethanol-acetone series, and then dried by critical point, mounted on aluminum stubs with adhesive tabs (Electron Microscopy Sciences, Hatfi eld, PA, USA), sputter coated with gold palladium in a Hummer 6.2 sputter coater (Anatech, Springfield, VA, USA), and examined and photographed in a Jeol JSM-5410 LV Scanning Electron Microscope operated at 10 kV. Seeds were cleaned in water prior to sputter coating and SEM.

Species plates
Both photographs of living plants and dissections of material in spirit are used to illustrate the species treated in this revision. For preserved material, fl oral structures were photographed under a Nikon SMZ1500 stereoscope equipped with a Nikon DXM1200F camera connected to a computer and using the software ACT-1. Th e plates were prepared with GIMP (Th e GNU Image Manipulation Program).

Results
Most species of Conostegia are in Central America (Fig. 5) with the highest density being in southern Central America (Fig. 6). I was fortunate to receive material and images of live plants from South American countries as well as the Caribbean Islands, which encompasses the whole range of Conostegia except for a disjunct population of C. icosandra in Bahia, Brazil. Th ey mostly occur in lowland rainforest and cloud forest habitats. Th e preferred elevational ranges are shown are shown in Figure 7.

Flowering phenology
Th e flowering phenology of species of Conostegia was tallied using herbarium label data. In total, the flowering time for 1420 specimens was recorded. To best visualize in a comparative manner the phenology of each species, circular histograms were produced . Th is assessment of phenology should be taken with caution since it is based on herbarium specimens which can often be the result of easy accessibility to collecting sites and other collecting biases such as sampling error and good weather conditions. Furthermore, realistic patterns of phenology are obscured for broadly distributed species as well as local patterns, which are best studied at the population level. Schnell (1996) reported a pattern present in species such as C. bracteata, C. montana, C. setifera, and perhaps C. balbisiana in which flowering begins around March and finishes in September. Schnell (1996) called this an example of the normal "out-ofseason low-frequency flowering" which might be favored in individuals in high light environments. Schnell (1996) also noted the year-long flowering of weedy species and hypothesized that these longer flowering times might be selected for longer fruiting seasons which might permit more colonization. A second possible non-exclusive explanation proposed by Schnell (1996) is that because of their weedy habit they tend to receive more light allowing them more energy for reproduction. Lastly, Schnell (1996) suggested the possibility that there are collection biases and that these habitats are more frequently visited by collectors. In addition, some species show very marked seasonality. For example, Conostegia caelestis and C. osaensis only flower between March and June. Conostegia oligocephala has a similarly narrow pattern being recorded in flower from May to July. Conostegia brenesii in the cloud forests of Costa Rica only flowers during the rainy season from July through September. Conostegia grayumii, C. incurva, C. pendula, C. povedae, and C. subpeltata appear to follow a similar pattern of rainyseason flowering from July through October. Another possible pattern includes C. allenii and C. calocoma both of which only flower from January through July.

Habit
Species of Conostegia are all terrestrial shrubs or trees. Some of the tallest trees in the tribe Miconieae are found in Conostegia, with C. osaensis reaching about 25 meters in height. Schnell (1996) noted a bimodal distribution of height classes and growth forms in which species were either large trees able to compete with canopy trees such as (e.g., C. volcanalis and C. rufescens), or fast-growing shrubs that seldom grow higher than 2-3 m. He further suggested that the shrubby growth habit had evolved more than once. In addition, I have observed C. rufescens growing both as a large tree and as a shrub. Th e latter was observed in Cerro Jefe, Panamá, and in the lower parts of Braulio   Carrillo National Park in Costa Rica. In Braulio Carrillo, which harbors forests over a wide elevational range, I also observed C. rufescens as a taller tree in the cloud forest. Th e pattern of tall trees is especially evident in the giant stigma group with a clear independent origin in C. osaensis and probably also in C. schlimii, which can also become a tall tree. Th e trunk of some species such as C. bernoulliana and C. oerstediana, can have flaky bark but in most taxa it is smooth. Th e smallest species are those previously described in Clidemia, such as C. subpeltata and C. trichosantha, which grow in the understory mostly of cloud forests. Twigs vary from terete and slender to somewhat White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. . Phenology in species of Conostegia. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. Figure 10. Phenology in species of Conostegia. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. Figure 11. Phenology in species of Conostegia. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. Figure 13. Phenology in species of Conostegia. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. Figure 14. Phenology in species of Conostegia. White slices denote fl owering specimens and gray slices fruiting specimens. Th e size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month. quadrangular and robust. Particularly the species with thick twigs such as C. bigibbosa and C. macrantha, tend to have lenticels on the nodes.

Leaves
Leaves are opposite and generally decussate as is the case in most species in the family. In one species, Conostegia henripittieri, they are always strongly anisophyllous. Most species of sections Australis and Conostegia have leaves with nerved venation and species in section Geniculatae tend to be strongly plinerved. Variation in leaf morphology among species is most evident in section Geniculatae with one peltate species (C. peltata) and a subpeltate species (C. subpeltata), as well as species with sessile leaves (C. dissitiflora) or strongly attenuate leaf bases (C. consimilis). In addition, many species of section Geniculatae tend to have asymmetric leaf venation. In C. grayumii for example, almost every leaf is asymmetrical at the base.

Leaf anatomy
Th irty leaves of 26 species within Conostegia were collected in the field, fixed in FAA, and then sectioned as explained above. Leaves of all Conostegia studied are hypostomatic. Th e cuticle is generally inconspicuous but sometimes can be relatively thick as in C. rhodopetala (Fig. 15). Th e mesophyll is dorsiventral (Figs 15,16), and druses were present in all species, usually near the adaxial or abaxial leaf epidermis. Th e size of the druses varies. Some species such C. tenuifolia have more space in the mesophyll. Leaf thickness varied but in general leaves were thinner in section Geniculatae (Fig. 17). At least two types of coriaceous leaves were found. Populations of C. montana from Cerro Jefe, Panama, have a very thick mesophyll and are the most evidently coriaceous leaves encountered in this survey. Th e other thick-leaved species were found to have a hypodermis one cell layer thick. Th is hypodermis is found only in C. montelagreana in section Australis and in the species of the giant stigma clade (species sampled were C. bigibbosa, C. icosandra, C. oerstediana, and C. pittieri) where it appears to have evolved in their common ancestor (Fig. 15). Mentink and Baas (1992) reported the presence of a hypodermis in C. subcructulata but sections done for this study failed to locate a hypodermis in that this species (Fig. 16). In the Melastomataceae the function of the hypodermis has been suggested to be related to water storage in epiphytic species (Reginato et al. 2009) or protecting the palisade parenchyma against solar radiation in terrestrial species (Ely et al. 2005). Th e spongy mesophyll was usually not lignified except in some species, such as C. ombrophila and C. schlimii.
Th e petioles in Conostegia bigibbosa and some Costa Rican and Guatemalan populations of C. montana have two protuberances at the apex of the petiole on the abaxial surface. Scanning electron micrographs of these protuberances evidenced a glabrous area with some stomata suggesting that they might be extrafloral nectaries. However, anatomical sections in C. bigibbosa did not indicate the presence of carbohydrates. Cross sections of petioles show variation in shape from rounded to somewhat heart shaped, grooved or flat adaxially as in C. consimilis, C. schlimii, C. tenuifolia, and C. bernoulliana (Fig. 16). Five to nine amphicribal bundles are present, forming an interrupted arc. Th e lowermost bundles tend to be larger as in C. rufescens, C. subcrustulata and C. schlimii, (Fig. 16). Smaller bundles are sometimes present inside the primary arc as in C. bernoulliana,C. bracteata,C. rufescens,and C. tenuifolia (Fig. 16). Petioles are mostly unlignifi ed but sclereids are present in some species, but besides the sclereids, petioles were mostly unlignified. Some lignification of the petiole was observed in C. rufescens and C. schlimii.

Leaf domatia
Th ree types of domatia occur in Conostegia. In two species, C. dentata and C. setosa, the domatia are of the formicarium type which house ants and are manifested as large swellings at the base of the leaf. Th e other two types are both mite domatia, present at the base of the leaf on the abaxial side at the point of divergence between the midvein and the primary lateral veins. Th e general classification of domatia used here follows Jacobs (1966). Th e first kind of mite domatium is the tuft mite domatium present in C. procera (Fig. 18). Th is species always has these structures, which are densely covered by stipitate stellate to dendritic trichomes. Th is type of domatium is also seen in some but not all specimens of C. hirtella. Th e second type of mite domatium is the pocket domatium which literally looks like a pocket formed by a membrane (Fig. 18). Some   pocket domatia are further called vesicular domatia because they are inflated. Pocket domatia are present in only the clade comprising C. ecuadorensis, C. hammelii, and C. ombrophila. Among the species having pocket domatia, C. ecuadorensis is unique in that there are domatia present on both the innermost pair of primary lateral veins as well as on the outermost pair of lateral veins.

Indument
Trichomes in Conostegia are quite diverse and variable (Fig. 19). Th is may not come as a surprise since trichomes in the Melastomataceae have been said to be the most diverse in the angiosperms (Wurdack 1986). In many cases trichome morphology is easy to describe, such as with simple or lepidote trichomes. Diffi culty in describing trichomes arises with the vast variation in dendritic trichomes, and with the gradations from dendritic to stellate. Wurdack (1986) noted that all types in the family are multicellular and, of the 46 trichomes types he recognized, 14 were recorded in species of Conostegia sensu Schnell (1996). Schnell (1996) discussed trichomes in Conostegia extensively noting that Conostegia species tend to have a mixture of trichomes and that variation within a species is extensive. I have noted extreme variation in C. icosandra, which can be densely hirsute in the northern part of its range. Conostegia superba has also caused confusion because in the mainland specimens tend to be glabrous or almost so, but in the Dominican Republic they can be quite pubescent except for the floral buds. As noted by Schnell (1996), and even now taking the phylogeny into account, it is evident that similar trichomes have evolved independently within Conostegia. One example involves the interdependent evolution of stipitate trichomes of C. brenesii and C. caelestis. Schnell (1996) divided the trichomes of Conostegia into the three general groups recognizable based on the work on Wurdack (1986): 1) tiny glands, 2) elongate simple hairs, and 3) a series of eight potentially intergrading kinds of stellate and branching hairs. Schnell (1996) stated that continuous variation among these eight groups made it diffi cult to describe them. I agree with Schnell and recommend that into the future, a quantitative method should be developed to describe and perhaps discretize these complicated dendritic to stellate trichomes. In the meantime, I have chosen to document the trichomes in a similar way as Wurdack (1986) did, with extensive Scanning Electron Micrographs . Apparently all taxa have minute glands on the abaxial leaf surface, and their shape is variable and may prove to be taxonomically informative. Th eir small size makes them diffi cult to describe, especially in species with dense pubescence on the leaf abaxial surfaces. To compliment Wurdack's (1986) initiative for Conostegia, I placed special attention on documenting these minute glands as much as possible.
All species with discolorous leaves are in section Geniculatae. However, this character has evolved independently a number of times within the section and not all discolorous leaves are the result of the same type of trichome. In some cases such as C. xalapensis the dense indument on the abaxial surface is made up of stellate trichomes with long, thin arms, whereas in closely related C. osaensis they are lepidote. Many species in section Geniculatae have an orange-colored indument of dendritic to stellate trichomes on the stems, especially towards the apex, which are less common in the other sections or if present do not tend not to form a conspicuous orange covering.

Inflorescence
Inflorescences in Conostegia are variable among species as in most clades of Miconieae (Michelangeli et al. 2004). Most species have terminal erect panicles with many flowers, but the infl orescence can be defl exed. In some species previously described in Miconia as well as in some species traditionally placed in Conostegia such as C. cinnamomea and C. muriculata, the inflorescence can be evidently deflexed. Axillary or pseudolateral inflorescences tend to have fewer, smaller fl owers and are present mainly in those taxa previously described in Clidemia. Some small-flowered taxa do have many flowers such as C. consimilis. Bracts are usually early deciduous in sections Australis and sections Conostegia except in species such as C. monteleagreana which has persistent bracts subtending the floral glomerules. On the other hand, many species in section Geniculatae have bracts and bracteoles that are persistent and fused at the base, forming an inconspicuous nodal collar. Inflorescence branches are particularly thin, wiry, and delicate in several species within section Geniculatae such as C. cinnamomea, C. grayumii, and C. ombrophila. It was the presence of this type of infl orescence, as well as the similar bracteoles amongst other characters that prompted Schnell (1996) to state that the similarities between C. cinnamomea and C. brenesiana are "little short of uncanny, involving almost every character, even to minute details".
Accessory branches in inflorescences are common in Conostegia, especially in taxa with relatively long terminal inflorescences. Although it is tempting to use the presence of these accessory branches in the systematics of the group, they are absent in some specimens, and in such cases it is diffi cult to tell if they fell off or were not there to begin with. Accessory branches are consistently absent in most species of section Geniculatae Pedicels in Conostegia are also variable among species and major clades. In particular, species within section Conostegia can have quite long pedicels, as previously noted by Judd and Skean (1991). Th ese pedicels can further elongate as the flowers develop into fruits. Pedicels are absent (fl owers sessile) in some species such as C. monteleagreana in section Australis and C. colliculosa and C. povedae in section Geniculatae. Schnell (1996) noted that in certain species the pedicels are clustered at the end of the inflorescence branches. Th is clustering is such that the pedicels appear to arise from the axils of other pedicels hence naming these taxa in his section Axilliflora Schnell -ined (see figure of C. superba for an example). Species in this section do not form a clade as evidenced by the molecular phylogeny where C. monteleagreana falls in a diff erent clade than C. cuatrecasii, C. rhodopetala, and C. superba ( Fig. 1; see also Kriebel et al. 2015). Nonetheless, Schnell (1996) himself noted that the species of section Axilliflora did not sort out into clear groups.

Flowers
Floral buds in calyptrate species of Conostegia are noticeable because of their calyx tends to fall as a unit at anthesis. Calyptrate calyces have arisen independently at least 15 times just in the Neotropical genera of the family and can be variously shaped ( Fig. 34) (Schnell 1996). In the Miconieae, five genera have at least some species with a calyptra namely: Conostegia, Mecranium, Miconia, Tetrazygia and Tococa (Schnell 1996). Schnell (1996) considered floral buds to be the most useful structures not only to identify species at the generic level but also within Conostegia. In general terms their size and shape can be taxonomically useful, as well as the presence of warts or diff erent types of indument on the hypanthium. Th e apex of the buds can also be helpful for identifying some species. An apiculate calyptra for example, is present in several distinct species from diff erent geographical areas. In South America, the aptly named C. apiculata is readily distinguished. In northern Central America, the distinctively apiculate calyptra of C. arborea is not easily confused with others; the same is true of C. pittieri in the mountains of Costa Rica and western Panama, but in this area there are other species that can have somewhat apiculate calyptras (e.g. C. tenuifolia and C. rhodopetala).
Phylogenetic and anatomical analyses have revealed that the calyptra evolved at least three times within Conostegia . Th is is a surprise because most workers previously thought the calyptrate calyx was shared by all species in the genus (Judd and Skean 1991;Schnell 1996) even referring to Conostegia as monophyletic without phylogenetic analyses of any kind (Judd and Skean 1991). Schnell (1996) did point to the possibility of the independent origin of the calyptra in C. cinnamomea, and stated that because these structures had evolved in the family there was possibly a selective force driving their evolution. Th e adaptive value of the calyptra remains unknown.
Th e calyptra of species in sections Australis and Conostegia are very similar in that they have conspicuous sclereids and no calyx teeth nor appendages (Fig. 35). Th e sclereids for the most part form a layer or two throughout the hypanthium and calyptra (Fig. 36). Th e main diff erence in the calyptra of these two sections is that in section Australis, the calyptra ruptures at one side and/or breaks into pieces in all species observed in the field (Fig. 37). In section Conostegia all evidence points to a cleanly circumscissle calyptra which Judd and Skean (1991) and Schnell (1996) believed was the case in all species of Conostegia. Independent origins of the calyptra are also seen in C. cinnamomea, which has a very thin calyptra, and in the clade comprised of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis. Th e calyptra in this latter clade lacks evident sclereids (Fig. 35). Although most species have a glabrous calyptra on the inside, three closely related species, i.e. C. plumosa, C. speciosa and C.xalapensis tend to have stellate trichomes inside the calyptra.
Within section Geniculatae, sclereids are mostly absent from the hypanthium and calyx. Some scattered sclereids are seen in the hypanthium of C. schlimii. In the fl oral anatomy of this section there are two general patterns that are absent in sections  Australis and Conostegia. First, in several species of Geniculatae there is a lining of druses around the ovary (Fig. 38). Second, several species have a distinctly bicolored anatomy where the ovary tissue stains red, indicating some lignification but the rest of the hypanthium stains blue. In fewer specimens of section Geniculatae the hypanthium stains either totally red or entirely blue. For further anatomical diff erences between the ovaries in section Geniculatae and the other two sections Australis and Conostegia, see below under the Gynoecium section.
Another unusual characteristic of the calyx of non-calyptrate species in section Geniculatae is the presence of an irregularly rupturing calyx in several of them (Fig. 39). Th is type of calyx is usually translucent. An example of the anatomy of the translucent calyx of C. friedmaniorum is presented next to the calyx of C. schlimii (a species without a translucent calyx) (Fig 35). In all cases of species with this type of calyx, because it ruptures, the calyx lobes are irregularly shaped. Th e calyx teeth in calyptrate taxa are diffi cult to assess. In sections Australis and most calyptrate species of Geniculatae they appear to be totally absent and anatomical sections failed to reveal them. In C. cinnamomea, anatomical sections did evidence their presence. On the other hand, within the clade of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis one sees what seem to be inconspicuous calyx teeth in C. subcrustulata. Schnell (1996) included the aforementioned clade (except C. osaensis which was undescribed at the time) in his section Tomentostegia. In his description of that section, Schnell (1996) described the calyx lobes as fused and usually with free teeth or prolonged appendages. Assessing if these teeth or appendages are homologous to calyx teeth is diffi cult. To complicate things even more, in C. osaensis, which is the sister clade, the calyx teeth appear to be at the level of the torus as is common in species of section Geniculatae and more generally in most species of the tribe Miconieae.
Flowers in the species traditionally treated in Conostegia also stand out among genera of Miconieae because most of them are pleiostemonous, meaning that they have more than double the number of stamens than petals (Fig. 40). Other pleiostemonous taxa do exist within the Miconieae, but they are not thought to be closely related to Conostegia. Th ere is one pleiostemonous species that has been named in Conostegia, C. inusitata, which appears not to be closely related to the other named taxa in the genus. Th is suggestion comes from a nuclear ribosomal DNA sequence (Kriebel, unpublished data) of an undescribed and closely related species to C. inusitata, which Schnell (1996) proposed as the new species Florbella wurdackii. Th e genus Florbella was proposed by Schnell to accommodate these two species, but it has yet to be published. Inclusion of this DNA sequence in phylogenetic analyses of Miconieae suggests it is more closely related to a clade of mostly Peruvian species of Miconia than to species of Conostegia. Th e concept of Conostegia included in the present treatment includes many diplostemonous species, particularly those previously described as Clidemia and Miconia. Th ese diplostemonous taxa are almost completely restricted to section Geniculatae (Fig. 41), with a few diplostemonous taxa also found in section Conostegia (e.g. C. setosa).

Corolla
Th e corolla in Conostegia varies dramatically in size, shape and number of parts among species (Figs 42,43) and sometimes also within species. Petal number can range from 4 to 12. Petal shape is quite variable among species and in general they tend to be asymmetric with notable exceptions e.g. in some species in section Geniculatae. Many species especially in section Australis and section Conostegia tend to have strongly asymmetrical petals. Th e posture of the petals is usually spreading with some species (particularly small-flowered ones) having reflexed petals. Petals are always imbricate in bud and overlap in a counter-clockwise fashion (Schnell 1996). At anthesis petals are usually imbricate in large flowered taxa but the overlap tends to decrease as the flowers get smaller. Petal margins are for the most part entire but in many large-flowered taxa they can have a more membranous texture on one side (Fig. 44). Petal apices are mostly rounded, truncate, or emarginate, and in a few species they can be narrowly rounded to acute. One species (C. consimilis) was originally described in the genus Leandra because of its acuminate petals. Petal bases are not evidently clawed, except in C. schlimii.
White is the most common petal color found in Conostegia with a few species having pink to purple petals such as C. bigibbosa, C. cuatrecasii, and C. muriculata. In a few taxa like C. fragrantissima the white petals can have a red band at the base. Translucent petals can be found in diff erent clades as well, and although it is tempting to think this might be related to floral size because several small-flowered taxa have them, some species with large petals have translucent petals as well (e.g. C. lasiopoda). Most species have glabrous petals but at least one has conspicuously papillose petals (C. papillopetala). Petal surfaces tend to be smooth in translucent petaled taxa and with rounded papillose cells in white-flowered species (Fig. 45). Although this diff erence in petal cells is evident also in micrographs (Fig. 46), intermediates exist and further study is needed to determine their possible systematic utility. In a few species, the petals persist after all other floral parts have fallen (of which C. pittieri might be the most notable example).

Androecium
Th e androecium in Conostegia consists of 8-52 isomorphic stamens. Th e basic arrangement of the stamens consists of five of them inserted opposite the sepals and five of them opposite the petals like most species in the Melastomataceae. Increase in the number of stamens are common, even predominant in sections Australis and Conostegia. Th ese increases result in pleiostemony, and were the subject of a recent floral developmental study (Puglisi 2007;Wanntorp et al. 2011) which found two ways in which Conostegia species increase in stamen number: 1) by having a large stamen in alternipetalous position with small ones in antipetalous position; or 2) by a process called dedoublement where the stamens split in two (Puglisi 2007;Wanntorp et al. 2011). Th ere appears to be no clear pattern indicating evolutionary relationships, with taxa with either of these developmental pathways being more closely related to another with the opposite pathway.
Th e posture of the stamens can go from erect, to forming a more or less a 45-degree angle between the filament and the anther (Fig. 47). Species with stamens that form an angle tend to have at least slightly bilaterally symmetric androecia. Th is condition is common in sections Australis and Conostegia as well as in the clade composed of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis within section Geniculatae. Bilateral symmetry in Conostegia appears also to be related to interactions between the stamen and style during development at least partly the result of the increase in number of stamens. Species in section Geniculatae, excluding the clade mentioned above, have stamens that cleanly surround the style and are radially symmetric, yet the flowers of these species might still have slight bilateral symmetry as a result of a gently curving style. In at least two species, C. fragrantissima and C. pittieri, the stamens are bent away and opposite the style.   Th e filaments are mostly white to translucent. Sections Australis and Conostegia lack a clear geniculation towards the apex of the filament, whereas in section Geniculatae the geniculation is present in almost every species, except perhaps in C. fraterna . It is significant that the molecular phylogeny of Kriebel et al. (2015) placed species traditionally recognized in Conostegia such as C. plumosa, C. speciosa, C. subcrustulata and C. xalapensis in Geniculatae and that the latter three species which     were studied from field-collected floral material have an evident filament geniculation. Th is morphological character thus supports their placement within section Geniculatae. It has to be considered that some variation does exist. For example, the micrograph of C. xalapensis (Fig. 49) does not show the evident geniculation very well whereas the photographs of the pickled stereoscope images of the same specimen do. In general, C. xalapensis shows an evident geniculation. Although most species which have fi lament geniculations present them towards the apex of the filament, in one species, C. brenesiana, the geniculation is in the middle of the filament.
Anthers in Conostegia are almost exclusively yellow to sometimes orange (i.e. C. fragrantissima) with some taxa having hues of pink towards the apex in some populations (i.e. C. rufescens). In some species particularly in section Australis the anthers can be cream colored. Anthers in Conostegia lack evident staminal appendages with at the most, some species (e.i., C. bernoulliana) having an inconspicuous dorsal bump on the anther connective. Th e only structure that resembles an anther appendage in Conostegia is present in most species of section Australis. In these taxa, the anther base is briefly prolonged below the junction of the filament and anther thecae (Fig. 50). Th is      Kriebel 515). Scale bar: 100 um. character was recently documented in some species of Tococa (Michelangeli 2005) and Pachyanthus (Bécquer 2008). Th e documentation of this type of anther base in Conostegia is relevant because it had not been used before in the systematics of the group, and although lost in some species, helps to identify species in section Australis.
Anther shape in Conostegia varies a lot even when considering only the species treated traditionally in the genus. Th is is in contrast to what has been previously stated in the literature. In their taxonomic studies of the Miconieae, Judd and Skean (1991) remarked that the non-appendaged anthers characteristic of Conostegia are ovoid, tapered to the apex, and open by a small apical pore. Th ese anther characters lead to conclude that the genus did not evolve from within Miconia, since members of the latter genus show various modifications of either pore or connective structure. We now know that Conostegia is phylogenetically nested within the Miconieae, and furthermore that anthers of its species can be ovoid, oblong, linear, or variations on these shapes (Schnell 1996;Fig. 49). Th ey are definitely not all ovoid. Also, they can be recurved or arcuate making them diff ciult to describe. In addition, it is not very useful to say that a small pore characterizes the anthers of Conostegia. In fact, most species of Miconieae have small anther pores corresponding to their buzz pollinated syndrome, and for this reason this type of pore was coded as plesiomorphic by Judd and Skean (1991). Even if the size of the pore proves useful in the systematics of Conostegia, continuous measurements would have to be employed to determine their utility. Th e shape of the anther apices and anther pores is quite variable in Conostegia and merits further research. Th is will require careful placement of critically point dried anther apices on stubs and Scanning Electron Microscopy. Some of the variation seen in the pores has to do with the deeply channeled anthers in the ventral surface. In some taxa this deep channel reaches the pore area but in others it does not. Conostegia brenesiana stands out because of its very small anther thecae with broad pores which can resemble anthers in Miconia section Cremanium. In most species the pores are terminal to ventrally inclined. Dorsally inclined pores can be found in some species of section Geniculatae, and are especially evident in C. cinnamomea (Figs 50,51).
With respect to the anatomy of the anthers, the sporangia in each anther theca were found to have two general arrangements. In species of section Australis and Conostegia they tend to be positioned side by side. On the contrary, in section Geniculatae they tend to be more-or-less superposed. All anthers were found to have druses. In some species such as Conostegia lasiopoda and C. oerstediana, the druses are present in the staminal connective, endothecium and anther septum (Fig. 52). In others, they are mostly restricted to the septum like in C. tenuifolia. Species in section Geniculatae tend to have druses on the proximal half of the outer side of the anthers (as in C. fraterna, C. friedmaniorum, C. ombrophila, and C. subcrustulata) (Fig. 52).
Pollen of species in the Melastomataceae has been rarely documented in taxonomic treatments. Th is may be because the few studies that have been done have revealed little variation (reviewed in Renner 1993). For the species studied, Renner (1993) summarized the pollen as small, tricolporate, radially symmetrical, and isopolar; occasionally grains may have 4, 5 or more colpi, and some grains are dicolporate or heteropolar. Faint to distinct pseudocolpi (or subsidiary colpi, in the terminology of Patel et al. 1984) are usually present. Th e surface sculpture is smooth and striate to rugulate or rugulate-verrucate. Polyads and tetrads are known from Tococa spadiciflora and Miconia melanotricha (Patel et al. 1984). In the case of Conostegia, Roubik and Moreno (1991) provided the only images known to me of the anatomy of pollen of Conostegia species (including C. shattuckii as Miconia shattuckii). I have found that the pollen characteristics of Conostegia agree with Renner's (1993) general description for the family; this is shown in micrographs provided for several species (Fig. 53).

