Monograph
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Monograph
A Monograph of Conostegia (Melastomataceae, Miconieae)
expand article infoRicardo Kriebel
‡ University of Wisconsin-Madison, Madison, United States of America
Open Access

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, C. 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.

Keywords

Conostegia, Melastomataceae, monograph, Neotropics

Introduction

Conostegia D. Don, a genus in the tribe Miconieae (Melastomataceae), is most famous for the calyptrate calyx of the flowers of its species. The 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 identified a core Conostegia clade (Kriebel et al. 2015). This Conostegia clade identified 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 different anatomical characteristics in each origin (Kriebel et al. 2015). The 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. This 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 (Ionta et al. 2012; Ionta and Judd 2012; Judd and Ionta 2013; Judd and Majure 2013; Majure and Judd 2013a, 2013b). These publications recommended and have started to divide the expanded Miconia into sections. Thus, 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. This problem also applies with the large number of genera that would have to be created to accommodate all species within Miconieae. The case of Conostegia is different 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 classifications in their organization. These 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. This 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. The last phylogeny of the Miconieae included two genes for 449 species (Goldenberg et al. 2008), 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. This 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).

The 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

The idea of separating a group of calyptrate species in the Melastomataceae was first suggested by Bonpland (1806-1816), 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. The 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 affinities of Conostegia to Miconia. Schnell (1996) informally designated M. procera as the type of Conostegia.

The 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. The first section Eriostegia DC. contained only one species that had been described as Melastoma mutisii Bonpl. and was proposed as Conostegia mutisii (Bonpl.) DC. The 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). The second section he called Euconostegia and characterized it based on the shape of the calyptra, which was abruptly acuminate and also hispid. The 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 (Axiliflora, Conostegia, Dasystegia, Notostegia, Parvistigma and Tomentostegia). Unfortunately Schnell’s work was never published. The genus had always been thought of as a morphologically distinct group (Bonpland 1806-1816; Don 1823; Almeda 1990; 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). The latter study followed very closely the work of Schnell (1996).

Phylogeny and infrageneric classification

The 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 (Kriebel et al. 2015). 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. These 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 identified 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 (flowers not herkogamous). This 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 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. These two sections also share the frequent pleiostemonous condition and the lack of an evident filament geniculation. The two groups differ 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. The calyptra of these species differs 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.

Figure 1.

Infrageneric classification in Conostegia next to the recently generated molecular phylogenetic hypothesis of Conostegia. Colors show the classification proposed by Schnell (1996). To the right of the phylogeny is the sectional classification proposed here. The phylogeny is a slightly modified version of the majority rule consensus tree derived from a Bayesian analysis of DNA sequences from four chloroplast regions and two nuclear ribosomal spacers (Kriebel et al. 2015) pruned to one accession per species.

Chromosome numbers

Chromosome counts have been reported for the following 13 species of Conostegia: C. arborea, C. colliculosa, C. consimilis, C. galdamesiae, C. hammelii, C. icosandra, C. montana, C. oerstediana, C. setosa, C. subcrustulata, C. superba, C. schlimii, and C. xalapensis (Solt and Wurdack 1980, Almeda and Chuang 1992, Schnell 1996, Almeda 2013). In all reported counts the haploid number was n=17, which is believed to be the base number of the tribe Miconieae (Almeda 2013).

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. This 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. The 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. The area with the most endemics is southern Central America (Costa Rica and Panama). The 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. The 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. The 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 Pacific.

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. The 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). This 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 different species (Fig. 3). The 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.

Figure 2.

Examples of galls in Conostegia. A A stem gall of a Lepidopteran in the family Momphidae in Conostegia oerstediana B A caterpillar of the family Momphidae that apparently can form ovary galls in C. macrantha.

Figure 3.

Beetles of the family Curculionidae in Conostegia flowers. A Conostegia oerstediana B Conostegia rhodopetala C Conostegia xalapensis D, F Scanning electron micrographs of two weevils found in Conostegia flowers E, G Two weevils from material in spirit.