Gynoecium
Th e gynoecium can have from 4 to 25 carpels (Schnell 1996). Placentation is axillary as in most Melastomataceae (Renner 1993). Th e placenta in most Conostegia protrudes into the locule, either in laminar fashion with the ovules attached at the sides, or it is peltate with a clear stipe (Fig. 54). Intermediates are present, and in a few cases (e.g. C. xalapensis), the placenta looks like the peltate kind but lacks a clear stipe. Species of sections Australis and Conostegia have inferior ovaries, and in some species of section Conostegia the ovary can be elevated into an evident collar around the style base. Species in these two sections all have glabrous ovary apices. In contrast, species of section Geniculatae tend to have ovaries about two thirds inferior and several species have glandular-pubescent ovary apices. A collar around the style base is usually absent in section Geniculatae. Species of sections Australis and Conostegia have a glabrous torus whereas species in section Geniculatae can have a glabrous or glandular-puberulent torus. In one species, C. osaensis, the torus is beset with lepidote trichomes.
A significant discovery of this study is the deep blue stain recorded in anatomical sections inside the ovary of species in sections Australis and Conostegia (Fig. 36). On the other hand, this staining is absent in section Geniculatae. Staining with Ruthenium red has provided evidence that the substance in question is mucilaginous pectin, and field tests and photographs by Reinaldo Aguilar of with C. bernoulliana in the Osa Peninsula have demonstrated that the berries are sticky as would be expected from the presence of mucilage.
Conostegia can be said to have the highest diversity in style and stigma morphology in the whole Melastomataceae (Fig. 55). Most notable is the presence of  crateriform lobed stigmas in several species (Figs 55,56). Th ese crateriform stigmas are unique to a subclade of Conostegia, although in some species the crater is reduced to almost absent but the lobes mostly remain present. Besides these obvious outliers, the diversity is still pronounced, ranging from short, stout, straight or apically hooked styles, to long, straight or gently curving styles. Stigmas can be punctiform, truncate, or variously enlarged (Fig. 55). Styles in Conostegia can be exserted or not. Biologically, this exertion corresponds to the maintenance of herkogamy (see Reproductive Biology section). Although the posture of the style appears to be taxonomically informative, this posture can vary through time within the same individual in some species. In species with exserted styles, two general trends are noticeable. Th e first trend, seen in species of section Australis as well as C. schlimii of section Geniculatae, is one in which the flower is either facing upward, or horizontally and thus the style is erect or horizontal. In both these similar postures, the style tends to surve gently towards the apex resulting in a more or less S shape. Th e other trend in species with exserted styles is almost the norm in species of section Geniculatae and involves deflexed styles. It should be noted that in this revision, exemplary flowers of all sections are presented in erect position for convenience (see Figs 40 through 43), but in the taxonomic treatment, images of live fl owering plants display the natural posture of the fl owers.
Th ere are two main clades where the style is not exserted beyond the stamens (not herkogamous). Th e first corresponds to section Conostegia and the other to the clade comprised of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis. Aside from the crateriform and or lobed stigma clade, which are derived from section Conostegia, all remaining taxa with a short style appear to have either a straight style or one that bends upward below the stigma. Within section Conostegia, style posture appears to change in most of its members, particularly in the crateriform or lobed stigma clade. Th e few observations of species in this group (Stratton 1989;Schnell 1996) suggest their flowers can last about two days. During this time the style bends downward usually blocking numerous stamens. Preliminary experiments (Kriebel unpublished data) show that if the style is removed, the stamens can unveil normally and that if the stamens are removed the style still bends (see photographs in section on Reproductive biology).
In the styles of the Conostegia clade, another outstanding aspect involves the presence of a stele within for their entire length (Figs 57,58). Specifically, section Conoste-  gia and convergently Conostegia ortizae in section Australis present this phenomenon. Th is stele is unknown in the rest of the Melastomataceae. Schnell (1996) was the first to document the stele in two species which are phenotypically divergent but belong to section Conostegia. Th is character is helpful to confi rm the hypothesis, derived by optimizing morphological traits on the molecular phylogeny, that the calyptra and short style seen in the clade composed of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis must have evolved independently from the similar morphology seen in section Conostegia. No species including the ones just mentioned of section Geniculatae were found to have a vascular cylinder inside the style. Th ese results also coincide with Schnell's (1996) observations of a lack of a cylinder in C. subcrustulata. Th e stele found in the style of C. ortizae, in general, diff ers from those in species of section Conostegia because it has fewer vascular traces.

Fruits
Fruits of most species traditionally recognized as Conostegia have a truncate apex produced by the dehiscence or breaking of the calyptra. In addition, some have a prominent ovary apex that is retained in the berry and can be highly elevated in fruit (Fig. 59). Fruits of some species such as those of C. oerstediana have a pleasant taste.

Seeds
Almost all species of Conostegia have small seeds averaging about 0.5 mm in length and varying in number from hundreds to thousands in a single berry. Th e most significant deviation from this pattern is observed in the clade comprised of C. osaensis, C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis, where seeds are fewer in number and average about double in length.
Seeds in Conostegia are mostly ovoid except for section Geniculatae where several species have pyramidal seeds (Figs 60-62) (Ocampo and Almeda 2013). In some cases like C. brenesiana most seeds are pyramidal but ovoid seeds can sometimes be found in the same fruit. Angles are absent except for some species in section Geniculatae that can be strongly angled. Th e lateral symmetrical plane is ovate in sections Australis and section Conostegia and frequently triangular in section Geniculatae. Tubercles on the seed surface are evident in in section Geniculatae and in C. monteleagreana of section Australis but are absent in section Conostegia.
Th e conclusion of this study is that despite the diffi culty in coding seed characters, species of section Australis and section Conostegia have very similar seeds in being ovoid and smooth, whereas there is a lot of variation in seed morphology is in section Geniculatae, with ovoid to pyramidal seeds that frequently have angles and tubercle-like cells present just on the angles or covering the whole seed. Description. Small shrubs to medium sized trees with slender and terete to stout and tetragonal branches; branches glabrous to variously pubescent with simple, stellate, stipitate-stellate, lepidote, or highly varying dendritic trichomes, usually the trichomes types not in combination. Twigs with or without nodal lines which can be obscured by the indument. Petiole absent or usually present, in one species (C. bigibbosa) and some populations of another (C. montana) with two tubercles near the apex abaxially. Leaves subisophyllous to isophyllous or rarely anisophyllous (most markedly in C. henripittieri), membranaceous, rarely coriaceous, several species with somewhat leathery texture,     particularly the ones with a hypodermis, nerved to strongly plinerved and if the latter, frequently asymmetric, entire to crenate, denticulate or serrate, adaxially usually glabrous, abaxially glabrous to variously pubescent with simple, stellate, stipitate-stellate, lepidote, or highly varying dendritic trichomes, the surface obscured by indument in a few species, all species with tiny glands on the surface, the apex acute to caudate, the base peltate, rounded, cordate, acute or long decurrent, with pouch-like formicaria at the base in two species (C. dentata and C. setosa), with pocket domatia at the base abaxially in three species (C. ecuadorensis, C. hammelii and C. ombrophila), and with evident tuft domatia in one species (C. procera). Inflorescences pseudoaxillary or terminal, erect or deflexed, small dichasia or small to large panicles, few to many flowered, branching at or above the base, in a few cases the branches terminating in bracteate glomerules (i.e. C. monteleagreana), bracts and bracteoles deciduous and sometimes appearing absent, or persistent and especially the bracteoles sometimes forming a nodal collar around the inflorescence branches, accessory branches present especielly in taxa with terminal, paniculate inflorescences. Flowers diplostemonous or pleiostemonous, 4-12-merous; calyx calyptrate or not, if calyptrate with or without calyx teeth and the calyptras varying from very thin to very thick, glabrous to pubescent like the hypanthium and with or without sclereids, if the calyx not calyptrate, fused in some species and rupturing irregularly at anthesis into irregular lobes, in non calyptrate species the calyx lobes mostly inconspicuous and similar to the calyx teeth, not conspicuous except in C. incurva; petals reflexed or generally spreading, linear-oblong to broadly obovate, the apex acute to rounded, evidently clawed at the base at least on one species (C. schlimii), mostly glabrous, in at least one species conspicuously papillose (C. papillopetala), translucent to mostly white, in a few species pink or purple, the petals cells on the adaxial surface mostly rounded in species with colored petals and flattened in species with translucent petals. Stamens 8 to ca. 52, arranged neatly around the style in diplostemonous taxa and less neatly but similarly in most pleiostemonous taxa, a few species with the stamens bent to one side of the flower (i.e., C. fragrantissima and C. pittieri), filaments translucent white to white, with or without an evident geniculation near the apex, the connective not prolonged nor appendaged, the filament anther insertion transitioning smoothly or with a "shoulder" or abrupt step, anthers linear, oblong, elliptic or ovate, apically rounded or acute, basally acute, rounded or sagittate, yellow, less frequently whitish or with pinkish hues, laterally compressed, with druses in the endothecium, the pore oriented totally upward, somewhat ventrally inclined or less frequently dorsally inclined, usually small, broad in one species (C. brenesiana), anther sporangia positioned laterally or more or less superposed. Ovary from almost superior to usually totally inferior, 4-25 locular, the placentas within each locule laminar, triangular or peltate, with or without mucilage inside; the style exserted beyond the stamens or not, cylindrical and linear, to shaped as an inverted crateriform cone, when linear, sometimes gently bending to abruptly bending below the stigma, hollow or with a stele within, the stigma lobed or not, papillose. Berries small to large, mostly purple, sweet and usually pleasant to human taste; seeds mostly numerous ad small (ca. 0.5 mm long), few and large in a few species, largest in C. osaensis (ca. 1.5 mm long), ovoid to pyramidal, sometimes evidently angular, the testa smooth in most species, tuberculate all over in some, and in fewer still, with the tubercles restricted to the angles. Species concepts in this revision for the most part follow the morphological species concept as defi ned by Cronquist (1978) in which species are "the smallest groups that are consistently and persistently distinct, and distinguishable by ordinary means." It is a problematic concept partly because defi ning "distinct" is subjective and varies across organisms. Nonetheless, it is the most common species concept applied to plants (Mc-Dade 1995), and an eff ort was made to recognize only species that appear distinct. Th is author views species as hypotheses that are subject to testing. Th us the reader should be aware that some species may someday prove to be synonymous with others, and some species complexes might prove to be made of cryptic species in need of recognition. Properties of other species concepts are desirable, such as defi ning species based on monophyly (e.g. the phylogenetic species concept), but the lack of enough representative specimens in the phylogenetic analyses of many species, as well as appropriate molecular markers to ascertain clades (species) with confi dence, and inherent biological processes such as incomplete lineage sorting, preclude much of its use in the present treatment. In this monograph when a negative measurement is given for the distance between the anther and the stigma, this means that the anthers are below the stigma for that length.

1
Sepals not fused into a calyptra, sometimes rupturing late at anthesis into irregular lobes, calyx teeth present at the level of the torus or slightly above; flowers diplostemonous (sect. Geniculatae, pro majore parte) 2 Leaves discolorous, the abaxial surface white to rusty and concealing the surface 3 Sepals fused and rupturing at anthesis into irregular lobes 4 Sepals the same texture as the hypanthium, persisting through flowering; anther apex rounded . Description. Trees 2-8 m tall with tetragonal, ridged and swollen stems which are covered with a mixture of sessile stellate and stalked-stellate hairs sometimes intermixed with simple hairs; the nodal line present but mostly obscured by indument.
Leaves of a pair equal to somewhat unequal in length. Petiole 1-7 cm. Leaf blades 8-26.9 (-30) × 6.6-11.5 (-15) cm, 3-5 plinerved with the innermost pair of veins diverging from the mid vein in sub alternate to alternate fashion up to 1.5 cm above the base, elliptic, the base acute to obtuse, the apex acute to acuminate, the margins dentate with gentle curves between the well separated teeth, adaxially glabrous, abaxi-ally with a a mixture of sessile stellate and stalked-stellate hairs sometimes intermixed with simple hairs. Inflorescence a terminal panicle 6-15 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, rachis flattened, bracts early deciduous or absent, bracteoles linear, ca. 2 mm long, early deciduous. Pedicel 3.5-6 mm long. Flowers 7-8(-12) merous, calyptrate. Flower buds 11-16 × 5-9 mm, not constricted in the middle, the base flat, the calyptra apiculate; the hypanthium 6-9 × 6-9 mm, smooth, glabrescent to evidently stellate pubescent. Petals 10-12.5 × 7-8 mm, white, obovate, glabrous, rounded-truncate to emarginate. Stamens 20-28, apparently slightly zygomorphic because the style is bent, the filament 4-5 mm, not evidently geniculate, white, anthers ca. 3 mm, oblong and somewhat recurved, sagittate at the base, the connective thickened, laterally compressed, yellow except for a hugh of rose at the base of thecae dorsally in one specimen, the pore ca 0.15 mm wide. Ovary 10-14 locular, inferior, glabrous, the apex elevated into a collar around the style. Style 5-6 mm, strongly bending downwards, vertical distance from the anther to the stigma ca. -1 mm, horizontal distance ca. 1-1.5 mm; stigma consisting of 10-15 laterally compressed lobes but not evidently crateriform, 2-3 mm wide. Berry 8-9 × 8-9 mm, purple. Seeds ca. 0.6 mm, pyramidal, smooth. Distribution (Fig. 63). Puebla and Veracruz, Mexico, 800-1600 m. Conostegia arborea can be recognized by its leaves with dentate margins, abaxial indument of sessile and stipitate stellate hairs and especially by its apiculate calyptra apices. Th e amount of indument on floral buds is variable. Th e flowers of C. arborea have been reported to have a good fragrance (Ventura 1141-NY).   Schnell (1996), K!, TCD!).

Conostegia balbisiana
Description. Shrubs and trees 2-12 m tall with thick strongly tetragonal glabrous stems; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petiole 1-6.6 cm. Leaf blades 5-18 × 2.9-9 cm, 3-5 nerved or if 3-5 plinerved, with the innermost pair of primary veins arising up to about 1 cm above the base, ovate, rounded at the base, acute to rounded and short acuminate at the apex, the margin entire, glabrous. Inflorescence a terminal panicle 8.5-19 cm branched well above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches apparently absent, bracts early deciduous, bracteoles 1-2 mm long, subulate, deciduous or persistent. Pedicels 7-10 mm. Flowers (5-)6(-7) merous, calyptrate. Flower buds 11-17 × 5.5-7 mm, elliptic pyriform, the base rounded, the apex acuminate and sometimes mucronate, constricted in the middle, hypanthium 4-4.5 × 5.5-7 mm, glabrous. Petals 12-20 × 7-11.5 mm, pink, obtriangular, emarginate, glabrous. Stamens 12-14, 7-9.5 mm long, androecium slightly zygomorphic, the filament 3.5-4.5 mm, not geniculate, apparently white to pink, anthers 2.85-4 × 1-1.5 mm, narrowly elliptic, sagittate at the base, laterally compressed, yellow, thickened dorsally and with a small hump in dried material, the terminal pore ca. 0.1 mm wide. Ovary 6-9 locular, inferior, glabrous, forming a collar around the style base; the style 6-8 mm, bent below the tip, vertical distance from the anthers to the stigma ca. -0.6 --0.2 mm, horizontal distance apparently very reduced to absent, stigma truncate, 0.5-0.75 mm wide. Berry ca. 6-7 × 8-9 mm, blue-black. Seeds ca. 0.8 mm, obliquely pyramidal, smooth. Distribution (Fig. 64). Endemic to Jamaica, 350-950 m elevation. Conostegia balbisiana can be recognized by its leaves which usually have tufted mite domatia on the leaf base abaxially, large pink flowers and linear anthers. Schnell (1996) discussed variation observed in the three Jamaican endemics and struggled with their recognition. Perhaps population genetic studies might help untangle how many species should be recognized within this Jamaican lineage. For the time being, the three species recognized by Schnell (1996), who studied the most specimens of these taxa, are here recognized. Th e short style characteristic of section Conostegia is evident in specimens Proctor 10432 and 10285 both at NY.  Description. Tree 6-16 m tall with grayish-brown bark peeling in large thick flakes, the somewhat tetragonal and ridged stems in newer branches glabrous or with inconspicuous underdeveloped stellate or dendritic trichomes; the nodal line evident mostly on young nodes. Leaves of a pair equal to somewhat unequal in length. Petioles 0.8-4 cm long. Leaves 5.5-18 × 2-6 cm, 3-5 plinerved, with the innermost diverging from the mid vein 0.5-1.5 cm above the base in opposite or sub opposite fashion, the outermost primary veins usually inconspicuous and resulting in a mostly 3 veined looking leaf, narrowly elliptic, acute at the base, the apex acuminate, the margin entire, glabrous on both surfaces. Inflorescence a terminal panicle 5-9 cm long branching above the base, accessory branches present, bracts to 4 cm, early deciduous absent on most specimens, bracteoles ca. 0.5 mm long, linear, early deciduous and appearing absent on most specimens. Pedicel 3.5-4.5 mm long. Flowers 6-9 merous, calyptrate; flower buds 7.5-9.3 × 5-7 mm, obovoid, rounded at the base, obtuse to rounded and apiculate at the apex, slightly constricted below the calyptra, the hypanthial and calycine portions not or only slightly diff erentiated, constricted above the torus, hypanthium 4.5-5 × 4.5-5 mm, smooth. Petals 6.75-8 × 6.5-7 mm, white, narrowly obtriangular, spreading, emarginate, glabrous. Stamens 18-25, 7.75-9.25 mm long, radially arranged, the filament 4.75-5.25 mm, lacking a geniculation, white, anthers 3-3.5 × 1.25-1.75 mm, ovoid, laterally compressed, yellow, the base sagittate, the connective thickened dorsally and with a small bump, the pore 0.15 mm wide, terminal or subterminal. Ovary 8-12 locular, inferior, apically glabrous and forming a stylar collar. Style 6-6.5 mm, strongly bending downwards resulting in a evidently zygomorphic flower, vertical distance from the anther to the stigma ca. -1 mm, horizontal distance ca. 1-1.5 mm; stigma crateriform, consisting of 8-12 laterally compressed lobes, 3-3.3 mm wide. Berry 6-8 × 6-8 mm, when dry; seeds not seen. Conostegia bernoulliana was synonymized under C. icosandra by Schnell (1996) and Almeda (2009). I have chosen to recognize C. bernoulliana after studying populations in the field of C. icosandra that correspond with the type and one that corresponds to C. bernoulliana in addition to many herbarium specimens of C. icosandra. Th e main diff erences between the two species are the presence of indument in the stems and leaves, the broader 3-5 plinerved leaves, the persistent ovate bracteoles and non crateriform stigma of C. icosandra. In contrast C. bernoulliana has glabrous stems and leaves, the latter narrow and usually three plinerved, bracteoles very early deciduous to apparently lacking, and an evidently crateriform stigma. Schnell (1996)    Description. Trees 3-15 m tall with tetragonal and ridged, glabrous to sparsely furfuraceous-puberulent stems; the nodal line conspicuous and frequently elevated, with conspicuous lenticels abaxially. Leaves of a pair equal to somewhat unequal in length. Petioles 2.7-6.5 cm, with paired projections on the abaxial surfaces at the petiole/laminar junction. Leaf blades 17-29 × 9.2-20 cm, 5-7 nerved, elliptic to elliptic ovate, obtuse to broadly rounded, abruptly acuminate, the margins undulate-denticulate, adaxially glabrous,abaxially glabrous or sparsely furfuraceous-puberulent. Inflorescence a terminal panicle 13-22 cm long branching well above the base, with accessory branches, bracts not seen, bracteoles subulate, 1-2 mm long, deciduous. Pedicel 4-10 mm long. Flowers 8 merous, calyptrate. Flower buds 7-9.2 × 6.4-9.6 mm, white, globose, rounded to flattened at the base, rounded to slightly flattened at the apex, the hypanthial and calycine portions undiff erentiated, the hypanthium 5-6 × 8-9 mm, glabrescent and tuberculate. Petals 7-11.25 × 7-11 mm, pink, broadly obovate, rotate at anthesis, rounded-emarginate, glabrous. Stamens 22-26, 9-10 mm, radially arranged around the style but frequently secondarily zygomorphic because the stamens that are below the side that the style bends get stuck below the downward bent style and giant stigma, the filament 5-5.5 mm long, not geniculate, white, anthers 4-4.5 × 1.5-1.6 mm, elliptic, laterally compressed, the base sagittate, yellow, the pore 0.2-0.35 mm, terminal to slightly dorsally inclined. Ovary 15-17 locular, inferior, fluted into a glabrous dome. Style 7-8.5 mm long, bending downwards, vertical distance from the anther to the stigma ca.-0.5 --0.25 mm, horizontal distance ca. 1 mm; stigma crateriform and lobed, 5.5-6 mm wide. Berry and seeds not seen. Conostegia bigibbosa was synonimized under C. oerstediana by Schnell (1996) and subsequently recognized by Almeda (2009). I here recognize it based on geographic distribution and morphological characters. Conostegia bigibbosa always has pink petals as well as two tubercles on the petiole apex which makes it unmistakable. At one point, Schnell annotated it as a subspecies of C. oerstediana which is allopatric and has white petals and lacks the two tubercles at the apex of the petiole. Schnell (1996) carried out some crossing experiments between C. bigibbosa and C. oerstediana and C. montana. He noted that fruit development was initiated when pollinating C. oerstediana with pollen of C. bigibbosa but not with pollen of C. montana or when left unpollinated. Unfortunately he could not finish his experiment because the material was vandalized. Th e two species are not sympatric however and maintain the petal color diff erence as well as the petiole tubercle diff erence in natural populations.  Description. Shrub to small tree 1.75-4.6 m tall with terete or nearly terete stems that are moderately to densely covered with an indument of simple or little branched hairs; the nodal line present but usually covered by indument. Leaves at a node equal to unequal in length. Petiole 0.4-2.1 cm. Leaf blades 5.5-18.5 × 2-7.2 cm, 3-5 nerved or slightly plinerved, narrowly elliptic to oblanceolate or narrowly obovate, the base attenuate to obtuse, the apex acuminate, margin entire to dentate, adaxially setose with spreading smooth hairs 1-2 mm long, abaxially moderately setose on the actual surface below with a mixture of smooth, barbed and stellulate hairs on the elevated primaries. Inflorescence a terminal panicle 3.5-9 cm long branched above the base, accessory branches present, the inflorescence rachis moderately to densely covered with an indument of simple or little branched hairs, elliptic to oblong, bracteoles 6-8 × 2-2.5 mm, persistent. Flowers sessile, (5-) 6 (-7) merous, calyptrate, buds 6-8 mm long, pyriform to ovoid or broadly ellipsoid, rounded at the base, short apiculate at the apex, the calyptra and hypanthium little diff erentiated, the hypanthium 3.5-3.75 × 3.5-4 mm moderately to densely covered with an indument of simple or little branched hairs. Petals 6-7 × 4-5 mm, white or pink, obtriangular, spreading, emarginate apically, glabrous on both surfaces, spreading. Stamens (12-)14-18, 5-6.5 mm, slightly zygomorphic, the filament 2.5-2.75 mm, white, anthers 2.5-2.75 × 0.5-0.75 mm, linear and somewhat laterally compressed, yellow, the base sagittate, the pore 0.1-0.16 mm, subterminal to ventral. Ovary 6-7 locular, inferior, fluted into a glabrous collar around the style. Style 3.5-4.25 mm, slightly curving upward, vertical distance between the anther and stigma ca. -1 mm, horizontal distance absent; stigma capitate, 1-1.5 mm wide. Berry 6-8 × 6-8 mm, dark purple-black. Seeds 0.4-0.6 mm, pyramidal, smooth. Distribution (Fig. 70). Nicaragua to Colombia, 0-1150 m in elevation.

Conostegia bracteata Triana
Conostegia bracteata is one of the most distinctive species in the genus based on the presence of a hirsute indument on most parts of the plant including adaxial leaf surface, and large, persistent bracts subtending the sessile flowers. Flowers have been observed to be buzzed by several types of bees including Euglossines (Fig. 4). Speci- mens of this species dry red when treated with ethanol. Schnell (1996) argued that C. bracteata provided an excellent example of convergent evolution between this species and Miconia barbinervis and M. cuneata in their "pubescence, foliage, and the general aspect of flowers, bracts and inflorescences". Th ese species look a lot alike in their leaf shape and pubescence but not in their flowers.     Schnell (1996).
Conostegia brenesii is a very distinctive and narrow endemic of middle elevation cloud forests in Costa Rica. It can be easily distinguished by its dense indument of stipitate stellate hairs on all plant parts. Because of its dense indument of stipitate stel-late hairs it is similar to C. caelestis which is allopatric occurring in northern Central America. In addition, C. brenesii tends to be a shrubby species whereas C. caelestis tends to be a larger tree. Th e flowering time diff ers with C. caelestis flowering in the first half of the year and C. brenesii flowering in the second half of the year consistently from July to September. Th e molecular phylogeny does not place these species as sister taxa, which suggests convergent evolution in the dense stipitate stellate indument (  Description. Small trees 2-12 m tall with tetragonal stems that soon become terete and are covered with simple and mostly stellate-stipitate trichomes to 1.5 mm long; the nodal line present but slight. Leaves of a pair equal to somewhat unequal in length. Petioles 0.6-3.8 cm long. Leaves 5.5-26 × 2-7.8 cm long, 3-5 nerved or 3-5 plinerved, if plinerved, the innermost pair of primary veins arising up to 1 cm above the in opposite or subopposite fashion, elliptical to obovate, the base acute or cuneate, the apex acute to acuminate, the margin entire to serrate, adaxial surface short-setose with simple bristles and sometimes stipitate stellate trichomes on the mid vein, abaxially covered with stipitate stellate hairs. Inflorescence a terminal panicle 3.9-9 cm long branched above the base, accessory branches absent, bracts linear, up to 5 mm long, deciduous, bracteoles 2-10 mm long, deciduous. Flowers 5-6 merous, calyptrate. Flower buds 5.8-7.6 × 2.9-3.5 mm, pyriform oblong, the base flat to rounded, the apex acute to apiculate, not constricted, hypanthium 3-3.25 × 3-3.25 mm, covered with stipitate stellate hairs. Petals 6-10 × 6.5-7 mm, white, obovate, not observed at anthesis, glabrous, 3 lobed. Stamens (14-)16-18, slightly zygomorphic, the filament 2.5-3 mm, not geniculate, anthers 2-2.5 × 0.3-0.6 mm, linear, reportedly white, somewhat laterally compressed, the base sagittate, the pore terminal, 0.1-0.15 mm. Ovary 7-10 locular, inferior, forming a glabrous collar around the base of the style. Style 3.5-4 mm, bent near the tip, vertical distance from the stigma to the anthers ca. -0.6 mm, horizontal distance absent; stigma subcapitate, 0.9-1.1 mm wide. Berry 6-7.5 × 6-7.5 mm, purple-black. Seeds 0.33-0.5 mm, pyramidal, smooth. Distribution (Fig. 74). Known from Mexico through Guatemala and Belize to Honduras from sea level to 1000 m in elevation.

Conostegia caelestis Standl.
Conostegia caelestis is easily distinguished from its congeners on the basis of its dense indument of stipitate stellate hairs especially on the leaf abaxial surface, inflorescence and hypanthia. As Schnell (1996) noted, this species has a very definite flowering time and is never fruiting at the same time that it is flowering, suggesting a long maturation time for the berries. Conostegia caelestis shares the dense indument of stipitate stellate hairs with C. brenesii but they are not closely related. See further discussion of their diff erences under C. brenesii.  Description. Trees 4-20 m tall and with tetragonal and ridged stems that are glabrous or sometimes with scattered sessile stellate trichomes; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 1-4.8 cm long. Leaf blades 6.2-16.5 × 3.2-7.7 cm, 3-5 nerved or slightly plinerved, elliptic, base obtuse to rounded, apex obtuse to acute and short acuminate, the margin entire to denticulate, essentially gla- brous on both surfaces. Inflorescence terminal, 5.7-13.5 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, bracts absent or very early deciduous, the bracteoles 1-5 mm, deciduous. Pedicels 3-6 mm. Flowers 7-11 merous, calyptrate. Floral buds 7.2-13 × 3.2-7.6 mm, mostly ellipsoid pyriform, constricted below the middle, the base flat to rounded, apiculate at the apex. Petals 10-12 × 5-6 mm, white to pale lavender, obtriangular, spreading, the apex rounded-truncate to emarginate, glabrous. Stamens 14-24, 8.5-10 mm long, androecium zygomorphic, the filament 4.25-5.25 mm, white but apparently turning red on some specimens perhaps when old, anthers 3.25-4.5 × 0.1-0.2 mm, subulate and slightly recurved, sagittate at the base, yellow except for a hugh of rose at the base of thecae dorsally in one specimen, the pore ventral-terminal, ca. 0.3 mm wide. Ovary 6-12 locular, inferior, glabrous and lacking a distinct apical collar. Style 9-12 mm, bent away from the stamens, vertical and horizontal distance not assessed, stigma barely expanded, made of lobes that are almost non discernible, ca. 1-2 mm wide. Berry 6-7 × 6-7 mm, blue-black or purple. Seeds 0.5-0.65 mm, pyramidal and smooth. Distribution (Fig. 75). Endemic to cloud forests in Costa Rica and Panama from 1000-2100 m. In Panama restricted to peaks near the Canal Area as well as Volcan Chiriquí. In Costa Rica common in Las Tablas Protected Zone on the Costa Rica-Panama border.

Conostegia chiriquensis
Conostegia chiriquensis is similar and possibly closely related to C. pittieri, especially in their glabrescence, apiculate calpytras and style lacking a conspicuous crater in the middle. Conostegia chiriquensis diff ers most notably in the more slender style that is not capitate like in C. pittieri. Schnell (1996) noted that the petals and particularly the style are persistent in C. chiriquensis and this diff ers from C. pittieri where only the petals tend to persist. In the protologue Gleason compared this species to C. rhodopetala and C. pittieri (Gleason, 1941). Conostegia rhodopetala can have an apiculate calyptra but has a noticeably short style lacking lobes. In Schnell's (1996) key C. chiriquensis is included in the group of species with large stigmas mostly because of the presence of lobes since it is barely expanded. Almeda noted "Gardenia like fragrance on flowers of this species NY). Schnell (1996)   Description. Shrubs to small trees 2-12 m tall with strongly tetragonal stems that are glabrous or with scattered sessile stellate trichomes; the nodal line inconspicuous or evident as a whitish line, not elevated. Leaves of a pair equal to somewhat unequal in length. Petioles 0.9-6 cm. Leaf blades 11-25.7 × 5-12.9 cm, 3-5 plinerved, with the innermost pair of vein arising just above the base in opposite or sub alternate fashion, elliptic to ovate, base acute to rounded, apex acuminate, margin entire to denticulate, glabrous on both surfaces. Inflorescence a terminal panicle 6-25 cm long branched above the base base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches absent or present, the bracts absent or early deciduous, bracteoles to 6 mm long, subulate, usually persistent at anthesis and deciduous in fruit. Pedicel 5-8 mm, lengthening in fruit. Flowers 6-8 merous, calyptrate. Floral buds 6-11 × 3-7 mm, slightly constricted at the torus, the base flat, the apex acuminate; the hypanthium 5-6 × 5-5.5 mm. Petals 7-12 × 7-11.5 mm, pink to lilac or white, obovate, spreading, apically emarginate, glabrous, spreading, the margin entire to undulate. Stamens 12-15(-17), 9-10 mm long, slightly zygomorphic, the filament 5-5.5 mm, white, anthers 4-4.5 × 0.8-1.2 mm, linear-oblong and recurved, somewhat laterally compressed, sagittate at the base, yellow, the pore 0.12-0.13 mm, slightly ventrally inclined. Ovary (6-)7-9(-11) locular, inferior, the apex glabrous and elevated into a pronounced collar around the style base. Style 4-6.7 mm, bent upward below the stigma, distance between the anther and the stigma -2 --0.5 mm, stigma broadly capitate, 0.3-0.5 mm wide. Berry 8-9 × 8-9 mm, purple-black. Seeds ca. 0.4 mm long, triangular in profile view, smooth. Distribution (Fig. 77). Ranging from Panama to Colombia, Venezuela and Ecuador, and the coastal range of Venezuela, 0-1000 m in elevation.
Th is species is particularly distinctive when found in the field with its large lavender flowers. Herbarium specimens with flower buds on the other hand can be hard to separate from some populations of C. superba. In general both can be separated on the basis of the larger flowers with usually lavender petals and more floral parts in C. cuatrecasii (versus smaller flowers with white petals in C. superba). Also, C. cuatrecasii tend to have a more markedly acute calyptra apex than C. superba. Schnell (1996) further separated the two stating that the stigma in C. cuatrecasii was lobed but I did not observe lobes in a specimen collected in spirit in Cerro Jefe, Panama. Also, Schnell (1996) stated the floral buds in C. superba are clearly constricted but I did not observe constriction in the floral buds of C. superba in Izabal, Guatemala. Good fragrance has been reported in the flowers of this species     Description. Shrubs to small trees 4-11 m tall with apically tetragonal glabrous stems; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 0.7-3 cm long. Leaf blades 3.5-10 × 1.2-4 cm, 3-5 plinerved, with the innermost pair of veins diverging from the mid vein up to 5 mm above the blade base, mostly elliptic, glabrous adaxially, glabrous or with some scattered minute trichomes abaxially, the base acute, the apex acuminate to caudate acuminate, margin entire. Inflorescence a terminal panicle 3-10 cm long branched well above the base, accessory branches apparently absent, the rachis glabrous or with some minute furfuraceous lepidote hairs, bracts apparently early deciduous, not observed, bracteoles 0.5-2 mm, narrowly triangular to subulate, early deciduous. Pedicel 2-7 mm long. Flowers 6-7 merous, calyptrate. Flower buds 5-7 × 2-3.25 mm, oblong-ellipsoid, the base rounded, the apex acute to apiculate, slightly constricted below the calyptra; the hypanthium 2.75-3.25 × 3.5-4 mm, glabrous or sparsely furfuraceous lepidote. Petals 4.5-5 × 4-4.5 mm, white with a red band at the base, obovate, reflexed to slightly spreading, glabrous, the apex three lobed. Stamens 14-19, 4-5.5 mm long, zygomorphic resulting from their bending all to one side, the filament 2.5-3.5 mm, white, non geniculate, anthers 1.5-2.3 × 0.65-0.75 mm, linear-oblong, sagittate at the base, yellow-orange, the connective thickened and forming a slight hump, the pore ventrally inclined, 0.25 mm wide. Ovary 6-7 locular, inferior, the apex glabrous and lacking an elevated collar.
Style 5-5.5 mm, bending opposite the stamens, vertical distance between the anther and the stigma absent, horizontal distance 2-2.3 mm, the stigma capitellate, with 6 or 7 lobes, 1.4-1.5 mm wide. Berry 3 × 3.5-4 mm, purple black. Seeds 0.5-0.75 mm, oblong or narrowly pyramidate, smooth. Distribution (Fig. 79). Known from south eastern Costa Rica and west Panama, 1200-1700 m elevation. Schnell (1996) and Almeda (2009) Kriebel and D. Solano 3174. in several living populations studied, or was there an evident style declination as is present in C. fragrantissima. My observations contrast with Schnell's because he stated most populations of C. montana do present a lobed stigma and a declinate style. Although Schnell (1996) makes a good point in stating that collectors tend to overlook fragrance, C. montana is one of the most frequently collected species of Conostegia and none of the specimens studied cited floral fragrance. On the other hand, of the only six specimens of C. fragrantissima, one describes evident fragrance MO,NY). Th e stigma lobes present in C. fragrantissima as well as the declinate style (Fig. 78) also suggest a possible relationship to C. pittieri. Unfortunately this species was not included in the phylogeny of Conostegia and thus its close relatives remain unknown.  Description. Shrubs to small trees 1.8-10 m tall with tetragonal stems becoming terete with age and that are finely puberulent with minute sessile stellate and stipitate-stellate trichomes; the nodal line present but slight. Leaves of a pair equal to somewhat unequal in length. Petioles 0.8-3.2 cm long. Leaves 6.5-16.1 × 1.75-5.7 cm, 3-5 nerved or if plinerved, with the innermost pair of primary veins diverging from the mid vein up to 2 cm above the base in opposite to sub opposite fashion, adaxially glabrous, abaxially with stipitate stellate hairs mostly on the primary veins and with minute stellate trichomes on higher order veins, some specimens with evident pocket domatia at the base abaxially, narrowly ovate to narrowly elliptic, acute at the base and apex, the margins undulate-dentate to entire. Inflorescence terminal panicle 4.3-11.8 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, bracts and bracteoles 0.6-4 mm long, subulate to linear-lanceolate, deciduous. Pedicels 2-5 mm long. Flowers 5-6 merous, calyptrate. Flower bud ca. 4-6 × 2.5-3.5 mm, obovate, obtuse or rounded at base, acute to short-apiculate at the apex, not constricted, the hypanthium 2.5-3 × 2.5-3 mm, smooth. Petals ca. 5 × 4 mm, white or pink, spathulate, emarginate, glabrous. Stamens 13-17, 3-4 mm long, the filament ca. 1.5-2 mm, white, anthers 1.8-2.0 × 0.4-0.5 mm, linear-oblong, straight or slightly recurved, laterally compressed, thickened dorsally, yellow, the pore ca. 0.2 mm, terminal. Ovary 5-6 locular, inferior, apically glabrous and forming a collar around the style base. Style ca. 3 mm, straight distance between the anther and the stigma -0.5 --0.1 mm; stigma punctiform, ca. 0.6 mm wide wide. Berry ca. 5 × 5 mm, dark purple. Seeds 0.4-0.6 mm long, obovoid, angulate or not, essentially smooth but frequently with the periclinal walls elevated to give a roughened look. Distribution (Fig. 80). Known from Guatemala, Honduras and Nicaragua, 700-1700 m.
Conostegia hirtella is quite similar to C. montana, a very variable species. It is recognized here on the basis of the presence of stipitate stellate hairs which C. montana lacks. Schnell (1996) considered C. hirtella to be close to C. caelestis on the basis of those stipitate stellate trichomes. Th e much sparser pubescence in C. hirtella compared to the dense indument in all the plant in C. caelestis makes it diff cult to confuse them. Unfortunately C. hirtella was not included in the molecular phylogeny to further help elucidate its close relatives.
Specimens examined. GUATEMALA. Description. Shrubs to trees 1-15.3 m tall with somewhat tetragonal and ridged stems that are glabrescent to hirsute with sessile and stipitate stellate as well as branching hairs; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 1-6.9 cm long. Leaves 4.6-25.2 × 1.5-11 cm, 3-5 plinerved, with the innermost diverging from the midvein just above the blade base in opposite or alternate fashion, elliptic to ovate, acute to rounded at the base, apex acute to acuminate, the margin entire to denticulate, adaxially generally glabrous, abaxially glabrous to densely hirsute with sessile or stipitate stellate and branching trichomes. Inflorescence a terminal panicle, 2.8-18 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches absent or present, the axis glabrous to hirsute with sessile and stipitate stellate, bracts linear to elliptic, up to 1.5 cm long, persistent or deciduous, bracteoles 1-10 mm long, oblong to ovate, mostly persistent. Pedicel 1-10 mm long. Flowers (5-)6-9(-11) merous, calyptrate. Floral buds 5-11 × 4.5-8.5 mm, rounded at the base, obtuse to rounded and apiculate at the apex, not constricted, the hypanthial and ca-lycine portions not or only slightly diff erentiated; the hypanthium 3.5-4.5 × 5-5.5 mm, glabrous to puberulent. Petals 7-8 × 7-8 mm, white, broadly obovate, spreading and rotate, emarginate, glabrous. Stamens (17)19-26 (30), 6.5-9 mm long, somewhat zygomorphic, the filament 4.5-5 mm, white, anthers 3.27-3.75 × 1.5-2 mm, oblong, laterally compressed, the base sagittate, yellow, the pore 0.1-0.15 mm, terminal or subterminal. Ovary 8-15 locular, inferior, apically glabrous and forming a stylar collar. Style 5.5-6 mm, slightly to strongly bent, vertical distance from the anther to the stigma ca. -0.25, horizontal distance 0.5-1.5 mm; stigma capitate, consisting of 8-15 lobes that are diffi cult to distinguish, not crateriform, the stigma 3-3.5 mm wide. Berry 9-12 × 7-10 mm, blue-black or purple. Seeds 0.4-0.8 mm long, obovoid, smooth. Distribution (Fig. 82). Ranging from Mexico throughout Central America and the Caribbean, reaching the coast of Venezuela, sea level to 1500 (-2000) m. Conostegia icosandra is variable in the amount of indument and degree of dentition of the leaf margin. Despite this variation, the species has usually been circumscribed as having conspicuous persistent bracteoles (Fig. 81). Th ese bracteoles are evident in populations mostly north of Nicaragua but in other populations of this species which were synonimized in Schnell (1996) and Almeda (2009) under C. icosandra, the plants are glabrous, the leaves narrower and the bracteoles missing altogether. Th ese specimens match the name C. bernoulliana which is here considered a distinct species. In addition, the style in C. icosandra can lack the large central hole that is evident in C. bernoulliana. Th e distinction of these two taxa is relevant for understanding the evolution of crateriform stigmas since the true C. icosandra falls as sister to the rest of the lobed stigma clade, suggesting the crateriform stigma was possibly not lost in C. icosandra whereas it was probably lost in C. pittieri. Th e specimen Davidse, González and León 18607 (NY) from Zulia, Venezuela, unlike any other specimens studied has extremely narrow leaves.   Description. Shrubs 2-4 m tall with young tetragonal stems which become terete with age that are glabrescent to finely and sparsely furfuraceous; the nodal line present. Leaves of a pair equal to unequal in length. Petioles 0.4-1.2 cm. Leaf blades 6-17 × 2-6 cm, 3-5 plinerved, with the innermost pair of veins diverging from the mid vein 1-2 cm above the base, elliptic, the base acute, the apex acute to acuminate, adaxially glabrous or sparsely ciliate, abaxially pubescent with stellate trichomes on the veins, the margins serrulate and ciliate. Inflorescence a terminal panicle 3-7 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches apparently absent, branches flattened; bracteoles linear to ovate, 2 mm long, early deciduous and appearing absent. Pedicels 2-5 mm. Flowers (5-)6-7 merous, calyptrate. Floral buds 10-15 × 4-7.1 mm, slightly ovoid to elliptic, subacute or rounded at the base, acute to acuminate at the apex, slightly constricted below the middle, the calyptra and base weakly diff erentiated; hypanthium 4.5-5.5 × 5-6 mm, glabrescent with inconspicuous sessile stellate hairs. Petals ca. 7-8 × 5-6 mm, white, broadly spatulate, probably spreading, glabrous. Stamens 15-20, 7 mm long, the filaments 3-4 mm long, white, anthers 3.5-4 mm long, linear-subulate, recurved, yellow, the pore not observed, terminal. Ovary 6-7 locular, inferior, glabrous, and elevated into a collar around the style base. Style 4-6 mm, curving and not widening below the stigma, bent beneath the tip, apparently no vertical or horizontal distance between the anthers and the stigma; stigma subcapitate, ca 1 mm wide. Mature berries not seen. Distribution (Fig. 83). Endemic to Mexico where it occurs in the Sierra Madre in the states of Jalisco and Guerrero from 700-1600 m in elevation.
Conostegia jaliscana is a rare species reported usually from alongside streams. It has few-flowered inflorescences and relatively long and acute to acuminate floral buds. Schnell (1996) placed C. jaliscana with the three Jamaican endemic in his section Conostegia but also hinted to the possibility of C. jaliscana being closely related to species in his subgenus Lobatostigma. Th is latter hypothesis is the one supported in the molecular phylogeny.
Specimens Description. Shrubs or small trees to 6 m tall, stems tetragonal but soon terete and densely pubescent with sessile and stipitate stellate hairs; the nodal line present yet slight. Leaves of a pair equal to subequal in length. Petiole 0.8-2.5 cm long. Leaf blade 6-18 × 2-6 cm, 3-5 nerved, narrowly ovate to elliptic or ovate, acute to rounded at the base, acute to acuminate or rarely obtuse at the apex,the margins entire or obscurely denticulate, adaxially glabrous, abaxially densely covered with stellate and stipitate trichomes. Inflorescence a terminal panicle 6-12 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches apparently absent; bracts and bracteoles to 3 mm long, linear, early deciduous. Pedicels 2-7 mm long, covered with stellate and branching trichomes. Flowers (4-)5(-6) merous, calyptrate. Floral buds 8-9 × 3-4  mm, pyriform to lachrimiform, truncate or rounded at the base, narrowly acute and acuminate at the apex, the apex with somewhat discernible lobes, the calyptra well differentiated from the hypanthium; hypanthium 4-5 × 3.5-4 mm, campanunlate, sparsely to densely beset with stellate hairs and minute brown glands. Petals ca. 5 mm long, white, obtriangular, spreading to a little reflexed, rounded apically, glabrous. Stamens 9-14, 3-3.5 mm long, radially arranged around the style but apparently becoming zygomorphic becasuse of the style bending to one side below the stigma, the filament ca. 1.5 mm long, white, lacking a geniculation, anthers 1.5-2 mm, linear and slightly recurved, yellow, the pore ca. 0.1 mm, terminal. Ovary 5-7 locular, inferior, apically glabrous and forming a low collar around the style base. Style 2-3 mm long, bent below the stigma, vertical and horizontal distance from the stigma to the anther absent; stigma truncate, ca. 1 mm wide. Berry 5 × 5 mm, blue black. Seeds 0.4-0.5 mm, ovoid, smooth. Distribution (Fig. 85). Cuba and the Dominican Republic, 750-1300 m elevation. Note that in Dominican Republic a pubescent morphotype of C. superba has been collected in the same place. Th ey are similar but when looking at the flower buds, C. lindenii has pubescent buds with the lobes somewhat evident in the calyptra apex. In C. superba on the other hand, the flower bud and calyptra are glabrous and the lobes not discernible at all.  Schnell (1996): C, GH, as well as a photo of a lost specimen from B at F!).
Description. Trees 3.5-15 m tall with thick tetragonal and ridged stems that are glabrous or furfuraceous on new growth with sessile stellate trichomes; the nodal line conspicuous and sometimes setulose in young branches, lenticellate abaxially. Petioles 1-7 cm long. Leaves of a pair equal to somewhat unequal in length. Leaf blades 6.7-30 × 2.3-15 cm, 5 nerved or slightly plinerved, ovate to elliptic, the base acute to obtuse, the apex obtuse and acute to acuminate, the margin entire or denticulate, adaxially glabrous, abaxially with branched and stellate hairs on the veins. Inflorescence a terminal panicle 6-21.2 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, rachis tetragonal, accessory branches present, bracts early deciduous or absent, the rachis glabrous or furfuraceous with sessile stellate trichomes these sometimes minute and inconspicuous, bracteoles linear, ca. 2 mm long, deciduous. Pedicels 4-15 mm, frequently nodding. Flowers 7-10 merous, calyptrate. Floral buds 8-15 × 7-13.5 mm, spherical, the base rounded, the apex rounded and mucronate, not constricted, the hypanthial and calycine portions little diff erentiated; hypanthium 9.5-10 × 13-14 mm, glabrous or beset with small sessile stellate trichomes, strongly tuberculate. Petals 9.5-16 × 12-14.5 mm, white, obovate, spreading, the apex retuse, glabrous. Stamens 28-45, 9-11 mm long, radially arranged around the style, occasionally secondarily zygomorphic resulting from some stamen getting stuck below the stigma, the filament 5.25-5.7 mm, white, anthers 4.5-5 × 1.75-2.25 mm, oval, yellow, the base sagittate, strongly laterally compressed, the pore 0.2-0.3 mm, terminal. Ovary 18-25 locular, inferior, the apex glabrous and forming a collar around the style; style 7-8 mm long, usually straight but sometimes slightly curving, distance from the anther to the stigma ca. -2 -0 mm, stigma crateriform, consisting of 18-25 laterally compressed lobes, 6.5-8 mm wide. Berry 14-18 × 10-12 mm, purple. Seeds 0.5-1 mm long, narrowly ellipsoid, the testa smooth. Distribution (Fig. 87). Endemic to Costa Rica, 1300-3000 m elevation. Conostegia macrantha can be recognized by its stout branches, stellate indument on abaxial leaf surface, and large flowers with crateriform, straight styles, and retuse petals. Flowers of this species have a good fragrance (i.e., Chavarría 817, Jiménez 626-both at MO). Th is species is also reported from northern Panama by Schnell (1996) but I have not seen any Panamanian specimens. Th e few specimens from Panama that resemble Conostegia macrantha are glabrous, the reason why I did not include them in the distribution of this species presented here. Schnell (1996) noted that the species is especially common in the volcanoes surrounding the central valley of Costa Rica and is mostly restricted to that area. Specimens collected near Barva and Poas volcano for this study have the typical straight styles of C. macrantha. Schnell (1996)  Description. Shrubs to trees 1.5-10 m tall with tetragonal to terete stem which are densely tomentose with sessile stellate hairs; the nodal line present yet slight. Leaves of a pair equal to subequal in length. Petioles 0.5-6 cm long. Leaves 5.7-21.5 × 2-10.5 cm, 3-5 nerved or slightly plinerved, ovate-elliptic to ovate, the base acute to obtuse, the apex acute to acuminate, the margins entire or remotely denticulate, the adaxial surface glabrous, the abaxial surface densely tomentose with sessile stellate hairs. Inflorescence a terminal panicle 5-12.5 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present or absent; bracteoles to 3 mm long, linear, early deciduous. Pedicels 0.5-3 mm long. Flowers (4-)5(-6) merous, calyptrate. Floral buds 2.5-6.5 × 1.5-3.5 mm, obovoid pyriform, the base rounded, the apex rounded to acute or or short apiculate, slightly constricted below the calyptra; the hypanthium 3-3.25 × 2.35-2.85 mm, with scattered stellate trichomes. Petals 3.5-5.25 × 2.5-3 mm, totally white with or white with pink or violet bases, oblong or broadly ovate, spreading to somewhat reflexed, glabrous, entire. Stamens mostly 12-18, 3.5-5.5 mm long, slightly zygomorphic, the filament 1.5-2.5 mm, white, 1.8-2.5 × 0.5-0.75 mm, linear to slightly sinuous, cream to yellow, the pore ca. 0.1 mm, subterminal and slightly ventrally inclined. Ovary 5-7 locular, inferior, glabrous, lacking an elevated collar around the style base. Style 3-4 mm, straight but bending slightly towards the apex, vertical distance between the anther and the stigma ca. -0.5 mm, horizontal distance absent; stigma capitellate, 1-1.5 mm wide. Berry 5-6 × 5-6 mm, purple black. Seeds 0.3-0.6 mm long, pyramidal, the testa smooth to slightly foveolate. Distribution (Fig. 89). Nicaragua to Panama and in Ecuador, 100-1200 m elevation.
Conostegia micrantha is very similar to C. montana from which it can de distinguished on the basis of the dense indument of stellate trichomes on the stem apices, the abaxial surface of the leaves, and the inflorescence. Some populations of C. montana on the Caribbean islands as well as in Central American highlands can have stellate trichomes, complicating their distinction. When sympatric, such as in La Selva Biological Station in Costa Rica, C. montana is glabrous and has narrower leaves. Schnell (1996) discussed the possibility that this species is derived from C. montana. Th is is indeed a possibility supported also by the molecular phylogeny where C. micrantha falls in a clade of several specimens of C. montana. Th e question does remain as to whether C. montana should be considered a single species.
Among the variation of this species there are populations in the mountains of Guatemala and Costa Rica that have two small knob-like structures at the apex of the petioles. In general this species tends to have some pubescence but never as dense and with as well defined stellate trichomes as typical C. micrantha. Other populations such as the one at La Selva Biological Station in Costa Rica are glabrous and symmpatric with typical C. micrantha. Other variants include leaves with conspicuously serrate margins such as a population in Cerro Hornito in Panama, and one population in Cerro Jefe, also in Panama, has very coriaceous leaves. For similarities to C. vulcanicola, see discussion under the latter. Th e molecular phylogeny  including several specimens of diff erent phenotypes of C. montana from Guatemala, Costa Rica, Panama, and Jamaica revealed a clade with good support which included all the latter accessions, but also included C. micrantha and C. setosa nested within them. Th is suggests C. montana is actually paraphyletic indicating this species complex is in need of more work. One species that is recognized in this revision and which Schnell (1996) considered a synonym of Conostegia montana is C. fragrantissima. For a discussion of the diff erences between both, see the discussion under the latter species. Schnell (1996)   Description. Shrubs to small trees to 3.5 m tall with slightly to evidently tetragonal stems towards the apex, essentially glabrous or with inconspicuous scales; the nodal  line present. Leaves of a pair somewhat unequal in size. Petioles 1.7-7.5 cm long. Leaf blade 18-27 × 6-13.5 cm, 3-5 nerved, elliptic to elliptic-ovate, the base acute or obtuse, the apex acute to abruptly acuminate, the margin entire, both surfaces glabrous but inconspicuously glandular puncticulate. Inflorescence a terminal, elongated and deflexed or arching panicle 8-30 cm long branching well above the base, accessory branches absent or present, rachis glabrous, bracts and bracteoles to 1 mm long, linear, early deciduous. Pedicels 1.5-2.5 mm. Flowers 5 merous, pyriform, calyptrate. Flower buds 5-9 × 3-4.5 mm, the base rounded to obtuse, the apex acute to acuminate, constricted below the calyptra, the calyptra and hypanthium diff erentiated; the hypanthium 2-2.5 × 2.75-3.15 mm, campanulate, glabrous and inconspicuously glandular puncticulate. Petals 5-7 × 4.5-6 mm, white to lavender, obovate, glabrous, emarginate. Stamens (9-)10, 4.5-5.5 mm long, slightly zygomorphic, the filament ca. 3 mm, white, anthers 2.25-2.75 × 0.6-0.9 mm, linear-oblong, yellow or pale yellow, the pore ca. 0.1 mm, subterminal and ventrally inclined. Ovary 5-6(-7), inferior, apically glabrous and lacking a collar around the style base. Style 3-3.5 mm, straight and bending below the tip, vertical and horizontal distance from the anther to the stigma ca. -1.25 --0.5 mm, stigma capitate, 1-1.5 mm wide. Berry 5-6 × 5-6 mm, dark purple. Seeds 0.4-0.6 mm, ovoid to pyramidal, the testa muriculate. Distribution (Fig. 94). Endemic to the Caribbean slope of south eastern Costa Rica and northern Panama, 40-1200 m in elevation.
Conostegia muriculata is a distinctive species recognized by being overall glabrous, pendant inflorescence, purple petals and muriculate seeds. Unfortunately the phylogenetic placement of this species remains unknown. Because of its small flower buds and being almost totally glabrous, it likely belongs in the C. montana complex in section Conostegia. As in all species in section Conostegia, the style is short (Fig. 93). Schnell (1996)  Description. Trees 4.5-18 m tall with whitish flaky bark and tetragonal and ridged stems that are glabrous or with sessile stellate trichomes, these usually minute and inconspicuous; the nodal line present but frequently inconspicuous, lenticels frequent at the nodes. Leaves of a pair equal to somewhat unequal in length. Petiole 1.3-9 cm. Leaf blade 5.2-25 × 2.9-15 cm, 3-5-plinerved, with the innermost diverging from the mid vein just above the blade or rarely up to 3 cm above the base in opposite or alternate fashion, ovate or elliptic, thick, the base acute to rounded, the apex rounded and abruptly acute to acuminate, the margin entire to denticulate, glabrous on both surfaces. Inflorescence a terminal panicle 6-18 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, bracts absent or if present elliptic to ovate and up to 3 cm long, accessory branches present, the rachis glabrous, bracts apparently lacking and bracteoles early deciduous. Pedicels 1-8 mm. Flowers 6-10 merous (mostly 8), calyptrate. Flower buds 5-12 × 5-10 mm, spherical, the base rounded to truncate, the apex rounded to truncate and usually short apiculate, not constricted and undiff erentiated; the hypanthium 5-5.5 × 7-8.25 mm, smooth to more frequently tuberculate, glabrous and frequently evidently white. Petals 7-11 × 7-11 mm, white, obovate, spreading and overlapping, emarginate, glabrous, spreading and overlapping, entire. Stamens (20-)24-28(-36), 7-9 mm long, radial but appearing bilateral because stamens often get stuck below the stigma, the filament 4-5 mm, white, anthers 3-4 × 1-1.5 mm, elliptical to linear-oblong, yellow, strongly laterally compressed, the base sagittate, the pore 0.2-0.3 mm, terminal. Ovary 14-20 locular, inferior, apically glabrous. Style 5.75-8 mm, bending downward to ca. 45 degrees throughout anthesis and protruding below the anthers, vertical distance from the anthers to the stigma ca. 0 mm, horizontal distance ca. -2-0; stigma crateriform, consisting of 14-21 laterally compressed lobes, 4-6 mm wide. Berry 11-14 × 8-10 mm, blue-black or purpleblack. Seeds 0.6-0.8 mm, pyramidal, the testa smooth. Distribution (Fig. 96). Nicaragua to west Panama where it occurs in the mountains from (550)1000-2400 m elevation.
Specimens from Chiriquí have been annotated as C. macrantha (White 194, Penneys and Olmos 1738-both at NY). I have included these specimens in the circumscription of C. oerstediana on the basis of their glabrous leaves as well as on the description of the style in the second above mentioned specimen where the style was described as curving downward whereas in C. macrantha it is straight. Most populations have tubercles on the hypanthium but in some populations these are reduced or lacking. When the tubercles are lacking, C. oerstediana looks similar to C. bernoulliana. Th e latter lacks the spherical floral buds of C. oerstediana and instead has a constriction in the middle of the bud. Also, the leaves in C. bernoulliana are narrower. Conostegia oerstediana is also similar to C. bigibbosa. Th e latter name was considered a synonym of C. oerstediana. See the discussion under C. bigibbosa for the diff erences between the two species. See also Schnell (1996) for details on crossing experiments between C. oerstediana and close relatives. Flowers of this species have good fragrance which has been reported on several specimens (e.i.,N. Zamora 5843,5844,MO). Th ese flowers have been observed to be visited by Bombus volluceloides and other large unidentified bees. Berries of this species have a good taste. Schnell (1996) notes the almost exclusivity of growing on volcanic soils of this species and Umaña Dodero (1988) documented peak flowering in December for populations in Grecia, Costa Rica. Phoradendron chrysocladon A. Gray has been reported as a parasite of C. oerstediana (Braby and Nishida, 2010).    . Th e latter collection is labeled Pittier and Tonduz 290 in US, a discrepancy common with the exsiccatae of these men and apparently due to erroneous use of the printed labels bearing both their names. Handwritten label with both names were never seen, nor evidence that the two collected in the same place at the same time (Schnell 1996).
Conostegia pittieri is one of the few species of Conostegia that retains its petals once the stamens and style have fallen. Th e functional significance of petal retention, if any, might have to do with continuing to make inflorescences attractive (Bertin 1982).Th e similar species Conostegia chiriquensis also retains the petals and the style after the stamens have fallen. See the discussion under the latter species for diff erences between C. chiriquensis and C. pittieri. Flowers of C. pittieri have been observed to be buzzed by female Bombus volluceloides (pers. obs.). As Schnell (1996) mentioned, this species can be quite abundant in pastures and second growth but is rare inside the forest where it occurs mainly in riparian habitats. Individuals within the forest produce much less flowers (pers. obs.).   Description. Trees to about 7 m tall with subtetragonal, stems stellate pubescent apically, glabrous to glabrescent with age; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petiole 1-5 cm. Leaf blades 4-15.5 × 3.1-7 cm, 3-nerved, elliptic, the base obtuse, the apex obtuse, acute or acuminate, the margin entire or weakly crenulate-dentate near the apex, the adaxial surface glabrous, the abaxial surface glabrous except for tuft domatia of stipitate branching hairs present at the base of the leaf and minute puncticulate glands throughout the surface. Inflorescence a terminal panicle 5-17.7 cm long branched above the base, rachis flattened accessory branches absent or present, bracts and bracteoles linear, 2-3 mm long, deciduous or appearing absent. Pedicels 5-12 mm. Flowers 6 merous, calyptrate. Floral buds 11-18 × 5-7 mm, elliptic pyriform, the base rounded, apex acuminate and apiculate, slightly constricted in the middle, the hypanthium 4-5 × 4.5-5.25 mm, glabrous and ribbed. Petals 10-14 × 6-9 mm, white or pinkish white, obtriangular, spreading, emarginate, glabrous. Stamens (17-)18(-20), 6-7.5 mm long, the filaments 3.5-4 mm long, reportedly yellow, anthers 3.5-4 × 0.5-1 mm, linear, yellow, slightly laterally compressed, the pore ca. 0.2 mm wide, terminal. Ovary (5-)6(-7) locular, inferior, the apex glabrous and elevated into a conspicuous collar around the style. Style 7-8 mm long, bending below the stigma, vertical and horizontal distance from the anthers to the stigma absent; stigma, punctate, ca. 0.5 mm wide. Dry berry 7-9 × 7-9 mm. Seeds 0.5-0.75 mm, ovoid to pyramidal, the testa angulate and smooth to slightly roughened. Distribution (Fig. 99). Endemic to Jamaica, 500-1300 m in elevation.
Conostegia procera can be recognized by its leaves with mite domatia, usually white flowers, and ribbed hypanthium. Schnell (1996) found this species does not overlap in flowering time with the other Jamaican endemics. Schnell (1996) thought C. procera highly resembled C. jaliscana and placed it in a section together with the rest of Jamaican endemics. Th e molecular phylogeny places both Jamaican endemics sampled (including C. procera) forming a well supported clade and belonging to a clade that also includes C. rufescens and C. cuatrecasii. Conostegia jaliscana on the other hand falls with species of the giant stigma clade, which is centered in Central America.  Description. Shrubs to small trees 1-5 m tall with subtetragonal stems that become terete and that are densely beset with simple roughened to dendritic trichomes near the tips, a small layer of inconspicuous underdeveloped dendritic trichomes also present, glabrescent to glabrous with age; the nodal line present, covered or not by indument on young branches. Leaves of a pair equal to somewhat unequal in length. Petiole 0.5-3.5 cm, adaxially pubescent like the stem apices. Leaf blade 4-13 × 2-5 cm, 3-nerved, oblong, linear to elliptic or ovate to obovate, the base acute to rounded, the apex rounded to acute or acuminate, the margin entire, the adaxial surface glabrous, the abaxial surface essentially glabrous, sometimes with short stipitate and sessile stellate trichomes on the mid vein and some secondary veins and inconspicuously glandular puncticulate throughout. Inflorescence a terminal panicle 3-8.5 cm long branched above the base, accessory branches apparently absent, bracts and bracteoles 0.5-2 mm, subulate, deciduous. Pedicel 7-13 mm. Flowers 5-7 merous, calyptrate. Floral buds 11-20 mm long, ovoid, the base obtuse to rounded, the apex acute and mucronate, not constricted, the hypanthium 6-7 × 6-7 mm, smooth and glabrous. Petals 6-17 mm long, white. Stamens 11-18, 6-8 mm long, the filaments 3.4-4.4 mm long, white, lacking a conspicuous geniculation, anthers 3-4 × 0.8-1 mm, subulate, recurved at the tip, yellow, somewhat laterally compressed, the pore ca. 0.1 mm wide, terminal. Ovary 5-7 locular, inferior, apically glabrous and forming a collar around the style base. Style ca. 8-9 mm, straight to slightly bent at the tip, vertical distance from the anther to the stigma -0-1 mm, horizontal distance absent, the stigma truncate, ca. 04-0.5 mm wide. Berry ca. 10 × 10 mm, red at first but turning purple black with maturity. Seeds ca 0.8 mm, pyramidal, smooth. Distribution (Fig. 100). Endemic to Jamaica, 200-1100 m in elevation. Schnell (1996) discusses at length that Conostegia pyxidata is very similar to C. balbisiana and C. procera. He chose to maintain the species on a "borderline" decision and gave three reasons for keeping the three taxa as distinct: 1) some morphological diff erences in a small area; 2) the disjunct range of C. balbisiana; and 3) diff erences in flowering time between the more similar species C. procera and C. pyxidata, and C. balbisiana overlapping somewhat in flowering time with the other two species, but having larger flowers with diff erent stamen morphology. Schnell (1996) argued that these morphological and phenological diff erences suggested the presence of biological isolation and a diverging lineage. For the time being I have chosen to follow Schnell's reasoning for recognizing three taxa. Schnell (1996) also suggested the possibility of introgression from C. rufescens. Th e type of Conostegia subprocera was not studied but Schnell (1996)    Description. Trees 3-12 m tall with tetragonal glabrous stems that sometimes have inconspicuous dendritic trichomes particularly noticeable when dry; the nodal line present but faint. Leaves of a pair equal to somewhat unequal in length. Petiole 1-7.1 cm. Leaf blades 7-23.1 × 3-9.8 cm, 3-5 plinerved, with the innermost pair of primary veins diverging from the mid vein 0.5-1 cm above the base in opposite or sub opposite fashion, elliptic to oblong or elliptic ovate, the base acute or obtuse, the apex acute to caudate, the margin entire, glabrous on both surfaces (except some specimens with tiny dendritic trichomes abaxially). Inflorescence a terminal panicle 11-25.7 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, rachis pink; bracts and bracteoles apparently lacking. Pedicel 2-15 mm. Flowers (5-)6(-7) merous, calyptrate. Floral buds 5-11 × 2.5-5 mm, obovoid pyriform, the base obtuse, the apex apiculate, slightly constricted in the middle, the hypanthium 3.5-4 × 4-4.25 mm, campanu- late glabrous. Petals 5-7(-8.3) × 6-8 mm, pink or rarely white, obovate, spreading, the apex emarginate, glabrous. Stamens 12-15(-17), 4.5-6.5 mm long, slightly zygomorphic, the filaments 2.25-3.75 mm, white, anthers 2.5-3.25 × 0.75-1 mm, elliptic to oblong, sagittate at the base, somewhat laterally compressed, yellow, the pore around 0.1 mm wide, terminal to slightly ventrally inclined. Ovary (5-) 6 (-7) locular, inferior, the apex glabrous, forming a collar around the style. Style 4.5-5 mm, bent below the stigma, vertical distance between anther pore and stigma ca.
Conostegia rhodopetala can be distinguished by being almost entirely glabrous, having pink inflorescences and floral buds, apiculate floral buds and pink petals. It is similar to C. superba, which as Almeda (2009) recently mentioned has wider leaves and white petals. Schnell (1996) noted a delay in of up to 5-6 months in fruit ripening and a much shorter fruiting than flowering season.  Fig. 103 Conostegia rufescens Naudin, Ann. Sc. Nat. Bot. ser. 3 16: 108. 1850. Type: Jamaica. no date, W. Purdie s.n. (Schnell (1996) cited holotype at P "fide Almeda pers. com." but annotated that of K as the holotype; isotypes: GH!, K!, fragment BR). Description. Shrubs to trees 1.5-20 m tall with tetragonal to terete slightly ridged stems that are usually densely covered with small brown dendritic trichomes sometimes intermixed with sessile stellate and stalked-stellate trichomes, sometimes glabrescent; the nodal line present (sometimes obscured by indument). Leaves of a pair equal to somewhat unequal in length. Petioles 0.4-5 cm, occasionally densely setose adaxially. Leaf blades 8-27 × 3-10.5 cm, 3-5 nerved or if plinerved, with the innermost diverging from the mid vein up to about 1 cm above the base in opposite or sub opposite or alternate fashion, elliptic, the base obtuse to acute and sometimes decurrent on the petiole, the apex acute, acuminate or short-caudate, the margin entire to denticulate, the adaxial foliar surface essentially glabrous, the abaxial surface densely or lightly furfuraceous or puberulent with mealy brown stellate or branching trichomes, sessile or short stipitate, thick bodied and short branched. Inflorescence a terminal panicle 5-23 cm branched well above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node and with the flowers frequently clustered at the end of the branches, accessory branches absent or present, the rachis pubsecent with brown stellate and branching trichomes, bracts early deciduous or absent, the bracteoles to 3 mm, deciduous. Pedicel 1-3 mm. Flowers 7-8(-12) merous, calyptrate. Floral buds 5.5-12.75 × 3-7 mm, broadly pyriform, the base and apex obtuse to acute, slightly constricted below the torus, the hypanthium 3.5-4.5 × 4-5 mm, ferrugineous. Petals 7-11 × 5-7.5 mm, white or pink, obovate, spreading, rounded-truncate to emarginate, glabrous. Stamens 20-28, 6-7 mm, slightly zygomorphic, the filaments 3.5-5.25 mm, white, anthers 2.75-3.5 × 0.5-0.75 mm, linear and often recurved, yellow or rarely pink or yellow with a pink tip, the base sagittate, somewhat laterally compressed, the pore ca. 0.1 mm, subterminal. Ovary 10-14 locular, inferior, apically glabrous and with a conspicuous collar around the style base. Style 4-7 mm long, straight or slightly bending just below the stigma, distance from the anthers to the stigma ca. -1.5-0 mm, horizontal ca. distance 0-2 mm; stigma capitate, 1.25-75 mm wide. Berry 9-15 × 9-15 mm, blue-black to purple. Seeds 0.5-0.7 mm long, obliquely pyramidal. Distribution (Fig. 104). In the mainland ranging from Nicaragua through Costa Rica and Panama to Colombia and Ecuador, in the Caribbean known from Dominican Republic, Jamaica, Haiti, and Puerto Rico, from sea level to 1700 m elevation.