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. This study included three species of Conostegia. This short floral life span is expected in flowers of the family because most are buzz pollinated (Renner 1989). Plants that are buzz pollinated tend to be homogamous (offer 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 offer a reward in both male and female phases. Because melastomes are generally not dichogamous, their flowers 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 The 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 flowers 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). The 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 flowers and is usually manifested within Conostegia by its species having exserted styles. This 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). The evolution of the giant stigma appears to correspond to a different 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. This 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. They 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). This 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).

Pollinators and breeding systems in Conostegia.

Species Family Genus Species Compatibility system Source
Conostegia bracteata Halictidae Augochlora Augochlora sp. ? R. Kriebel pers. obs.
Conostegia bracteata Apidae Euglossa Euglossa sp. ? R. Kriebel pers. obs.
Conostegia cinnamomea Apidae Melipona Melipona cf costaricensis ? R. Kriebel pers. obs.
Conostegia consimilis Halictidae Augochlora Augochlora sp. ? R. Kriebel pers. obs.
Conostegia grayumii Halictidae Augochlora Augochlora sp. ? R. Kriebel pers. obs.
Conostegia macrantha ? ? ? SIC Renner 1989
Conostegia montana ? ? ? SC Tanner 1980
Conostegia oerstediana Apidae Bombus Bombus volluceloides SC R. Kriebel pers. obs.; Schnell 1996
Conostegia ortizii Halictidae cf Augochlora cf Augochlora sp. ? D. Penneys 1857
Conostegia pittieri Apidae Bombus Bombus volluceloides ? R. Kriebel pers. obs.
Conostegia schlimii Apidae Melipona Melipona cf costaricensis ? R. Kriebel pers. obs.
Conostegia setosa Halictidae Augochlora Augochlora sp. ? R. Kriebel pers. obs.
Conostegia subcrustulata Apidae Melipona Melipona cf costaricensis ? R. Kriebel pers. obs.
Conostegia xalapensis Apidae Melipona Melipona beechei and Melipona fasciata SC J. A Reed pers. obs.
Conostegia sp. Apidae Melipona Melipona panamica ? D. Roubik pers. comm.
Conostegia sp. Halictidae Augochlora Augochlora sp. ? D. Roubik pers. comm.

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 flower type (Nesom 2012). This term derives from the Greek, plesios, meaning near, alluding to the proximity of the stamens and stigma (Nesom 2012). The evolution of plesiogamy has occurred at least twice within Conostegia (Kriebel et al. 2015) 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 1996). Throughout 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). These 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 flowers 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.

Figure 4.

Bee pollinators and pollen robbers in Conostegia. A Female of the genus Euglossa approching an inflorescence of C. bracteata B Euglossa sp. buzzing the flower of C. bracteata C Unidentified female of a species of Halictidae buzzing the flower of C. bracteata D Female of Melipona costaricensis buzzing flower of C. subcrustulata E Female of Bombus volluceloides buzzing the flower of C. oerstediana F, G Unidentified large bees buzzing the flowers of C. oerstediana H A species of Trigona robbing pollen from a flower of C. oerstediana I A species of Trigona robbing pollen from a flower of C. bigibbosa J Unidentified female of a species of Halictidae buzzing the flower of C. ortizae. Photograph by Darin Penneys K, L Female of Melipona costaricensis buzzing flower of C. schlimii. Photographs by Reinaldo Aguilar M Unidentified female of a species of Halictidae buzzing the flower of C. grayumii N, O Unidentified female of a species of Halictidae buzzing the flower of C. cinnamomea. Photos by the author if not specified.

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. The display of the berries in taxa with terminal inflorescences contrasts with that of taxa with axillary inflorescences. These two different types of fruit display result in different 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). Then, 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. This 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. The 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. This 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 identified by experts (mainly Charles Schnell and Frank Almeda) added to improve the distribution maps. The 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). The 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, Hatfield, 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, floral structures were photographed under a Nikon SMZ1500 stereoscope equipped with a Nikon DXM1200F camera connected to a computer and using the software ACT-1. The plates were prepared with GIMP (The 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. They mostly occur in lowland rainforest and cloud forest habitats. The preferred elevational ranges are shown are shown in Figure 7.

Figure 5.

Distribution of Conostegia.

Figure 6.

Species density of Conostegia per ten square kilometer grid cell size. A couple of known localities of Conostegia icosandra from Bahia, Brazil, not shown.