Conostegia formosa
Conostegia rufescens can be recognized by its dense but short indument of mainly small brown dendritic trichomes covering floral buds and veins on the abaxial leaf surface. Th is species is variable in habit, the amount of indument on the leaves and the shape of the leaf base. In Costa Rica for example, one can find shrubs to small trees in Braulio Carrillo National Park at 500 meters elevation with acute leaf bases and in the same park at 1500 meters tall trees with decurrent leaf bases. Conostegia rufescens can be easily confused with C. centronioides and C. rubiginosa on the basis of the rusty indument. Th e latter two species diff er from C. rufescens in their exserted styles. Schnell (1996) noted limited local flowering seasons and diff erentiation between population in flowering time due to local adaptation. Schnell (1996) also studied a population in Alto La Palma, Costa Rica, and noted brief and concentrated flowering peaks, more so than he observed in other species such as C. macrantha, C. montana, C. oerstediana, and C. rhodopetala. Lastly, Schnell (1996) noted the earlier flowering season of populations at lower elevations and a later flowering season for more higher elevation populations. Conostegia formosa Macfad. from Jamaica was considered by Schnell (1996) as conspecific with C. rufescens based on Macfadyen's description. Schnell (1996) argued that if Macfadyen's treatment were accepted, C. formosa would be the oldest name available.
Specimens examined. HAITI (fide Schnell). Riviere Glace, Holdridge 2120 (MICH, US).   Description. Trees 4-12 m tall with tetragonal and ridged stems that are sparsely to densely setose with stramineous hairs 2-5 mm long, with or sometimes replaced by a dense but inconspicuous puberulent understory; the nodal line present and bearing coarse setae. Leaves of a pair equal to somewhat unequal in length. Petiole 1.8-7.8 cm.

Conostegia setifera Standl.
Leaf blades 8-27 × 4-13.5 cm, 3-5 plinerved, with the innermost pair of veins arising 0.5-2 cm above the base in opposite to mostly alternate fashion, elliptic to obovate, the base acute or obtuse, the apex acute to obtuse or rounded, the margin denticulate and often ciliate, the adaxial surface glabrous, the abaxial surface glabrous except for some pubescence on the nerves. Inflorescence a terminal panicle 4-17.3 cm long with flowers frequently congested at the end of small branches, branching above the base, accessory branches present, the rachis covered mostly with stellate trichomes, bracts to 3 cm long, linear or setulose, deciduous or if setulose persistent, bracteoles replaced by clusters of setae subtending the buds, the setae persistent. Pedicels to 3 mm. Flowers 7-9 merous, calyptrate. Floral buds 6-10 × 6-10.5 mm, globose, the base rounded to truncate, the apex rounded and short-mucronate, not constricted, hypanthium 4.5-5 × 6-8 mm, stellate pubescent and tuberculate. Petals 8-11 × 8-11 mm, white, obovoid, spreading, apically emarginate, glabrous. Stamens 26-31, 6-8 mm long, radial to slightly bilateral resulting from stamens getting stuck below the stigma, the filaments 3.5-4.5 mm long, white, anthers 2.5-3.5 × ca. 1 mm, elliptic, the base sagittate, laterally compressed, yellow, the pore ca. 0.15 mm wide, terminal. Ovary 13-18 locular, inferior, apically glabrous and forming a collar around the style. Style ca. 5 mm long, bending downwards during anthesis and protruding below the anthers, vertical distance frome the anthers to the stigma ca. -2 -0 mm, horizontal distance ca. 0.5-2 mm; stigma capitate, consisting of 13-18 laterally compressed lobes, ca. 4 mm wide. Berry 10-12 × 10-12 mm, purple black. Seeds ellipsoid, the testa smooth. Distribution (Fig. 106). Ranging from Nicaragua to Costa Rica, 0-1350 m elevation. Conostegia setifera is distinguished by its setose petiole adaxial surface, setose bracteoles and sessile to subsessile, spherical flower buds. Specimens of C. setifera have been confused with C. lasiopoda based on the setose indument on the petioles. One way to diff erentiate them with infertile material is that C. lasiopoda is consistently basinerved whereas C. setifera is plinerved. With fertile material they are unmistakable since C. lasiopoda has a long exserted style with a capitate stigma whereas C. setifera has a large crateriform lobed stigma, and its style is not exserted. Th is species was reported from Panama from the specimen Mori et al. 3850 (at MO and reportedly at WIS) in both Schnell (1996) and Almeda (2009) but this specimen was also cited as Conostegia dentata in both of the latter publications. I have not been able to locate the specimen Mori et al. 3850 to confirm its identification. Th is species is usually encountered in the forest understory and Schnell (1996) noted that within this habitat, only well lit branches produce flowers.    Description. Shrubs to less commonly small trees 0.9-1.5(-3) m tall with terete to a somewhat tetragonal stems that are covered with long smooth spreading hairs and a sparse and inconspicuous ground layer of brown lepidote hairs; the nodal line obscured and covered by the setae as the rest of the node and internode. Leaves of a pair equal to unequal in length. Petiole 0.2-3.9 cm. Leaf blades 7.6-35.5 × 3.22-13.5 cm, usually clustered at the apex of the branches, 5-7 plinerved, with the innermost pair of primary veins diverging from the mid vein up to about 4 cm above the base usually after the formicarium mostly opposite fashion, elliptic to obovate, the base acute and attenuate or rounded and with formicarium 1.5-3 cm long entirely on the leaf blade when the base is decurrent ot half of the formicarium on the petiole when not, the apex acute to abruptly acuminate, the margin denticulate to dentate, setose on both sur- faces. Inflorescence a terminal panicle 3.4-16.3 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, the rachis setose with green or red trichomes, bracts subtending the nodes up to 3 cm long, persistent or deciduous, bracteoles up to 1 cm long, linear, persistent. Pedicel 0.5-3 mm. Flowers (4-)5(-6) merous, obovate to pyriform, calyptrate, the floral buds 4-7 × 2-4 mm, the base rounded, the apex apiculate, slightly constricted; the hypanthium 2.35-3.5 × 2-3 mm, setose with green or red trichomes and tiny brownish glands to rarely glabrescent. Petals 6-7 × 4-5 mm, white to pale pink, broadly obovate, spreading, eventually closing and persisting closed, emarginate, glabrous. Stamens (13-)15(-17), 4-5.5 mm long, radially arranged but sometimes bilaterally symmetric or asymmetric apparently from interactions with the style, the filament 2.45-2.75 mm long, white, anthers 2.25-2.75 × 0.5-0.75 mm, linear and sinuous, laterally compressed, the base sagittate, yellow, the pore ca. 0.15 mm wide, ventro terminal. Ovary (4-)5(-7) locular, inferior, apically glabrous and forming a low collar around the style. Style 4-5 mm long, straight and just slightly curved upward apically, vertical distance of the anther pore to the stigma -2 -0 mm, horizontal distance absent; stigma capitellate to subcapitate, 1-1.5 mm wide. Berry 5-6 × 5-6 mm, dark purple to black. Seeds 0.3-0.5 mm, ovoid, the testa smooth. Distribution (Fig. 108). From Nicaragua through Costa Rica and Panama to Colombia and Ecuador, 0-1400 m elevation.

Conostegia setosa Triana
Th is is one of the most distinctive species of Conostegia because of the densely setose indument on most parts and the presence of pouch formicaria at the base of the leaf. Only one additional species has this kind of structure within Conostegia and that is C. dentata. Th e latter taxon diff ers from C. setosa in its reduced inflorescences, larger flowers, and exserted styles. Two morphotypes exist in C. setosa, with typical plants having almost sessile leaves with mostly acute to attenuate bases in which the formicarium is almost all on the lamina. On the other hand plants described by Standley and Macbride as Cyprophysa setosa and given the new name of C. hirsuta by Gleason have long petioles with the formicarium placed half on the petiole and half on the lamina. Schnell (1996) considered these two morphotypes to be the same species because he saw intermediate morphologies. Th e latter author also observed no geographic pattern but noted the similarity between plants from the same locality. During the course of this study plants of both morphotypes were collected in the same locality in Santa Fé, Veraguas, Panama (short petiolate leaves with acute to attenuate bases in Kriebel and Burke 5731, petiolate leaves with rounded bases in Kriebel and Burke 5712-both at NY, PMA). Schnell (1996) studied the phenology of this species in detail and observed one or two flowers opening everyday for two to three months. He hypothesized that this behavior might result in a greater degree of outcrossing by forcing the bees to forage to other plants in these large clonal populations. He further noted that bee species that visit C. setosa are non traplining opportunistic species (Schnell 1996). I have observed one of these opportunistic bees in the Halictidae family visiting C. setosa at La Selva, Costa Rica. Perhaps as expected for a species that flowers over a long period of time, fruiting is also spread through large periods of time (Schnell 1996). Alonso (1998) studied populations of Conostegia setosa in Costa Rica and Panama and observed some populations in Colombia and Ecuador. She found that in the southern part of it distribution, C. setosa was inhabited by more specialized ants and one obligate inhabitant, Pheidole melastomae. Th e latter inhabitant was most common in South America so Alonso (1998) hypothesized that perhaps this pattern is due to the fact that because C. setosa is bird dispersed, plants have dispersed more rapidly than their obligate ant inhabitant P. melastomae. In general, Alonso (1998)  Description. Shrubs to small trees 1-8 m, tetragonal and sulcate stems that are glabrous or beset with sessile stellate trichomes or densely covered with stipitate stellate trichomes; the nodal line present but inconspicuous. Leaves of a pair equal to somewhat unequal in length. Petiole 0.7-10.8 cm. Leaf blades 7.1-36 × 2.3-16 cm, 5-plinerved, with the innermost pair of primary veins diverging from the mid vein up to 1.5 cm above the base in opposite to sub opposite fashion, narrowly ovate to broadly ovate, or oblong-elliptic, the base acute to rounded, the apex acute to obtuse and acuminate, the margin entire, undulate-ciliate, or dentate, the adaxial surface glabrous or with simple hairs in young leaves, the abaxial surface glabrous or glabrescent with sessile stellate trichomes, to evidently pubescent with stipitate stellate trichomes especially on the veins. Inflorescence a terminal panicle with the flowers disposed in umbels terminating the inflorescence branches, 7-27.5 cm long and branching above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, rachis often reddish, sulcate, bracts and bracteoles 1 mm or less, early deciduous. Pedicels thick, 1.5-5.5 mm long. Flowers (4-)5-7 merous, oblong to obovate-pyriform, calyptrate, floral buds 5-9 × 2.5-5 mm, the base rounded, the apex acute to slightly apiculate, slightly to not con- stricted at the middle, the hypanthium 2.5-3 × 2.25-2.75 mm, glabrous. Petals 4-6.5 × 3.5-6 mm, white or less commonly pink, obovate or obtriangular, spreading at anthesis, emarginate, glabrous. Stamens (10-)14-16(-17), anthers 5-7.5 mm long, slightly zygomorphic, the filament 3-4.25 mm long, not geniculate, white, anthers 2-3.25 × 0.5-1 mm, linear-oblong, slightly recurved, laterally compressed, briefly sagittate at the base, yellow, the pore 0.1-0.15 mm wide, subterminal and ventrally inclined. Ovary (4-)5-6(-9) locular, inferior, apically glabrous and forming a collar around the style. Style 3-5.25 mm, enveloped at the base by a collar, straight or slightly bent towards the tip, vertical distance between the anther pore and the stigma ca -1 -0 mm, horizontal distance absent, the stigma subcapitate, 1-1.5 mm wide. Berry 6-9 × 6-9 mm, purpleblack. Seeds 0.4-0.6 mm long, narrowly pyramidal, the testa smooth. Distribution (Fig. 110). In the mainland from Mexico through most of Central America, restricted to the Pacific coast of Costa Rica and in Panama to the western portion. In South America in Colombia, Ecuador, and Venezuela. Schnell (1996) also reports this species from Peru, 0-1700 m in elevation.
Th e recent molecular phylogeny of Conostegia included three samples of C. superba. One, from the Dominican Republic, the second from Guatemala, and the third from Ecuador. Two of the samples, the one from Guatemala and the one from the Dominican Republic fell sister to each other in a clade that also includes C. bracteata and C. caelestis. Th e third specimen from Ecuador, falls sister to C. rhodopetala. On the one hand, this confirms the position of Schnell (1996) of treating C. clidemioides as a synonym of C. superba but on the other hand underlines the species delimitation problems between C. superba and C. rhodopetala. Also, although genetically C. cuatrecasii appears to be a distinctive species based on the Panamanian and Ecuadorian samples included in the phylogeny, morphologically the populations of pink flowers of C. superba can resemble C. cuatrecasii. Schnell (1996) notes more limited flowering seasons in local populations than are reflected by specimens. A Brazilian infertile specimen which corresponds to a syntype of C. poeppigii cannot be positively identified (Schnell 1996) and is thus excluded from the distribution circumscription until it is collected again. Schnell (1996)    Description. Trees 2-20 m tall with tetragonal and ridged branches that are generally sparsely to copiously covered with a mixture of caducous, sessile stellate and stalkedstellate hairs; nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 0.7-7 cm. Leaf blades 6-32 × 2.6-20 cm, 3-5 plinerved, with the innermost pair of primary veins diverging 1-3.5 cm from the mid vein in opposite to alternate fashion, ovate to elliptic, the base acute or obtuse, the apex acute or obtuse and short acuminate, the margin undulate dentate, the adaxial surface glabrous or glabrescent with sessile or stipitate trichomes which are branching or stellate,the abaxial surface with with sessile or stipitate trichomes which are branching or stellate especially on the veins. Inflorescence a terminal panicle 3.7-16 cm long branching above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present or absent, the rachis glabrescent with few scattered stellate trichomes, bracts and bracteoles to 5 mm long, linear, early deciduous. Pedicel 1.5-15 mm. the hypanthium 2.25-3 × 2.5-3 mm, smooth and mostly glabrous. Flowers 6-10(-12) merous, calyptrate. Floral buds 6-14 × 4-9 mm, spherical, the base rounded or flattened, the apex obtuse to flattened and apiculate, not constricted. Petals 7.5-15 × 4.5-10 mm, white, obovate, spreading, rounded-truncate to emarginate, glabrous. Stamens18-30, 7-8 mm long, radially arranged, to slightly bilateral apparently because of the downward bending style, the filaments 3.75-4.5 mm, white, lacking a geniculation, anthers 2.75-4 × 1-1.25 mm, oblong, straight or recurved, laterally compressed, yellow, the pore 0.1-0.3 mm wide, terminal. Ovary 9-16 locular, inferior, apically glabrous and forming a collar around the style. Style ca. 7 mm long, curving downward, vertical distance from the anthers to the stigma ca. -0.5 --0.25 mm, horizontal distance ca. 1-2 mm; stigma crateriform, consisting of 9-16 laterally compressed lobes, ca 3-4 mm wide. Berry 10-13 × 8-10 mm, blue-black or purple. Seeds 0.5-0.75 mm, obliquely pyramidal, the testa smooth. Distribution (Fig. 112). From Mexico through Guatemala to Honduras, 500-2200 m in elevation.
In general, Conostegia volcanalis can be recognized on the basis of its mostly spherical flower buds and broad leaves with undulate dentate margins. Schnell (1996) discussed this species as having "three well defined allopatric races". Th ese morphotypes were considered on the "borderline" of deserving species status (Schnell 1996). Th e three morphotypes diff ered in their distribution, habitat preference, indument density, and floral part size. Th e first morphotype is found in Guerrero and Jalisco and (Schnell 1996) noted that as C. jaliscana. Th ese plants prefer streamsides in pine forests. Th e leaves are is more pubescent and have larger floral parts. Th ese trees flower January through April. Th e second morphotype recognized by Schnell (1996) occurs in cloud forests of Chiapas, San Marcos and Quetzaltenango in Mexico. Th e plants from this morphotype are larger trees with less pubescent leaves and flowering November through March. Th is morphotype is reminiscent of C. icosandra and had been described as C. sphaerica Triana. Th e third race occurs in cloud forests of central Guatemala, Honduras, El Salvador and Nicaragua. Th is morphotype Schnell (1996) characterized as being like morphotype one in pubescence but morphotype two in inflorescence structure. Th e southernmost population within the third morphotype occurs in Costa Rica and Panama and was described as C. orbeliana Almeda. Very few specimens have been collected of this species and they tend to look dif- ferent than C. volcanalis because of their few-flowered inflorescences. When it was only known from Panama, Schnell (1996) noted its flowering time as in January, diff ering from populations from Guatemala to Nicaragua that flower May through October. Very few specimens are known of this species and Costa Rican specimens remeniscent of C. orbeliana have been collected. One of them, was flowering in August, undermining the possible phenological gaps between these populations or possible species. I have chosen not to include C. orbeliana under the synonymy of C. volcanalis not only because they are allopatric, but also because it might also be a variant of for example C. oerstediana or C. macrantha which occur in nearby areas. In general, C. volcanalis is a complex in need of study to assess habitat and phenological specialization. Perhaps more species can be recognized within this complex in the future.  Description. Shrubs or small trees 3 to 5 m tall with flattened stems that become terete with age and are densely puberulent with sessile stellate trichomes; the nodal line present yet slight. Leaves of a pair equal to somewhat unequal in length. Petioles 1-3.8 cm.

Conostegia vulcanicola
Leaf blades 5-15 × 2-7.1 cm, 5-plinerved, with the innermost pair of primary veins diverging from ca. 1 cm from the blade base the midvein in opposite to subopposite fashion, elliptic-ovate to elliptic, the base acute, the apex acute to acuminate,the margin entire or ciliate, the adaxial and abaxial foliar surfaces covered with small stellate hairs. Inflorescence a terminal panicle 3-10 cm, accessory branches apparently absent, the rachis with small stellate hairs, linear, the bracteoles to 3 mm, deciduous. Pedicel 5-20 mm. Flowers 5-6 merous, calyptrate. Floral buds 4-7 × 2-3.5 mm, oblong-pyriform, the base rounded, the apex acute to short apiculate, slightly constricted below the calyptra. Petals 5-7 mm long, white or pinkish, broadly obovate, spreading, glabrous. Stamens 10-12 (-13), 4-5 mm long, possibly slightly bilateral, the filaments ca. 2.5 mm, white, anthers ca. 2 mm long, linear oblong, yellow, the pore ca. 0.2 mm, terminal. Ovary 4-5-locular, inferior, apically glabrous and forming a low collar around the style. Style 3-5 mm, bending below the stigma, apparently no vertical or horizontal distance between the anthers and the stigma, truncate or subcapitate. Berries 5-7 × 5-7 mm, purple-black. Seeds ca. 0.7 mm long, obliquely pyramidal, the testa smooth. Distribution (Fig. 113). Costa Rica and reported from Panama by Schnell (1996), 1550-2400 m in elevation. Considering the lengthy discussions provided by Schnell (1996) in problematic species, it is interesting he did not discuss anything under Conostegia vulcanicola, although he did mention that C. micrantha and C. vulcanicola were probably derived from C. montana. I have gone to several volcanoes in Costa Rica looking for C. vulcanicola and I found populations that resemble the type specimen except mostly for the reddish hairs on the newest vegetative growth. In general, these specimens also resemble and I have annotated as C. montana. Th e isolectotype of C. vulcanicola was previously identified as C. montana apparently by A. Tonduz and another specimen identified by H. A. Gleason as C. montana was annotated as C. vulcanicola by Schnell. Th e main diff erences between the two taxa at this time are the indument mentioned above as well as leaf venation. C. vulcanicola tends to have plinerved leaves whereas C. montana lack the reddish indument and usually has nerved leaves. Further investigation into this species complex is needed.
Conostegia sect. Australis Kriebel, sect. nov. urn:lsid:ipni.org:names:77156240-1 Diagnosis. A mostly South American group distinguished by the following combination of characters: pleiostemonous flowers, calyx calyptrate, lacking calyx teeth altogether, and with conspicuous sclereids internally exserted styles, stamens lacking a filament geniculation but frequently bearing an evident distinction between the filament and the anther, exserted style, most species lacking a stele inside the style, mucilage inside the ovary present, and seeds ovoid and smooth.
Type. Conostegia lasiopoda Benth. Description. Small tree to 7 m tall (one label had this information) with tetragonal and ridged young stems that are covered with a rusty indument of sessile stellate trichomes intermixed with dendritic trichomes which have a thick axis and appear stipitate stellate; nodal line absent and/or covered by indument. Leaves equal or unequal at each node. Petiole 0.8-3.5 cm. Leaf blades 8-22 × 2.-8 cm, 3-5-plinerved, with the innermost pair of veins arising just above the blade base or up to 1 cm above the base in opposite to alternate fashion, narrowly elliptic to elliptic, the base acute, the apex acuminate to caudate, entire to inconspicuously denticulate, glabrous adaxially, with small stellate trichomes and branched trichomes mostly on the veins abaxially. Inflorescence a terminal panicle 4-11 cm long, accessory branches absent, branches rusty from the indumenta like the stems, bracteoles linear to lanceolate, 3-10 mm long, persisting but ultimately deciduous. Flowers mostly 6 merous, calyptrate. Flower buds (7.5-) 10-18 × 4-8 mm, slightly constricted in the middle, flattened at the base and apiculate at the apex, the calycine and hypanthial portions weakly diff erentiated, the hypanthium 4.5-6.25 × 5-6.5 mm, covered with stellate hairs. Petals 8-11 × 6-8.8 mm, white, obovate to obovate-spatulate, emarginate, and glabrous. Stamens 18-24, ca 6-8 mm long, the filament 3-4.5 mm long, white, without an abrupt geniculation but curving below the anthers, anthers 3-3.5 × 0.5-0.7 mm, linear, slightly recurved, the anther connective thickened but without an evident shoulder, yellow, the pore ca. 0.14 mm wide. Ovary mostly 6 locular, glabrous, inferior and elevated into a small collar around the style base. Style ca. 7.5-11 mm, vertical distance from the anther to the stigma ca. 4 mm, horizontal distance absent; stigma capitate, 1.4-1.6 mm wide. Berry not seen. Distribution (Fig. 115). Known from few collections in the Pacific coast of Colombia to Central Ecuador, 600-1800 m in elevation.

Conostegia apiculata
Th is rare species can be recognized based on its indument of stellate hairs, long apiculate calyptras, and stout, exserted styles. Th e drawing made by Wurdack deposited at NY shows the lack of a filament geniculation in this species. Floral measurements made of an Ecuadorian specimen (Toasa et al. 11350-NY) show longer styles at least in that population than in Wurdack's drawing.    Description. Shrub to small tree 0.5-3 m tall with terete glabrescent stems that are beset with an inconspicuous indument of minute sessile stellate or branching trichomes; the nodal line present but inconspicuous. Leaves equal or unequal at each node. Petiole 0.5-2.4 cm. Leaf blades 2.5-15 × 1-3.5 cm, 3-nerved, linear to ellip-tic, the base attenuate, the apex acuminate, adaxially glabrous and inconspicuously glandular puncticulate, abaxially with small stellate trichomes mostly on the primary and higher order veins, entire to more commonly denticulate. Inflorescence terminal panicles 3.5-7.9 cm long branching above the base, but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present or absent, the rachis glabrescent with few scattered stellate trichomes, bracts and bracteoles 1-5 mm long, linear, persistent. Flowers (5-)6-8 merous, calyptrate. Flower buds 3.9-8.6 × 2.3-4.6 mm, not to slightly constricted in the middle, pyriform to obovate, flattened at the base, acute to short apiculate apically, the calycine and hypanthial portions weakly diff erentiated, the hypanthium 3-4.5 × 3-5 mm, inconspicuously stellate. Petals 5-9 × 4-7.2 mm, white or pinkish, broadly obovate-spatulate, glabrous, the apex emarginate. Stamens 16-21, 4.1-5.5 mm long, the filament 2.3-3 mm long, anthers 1.8-2.5 × mm, linear or narrowly oblong, straight to slightly recurved, laterally flattened, the base sagittate, with a small bump on the anther connective dorsally at the filament insertion, yellow, the pore ca. 0.1 mm wide. Ovary (4-)5-6 locular, glabrous, inferior and elevated into a conspicuous collar around the style base. Style 4.7-5.7 mm long, vertical distance from the anther to the stigma ca. 1.5 mm, horizontal distance absent; stigma capitate, 0.6-1 mm wide. Berry 4-5 × 4-5 mm, dark purple to black. Seeds ca. 0.4 mm long, pyramidal, the testa smooth. Distribution (Fig. 116). Colombia to northern Ecuador, 0-600 m in elevation. Also reported from Panama by Schnell (1996). Conostegia attenuata var. peruviana is known only from a single infertile specimen in Peru and cannot be positively identified at this time. Th us, Peru is excluded in the present distribution of the species.

Conostegia attenuata
Floral measurements of Schnell are quite small in comparison to those of Gleason. Th e latter author measured three specimens of which the drawings are deposited at NY. With regards to staminal morphology, Gleason also noted in his drawings that "thecae are prolonged below the insertion of the filament. Connective minutely gibbous on the back just above apex the filament." Th is morphological feature is common and only present in section Australis. Schnell (1996) and Almeda (2009) reported C. attenuata from Costa Rica. Th e Costa Rican specimens cited by these authors (Skutch 2734, 4941-NY) diff er quite a bit from typical C. attenuata material. Th e main diff erences are that the Costa Rican material is glabrous to inconspicuously lepidote, has plinerved leaves, and most notably has short styles. Th e typical C. attenuata has stellate trichomes on the branch apices and floral buds, nerved leaves and exserted styles. Both specimens were also annotated as a probable new species on the NY specimens by Schnell. Given that they are highly reminiscent to C. montana and have not been recollected, I consider them a narrow leaf variant of C. montana for the time being.  Fig. 117 Conostegia centronioides Markgraf, Notizbl. Bot. Gart. Berlin-Dahlem 14: 25-44. 1938 Description. Shrubs to small trees 2-11 m tall with flattened or tetragonal that are finely and densely pubescent with tiny sessile stellate trichomes; the nodal line present yet slight. Leaves equal to unequal at each node. Petiole 1-5 cm long. Leaves 5-21 × 1.9-10 cm, 3-5 nerved or if 3-5 plinerved, with the innermost pair of primary veins diverging from the mid vein up to about to about 2 cm above the base, elliptic, the base acute to obtuse, the apex abruptly acuminate, usually conspicuously undulate denticulate, adaxially glabrous except for inconspicuous stellate trichomes when young, abaxially stellate pubescent with small trichomes on the veins to almost glabrous. Inflorescence terminal a panicle 3.7-22 cm long branching above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches present, rachis pubescent with small stellate trichomes; bracts and bracteoles 0.5-2 mm, linear to lanceolate, usually deciduous. Flowers 5-6(-7) merous, calyptrate, the calyptra apparently consistently breaking in pieces. Flower buds 4.7-11 × 2.5-6.5, pyriform, the base flat to rounded, the apex rounded to short apiculate, the calycine and hypanthium portions weekly diff erentiated, constricted below the calyptra; the hypanthium 3-6 × 3-6 mm, pubescent with rusty stellate hairs. Petals 6-9 × 6-9 mm, white to translucent white, obtriangular, glabrous, emarginate to three lobed apically. Stamens 15-24, 4.8-7 mm long, slightly zygomorphic, the filament 3-4 mm, white, not geniculate, anthers 1.8-3, linear-oblong, pale yellow, laterally compressed, dorsally thickened, basally sagittate, the pore ca. 0.1 mm wide, terminal. Ovary 5-6(-7), inferior, glabrous, the apex forming a conspicuous collar around the style base; style ca. 6-8 mm, straight to gently curving near the apex, vertical distance from the anther to the stigma 1.5-3 mm, horizontal distance absent, the stigma capitate, ca. 1 mm wide. Berry 6-7 × 6-7 mm, purple. Seeds 0.4-0.5 mm long, ovoid and smooth. Distribution (Fig. 118). Mostly in northwestern Ecuador and probably overlooked in south-western Colombia, 20-900 m in elevation. Schnell (1996) reported that the holotype of this species (as well as the isotypes which had not been distributed) were lost at Berlin. As mentioned by Schnell (1996), the neotype chosen here (the same chosen by Schnell's unpublished work) fits the original description. Conostegia centronioiodes is quite variable in the sizes of leaves, inflorescences and flower buds as well as in the flower bud apex which can be more or less rounded to short apiculate. Perhaps because of its small flower buds, this species has been confused with C. montana. An easy way to tell these species apart in bloom is that C. montana has a short style whereas C. centronioides has a long style. Th e same character can be used to separate C. centronioides from C. rufescens, another species with which it gets confused. Lastly, C. centronioides is similar to C. lasiopoda, particularly when in the latter the large bracts that cover the flower buds have fallen off . C. lasiopoda tends to have conspicuously setose adaxial petiole surfaces which aid in its distinction from C. centronioides.

Conostegia centronioides Markgraf
Specimens  Description. Shrub to small tree 1.3-8 m with sub-terete stems that are densely setose with simple bristles up to 4 mm long; the nodal line hard to see and covered with setae as the rest of the node and internode. Leaves at a node equal to sub equal in size. Petiole 0.5-4.9 cm long. Leaves 11.9-35 × 5-12.9 cm, 3-5 plinerved, with the innermost pair of veins arising up to about 3 cm above the base and diverging mostly in opposite or sub opposite fashion from the mid vein, obovate to nearly elliptical, the base acute to decurrent on the petiole and with paired formicaria ca. 2-4 cm long on the leaf surface or extending to the petiole, the apex rounded to obtuse and abruptly acuminate, the margin dentate, undulate-dentate or denticulate, adaxially sparsely setose, abaxially setose. Inflorescence a terminal compact panicle 1.5-4(-7) cm long, inflorescence rachis obscured by the dense setose indument; bracts and bracteoles 1.5-4 mm. Pedicel ca. 2 mm long, obscured by the indument. Flowers (5-)6(-8) merous, calyptrate. Flower buds ca. 9-15 × 5-7 mm, narrowly ovate not constricted about the middle, rounded a the base, acute and long attenuate at the apex, not constricted in the middle, the calycine and hypanthial portions undiff erentiated; the hypanthium 5-6.25 × 5.5-7.5 mm, densely hirsute with trichomes with swollen bases. Petals ca. 9-13 × 9-12 mm, white or pink, obtriangular, spreading, glabrous, the apex emarginate. Stamens 19-30, ca. 7-8 mm long, their posture at anthesis not seen, the filament 4-5 mm long, lacking a conspicuous geniculation, anther 2.8-3.5 mm, linear-oblong, recurved near the base, yellow, laterally compressed, the connective thickened and with a small bump dorsally, the pore ca. 0.1 mm, subterminal and slightly ventrally inclined. Ovary 6-10 locular, inferior, the apex glabrous and forming a collar around the style base. Style 6.5-7.5 mm long, gently curved, vertical distance from the anthers to the stigma ca. 1-2 mm, horizontal distance absent, the stigma sub capitate, ca. 1.3 mm wide. Berry ca. 7 × 7 mm when dry. Seeds not seen. Distribution (Fig. 119). Ranging from Panama to Colombia and Ecuador on the Pacific coast, 0-680 m in elevation.
Conostegia dentata is easily recognized because of its setose indument on stems, inflorescences and flower buds, leaves with formicaria at the base and dentate margins, and compact inflorescences. Th e study of flowers in herbarium specimens revealed an exserted style as is typical of species of section Australis. Almeda (2009) was unable to locate the type specimen Triana 4113 of this species at BM and when checking Triana's herbarium list noted that that number is stated to be from Antioquia instead of Chocó, but no genus or species are stated. Almeda did locate Triana 4112 which corresponds to C. dentata but that specimen is from Barbacoas province instead of Chocó province where Triana 4113 was cited to have been collected. I agree with Schnell (1996) that the original description leaves little doubt on its identity namely because of the description of the setose indument, congested inflorescences and long petals. Because the setae of the fruiting hypanthia are white instead of brown as in the rest if the plant, Schnell (1996) hypothesized that this color diff erence might function to attract dispersal agents.  Description. Shrub to small tree 2.5-7(-12.5) m tall with flattened stems that are furfuraceous on new growth with stellate trichomes; the nodal line present yet slight. Leaves at a node equal to sub equal in size. Petiole 0.7-2.8 cm long. Leaves 6-24.8 × 3-10.1 cm, 5 plinerved, with the innermost pair of veins diverging up to 2.5 cm above the base mostly in sub opposite fashion, elliptic to narrowly ovate, the base subacute to obtuse, acute or acuminate at the apex, the margins remotely undulate to denticulate, adaxially glabrous or glabrescent, abaxially with numerous underdeveloped stellate trichomes. Inflorescence a terminal panicle 6-12.3 cm long branched above the base, accessory branches present, the rachis covered with stout stellate or dendritic trichomes. Pedicel 2-4.4 mm. Flowers (4-)5-6(-7) merous, calyptrate. Floral buds 3-5.5 × 3.75-4.25, oblong-pyriform, acute at the base, acute and apiculate at the apex; hypanthium ca. 3-4 × 3.25-4.25 mm, not constricted in the middle, stellate pubescent, the calyptra and hypanthium not diff erentiated. Petals 4-5 × 3-3.5 mm, white, broadly obovate, spreading, glabrous, rounded with a lobe to one side of the flower. Stamens (16)27-36, 4.75-6 mm long, apparently slightly zygomorphic, the filament 2.5-3 mm, white, anthers 2.25-2.75 × 0.25-0.75 mm, linear, white to yellowish, not evidently sagittate at the base, the pore 0.1-0.12 mm, ventro-terminal. Ovary (5-)6-7(-8) locular, inferior, glabrous. Style 7-8 mm, straight, lacking a prominent a collar at the base, vertical distance from the anthers to the stigma ca. 1-1.5 mm, horizontal distance absent; stigma subcapitate, 0.75-1 mm wide. Berry 4-5 × 4-5 mm, purpleblack. Seeds 0.3-0.4 mm, ovoid, the testa smooth to a little roughened. Distribution (Fig. 121). From southern Costa Rica to Colombia, with a disjunct population in the Cordillera de Vilcabamba in Peru, 700-1350 m in elevation. Th e inclusion of the Peruvian locality is based on a specimen cited by Schnell (1996).
Conostegia extinctoria is one of the smallest flowered species in section Australis. Th e style of this species is exserted as in other species of section Australis (Fig. 120). Schnell (1996)     Description. Shrubs or small trees 1-5 m tall with tetragonal to rounded stems which are covered predominantly by stipitate stellate trichomes; nodal line present, somtimes obscured by indument. Leaves at a node equal to unequal in size. Petiole 0.4-1.8 cm. Leaf blades 5-17 × 2-6 cm, 3-5 slightly plinerved, elliptic to lanceolate, rounded to subacute at the base, the apex and acuminate, margins entire to undulate-denticulate, glabrous adaxially, pubescent mainly on the primary veins abaxially with indument like that of the stems. Inflorescence a terminal panicle 4-9 cm long branched above the base but sometimes appearing branched at the base because of multiple inflorescences arising at opposing meristems at the terminal node, accessory branches absent or present. Bracts and bracteoles 1.5-3 mm long, linear. Pedicels 1-3 mm long. Flowers 7-9 merous, calyptrate. Floral buds 6-9.5 × 3-6.5 mm, ovoid to obovoid, slightly constricted below the calyptra, the hypanthium and calyptra undifferentiated; hypanthium 3-3.5 × 3.75-4.25 mm, pubescent with sessile and/or stipitate stellate trichomes. Petals 8-10 × 5-6 mm, white, obtriangular, posture not seen, glabrous on both surfaces, emarginate apically. Stamens 30-42, 7-8 mm long, the filament 4-4.5 mm, apparently white, anthers 3-3.5 × 0.5-0.75 mm, linear subulate, pale yellow (in rehydrated specimen), the pore ca. 0.18 mm, terminal to slightly ventrally inclined. Ovary 11-12-locular, inferior, glabrous and forming a collar around the style base; style ca. 7.5-8 mm, gently bending from the base, glabrous, the style appears to be exserted but diffi cult to assess in the rehydrated material as well as if it is bent opposite the stamens or not, stigma capitate, ca. 2 mm wide. Mature berry not seen. Distribution (Fig. 122). Southeastern foothills of the Andes in Ecuador at 800-1200 m in elevation. Schnell (1996) noted that as like for Conostegia centronoides, the holotype of Conostegia centronoides var. lancifolia was lost during the war in Berlin. I concur with Schnell (1996) in that the description of the latter including stipitate stellate hairs and narrowly ovate leaves matches topotypical material which Schnell cited as the neotype (E. Asplund 18868). For this reason and because the work of Schnell (1996) wasa not published, the neotypification is made here. Schnell (1996) included in his circumscription of C. lancifolia the decurrent leaved and large leaf blade plants here referred to as C. ortizae. Schnell pointed out that if intermediates between the two morphotypes did not arise they might as well be called subspecies or species. Considering they are easy to distinguish they are here proposed as diff erent species.  Description. Shrubs to small trees 1.5-6 m tall with stems that are first tetragonal then terete and setose with appressed single bristles 1-2 mm long and many minute sessile stellate hairs; nodal line present yet slight. Leaves at a node equal to subequal in size. Petioles 0.7-3.6 cm long, densely hirsute adaxially. Leaves 8.3-30 × 3.6-11.7 cm, 3-5 nerved, ovate to obovate, cuneate to obtuse at the base, the apex acute to rounded and attenuate to abruptly acuminate,the margins entire to undulate-dentate, the adaxial surface glabrous, slightly concave and with deeply impressed tertiary venation, abaxially covered only with a layer of these tiny sessile stellae. Inflorescence terminal, 5-19.1 cm long, accessory branches absent or present, the branches subtended by deciduous, setose linear bracteoles 2-20 mm long, the clusters of flowers subtended by puberulent foliose, persistent to deciduous bracteoles 2-7 mm long. Pedicels of 0.5-2.0 mm. Flowers (5-)6(-8) merous, calyptrate. Floral buds 4-9 × 2-5 mm, ovate to oblong-pyriform, the base rounded, the apex acuminate, constricted below the middle, the upper and lower portions undiff erentiated, the hypanthium 3.5-4 × 3.25-3.5 mm, pubescent. Petals 7-8 × 7-8 mm, translucent white, obtriangular, spreading, glabrous, and slightly asymmetrical apically. Stamens 17-25, 4-6 mm, slightly zygomorphic apparently from the movement of the style, the filament 2.5-3 mm, white, anthers 1.5-2 × 0.5-1 mm, linear-oblong, straight or recurved, light yellow., the pore terminal, less than 0.1 mm. Ovary 6-9 locular, inferior, apically glabrous and forming a low collar around the style. Style 5-8 mm, straight for most of its length and bending gently near the apex, vertical distance from the anther to the stigma 2-3 mm, the stigma capitate, 1-1.25 mm wide. Berry 8-10 × 8-10 mm, light purple to purple-black. Seeds 0.3-0.5 mm long, ovoid, the testa smooth to roughened. Distribution (Fig. 124). Distributed in southeastern Nicaragua, wet forests of Costa Rica, somewhat restricted in Panama to mostly western Colombia, with a few specimens collected in northwestern Ecuador, also in Cocos Island, from sea level to 2100 m in elevation. Conostegia lasiopoda is usually a quite distinctive species. Th e leaf veins are ele vated and arise exactly at the base ("perfectly nerved"), the petioles are setose adaxially and the flowers have foliaceous bracts covering them. When these bracts fall, distinguishing C. lasiopoda from C. centronioides can be diffi cult. Th e latter tends to be at least somewhat plinerved and lack the adaxial setose petioles. C. rubiginosa is also similar but lacks the foliaceous bracts covering the floral bracts and the indument in flower bud is more evident. Also, C. rubiginosa tends to have smaller coriaceous leaves. Th e stamens of C. lasiopoda have a conspicuous anther shoulder (Figs 49, 123) which has not been confirmed in C. centronioides or C. rubiginosa and the style is exserted (Fig. 123) like the latter two taxa and as usual in section Australis. Lastly, the calyptra has been observed to rupture into pieces at anthesis (pers. obs. and photographs by Reinaldo Aguilar).   Schnell (1996)).
Conostegia monteleagreana is easy to distinguish because of the foliaceous bracts that subtend the glomerulate inflorescences. Th e seeds of this species are also distinctive because of their particular tubercles. It is the only species in section Australis to have this testa ornamentation. Th is species has two morphotypes, one in Colombia with larger leaves and inflorescences than the one present in Costa Rica and Panama. It also appears to vary in the degree of herkogamy. Th e distinction between the insertion of the anther and filament ("anther shoulder") is very conspicuous as in other species of section Australis. Schnell (1996)   Description. Shrubs or small trees to 3-6 m tall with flattened and grooved stems that are covered with an orangish mixture of long and shorter-branching stellate trichomes, the long branching ones graceful and sessile, sometimes also becoming dendritic; the nodal line present yet slight. Leaves at a node equal to subequal in size. Petiole 0-1.5 cm, grooved above, usually entirely covered by the decurrent leaf base. Leaf blades 17-37 × 8-15 cm, 3-5 plinerved, with the innermost pair of veins diverging above the base in fashion, broadly elliptic to obovate, decurrent at the base, the apex long-caudate up to 3.5 cm long, margins entire to undulate-denticulate, glabrous adaxially, abaxially with indument like that of the stem. Inflorescence a terminal panicle 4-13 cm long with accessory branches; bracteoles 2-3 mm long, linear, tending to persist. Pedicels 1-2 mm. Th e hypanthium 3-4 × 4.5-5 mm, pubescent with long-and shorter-branching stellate trichomes. Flowers 6-8 merous, calyptrate; floral buds 7-11.5 mm, not constricted below the calyptra. Petals 10.5-12 × 8.25-8.75 mm, white, obtriangular, spreading, glabrous on both surfaces. Stamens (26-)34-52, slightly zygomorphic, the filament 4-4.5 mm, white, anthers 3.5-4 × 0.5-0.75 mm, linear subulate, pale yellow, the pore ca. 0.15 mm, slightly ventrally inclined. Ovary 12-13 locular, inferior; style ca. 11-12 mm long mm, straight but gently bending from the base, glabrous, distance from the anther to the stigma ca. 3.5-4 mm, stigma subcapitate, 1.4-1.7 mm wide. Mature berry ca. 8-9 × 8-9 mm when dry. Distribution (Fig. 128). Northeastern foothills of the Ecuatorian Andes with one more southern population in Ecuador, at 250-1250 m in elevation.
Conostegia ortizae can be easily recognized by its large sessile leaves with an evidently decurrent leaf base and strongly plinerved venation. Of the species for which the anatomy of the style has been studied, this is the only one in section Australis to have a stele within the style.
Etymology. Th is species is dedicated to the Ecuadorian biologist Patricia Ortiz who tragically passed away in her second home of Monteverde, Puntarenas, Costa Rica. Pati was one of the best and most passionate naturalists I have had the fortune to meet.    Description. Small trees 1-13.6 m tall with flattened stems that become terete with age and are sparsely to densely puberulent with sessile stellate hairs; the nodal line present yet slight. Leaves at a node equal to subequal in size. Petiole 0.4-3.5 cm long. Leaf blade 4-14 × 2-7.3 cm, 3-5 nerved, usually elliptic to elliptic ovate, the base rounded to obtuse, the apex acute or short acuminate, the margin serrulate and ciliate, the adaxial sur- face glabrous, the abaxial surface sparsely to densely puberulent with sessile stellate hairs and minute white rounded and roughened secretions. Inflorescence a terminal panicle 3-15 cm long, accessory branches absent or less frequently present, the rachis with sessile stellate and stalked stellate hairs, the bracteoles 1-8 mm long, linear to ovate, persistent or deciduous. Pedicel 1-5 mm long. Flowers 5-8-merous, calyptrate, floral buds 6.5-10 × 3.25-6.25 mm, elliptic-pyriform, the base obtuse to rounded, the apex apiculate, slightly constricted below the calyptra, the calyptra and hypanthium not diff erentiated; the hypanthium 3.5-4 × 5.25-5.75 mm, stellate puberulent. Petals 7-9.5 × 5.25-9 mm, translucent white to white or pink, obtriangular to obovoid, spreading, glabrous, apically slightly bilobed to emarginate. Stamens 26-36, 5-7 mm long, radially arranged around the style, the filaments 3-4 mm, white, without an evident geniculation, anthers 2-3 × 0.4-0.6 mm, yellow, the pore ca. 0.25 mm. Ovary 6-8 locular, inferior, apically glabrous, forming a very low to absent collar around the style base. Style 8-10 mm, basically straight but slightly bending along its length, vertical the distance from the anthers to the stigma ca. 2.5-3.5 mm, horizontal distance absent, stigma capitellate, 0.5-0.75 mm wide. Berry 6-8 × 6-8 mm, dark purple. Seeds 0.45-0.7 mm long, obliquely pyramidal and somewhat angulate, the testa smooth to roughened. Distribution (Fig. 130). On the Caribbean coast of Nicaragua, both coasts of Panama and on the Pacific coast of Colombia and Ecuador and a disjunct population in Peru, sea level to 200(-600) m elevation. Conostegia polyandra has white secretions on the leaf abaxially. Since this species grows in mangrove habitats, these secretions are probably salt. Th is is the only species of Neotropical Melastomataceae known to grow in mangroves. It can further be recognized by its 3-4 nerved leaves which tend to have a consistent elliptic shape and serrulate and ciliate margins. Flowers in this species are also noticeable for their great number of stamens and strongly exserted styles. Schnell (1996) and Almeda (2009) commented on the phenological patterns of this species citing its continuous flowering and fruiting throughout the year. In addition these authors stated that the this phenological behavior coupled with few large seeds per fruit might help explain its weedy tendencies. I note here that the specimens I studied did not have particularly few large seeds per fruit. Schnell (1996) suggests C. polyandra is aggressive and somewhat weedy, reminiscent of C. speciosa and C. xalapensis. Its apparent absence in coastal areas in the Caribbean of Costa Rica might be explained by its replacement in these areas by C. xalapensis (Schnell 1996). Duke and Idrobo 11567 (MO, NY) reported the berries as "quite edible".
Description. Shrubs to trees 3-6 m tall with tetragonal to terete and slightly ridged stems that are covered with a mixture of sessile stellate and stalked-stellate hairs sometimes intermixed with simple hairs or sometimes glabrescent and becoming glabrous with age; the nodal line present (sometimes obscured by indument). Leaves at a node equal to subequal in size. Petioles 0.4-2.5 cm. Leaf blades 5.5-25 × 3-9 cm, 3-5 nerved or less frequently plinerved, if plinerved with the innermost veins diverging from the midvein up to ca. 1 cm above the base in opposite or alternate fashion, elliptic to ovate, the base obtuse to acute, acuminate to caudate, the margin entire to denticulate, the adaxial foliar surface essentially glabrous, the abaxial surface densely pubescent with rusty brown stellate and branching trichomes, sessile or short stipitate, especially on the main veins and lightly pubescent to glabrescent on the surface. Inflorescence a terminal panicle 5.5-13 cm long, accessory branches present or absent, the rachis pubsecent with brown stellate and branching trichomes, the bracteoles to 4 mm long, lanceolate to ovate, persistent or caducous. Flowers sessile or with pedicels up to 2 mm long, 6-7 merous, calyptrate. Floral buds 5.5-11 × 3.7-5.5 mm, obovoid, the base flat to rounded, and apex broadly acute, slightly to not constricted near the torus; the hypanthium 4.5-6 × 4.5-6 mm, rusty brown pubescent and sometimes sparsely tuberculate. Petals 7-9 × 6-8.5 mm, translucent white, obovate, spreading, glabrous, asymmetrical. Stamens 15-24, 5.5-6.5 mm, slightly zygomorphic, the filaments 2.75-3.25 mm, white, anthers 2.5-3 × 0.5-0.75 mm, linear to elliptic, laterally compressed and with an anther shoulder, yellow, the pore ca. 0.1 mm, subterminal and slightly ventrally inclined. Ovary 6-7 locular, inferior, apically glabrous and with a low collar around the style base. Style 6-7 mm long, straight to bending sideways, vertical distance from the anthers to the stigma 1.75-2.5 mm, horizontal distance 0-1 mm; stigma capitate, ca. 1 mm wide. Berry and seeds not seen. Distribution (Fig. 132). West coast of Colombia, sea level to 100 m elevation. For the few specimens studied, Conostegia rubiginosa is quite variable in leaf size and consistency. Conostegia rubiginosa can be confused with C. centronioides, C. lasiopoda and C. rufescens. From C. centronioides it can be distinguish by the larger flowers without a constriction in the hypanthium, and denser reddish indument. From C. rufescens it can be distinguished by its exserted style. For diff erences between this species and C. lasiopoda, see the discussion under the latter.    Description. Shrub or small tree 1.5-4 (-8) m tall with young tetragonal stems that then become terete and which are glabrescent with some minute stellate trichomes scattered throughout; the nodal line present yet slight. Leaves at a node equal to subequal in size. Petioles 0.3-3 cm. Leaf blades 5.9-19 × 2.1-9 cm, 3-5 plinerved, with the innermost pair of primary veins diverging from the midvein up to 1.5 cm above the base in opposite to alternate fashion, the secondaries obscure, elliptic to obovate, the base acute ro rounded, the apex obtuse to acute and caudate, the mar-gin entire, the adaxial surface glabrous, the abaxial surface glabrous except for tiny stellate hairs on the veins. Inflorescence a terminal few flowered panicle 2.2-8 cm long, accessory branches absent or if present frequently reduced to single flowers the rachis glabrescent with minute stellate hairs, the bracteoles up to 1mm long, linear to ovate, deciduous. Pedicels 1-2.2 mm. Flowers (5-)6(-7) merous, calyptrate. Floral buds 6-9 × 3-5 mm, globose, wider below the torus, the base truncate to rounded, the apex apiculate, constricted in the middle, the hypanthium 4-5 × 4-5 mm, sparsely stellate with minute trichomes. Petals 6-8.25 × 6-7.25 mm, translucent white, obtriangular, spreading at anthesis, apically asymmetrical, glabrous. Stamens 18-24, 3.5-5 mm long, radial to slightly bilateral apparently from interactions with the style, the filament 2-3 mm long, white, anthers 1-2 × 0.5-0.75 mm, oblong, forming an anther shoulder at the filament anther junction, yellow, the pore 0.1-0.2 mm, terminal. Ovary (6-)8-12 locular, inferior, apically glabrous and forming a low inconspicuous collar around the style. Style 5-6 mm long, straight with the stigma turning slightly upward, vertical distance from the anther pore to the stigma ca. 3-3.5 mm, horizontal distance absent; stigma capitellate, 1-1.25 mm wide. Berry 5-7 × 5-7 mm, purple black. Seeds 0.25-0.45 mm long, ovoid, the testa smooth. Distribution (Fig. 134). Nicaragua, Costa Rica, Panama, Colombia and reaching Ecuador, most common below 1000 m in elevation but reaching 1600 m.

Conostegia tenuifolia Donnell Smith
Th is species is easy to recognize because of its glabrosity, plinerved leaves with entire margins and caudate apices, few flowered inflorescences, apiculate calyptra, and exserted styles with a capitate stigma. In addition, C. tenuifolia has an evident anther shoulder. Th e calyptra of Conostegia tenuifolia has been observed to rupture longitudinally at one site in San Vito de Coto Brus, Puntarenas, Costa Rica (Fig. 133). Th e specimens from Peru (Kayap 955 and Berlin 723, both at NY), resemble this species, but more flowering specimens are required to assess their identity.   Diagnosis. Leaves frequently conspicuously plinerved, calyx not calyptrate or the few species with a calyptra then the latter lacking sclereids. Several species have fused calyces in bud that rupture irregularly. Flowers mostly diplostemonous and herkogamous, with the stamens bearing a conspicuous geniculation towards the apex, anthers radially arranged around the exserted style except for the calyptrate species which have short styles. Th ese short styled species resemble species of section Conostegia but lack a stele in their internal anatomy. Species in this section are almost entirely endemic to Costa Rica and Panama with a few species reaching South America and a few others northern Central America. Only Conostegia xalapensis reaches the Caribbean.
Type. Conostegia cinnamomea (Beurl.) Wurdack ( Description. Shrub to small trees 1.5-5 m tall with terete to flattened internodes that are moderately to copiously covered with smooth spreading hairs 1.5-3 mm long, sometimes underlain with a sparse to moderate understory of sessile stellate and stipitate-stellate hairs; the nodal line not evident. Leaves at a node equal to somewhat unequal in length. Petiole absent or up to 1.2 cm long. Leaf blades 3.5-23 × 1.8-10 cm, 5-7 plinerved with the innermost pair of primary veins diverging from the midvein 0.7-1.5 cm above the base and arising in alternate fashion, elliptic to elliptic-ovate, oblique and rounded, acuminate to attenuate, entire and ciliate varying to obscurely denticulate distally, adaxially moderately covered with spreading hairs 1-2.5 mm long to nearly glabrous, abaxially moderately to sparingly covered with spreading simple hairs 1-3 mm long on the median vein and innermost primaries and sparsely underlain with sessile stellate and stipitate-stellate hairs varying to glabrate. Inflorescence a pseudolateral modified dichasium 2.6-9 cm long sometimes divaricately branched from the base, accessory branches absent, the branches reddish green to red, moderately to copiously covered with smooth spreading hairs, sometimes underlain with a sparse to moderate understory of sessile stellate and stipitate-stellate hairs, bracteoles 1-6 mm long, subulate to narrowly triangular, persistent. Pedicel 0.5-1 mm long. Flowers 5 merous, calyx not calyptrate. Floral buds 3.75-4.25 × 3-3.5 mm, cupulate to campanulate, the hypanthium 3-4 × 2.5-3.5 mm, moderately to sparingly covered with spreading hairs 1-3 mm long and sparsely underlain with deciduous sessile stellate and stipitate-stellate hairs; calyx tube 0.5 mm long, calyx lobes hyaline, rounded-triangular, 0.25-1.5 mm long, calyx teeth linear to subulate, 2-4 mm long. Petals 4.5-5 × 1-2 mm, white or reportedly pink, oblong to oblong-ovate, reflexed, glabrous, truncate or rounded. Stamens 10, radially arranged around the style, the filament 2-2.5 mm long with a geniculation near the apex, yellow, anthers 1.5-2 × 0.5 mm, linear-oblong, yellow, the pore ca. 0.1 mm, terminal and slightly ventrally inclined. Ovary 5-locular, 2/3 inferior, apex elevated into a low ringlike collar with or without smooth hairs that surround the stylar scar. Style ca. 7 mm long, straight to slightly curving, vertical distance from the anther apex to the stigma ca. 1.5-2.5 mm, horizontal distance absent, stigma punctiform, ca. 0.25 wide. Berry 5 × 5 mm, purple black. Seeds ca. 0.5 mm long, more or less triangular in profile, angulate and somewhat muriculate to papillate on the convex face. Distribution (Fig. 136). Primary forest understory in the Golfo Dulce and Osa Peninsula region in southern Costa Rica, and a population in Panama on the Santa Rita Ridge, from sea level to 400 m elevation.

Conostegia allenii
Conostegia allenii can be distinguished by its sessile to subsessile leaves that usually have oblique bases, pseudolateral inflorescences, and reddish hirsute indument on inflorescence branches and hypanthium. In the protologue Almeda (2004)  ing glabrous adaxial foliar surfaces. An additional specimen from this locality (Aguilar 13243-NY) also has glabrous leaves adaxially. Th e latter specimen was accompanied by photographs of the flowering plant (Fig. 135) which evidenced the white petals of the flowers. Other Costa Rican specimens reportedly have pink flowers, and the only other report of white petals in this species are the only known Panamanian specimen cited above. Conostegia allenii represents a strong case of vegetative convergence with taxa such as Clidemia costaricensis and Clidemia petiolaris. Almeda (2004)  Description. Small trees 1.5-10 m tall with rounded-quadrate stems that are moderately to sparsely ferrugineous scurfy or stellulate-puberulent, to almost glabrous; the nodal line present but inconspicuous. Leaves at a node equal to unequal in length. Petioles 0.4-2 cm. Leaf blades 4-14.4 × 1.2-4.7 cm, 3-5 plinerved, with the innermost pair of primary veins diverging from the midvein 0.4-2 cm above the base in opposite or generally alternate fashion, elliptic, the base acute to obtuse and typically asymmetrical, the apex attenuate to acuminate, the margin entire to distally crenulate, the adaxial surface glabrous, the abaxial syrface glabrous on the surface and scurfy puberulent to glabrous on the veins, frequently with white dots irregularly spread on the lamina. Inflorescence a terminal panicle 2.8-7.5 cm long branching at or above the base, the rachis glabrous to scurfy puberulent, the branches very thin, the bracteoles 0.5-1 mm, narrowly triangular to subulate, persistent and fused basallly forming a shallow inconspicuous nodal collar or elevated ridge. Pedicels 0.5-2 mm. Flowers 5 merous, not calyptrate, floral buds 1.5-3.75 × 0.9-2.0 mm, the hypanthuim 1.5-2 × 1.25-1.75 mm, scurfy puberulent, calyx tube ca. 0.1 mm long, the calyx lobes depressed undulate, 0.5 × 1 mm, the calyx teeth bluntly triangular, 0.25 mm long, equaling or barely exceeding the calyx lobes. Petals 1.5-2.5 × 1-1.5 mm, white, oblong, glabrous reflexed, emarginate. Stamens 10, 2-2.5 mm long, actinomorphic, the filaments 1.25-1.75 mm, with a geniculation near the middle, white, anthers 0.5-1 × 0.25-0.55 mm, cuneate in outline and widest at the apex, white or yellow, the connective prolonged briefly below thecae, the pore subterminal ca. 0.3 mm wide, dorsally inclined. Ovary 5 locular, inferior, glabrous or scurfy puberulent apically. Style 3.25-4 mm long, straight, vertical distance from stamens to stigma ca. 0.5-1 mm, horizontal distance absent, the stigma punctiform to truncate, ca. 0.35 wide. Berry 4.25-5.25 × 4.25-5.25 mm, purple black. Seeds 0.34-0.5 mm long, more or less pyramidal, frequently asymmetrical. Distribution (Fig. 138). Endemic to cloud forests in Costa Rica from 550-2100 m elevation.
Th e epithet brenesiana was chosen for this species because brenesii is preempted by Conostegia brenesii Standl. Conostegia brenesiana can be recognized because of its almost glabrous vegetative parts, plinerved leaves frequently with asymmetric venation, wiry inflorescences with persistent bracteoles and small five merous flowers. In addition, this species has the widest anther pores in Conostegia and some of the smallest anthers. Almeda (2009) reports the ovary of this species as glabrous. Th e specimen I studied from spirit had small glands suggesting intraspecific variation. Further observations should be made in this species considering it is also variable in anther color.  Some specimens report the flowers as totally white while others report the anthers as yellow. An interesting aspect of this variation concerns the possible transition of pollination systems in this species. C. brenesiana inhabits cloud forests mainly at middle elevation but spanning a broad range. An interesting hypothesis to test is if the more white flowers occur at higher elevations where the buzz pollinated species tend to decline. Th e species already presents the broad anther pores which in themselves hint to a transition that might be happening. One specimen from Cerro Hornito in western Panama (Kriebel and Burke 5752) might represent this species but the inflorescence is too immature to tell.  Description. Shrub to small tree 2.5-5 m tall with terete stems which are covered with a rusty mixture of stellate and stipitate stellate trichomes; the nodal line covered by indument. Petiole 0.4-1.4 cm. Leaf blade 4.5-19.5 × 2.5-9 cm, 5(-7) plinerved, with the inner pair of primary nerves diverging from the mid vein 0.4-1.5 cm above the blade base in opposite, sub opposite or alternate fashion, elliptic to elliptic ovate, the base typically obtuse to rounded but varying to asymmetrical and briefly decurrent, the apex acuminate, the margin undulate to undulate-dentate, the adaxial surface sparingly stellate to glabrous, abaxially copiously stellate on the elevated primary veins with a sparser cover on the transverse secondary and higher order veins. Inflorescence a terminal, erect or deflexed panicle 2-8 cm long, branched above the base, with flowers in terminal congested glomerules, accessory branches absent; bracteoles 0.5-1.5 × 0.25-5 mm, linear-oblong, persistent. Pedicel 0.5 mm long. Flowers 4-merous, not calyptrate but flower bud closed and crowned by an apiculum ca. 0.25 mm long that ruptures at anthesis into 2 to 4 deltoid and hyaline lobes mostly 1 × 1-1.5 m the calyx teeth linearoblong, 1 mm long, the hypanthium 2-3.75 × 2-2.5 mm, densely stellate pubescent. Petals 3-4 × 1.5-2.5 mm, white, obovate to oblong-obovate, apparently reflexed at anthesis, glabrous, rounded to emarginated apically. Stamens 8, 3-3.5 mm, actinomorphic, the filaments 1.5-2 mm, with a geniculation near the apex, white, anthers 1-1.5 × 0.5-0.75 mm, linear-oblong, laterally compressed, the connective not prolonged nor appendaged, yellow, the pore ca. 0.17 mm, omewhat ventrally inclined. Ovary 4-locular, 3/4 inferior, the apex glabrous and not forming a collar around the style base. Style 5-7 mm long, exserted and straight to slightly curving towards the apex, vertical distance from the anthers to the stigma ca. 1.5-2 mm, horizontal distance absent; stigma punctiform, 0.25-0.5 mm wide. Berry 6-8 × 6-8 mm, purple black. Seeds ca. 1 mm long, obovoid to pyriform, angulate and with a densely tuberculate testa. Distribution (Fig. 140). Endemic to Caribbean lowlands and foothills of Costa Rica, from sea level to 600 m in elevation. To be expected in the south eastern lowlands of Nicaragua.
Conostegia calocoma can be recognized on the basis of its rusty stellate pubescence, short petiolate leaves with plinerved venation and undulate margin, fused floral buds in which the calyx ruptures irregularly, 4 merous flowers, and bright yellow anthers. Th e molecular phylogenetic study revealed strong support for a clade comprised of C. calocoma as sister to the species pair C. colliculosa and C. subpeltata. A high quality line drawing was provided in the protologue of this species (Almeda 1989a).  Description. Small trees 3-7 m tall with flattened and two edged stems when young that become terete with age and which are reddish brown and densely stellulate-lepidote on young branches; the nodal line nodal line evident. Leaves at a node equal to subequal in length. Petiole 0.5-2 cm. Leaf blades 5.25-12.5 × 1.5-4.5 cm, 3-plinerved, with the inner pair of primary nerves diverging from the mid vein 0.4-3 cm above the blade base in opposite, subopposite or alternate fashion, elliptic to elliptic lanceolate, the base acute to decurrent on the petiole, the apex acuminate, the margin entire or slightly denticulate, the adaxial surface glabrous, the abaxial surface reddish brown and densely stellulate-lepidote. Inflorescence a terminal little branched panicle 1.6-4.8 cm long with the branches terminating in three sessile-flowered clusters, accessory branches mostly absent, bracteoles 3-4 mm long, linear-oblong, early deciduous. Floral buds ca. 4-5 × 2-3 mm, campanulate to urceolate. Flowers 5-merous, not calyptrate, but with the calyx lobes fused in bud and rupturing at anthesis (with the style sticking out initially) into irregular persistent lobes ca. 2 mm long, the calyx teeth obsolete or evident as blunt protuberances at or near the torus, the hypanthium 2-3 × 2-3 mm, densely reddish brown and densely stellulate-lepidote. Petals 3.75-4.25 × 2-2.5 mm, white, oblong-obovate, spreading, glabrous, truncate to asymmetrical apically. Stamens 10, 4.5-5.25 mm long, radially arranged around the style, the filament 2.5-3.25 mm, with a geniculation near the apex, white, anthers 1.5-2 × 0.5-0.75 mm, linear oblong, yellow, laterally compressed, the pore ca. 0.2 mm, terminal. Ovary 5-locular, inferior, costate apically and forming a setose collar around the style. Style 6-6.5 mm, exserted and straight, vertical distance from the anther to the stigma ca. 1.75-2 mm from the stigma, horizontal distance absent, the stigma truncate to slightly expanded, 0.65-0.85 mm wide. Berry 3-4 × 3-4 mm, purple black. Seeds ca 0.5 mm, cuneate, angularly ridged with a conspicuous spur on the distal truncate surface. Distribution (Fig. 142). Endemic to central Panama where it has been collected in the foothills of Cerro Jefe and El Llano Cartí Road, 200-750 m elevation. Almeda (1983) compared Conostegia centrosperma to C. xalapensis in the protologue, a species from which C. centrosperma diff ers by its non-calyptrate calyx, more lepidote than stellate indument, exserted style, and setose rim around the style base. Almeda (1983) also compared C. centrosperma to C. fulvostellata and C. oligocephala which interestingly also belong to section Geniculatae. Th ese two species diff er from C. centrosperma in their non rupturing calyx and stellate indument. When Almeda described C. centrosperma, C. dissitinervia had not been described. When C. dissitinervia was described (Kriebel et al. 2005) its possible relationship to C. centrosperma was discussed and both taxa were separated in a dichotomous key. Th ese two species form a strongly supported sister pair in the molecular phylogenetic analyses. Th ey can be distinguished by the larger leaves with stellate indument abaxially and longer inflorescences in C. dissitinervia. In addition to the hyaline calyx of the latter when rupturing, does not leave the evident calyx lobe pieces of C. centrosperma. Lastly, C. dissitinervia lacks the setae around the style base present in C. centrosperma and has an apiculate anther apex that C. centrosperma lacks.   Fig. 143 Conostegia cinnamomea (Beurl.) Wurdack, Phytologia 38: 287. 1978 Description. Shrubs to small tress 1.5-6 m tall with terete, sometimes slightly rectangular stems especially towards the apex which are moderately to densely covered on new growth with short dendritic hairs; the nodal line present yet slight. Leaves at a node equal to unequal in length. Petioles absent or to 1.3 cm long. Leaves 4.5-20.1 × 1.9-6.9 cm long, 5-plinerved, with the innermost pair of veins arising up to 4 cm above the blade base in opposite or more commonly strongly alternate fashion, elliptic, acute to cuneate, attenuate to acuminate, margin entire to crenate, glabrous adaxially, abaxially glabrous or with a few scattered stellate furfuraceous trichomes on the main veins. Inflorescence a terminal reflexed and pseudoaxillary panicle 2.6-7.6 cm long branching at the base, accessory branches absent, bracteoles lanceolate, to about 2 mm, persistent. Pedicel 0.5-1 mm. Flowers (4-)5 merous, calyptrate. Flower buds 2.3-4 × 1.1-1.5 mm, elliptic to slightly pyriform, rounded at the base, apiculate at the apex, calyx teeth present but minute and not discernable to the naked eye, the calycine and hypanthium diff erentiated with the calyptra tending to dry white, slightly to not constricted below the calyptra; the hypanthium 2-2.5 × 2-2.5 mm, glabrescent. Petals 3-4.25 × 1.75-2.5 mm, white, turning pink and closing with age, narrowly ovate, reflexed, glabrous, apically acute. Stamens (8-)10, 3.25-4.25 mm long, radially arranged around the style, the filament 1.75-2.25 mm, with a geniculation near the apex, white, anthers 1.5-2 × 0.5-0.75 mm, oblong, yellow, ventrally wrinkled, the pore 0.18-0.2 mm, dorsally inclined. Ovary 5 locular, inferior, apically glabrous and not forming a collar around the style base. Style 5.75-6.25 mm, straight to gently curving, vertical distance from the anther apex to the stigma 1.5-2 mm, horizontal distance absent, stigma punctiform, 0.35-0.45 mm wide. Berry 4-5 × 4-5 mm, blueblack. Seeds 0.4-0.5(-0.8) mm, roughly pyramidal, the testa tuberculate. Distribution (Fig. 144). Nicaragua, Costa Rica, Panama, Colombia and northern Venezuela, from sea level to 1450 m elevation.

Conostegia cinnamomea (Beurl.) Wurdack
Conostegia cinnamomea is easily recognized its glabrous appearance, evidently plinerved leaves, deflexed inflorescences and calyptrate calyx. Its bracteoles are also helpful when flowers are lacking because they are persistent and form a nodal collar around the inflorescence branches. Th e style of this species is exserted and one of the bees observed to visit its flowers, Melipona costaricensis, curls its abdomen over the style before buzzing the flowers (Fig. 143). Th is same behaviour was also observed by another small halictid bee in a species with similar floral morphology, Conostegia grayumii. It is possible most of the species in section Geniculatae are buzzed in a similar way since they have floral morphology very similar to the above mentioned taxa. Th e anther pore in this species is perhaps the most strongly dorsally inclined in any species of the genus.  Description. Shrub to small trees 1.5-6 m tall with terete branches that are apically densely covered with brown penicillate-stellate hairs, older branches glabrous; the nodal line present yet slight and covered by indument. Leaves at a node equal to subequal in length. Petiole 0.4-1.5 cm. Leaf blades 9.5-20.5 × 4.2-10.6 cm, 5(-7) plinerved, with the inner pairs of primary nerves diverging from the mid vein up to 2 cm above the base in sub opposite or alternate fashion, elliptic-ovate, the base obtuse to rounded and frequently oblique, slightly decurrent on the petiole, the apex acuminate, the margin undulate-denticulate to almost entire, the adaxial surface glabrous at maturity, the abaxial surface with penicillate stellate hairs on the elevated primary veins and stellate hairs on the secondary and higher order veins. Inflorescence a terminal panicle with ultimate branches termi-nating in simple dichasia appearing pseudolateral from elongation of lateral branches, sometimes deflexed, 3.5-8 cm long, branching above the base, accessory branches absent, densely covered with brown penicillate-stellate and / or coarse dendritic hairs, bracteoles 1-4 mm × 0.25-1 mm, linear-oblong, persistent. Pedicel lacking or to 0.25 mm long. Flowers 5-merous, calyx not calyptrate. Flower buds not seen; the hypanthium 2-2.5 × 1.75-2.25 mm, densely covered with brown penicillate-stellate and/or coarse dendritic hairs; the calyx tube 0.5 mm long, the calyx lobes ovate to suborbicular, often mucronate at the apex, stellate pubescent on both surfaces, the calyx teeth subulate, 1.5-2 mm long. Petals 3-4 × 1.5-2 mm, white, oblong-obovate, petal posture uncertain, the apex mostly rounded, glabrous. Stamens 10, ca. 2.5-3.5 mm long, radially arranged at anthesis, the filament 1-2 mm, with a geniculation near the apex, white, anthers 1-1.5 × 0.5-0.75 mm, elliptic, yellow, the pore 0.1 mm wide, dorsally inclined. Ovary 5-locular, inferior, costate apically and with a lobed glandular puberulent crown around the style base. Style 3.15-3.45 mm, exserted and straight to slightly curving, vertical distance from the anther to the stigma 1.45-1.65 mm, horizontal distance absent, stigma punctiform, 0.25-04 mm wide. Berry 4-5 × 4-5 mm, not seen at maturity. Seeds 0.5 mm long, triangular in outline, testa colliculose. Distribution (Fig. 146). Restricted to the Caribbean slope of Cost Rica and Panama, 150-650 m in elevation.
Th is rare species was assigned to Miconia section Ambyarrhena by Almeda (2000). Th e drawing in the protologue and study of the type specimen confirmed that the style is exserted and that the anthers are yellow, like most of its relatives, including Conostegia calocoma and C. subpeltata. With the latter two species it also shares an irregularly rupturing calyx but in the case of C. colliculosa the lobes are thicker and the calyx teeth longer. In addition, C. colliculosa has five merous sessile flowers disposed in bracteate clusters whereas C. calocoma and C. subpeltata have four merous fl owers that are not clustered in the infl orescences. C. supeltata is further distinguished from C. colliculosa in having subpeltate leaves and axillary inflorescences. Almeda (2000)   Conostegia consimilis (Gleason) Kriebel, comb. nov. Basionym: Leandra consimilis Gleason, Ann. Missouri Bot. Gard. 45(3): 268. 1958. Type: Panama. Panamá: Las Minas, 9 January 1941 Allen 2702 (holotype: NY!). Miconia ligulata Almeda, Proc. Calif. Acad. Sci. 46(9): 216. 1989.
Conostegia consimilis is a common species especially in lowlands and middle elevation in Costa Rica and Panama. It can be recognized by its large leaves with decurrent bases and strongly plinerved venation. Its flowers are quite small and its petals are quite narrow and somewhat acute at the apex which prompted in its original description in the genus Leandra. Other useful characters are its yellow anthers, exserted styles, and angulate seeds with verruculose angles. In general, C. consimilis shares its rusty indument and petal shape with its close relatives which include C. iteophylla, C. jefensis, C. peltata, and C. trichosantha. Of the latter taxa, it is most similar to C. iteophylla and the common morphotype of C. jefensis. From C. iteophylla it can be distinguished by its much larger leaves, and from C. jefensis by its white petals (vs. magenta). Th e drawing in the protologue (Almeda 2000) evidences the exserted style which is common in section Geniculatae.  Description. Sparingly branched shrub 1-3 m tall with terete and glabrous internodes; the distal branches moderately to sparingly stellate-furfuraceous; the nodal line present. Leaves of a pair somewhat unequal in size, sessile and clasping or with petioles 1-9 mm long. Leaf blade 4-22 × 1.3-10 cm, 5-7 plinerved, with the innermost pair of veins diverging 0.4-3.5 cm above the blade base in sub opposite or alternate fashion, elliptic to elliptic ovate, the base rounded to subcordate and sometimes oblique, the apex acuminate, the margin undulate-dentate to subentire, adaxially glabrous, abaxially glabrous or with some stellate hairs on the elevated primary veins. Inflorescence a terminal, erect or reflexed divaricately branched, paniculiform dichasium, 6.4-20.5 cm long, branching at or above the base, accessory branches absent, branches glabrescent, thin and wiry, bracteoles paired, sessile and fused into a short nodal collar forming an elevated internode ridge, 0.5 mm long, lance triangular to subulate, persistent. Pedicels 1.5-2.5 mm long. Flowers 5 merous, calyx in bud not calyptrate but closed in bud and crowned by an apiculum, rupturing irregularly at anthesis into 2-5 hyaline lobes 1-1.5 mm long. Flower buds 1.75-2.25 × 1.5-1.75 mm, hypanthium 1.5-1.85 × 1.5-1.65 mm, campanulate, glabrous to stellate puberulent, calyx tube ca. 0.5 mm long, the calyx teeth subulate, 0.5 mm long. Petals 3.25-3.75 × 1.5-2 mm, translucent white, oblong-lanceolate, reflexed at anthesis, the apex acute to retuse, glabrous. Stamens 10, alternately unequal with the larger stamens inserted on the torus opposite the petals and the small ones opposite the calyx lobes, radially arranged around the style, the filaments 1.5-2 mm, with a geniculation near the apex, white, anthers 1.5-2 × 0.5 mm, linear-oblong, yellow, laterally compressed, the pore 0.15 mm, somewhat dorsally inclined. Ovary 5-locular, inferior, minutely puberulent at the apex. Style 2.5-3 mm, straight, vertical distance from the anther to the stigma 1.25-1.5 mm, horizontal distance absent, stigma truncate, 0.3-0.5 mm wide. Berry 3-4 × 3-4 mm, purple black. Seeds 0.36-0.7 mm, mostly ovoid, with densely muricate or verrucose testa. Distribution (Fig. 150). Endemic to south Pacific Costa Rica, mainly in the Golfo Dulce region and the Osa Peninsula but also towards to mountains on the road from San Vito de Coto Brus to Ciudad Neily, from sea level to 660 m elevation.
Conostegia dissitiflora is a distinctive species on the basis of its overall glabrous, plinerved, sessile leaves, and 5-merous flowers with an irregularly rupturing calyx. Almeda (1989a) discussed an unusual feature of this species which is the presence of "anisomorphic" stamens, with the larger set of stamens positioned opposite the petals. Th e    Fig. 152). Restricted to central and south Pacific slope of Costa Rica and barely getting into Panama, 0-600 m in elevation.
Conostegia dissitinervia can be recognized by its large leaves with acute to decurrent leaf bases that are abaxially covered with stellate trichomes and are strongly plinerved. In addition, the flowers have a fused calyx that ruptures irregularly and this species tends to have a very exserted style as many other species in section Geniculatae. Th e acute anther apex is distinctive. Th is species has been confused with Miconia argentea in herbaria because of the similar colored abaxial leaf surface as a result of the leaf indument. Th ey are easily distinguished because M. argentea has nerved leaf venation whereas C. dissitinervia is strongly plinerved. In the molecular phylogeny this species forms a well supported sister pair with C. centrosperma which it resembles. See the discussion under the latter species for the diff erences between the two. Specimens examined. COSTA RICA. Puntarenas: Península de Osa, about 5 km west of Rincón de Osa, Burger and Liesner 7253 (NY); Península de Osa, Cerca del Río Piro, Kriebel et al. 5046 (INB,NY); Península de Osa, Guadalupe La Tarde, Kriebel et al. 5377 (INB,NY).

Conostegia ecuadorensis (Gleason) Kriebel, comb. nov.
On the other hand, he did not place C. ombrophila in any section or compare to any other species, stating that its position in Clidemia was uncertain.  Description. Shrub 1-1.5 m tall with terete branches that have a sparse to dense covering of smooth spreading hairs (2-3.5 mm) intermixed with inconspicuous, early deciduous, asperous headed underlain by a moderate to dense understory of stellulate-furfuraceous or short asperous headed hairs but that become glabrate at maturity; the nodal line absent. Leaves at a node equal to somewhat unequal in size. Petiole 0.5-1.5 cm. Leaf blade 6-10 × 2.5-4 cm, 5-nerved to slightly plinerved, elliptic, the apex acute to acuminate, the margin entire to crenulate, adaxially moderately strigose to subhirsute with hairs mostly 1-2 mm long, abaxially moderately hirsute with a mixture of smooth hairs (1-2.5 mm long) and minute glandular hairs essentially restricted to the primary and higher order veins. Inflorescence a pseudolateral modified dichasium 3-5 cm long divaricately branched at the base and appearing axillary on older nodes, accessory branches absent, rachis sparingly setulose, bracteoles lanceolate to ovate, 1.5-3 × 1-1.5 mm, paired at each node and persistent, setose in between the bracteoles. Pedicels 2-3 mm long. Flower buds 2 × 1.5 mm, the hypanthium campanulate, ca. 1.5-2 mm long, densely covered with spreading smooth hairs 1-2 mm long and a sparse understory of sessile stellulate furfuraceous hairs.
0.15 mm wide, dorsally inclined. Ovary 5-locular, totally inferior, apex flat and glabrous except for s few minute glands. Style ca. 5.7-5.9 mm long, straight to slightly bending, vertical distance from the anther to the stigma 1.9-2.15 mm, horizontal distance absent, stigma truncate, 0.25-0.37 mm wide. Berry 3.5 × 3.5 mm when dry, purple-black. Seeds ca. 0.5 mm long, pyramidal, the testa asperulate or undulate. Distribution (Fig. 157). On the foothills of the Pacific slope of Talamanca Cordillera in Valle del General and San Vito de Coto Brus and barely reaching neighboring Panama, 950-1300 m in elevation. One outlying population collected on the south side of Lake Arenal in northern Costa Rica.
Conostegia fraterna is a distinctive species that can be identified on the basis of its short petiolate, narrow, plinerved leaves, and unusual inflorescence which consists of pedunculate mostly 3-flowered glomerules. Th is kind of inflorescence is not seen in any other species of Conostegia. Th e sessile flowers with fused buds that rupture at anthesis, herkogamous flowers, and broad anthers are also helpful to place it in section Geniculatae. C. fraterna did not fall as sister to any species with support in the recent molecular phylogeny. Nevertheless, it fell in a clade that includes similar looking species such as C. cinnamomea, C. dissitiflora and C. grayumii. All these species are glabrous or just barely pubescent and plinerved like C. fraterna. Gleason (1937) compared this species to Clidemia conglomerata and Clidemia involucrata. Of the latter two species, he believed C. involucrata to be the closest relative to C. fraterna. C. involucrata diff ers in its glandular pubescence, ciliate-serrulate margins and 3-locular ovary. Based on the molecular phylogeny available , we know they are not closely related at all. A seed micrograph of this species is provided in Ocampo and .  Description. Shrub to small tree 2-5 m tall with somewhat tetragonal and ridged stems in newer branches that are densely covered with inconspicuously stalked reddish-orange asperous-headed hairs usually also with some simple multicellular hairs on the distal internodes; the nodal line nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 0.5-4.8 cm. Leaf blades 7.5-18.6 × 3.4-7.7 cm, 5-plinerved, with the innermost pair of veins arising up to about 4 cm above the base in opposite or frequently in strongly alternate fashion, elliptic to elliptic-ovate, the base obtuse to oblique, the apex acuminate,the margin entire to inconspicuously denticulate, the adaxial surface mostly glabrous, moderately to sparsely covered with a mixture of inconspicuously asperous headed and scalelike multicellular hairs on the secondary and higher order veins abaxially. Inflorescence a mostly deflexed terminal modified cyme branching at the base, 3.6-9.8 cm long, accessory branches absent, rachis covered with stalked asperous headed hairs, linear oblong to triangular, bracteoles 1.5-4 mm, persistent. Pedicel 1-1.75 mm long. Flowers 5-merous, calyx in bud not calyptrate but closed in bud and crowned by an apiculum, rupturing irregularly at anthesis into 4-5 hyaline irregular lobes 0.3-1.5 mm long, flower buds 3-4 × 2-2.5 mm, the hypan- thium 1.5-2.5 × 1.75-2.25 mm, campanulate, covered with orangish asperous headed hairs; exterior calyx teeth conspicuous, triangular, 0.5-0.6 mm long, torus glabrous. Petals 3-3.5 × 0.9-1.25 mm, translucent pink, linear-oblong, reflexed at anthesis, glabrous, the apex acute. Stamens 10, 2.2-3 mm, radially arranged around the style, the filaments 1-1.75 mm, geniculate near the apex, white, anthers 1-1.25 × 0.25-0.75 mm, linear-oblong, yellow, laterally compressed, the pore 0.1-0.14 mm wide, terminal and slightly dorsally inclined. Ovary 5 locular, 3/4 inferior, slightly fluted and with glandular hairs on the apex. Style 5.25-5.5 mm long, slightly to strongly bending, vertical distance from anther pore to stigma 1.75-2.25 mm, horizontal distance absent or up to1.3 mm, stigma truncate and somewhat dilated, 0.25-35 mm wide. Berry 3-4 × 3-4 mm, red turning purple black when mature. Seeds 0.39-0.6 mm, angular pyramidate to somewhat cescent shaped in profile, the testa smooth. Distribution (Fig. 159). Endemic to the Caribbean slope of the Cordillera de Tilarán and Volcán Tenorio National Park in Costa Rica, 1250-1700 m in elevation.
Conostegia friedmaniorum can be recognized by its rusty pubescence, leaves which are strongly plinerved with the veins frequently arising in asymmetric fashion, deflexed inflorescences and linear-oblong petals. Vegetatively it resembles another Costa Rican endemic, C. pendula, as well as the Panamanian endemics C. galdamesiae and C. papillopetala. From C. pendula it can be distinguished because the latter has lanate indument on the stems which is absent in C. friedmaniorum. From C. galdamesiae and C. papillopetala it can be distinguished by its deflexed inflorescence and linear-oblong petals.
Almeda (2009)  petals. More specimens are needed to better establish the diff erences noted by Almeda. Almeda (1989a) compared C. fulvostellata in the protologue of M. ibarrae and noted diff erences in pubescence and notably in style curvature and seed characteristics. C. fulvostellata has a dense indument in abaxial leaf surface reminiscent of C. oligocephala. With the latter C. fulvostellata also shares an overall geographic range. C. fulvostellata diff ers from C. oligocephala in its small calyx teeth, four locular ovary and usually papillose petals (vs. evident calyx teeth, five locular ovary and glabrous petals in C. oligocephala).
Conostegia galdamesiae is a Panamanian cloud forest endemic that can be recognized on the basis of its reddish indument, broad leaves with extremely plinerved leaf veins which are frequently asymmetric, and inflorescences with clustered, sessile flowers that have a fused calyx in bud. It is similar to C. brenesiana, C. friedmaniroum, and C. papillopetala. From all of these species C. galdamesiae can be recognized by its inflorescences with the flowers clustered at the end of the branches. From C. brenesiana it can be further distinguished by its evident indument on branch apices and veins on the abaxial leaf surface. C. brenesiana also has a unique broad anther pore. From C. friedmaniroum it can be distinguished because the latter has deflexed inflorescences and linear-oblong petals. Lastly from C. papillopetala it can be distinguished because the latter has narrower, somewhat bullate leaves, and papillose pink petals.  Description. Shrub to small tree 1-5 m tall with subquadrate to terete stems which are sparsely to moderately ferrugineous scurfy-pulverulent to glabrous; the nodal line present but inconspicuous. Leaves of a pair somewhat unequal in size. Petioles 0.2-2.1 cm. Leaf blades 3.4-13 × 1.1-3.7 cm, 3-plinerved, with the inner pairs of subparallel veins arising up to 2 cm above the base opposite or generally alternate fashion, narrowly elliptic, the base broadly acute to obtuse and typically asymmetrical, the apex acuminate, the margin undulate denticulate to entire, glabrous above when mature, abaxially glabrous or scurfy pulverulent or glandular-pulverulent. Inflorescence a terminal and deflexed paniculiform cyme 2.4-5.7 cm long, sometimes becoming pseudolateral because of axillary bud elongation, accessory branches absent, rachis thin and glabrous, bracts and bracteoles to 3 mm long, lance-triangular to subulate, persistent and forming a nodal collar. Pedicel 2-2.5 mm. Flowers 5-merous, not calyptrate, nor the sepals fused in bud, floral buds ca.  Fig. 164). South eastern Nicaragua to the Caribbean lowlands of Costa Rica and one outlying population in the Osa Peninsula of Costa Rica on the Pacific side, 0-300 m in elevation.
Conostegia grayumii is a small tree restricted to lowland rain forests which can be recognized by its glabrosity, small plinerved and asymmetric leaves, and small flowers on wiry inflorescences. Th is species was identified before its recognition as a distinct taxon as C. brenesiana (as Miconia brenesii). Almeda (1989b) noticed important differences between both such as the fact that C. brenesiana has noticeably broader anther pores and lacks seed testa ornamentation present in C. grayumii. Lastly, Almeda (1989b) noted the diff erence in habitat occupied by both taxa. Conostegia. grayumii is restricted to the lowlands whereas C. brenesiana is mostly restricted to middle elevation cloud forests. Th e molecular phylogeny shed further light on the relationships of these taxa. In fact, they did not form a sister species pair. Conostegia grayumii appears more closely related to C. cinnamomea, C. dissitiflora, C. fraterna, and C. povedae than to C. brenesiana.
Specimens    Description. Shrub 1.5-3.25 m tall with slightly tetragonal branches at the apex that have a sparse to dense covering of smooth spreading hairs (2-3.5 mm) intermixed with inconspicuous, early deciduous, asperous headed hairs that are underlain by a moderate to dense understory of stellulate-furfuraceous or short asperous headed hairs but becoming glabrate at maturity; the nodal line absent. Leaves at each node equal to somewhat unequal in size. Petiole 0.5-4 cm. Leaf blade 7.4-29 × 3.7-14.5 cm, 5-7 nerved or if 5-7 plinerved, with the innermost pair of primary veins diverging from the midvein up to 2 cm above the base in opposite or alternate fashion, elliptic, the apex long acuminate, the margin entire to crenulate, adaxially moderately strigose to subhirsute with hairs mostly 1-2 mm long, abaxially moderately hirsute with a mixture of smooth hairs (1-2.5 mm long) and minute glandular hairs essentially restricted to the primary and higher order veins and with pocket mite domatia at the base. Inflorescence a pseudolateral modified dichasium 2.1-6 cm long divaricately branched at the base and appearing axillary on older nodes, accessory branches absent, bracteoles 1.5-3.5 × 0.5 mm, paired at a node and persistent, sometimes fused forming a nodal collar, lanceolate to subulate. Pedicels 1-2.5 mm long. Flower buds 2-2.5 × 1.9-2.4 mm. Flowers 5 merous, calyx not calyptrate but closed in bud and crowned by an apiculum 0.5 mm long and rupturing irregularly at anthesis into 3-5 hyaline, persistent lobes 0.5 mm long, 2-2.5 × 1.9-2.4 mm, ca. 0.25 mm, external calyx teeth triangular-subulate, 0.5-1 mm long, the hypanthium campanulate, 0.5-2 mm long, moderately to sparsely covered with spreading smooth hairs 0.5-2 mm long and a sparse understory of sessile stellulate furfuraceous hairs, the torus not evidently glandular puberulent but the inner hypanthium wall sparsely so. Petals 3.5-4 × 1.5-2 mm, white or translucent white, oblong, rounded apically, spreading at anthesis. Stamens 10, 2.5-4 mm long, radially arranged around the style, the filament 1.5-2.5 mm with a geniculation near the apex, translucent white, anthers 1-1.75 × 0.35-0.85 mm, oblong and laterally compressed, yellow, the pore 0.15 mm wide, somewhat dorsally inclined. Ovary 5 locular, totally inferior, apex fluted and elevated into a glandular-puberulent lobulate collar. Style 5-6 mm long, straight to slightly bending, vertical distance from the anther to the stigma 1.25-1.75 mm, horizontal distance absent; stigma truncate, 0.3-0.5 mm wide. Berry 6-7 × 5-6 mm, purple-black. Seeds 0.5-0.7, galeiform to deltoid, irregularly angulate with a densely papillate testa and a lateral flattened or somewhat convex raphe. Distribution (Fig. 166). Nicaragua, Costa Rica, Panama and reaching Colombia, 0-950 m in elevation.
Conostegia hammelii is easy to distinguish by the presence of pocket domatia on the base of the abaxial leaf surface, its pseudolateral inflorescences branched at the base, sepals that are fused in bud into an apiculum, ovary apex with a glandular-puberulent lobulate collar and papillose seed testa. Almeda (1989a) noted that some specimens have stamens that are somewhat unequal in length. Th is phenomenon is only present in C. dissitiflora within Conostegia. I have not noted this staminal length diff erence in my collections in spirit from Costa Rica and Panama of this species. In the molecular phylogeny, C. hammelii forms a relatively well supported clade with C. allenii, C. ecuadorensis, and C. ombrophila. Th is clade shares the pseudolateral inflorescences as well as the papillose seed testa. It is also noteworthy that three species of this clade, namely C. ecuadorensis, C. hammelii, and C. ombrophila are the only ones in section Geniculatae to have domatia. Almeda (1989a) reported on the presence of mites of the genus Ololaelaps found in the domatia of this species.     3-4 × 3-4 mm, pink, broadly obovate to subrotund, rounded to emarginated apically, glabrous. Stamens 10 or 12, 2.25-3 mm long, the filament 1.25-2, apparently lacking a geniculation near the apex, anthers 1 × 0.6-0.8 mm, yellow, laterally compressed, connective thickened, the pore 0.18-0.2 mm wide, terminal and somewhat dorsally inclined. Ovary 5-6 locular, inferior, apically glabrous and flat, with five or six callose thickenings around the style base. Style not seen. Berry 4-6 × 4-6 mm, purple black. Seeds 0.45-0.6 mm long, triangular in sideview, the testa angulate and smooth. Distribution (Fig. 168). Endemic to western Panama at 1300-2000 m. Th e epithet henripittieri was chosen for this species because pittieri is preempted by Conostegia pittieri Cogn. ex T. Durand. Conostegia henripitteri is unmistakeable because of its glabrous, subsessile, anisophyllous leaves. Although I was not able to see flowering material of this species, the isotype at F shows the evidently exserted style that is almost universal in section Geniculatae. Description. Trees 3-10 m tall with rounded-quadrate stems that are densely covered with a ferrugineous stellate indument; the nodal line not evident from the consipucous indument. Leaves of a pair equal to unequal in length. Petiole 0.5-2.8 cm. Leaves of a pair equal to somewhat unequal in length; leaf blade 4.3-18 × 1.9-5.9 cm, 5-plinerved, with the innermost pair of veins diverging from the midvein ca. 1-2 cm above the base in opposite or alternate fashion, elliptic to oblong-elliptic, the base acute to obtuse, the apex acuminate, the margin entire, the adaxial surface glabrous, the abaxial surface covered with a ferrugineous stellate indument. Inflorescence a terminal paniculiform dichasium, 2.5-7 cm long, accessory branches absent, rachis densely covered with a ferrugineous stellate indument; bracts and bracteoles triangular subulate, 1.5-3 × 0.5-1 mm, persistent. Flowers 5-merous, sessile or subsessile at anthesis, not calyptrate, nor the sepals fused in bud, floral buds ca. 8-9 mm, the calyx truncate and forming a flange ca. 1.5 mm long, exterior calyx teeth conspicuous, linear and more or less hooked, 2-7 mm long, torus glabrous. Petals 0.6-1.1 × 0.6-0.8 mm, white, broadly obovate, asymmetrical apparently spreading an anthesis, glabrous, emargiante apically. Stamens 10, 6.5-7 mm long, radially arranged around the style, the filament 3-3.25 mm, with a geniculation near the apex of the filament, white, anthers 3.25-3.75 × 0.75-1 mm, oblong, yellow, laterally compressed, the connective thickened, the pore ca. 0.15 mm wide, ventrally inclined. Ovary 5 locular, 4/5 inferior, glabrous and elevated into a collar around the style base. Style 8-8.5 mm long, vertical distance from the anther pore to the stigma ca. 1-2 mm, horizontal distance absent, the stigma capitate, ca. 1 mm wide. Berry 6-10 × 6-10 mm, reportedly light red. Seeds ca. 0.5 mm long, triangular to angulate-ovoid, the testa muriculate.
Distribution (Fig. 170). Endemic to the Pacific slope of the Tilarán, Central and Talamanca Cordilleras in Costa Rica, 500-1500 in elevation. Conostegia incurva is a rare species that can be recognized by the ferrugineous indument that covers the abaxial leaf surface, the strongly plinerved leaves with frequently asymmetrical primary veins, and large flowers with linear and evident calyx teeth. It resembles C. schlimii, but the latter lacks the linear calyx teeth. Interestingly, C. incurva forms a strongly supported species pair with the northern Central American endemic C. oligocephala in the molecular phylogeny. Th e latter has much smaller flowers and more whitish indument on the leaves abaxially. Th e specimen Herrera 627 (CAS, CR) mentions the style separate from the stamens like most other species in section Geniculatae. Description. Shrubs 0.4-1.5 m tall with sub quadrate to terete branches moderately covered with a brown scurfy-pulverulent indument; the nodal line nodal line present. Leaves of a pair equal to somewhat unequal in size. Petiole 2.5-1.1 cm. Leaf blade 1.7-9.5 × 0.4-1.7 cm, 3-plinerved, with the innermost pair of veins diverging from the midvein 0.2-0.7 mm above the base in opposite or sub opposite fashion, narrowly elliptic, the base acute to attenuate, the apex acuminate, the margin entire, deciduously pulverulent adaxially when young but soon becoming glabrous, the abaxial surface deciduously scurfy pulverulent on the veins. Inflorescence a terminal paniculate cyme 2-7 cm long, branching above the base, accessory branches absent, rachis brown scurfy, bracteoles 0.5-3.5 × 0.25-0.5 mm, linear-oblong to triangular, sessile and persistent, paired and fused laterally into a short nodal collar. Pedicel 1-2 mm. Flowers 5-merous, calyx in bud not calyptrate nor fused in bud, hypanthium 1-1.5 mm, campanulate, deciduously scurfy pulverulent, calyx tube 0.25 mm long, the lobes barely discernible as triangular undulations, calyx teeth subulate ca. 0.25 mm long. Petals 2-2.5 × 0.6-0.8 mm, white, elliptic oblong, their posture not observed at anthesis, glabrous and entire, narrowly rounded to acute apically. Stamens 10, 2.5-3 mm long, apparently radially arranged around the style, the filament 1.25-1.5 mm, the geniculation near the apex of the filament apparently present, white, anthers 1.25-1.5 × 0.25-0.5 mm, linear-oblong, yellow, laterally compressed, the connective thickened dorsally, the pore ca. 0.1 mm wide, dorsally inclined. Ovary 5 locular, 2/3 inferior, the apex fluted and glandular puberulent. Style 3.5-4 mm long, distance from the anther pore to the stigma ca. 1 mm, the stigma capitate, ca. 0.3 mm wide. Berry 2-4 × 3-4 mm, red turning purple black. Seeds ca. 0.5 mm long, irregularly triangular to angulate-ovoid, the testa smooth and polished on the convex face.
When Almeda (1989a) described Miconia iteophylla he noted the strong similarity between it and M. ligulata (here combined to Conostegia consimilis). Indeed it appears to be a narrow leaf C. consimilis. Unfortunately I have not been able to place M. iteophylla in the molecular phylogeny and test this possible close relationship. A line drawing of this species is available in the protologue (Almeda 1989a (Fig. 173). Endemic to Cerro Jefe in Panama from 750-1000 m elevation. Conostegia jefensis can be recognized by its reddish scurfy pubescence, strongly plinerved leaves and magenta petals. Th is is a puzzling species because it includes two morphotypes endemic to Cerro Jefe. Th e common morphotype resembles C. consimilis in the decurrent leaf base but diff ers from it in its coarse habit, somewhat coriaceous leaves and magenta petals. Th e second morphotype which corresponds to the mor- phology of the holotype is apparently much less common and does not have the decurrent leaf base but rather a rounded to cordate leaf base. Th is morphotype also has the typical magenta petals of the other morphotype. Unexpectedly, the two morphotypes did not form a clade in the molecular phylogeny. Th e holotype matching morphotype formed a strong sister relationship to another Cerro Jefe endemic, C. peltata. Lack of resolution and more samples of each morphotype are needed at this time to further confirm the possibility that they represent diff erent taxa.
Specimens Description. Shrubs to small trees 2-12 m tall with somewhat flattened cauline internodes that are densely covered with a whitish and ferrugineaous stellate indumentum; the nodal line not evident from the indument. Leaves of a pair equal to unequal in length.Petioles 0.5-3 cm. Leaf blade 3.5-14.2 × 1.5-5 cm, 5-plinerved, with the innermost pair of veins diverging in from the midvein 0.5-2 cm above the base in opposite or usually alternate fashion, elliptic, the base acute to obtuse, the apex acuminate, the margin denticulate to subentire, the adaxial surface glabrous, the abaxial surface densely covered with white stellate trichomes in between the veins and white intermixed with ferrugineus trichomes on the main veins. Inflorescence a terminal panicle 3.6-8 cm long, branching at or above the base, accessory branches absent, the axes covered with stellate hairs; bracteoles 2-5 × 0.5-3 mm, elliptic to linear-oblong, persistent. Flowers sessile or subsessile, 5 merous, not calyptrate or with the sepals fused in bud and rupturing, hypanthium 2.75-3.25 × 2.5-3 mm, campanulate, covered with stellate hairs, calyx tube 0.5 mm long, calyx lobes rounded-triangular to oblong, 1.5 × 1.5-2 mm, oblong, 1-1.5 mm long. Petals 3.5-4 × 3-4.5 mm, white, obovate, spreading, glabrous the apex rounded to emarginate. Stamens 10, 5-6 mm long, radially arranged around the style, the filaments 2.5-3 mm, with a geniculation near the apex, anthers 2.5-3 mm, linear-oblong, yellow, laterally compressed, the pore ca. 0.2 mm wide, ventrally inclined. Ovary 5-locular, half inferior, densely sericeous with trichomes surrounding and overtopping the stylar scar. Style ca. 5-7 mm long, straight but somewhat bent in some specimens, vertical distance from the anther to the stigma ca. 2.5-3 mm, the stigma punctiform, Conostegia oligocephala can be recognized by its leaf abaxial surface which is obscured by whitish and ferrugineaous trichomes, sessile flowers subtended by persistent bracteoles, non calyptrate calyx, and linear calyx teeth. Th is species has been confused with C. plumosa and C. xalapensis but can be distinguished from them by the lack of a calyptrate calyx and presence of evident calyx teeth. It is also similar to another non-calyptrate specie, C. fulvostellata. See the latter species for diff erences between the two. Th e molecular phylogeny places C. oligocephala as sister to C. incurva with strong support. Th e two share a similar leaf indument and calyx teeth but are easy to separate based on inflorescence and flower size.
Conostegia ombrophila can be recognized by its almost entire glabrosity, relatively small plinerved leaves with caudate apices and pocket domatia usually present at the base of the lamina abaxially. It also has pseudolateral, divaricately branched and wiry inflorescences with persistent bracteoles fused at the base to form a small collar. Its anthers are laterally compressed and their shape is unusual in that they are somewhat ovate and conspicuously flatted at an angle towards the apex. C. ombrophila in general is a very uniform species morphologically but two morphotypes of the species are recognizable. Th e common morphotype occurs more commonly at middle elevation cloud forests and has leaf domatia at the base of the blade abaxially. Th e lowland morphotype which occurs for example at La Selva Biological Station in Costa Rica does not tend to have domatia. Suprisingly, the two morphotypes did not form a sister pair in the molecular phylogeny. Further morphological studies should be made and more populations collected for genetic analyses. Th e non domatia bearing morphotype can also resemble C. grayumii in places where they are sympatric like in the Sarapiquí area in Costa Rica. Th e latter can be distinguished by the more glabrous and smaller leaves of C. grayumii as well as the five merous flowers and seeds with tubercles on the edges in the latter. In the lowlands, it is not uncommon to find several fruit with galls.  Description. Trees 9-25 m tall with the uppermost flattened and two sided branchlets completely covered with a stellulate-lepidote indumenta; the nodal line present. Leaves of a pair equal to subequal in size. Petioles 1.5-4.5 cm long. Leaf blades 7.5-21 × 3-7 cm, 5-plinerved, with the innermost pair of primary veins diverging ca. 0.5-1.5 cm above the blade base in opposite, alternate or subalternate fashion, elliptic to elliptic-oblong or slightly elliptic-lanceolate, the base acute, the apex acute to acuminate, the margin entire to inconspicuously crenulate-denticulate, the adaxial surface glabrous and inconspicuously glandular puncticulate, the abaxial surface completely covered by peltate scales. Inflorescence a terminal panicle 10.8-15 cm long branching above the base, accessory branches present, reddish stellulate-lepidote indument throughout; bracts and bracteoles 1-1.5 × 0.25-0.5 mm, triangular, persistent. Pedicels 0.25-0.5 mm. Flowers 5-merous, calyx calyptrate, flower buds 3-4.5 × 2-2.75 mm, elliptic-ovate, calyx teeth, the base rounded, the apex acute, not constricted, the hypanthial and calyptrate portions diff erentiated in texture, color and indument diff erent, the calyptra being white or translucent, very thin and more sparsely pubescent than the hypanthium, the latter campanulate to urceolate, 2.5-3 × 2.25-2.75 mm, densely covered with peltate scales that grade into stellate hairs, the torus also beset with scales. Petals 5-8 × 3-6 mm, white, obtriangular, spreading at anthesis, emarginate at the apex, glabrous. Stamens 10, Stamens ca. 5-8 mm long, androecium slightly bilaterally symmetric, the filaments 2.5-4.5 mm, white, 3.25-3.75 × 1-1.5 m, oblong, yellow, laterally compressed, the connective dorsally thickened, the pore ca. 0.2 mm wide, slightly ventrally inclined. Ovary 5-locular, inferior, apically glabrous and slightly ribbed, forming a low collar around the style base, with pronounced elevated lines. Style ca. 5 mm long, straight and abruptly curved near the apex, vertical distance from the anther to the stigma ca. 0-0.5 mm, stigma punctiform to truncate, ca. 0.8 mm wide. Berry ca. 6 × 6 mm, purple black. Seeds 1.3-1.9 mm long, broadly pyramidate, rounded to bluntly angled on the convex face, the testa smooth. Distribution (Fig. 179). Endemic to the Osa Peninsula in southern Costa Rica, 40-200 m in elevation.
Conostegia osaensis is endemic to the Osa Peninsula of Costa Rica and quite distinctive as it tends to be a large tree up to about 25 m tall. In addition, the leaf abaxial surface is covered with a stellate lepidote indument and the flowers are calyptrate, not pleiostemonous, and have a short style. In the molecular phylogeny C. osaensis falls sister to the clade comprised of C. plumosa, C. speciosa, C. subcrustulata, and C. xalapensis. With these taxa C. osaensis shares the calyptrate calyx and the short style. C. osaensis has the largest seeds of any species in Conostegia.
Specimens examined. PANAMA. Panamá: Cerro Jefe, Aranda 185 (CAS); P. N. Chagres, Cerro Jefe, subiendo por calle principal hacia la cima, Kriebel  Description. Shrub to small tree 1.5-5 m tall with somewhat tetragonal stems in newer branches densely covered with a lanate indument of variable dendritic trichomes; the nodal line present. Petioles 1-7 cm. Leaves at a node equal or somewhat unequal at a node. Leaf blades 6.5-28 × 3.5-13.5 cm, 7-plinerved, with the inner pairs of innermost veins arising 1-5 cm above the base mostly in alternate fashion, elliptic to elliptic-ovate, the base obtuse to asymmetrical, the apex acuminate, the margin entire to inconspicuously denticulate, the adaxial surface strigose or lightly lanate to glabrous, lanate on the primary veins abaxially, and with furfuraceous hairs on higher order veins. Inflorescence a pendant modified cyme branching well above the base, 4-12 cm long, the flowers clustered at the end of the branches, accessory branches absent, rachis lanate, bracteoles 0.2-0.3 mm, persistent. Pedicel 0.5-2.5 mm. Flower buds 3-4 × 2-3 mm.
Flowers 5-merous, calyx not calyptrate but closed in bud and crowned by an apiculum, rupturing irregularly at anthesis into 5 hyaline irregular lobes 0.25-0.5 mm long, exterior calyx teeth narrowly triangular, 0.9-1.3 mm long, torus glabrous; the hypanthium 2-3 × 1-2 mm, campanulate. Petals 2.9-3.5 × 1 mm, pale pink, linear-oblong, not observed at anthesis, glabrous, the apex obtuse to rounded. Stamens 10, ca. 3-4 mm, radially arranged around the style, the filaments ca. 2 mm long, geniculate near the apex, white, anthers 1-1.5 × 0.35-0.5 mm, linear-oblong, somewhat recurved, yellow, the connective thickened dorsally, the pore 0.1 mm wide, slightly dorsally inclined. Ovary 5 locular, 2/3 inferior, slightly fluted and with glandular hairs on the apex. Style reportedly 2.5-5 mm long, not observed at anthesis but notes on the type suggest it is exserted, stigma truncate and somewhat dilated. Berry 4-5 × 4-5 mm, pink turning purple black when mature. Seeds 0.4-0.6 mm long, angular pyramidate to somewhat cescent shaped in profile, the testa smooth. Th e description of this species is for the most part based on the original one by Almeda and Umaña Dodero (1993). Few recent collections have been made since its description but as Almeda and Umaña Dodero (1993) predicted, the species has been collected in the Cordillera de Talamanca in Fila Matama, Limón. Th e notes on the type specimen describe the anthers around the style which suggests the style is exserted as in its close relative like C. friedmaniorum. Almeda and Umaña Dodero (1993) also noted the filament geniculation in this species, something not commonly done. Th ey mention strong herbivory on these plants as is frequently noted in species of section Geniculatae. Th e label on Almeda et al. 7245 (CAS) records a pleasant acidic flavor of the berries of this species.
Specimens examined. COSTA RICA. Cartago: Refugio Nacional de Vida Silvestre Tapantí, sendero Palmito about 6 km from main gate to refuge, Almeda et al.
Conostegia plumosa is a rare species and one of the few in the genus to have the abaxial leaf surface tan colored from the dense indument covering it. Th is characteristic leaf underside is shared with C. dissitinervia, C. centrosperma, C. incurva, C. oligocephala and C. xalapensis. In particular, C. plumosa has been confused with C. oligocephala and C. xalapensis, both of which occur in the same general area of northern Central America. Th e other species mentioned are endemic to southern Central America. C. plumosa shares the calyptrate calyx and short style with C. xalapensis but not with C. oligocephala which lacks the calyptrate calyx and has an exserted style. C. plumosa can be distinguished from C. xalapensis on the basis of its evident calyptra appendices which are absent in C. xalapensis.   Description. Shrub to small tree to 2.5-4 m tall. Young branches tomentose with golden to orange indument consisting of dendritic to stellate trichomes. Th e interpetiolar line inconspicuous and covered by indument as in the stems. Leaves sessile or subsessile, somewhat anisophyllous. Leaf blades 9-24 × 5-9 cm, 3-5-plinerved, with the innermost pair of veins diverging from the midvein 1.5-7 cm above the base generally in asymmetric fashion, elliptic to obovate, the base attenuate, apex acuminate, the margin subentire to denticulate, adaxially glabrous, abaxially moderately stellate pubescent on tertiary and higher order veins. Inflorescence a terminal and deflexed panicle with dichasial branches, 5-10 cm long, with about 3-7 sessile flowers clustered at the end of the branches; bracts 2, 3-10 mm long, elliptic to ovate, persistent; bracteoles 2, ovate, ca. 1-2 × 0.5-1.25 mm, persistent. Flower buds 4-5 mm long, with the petals forming an acuminate cone when mature. Hypanthium narrowly campanulate, sparsely stellate. Flowers 5(-6) merous, calyx not calyptrate but with the calyx lobes fused and irregularly rupturing in their early stages, the calyx tube 0.3-0.5 mm, the teeth 0.25-0.4 mm long, linear-oblong. Petals 4.5-5.5 × 2-2.5 mm, white, spreading at anthesis, elliptic to elliptic-ovate, apically acute, glabrous. Stamens 10(12), 4-5 mm long, radially arranged around the style, the filament 2.25-2.75 mm long with a geniculation near the apex, white, anthers 1.5-2 × 0.6-0.8 mm, elliptic, somewhat laterally compressed, lacking an appendage or collar at the anther filament junction, yellow, pore ca. 0.15 mm wide, terminating the truncate apex. Ovary 5-locular, half superior, the apex beset with small brown glandular trichomes and elevated into a collar ca. 0.25 mm high around the style. Style ca. 6.4 mm long, straight to slightly curving, the stigma truncate, ca. 0.38 wide, the distance between the anther apex and the stigma ca. 1.5 mm, stigma truncate to capitellate, ca. 0.3 mm wide. Berry and seeds not seen. Distribution (Fig. 187). Endemic to the remaining primary forest in San Vito de Coto Brus, Puntarenas, Costa Rica at 1100-1200 m elevation.
Conostegia povedae is a very narrow endemic easy to recognize because of its rusty indument, sessile leaves with attenuate leaf bases and which are strongly plinerved, and sessile flowers in clusters that when in bud have a fused calyx that ruptures irregularly. In their evidently yellow anthers, exserted styles, and geniculate filaments, C. povedae shows the typical floral morphology present in section Geniculatae.

m in elevation.
Conostegia schlimii is one of the most common species of Conostegia and an easy one to recognize. It has a conspicuous rusty indument on branch apices, the leaves are evidently plinerved and frequently with asymmetrically arising primary veins, as well as an undulate-denticulate margin. Th e flowers are quite large and have distinctive basally clawed petals, large bright yellow anthers and a capitate stigma. It is one of the largest flowered species in the genus and one of the few large flowered species not to be pleiostemonous. Its berries are also quite large. Flowers of this species have been observed being buzzed by Melipona costaricensis in the Osa Peninsula of Costa Rica. For such a common species in the Pacific slope of Costa Rica as well as being present towards the Caribbean slope in Nicaragua, Honduras, Guatemala and Belize, it is suprising it is absent in the Caribbean slope of Costa Rica. Based on morphology, C. schlimii is hard to confuse with any other taxon. Perhaps the one that most closely resembles C. schlimii, based on floral size and general morphology is C. incurva. Th e latter diff ers in its leaf abaxial surface being covered with indument and most noticeably in its claw like, conspicuous, calyx teeth.   Description. Shrubs 1-4 m tall with terete stems that are moderately to densely covered with rusty barbellate or plumose hairs and moderately underlain with minute oblong glands; the nodal line not evident from the copious indument. Leaves of a pair mostly equal in length. Petiole 0.5-3 cm. Leaf blades 9.5-40 × 7-20, 5-7 nerved, elliptic-ovate to ovate, base cordate, apex acuminate, the margin undulate-denticulate, adaxially with simple hairs on the veins when young but becoming glabrous with age, abaxially with a mixture of simple, barbellate and minute glandular hairs. Inflorescence a terminal panicle 6-15 cm long, accessory branches absent, rachis copiously covered with rusty barbellate hairs, bracteoles 0.5-1 mm long, subulate, persistent. Pedicels about 1.5-2.5 mm. Hypanthium campanulate 2-2.25 × 1.25-1.75 mm, sparingly covered with barbellate, plumose and glandular hairs, the torus and inner hypanthium walls glandular puberulent. Flowers 5-merous, calyx not calyptrate nor fused in bud, the undulate apiculate lobes 0.25-0.5 mm long, the exterior calyx teeth tuberculate, 0.25 mm long. Petals 2.5-5 × 1.5-3 mm, ovate-oblong, translucent white, glabrous, not observed at anthesis but petals apparently closing after anthesis, emarginate. Stamens 10, 3.75-4.5 mm long, radially arranged around the style, the filament 1.75-2 mm long with a geniculation near the apex, translucent white, anthers 2-2.25 × 0.4-0.6 mm, linearoblong, somewhat laterally compressed, cream yellow, the pore ca. 0.17 mm, truncate.
Conostegia speciosa is a distinctive species based on its dense hirsute indument of stipitate stellate and simple trichomes. Also, the indument on its inflorescences tends to be bright purple. Its style is short like that of its close relatives C. subcrustulata and C. xalapensis and like the latter taxa lacks a stele inside the style. Other species with short styles which are restricted to section Conostegia have a stele inside the style. Schnell (1996) argued that C. speciosa was the most abundant weed of the genus in Central Panama in habitats that would probably be occupied by C. icosandra, C. poly- andra, C. subcrustulata, or C. xalapensis. For this reason, Schnell (1996) suggested they might be ecological replacements of each other. Of all these taxa, C. speciosa is the one that inhabits the driest areas. Schnell (1996) also reported possible hybrids between C. speciosa and C. xalapensis form the Canal Zone in Panama and from Santa Marta, Colombia, and tested their pollen viability. He found that the purported hybrids in fact had a high percentage of non viable pollen grains. Lastly Schnell (1996)   Description. Shrub to small tree 1-5 m tall with tetragonal stems at first but soon becoming terete that are tomentulose, covered with small stellate hairs and sometimes also more developed somewhat branched or roughened trichomes; the nodal line present but slight and obscured by indument. Leaves of a pair equal to somewhat unequal in length. Petioles 0.5-12 cm. Leaf blades 5-20.7 × 3.1-12 cm, 5-9 plinerved, with the innermost pair of primary veins diverging from the mid vein 0.5-3 cm above the base in mostly opposite to sub opposite fashion, ovate, the base rounded to cordate, the apex acute or short acuminate, the margin serrate and short ciliate, the adaxial surface sparsely hirsute to glabrous,the abaxial surface tomentulose covered with small stellate hairs and sometimes with also more developed somewhat branched or roughened trichomes intermixed. Inflorescence a terminal panicle 6.7-30 cm long branching above the base, accessory branches present, the rachis covered with stellate hairs, bracteoles to 1.5 mm, linear, deciduous. Th e pedicels 0.7-2 mm. Flowers 5(-6) merous, calyptrate. Floral buds 3-6 × 2-3.5 mm, pyriform, the base rounded, the apex acute to apiculate and with teeth at the tip, constricted at the middle, hypanthial and calyptrate portions well diff erentiated, the hypanthium 2-3.5 × 2-3 mm, glabrescent, with stellate trichomes. Petals 4-5 × 2.5-3.25 mm, pink, narrowly obovate, spreading, glabrous, apically acute. Stamens 10 (-12), 4-5 mm, somewhat zygomorphic, the filament 2-2.75 mm, white, geniculate near the apex, anthers 1.8-2.5 × 0.5-1 mm, linear to elliptic, yellow, laterally compressed, the pore 0.2-0.3 mm wide, ventrally inclined. Ovary (4-)5(-6) locular, inferior, apically glabrous and not forming an evident collar around the style. Th e style 4-4.5 mm, straight but abruptly bent just below the stigma, vertical distance of the anther pore to the stigma -1 --0.25 mm, stigma clavate to capitellate, 0.65-1 mm wide. Berry 4-6 × 4-6 mm, at first pink but turning dark purple to black. Seeds 1-1.5 mm long, narrowly wedge shaped, glossy on one side and smooth on the other.
Distribution. Nicaragua, Costa Rica, Panama, Colombia, and a locality in central Ecuador, mostly at low elevations but occasionally to 1500 m in elevation. Fig. 195.
Conostegia subcrustulata is a weedy species frequently found in open areas and on roadsides. It is easy to recognize because of its membranaceous, broad leaves with serrate and ciliate margins, and whitish stellate trichomes on twigs and inflorescences. Th e latter are generally pink, as are the hypanthia and calyptras. Th e calyptra in this species tends to have small appendage or calyx teeth at the apex. Th e calyptra is quite thick but lacks sclereids. Th e filaments are evidently geniculate which agrees with its phylogenetic placement in the Geniculatae clade. In the Osa Peninsula, Costa Rica, flowers of this species were found to be quite variable. In particular, the positioning of the stamens with respect to the style varies almost flower to flower. Th e short style slightly bent apically might contribute to this apparent intrafloral interference.
Schnell reports on observation by Dent-Acosta who at La Selva observed a species of Melipona as the most common visitor of C. subcrustulata flowers between 8:30 and 10:30 am. Dent-Acosta also reported on pollination of all flowers but fruit set of a couple of berries a day per inflorescence. Lastly, Dent-Acosta noted that ovaries have about 200 ovules and stigmas received more than 200 grains but only up to seventy seeds were pro- duced in each one. Schnell (1996) suggests this is either a case of a self-incompatibility mechanism or self limitation of seed and fruit development. He further suggests it might be an example of fruit and or seed abortion. I have also observed Melipona costaricensis as a common visitor of C. subcrustulata at Los Charcos, Osa Peninsula, Costa Rica.
Hybrids with C. xalapensis were reported by Schnell (1996) from El Valle, Coclé, Panama. Schnell tested the viability of these potential hybrids and found very low viability suggesting they were indeed hybrids. I have collected a specimen which appears to be a hybrid with C. xalapensis in Sarapiquí, Costa Rica. Th e specimen looks like C. subcrustulata but has an abaxial leaf surface more like C. xalapensis in its heavy indument. Schnell also reports possible hybridization with C. speciosa from two Costa Rican specimens, but the testing of pollen viability resulted in normally filled grains. Schnell (1996) reported the findings of Ellison et al. (1993) and colleagues who studied the germination of seedling ecology. Th ey reported that the seeds handled dry storage and had about a 50% germination rate. Description. Shrubs to small trees with adventicious roots 1.5-2 (reportedly to 5) m tall with quadrisulcate young nodes that become terete with age and that are moderately to copiously covered with pinoid hairs intermixed or replaced with stellate and asperous hairs; the nodal line not evident. Leaves of a pair equal to somewhat unequal in length. Petiole 0.7-3.1 cm. Leaf blades 5.75-16.5 × 2.8-8.9 cm, 3-5-plinerved, diverging from the midvein 0.8-1.4 cm above the blade base in opposite to alternate fashion, elliptic to elliptic-ovate, base rounded to subcordate and generally slightly peltate, apex acuminate, margin subentire to denticulate or crenulate, adaxially glabrous, abaxially stellate pubescent on tertiary and higher order veins. Inflorescences laxly branched dichasia in the axils of the upper leaves or lateral, usually arising on branchlets below the leaves and paired mainly at defoliated nodes, 1.5-4 cm long, branching above the base, accessory branches absent, rachis moderately to copiously covered with pinoid hairs intermixed or replaced with stellate and asperous hairs, bracteoles 0.5-1 mm long, subulate. Pedicels 1-2 mm long. Floral buds 4 × mm. Flowers 4 (-5) merous, calyx not calyptrate but fused in bud, shortly apiculate and rupturing into 4 irregular, broadly rounded hyaline lobes 0.25-0.75 mm long and 0.5-0.75 mm wide at the base, the exterior calyx teeth 0.25-0.5 mm long, linear oblong, hypanthium 2-2.75 × 1.25-2 mm, narrowly campanulate to urceolate, sparsely stellate. Petals 2.5-3.5 × 2.5-3 mm, translucent white, ovate to orbicular, glabrous m, reflexed at anthesis, emarginated and strongly asymmetrical. Stamens 8, 3-3.75 mm long, radially arranged around the style, the filament 1.5-2 mm long with a geniculation near the apex, translucent white, anthers 1.25-1.75 × 0.5-0.75 mm, linear-oblong, laterally compressed, yellow, the pore 0.1-0.15 mm, truncate to somewhat ventrally inclined. Ovary 4-locular, inferior, the apex elevated into glabrous collar around the style base. Style 5-5.75 mm long, straight to slightly curving, vertical distance between the anther apex and the stigma 1.9-2.25 mm, horizontal distance absent, stigma truncate to capitellate, ca. 0.3 mm wide. Berry ca. 4-5 × 4-5 mm. Seeds 0.4-0.6 mm, triangular, regulate or muriculate. Distribution (Fig. 197). Endemic to Costa Rica where it is known from the Caribbean slope of the Guanacaste and Talamanca mountain ranges at (100-)900-1200 m.
Conostegia subpeltata is a very distinctive species because of its slightly peltate leaves, well defined stellate trichomes on the leaves and hypanthia, axillary inflorescences, and 4-merous flowers with a calyx that is fused in bud and ruptures irregularly into unequal lobes. Th e flowers of this species resemble the common morphology on section Geniculatae with bright yellow anthers and exserted styles. Th is species is known from very few specimens but surprisingly these few specimens come from two mountain ranges, one in Volcan Miravalles in Guanacaste and the other from the Talamanca mountain range (Tapanti National Park and Hitoy Cerere Biological Reserve). C. subpeltata forms a well supported species pair with C. colliculosa in the molecular phylogeny. Th is species pair is in turn sister to C. calocoma. C. subpeltata can be easily distinguished from both of these close relatives by the subpeltate leaves and axillary inflorescences. It shares the 4-merous flowers with C. calocoma.  Calif. Acad. Sci, Series 4, 43 (17) Description. Shrub 1-2.5 m tall with terete stems that are moderately to densely covered with smooth e spreading hairs mostly 1-3 mm long; the nodal line inconspicuous and not evident from the indumenta. Leaves of a pair equal or usually unequal in size. Petiole 0.1-2.2 cm. Large leaf blade 5-14.5 × 2.5-6 cm, 5-7 plinerved with the innermost pair of primary veins diverging 0.6-2.2 cm above the blade base and in alternate fashion, elliptic, acute to obtuse, rounded or oblique, acuminate, the margin conspicuously serrate denticulate, adaxially moderately to sparely strigose to hirtellous, abaxially hirsute. Inflorescence a pseudolateral, modified and usually deflexed dichasium with flowers borne in pedunculate terminal glomerules 1.2-3 cm, branched above the base, accessory branches absent, the rachis densely hirsute, lanceolate to naviculiform, 1.5-3 × 0.5-1.5 mm. Flower buds 1.75-2.5 × 1.25-1.75 mm, more or less ovoid, copiously covered with smooth spreading trichomes. Flowers 5 merous, not calyptrate, the hypanthium 1.75-2.2 × 1.5 mm wide, calyx lobes broadly deltoid, 1 × 1.5 mm long, calyx teeth setiform, 1-2 mm long. Petals 4-4.5 × 1.5-2 mm, translucent white, elliptic-lanceolate, spreading to somewhat reflexed, apex acute, glabrous. Stamens 10, radially arranged around the style, the filaments 2-2.5 mm, with a geniculation near the apex, translucent white, anthers 1.25-1.75 × 0.5-0.75 mm, linear-oblong, yellow, somewhat laterally compressed, the pore ca. 0.15 mm wide, truncate. Ovary 5-locular, 2/3 inferior, apically mostly glabrous but with some scattered sessile glands and elevated into a low lobulate collar around the style. Style 5-6.5 mm, gently curving, vertical distance from the anther apex to the stigma 2-2.5 mm, horizontal distance ca. 0.5 mm, stigma punctiform and conspicuously papillose, 0.3-0.4 mm wide. Berry 4-6 × 4-5.5 mm, dark purple; seeds 0.3-0.5 mm, cuneate to triangular, smooth with verruculose angles. Distribution (Fig. 199). Endemic to Panama where it grows mostly in cloud forests, 250-1250 m in elevation.
Conostegia trichosantha is a very distinctive species because of its dense indument of smooth hairs, small leaves with very plinerved and usually asymmetric venation, pseudolateral, short inflorescences, 5-merous flowers with narrow and acute white petals and very exserted styles. It is perhaps not surprising that in the molecular phylogeny, C. trichosantha appears as closely related to other Panamanian endemics such as C. jefensis and C. peltata as well as with the more widespread C. consimilis. With these taxa it shares narrow petals and with most of them angulate seeds with verruculose angles. Th is species accumulates water in the hypanthium indument.