Figure 7.

Boxplots of elevation (in meters above sea level) for species of Conostegia.

Flowering phenology

The 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 (Figs 814). This 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-of-season 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 rainy-season flowering from July through October. Another possible pattern includes C. allenii and C. calocoma both of which only flower from January through July.

Figure 8.

Phenology in species of Conostegia. White slices denote flowering specimens and gray slices fruiting specimens. The size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month.

Figure 9.

Phenology in species of Conostegia. White slices denote flowering specimens and gray slices fruiting specimens. The 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 flowering specimens and gray slices fruiting specimens. The 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 flowering specimens and gray slices fruiting specimens. The size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month.

Figure 12.

Phenology in species of Conostegia. White slices denote flowering specimens and gray slices fruiting specimens. The 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 flowering specimens and gray slices fruiting specimens. The 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 flowering specimens and gray slices fruiting specimens. The size of the slice represents the relative amount of specimens from the total which was blooming or fruiting in that month.

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. The 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. The 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. The trunk of some species such as C. bernoulliana and C. oerstediana, can have flaky bark but in most taxa it is smooth. The 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 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

Thirty 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. The cuticle is generally inconspicuous but sometimes can be relatively thick as in C. rhodopetala (Fig. 15). The mesophyll is dorsiventral (Figs 15, 16), and druses were present in all species, usually near the adaxial or abaxial leaf epidermis. The 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. The other thick-leaved species were found to have a hypodermis one cell layer thick. This 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). The spongy mesophyll was usually not lignified except in some species, such as C. ombrophila and C. schlimii.

The 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. The 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 unlignified 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.

Figure 15.

Leaf anatomy in Conostegia A C. lasiopoda (R. Kriebel 5780) B C. monteleagreana (R. Kriebel s. n.) C C. tenuifolia (R. Kriebel 5773) D C. bigibbosa (R. Kriebel 5771) E C. bracteata (R. Kriebel 5816). F C. brenesii (R. Kriebel 5546) G C. caelestis (R. Kriebel 5617) H C. bernoulliana (R. Kriebel 5772). I C. montana (R. Kriebel 5548) J C. montana (R. Kriebel 5662) K C. oerstediana (R. Kriebel 5627) L C. pittieri (R. Kriebel 5543) M C. rhodopetala (R. Kriebel 5542). N C. rufescens (R. Kriebel 5524) O C. setosa (R. Kriebel 5813) P C. fraterna (R. Kriebel 5774) Q C. hammelii (R. Kriebel 5539) R C. ombrophila (R. Kriebel s.n.) S C. subcrustulata (R. Kriebel 5808). T C. xalapensis (R. Kriebel 5817) U C. calocoma (R. Kriebel s. n.). Scale bar: 100 µm.

Figure 16.

Leaf and petiole anatomy in Conostegia. A C. grayumii (R. Kriebel 5807) B C. consimilis (R. Kriebel 5811) C C. oligocephala (R. Kriebel 5575) D C. schlimii (R. Kriebel 5776) E C. lasiopoda (R. Kriebel 5780) F C. tenuifolia (R. Kriebel 5773) G C. monteleagreana (R. Kriebel s. n.) H C. bigibbosa (R. Kriebel 5771). I C. bracteata (R. Kriebel 5816). J C. bernoulliana (R. Kriebel 5772) K C. rufescens (R. Kriebel 5524). L C. setosa (R. Kriebel 5813) M C. subcrustulata (R. Kriebel 5808). N C. grayumii (R. Kriebel 5807) P C. consimilis (R. Kriebel 5811) O C. schlimii (R. Kriebel 5776). Scale bar: 100 μm (A–D); 1 mm (E–O).

Figure 17.

Boxplots of leaf thickness by section of Conostegia. The outlier in section Conostegia is a population of C. montana from Cerro Jefe, Panama. Units in microns.

Leaf domatia

Three 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. The 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. The general classification of domatia used here follows Jacobs (1966). The first kind of mite domatium is the tuft mite domatium present in C. procera (Fig. 18). This species always has these structures, which are densely covered by stipitate stellate to dendritic trichomes. This type of domatium is also seen in some but not all specimens of C. hirtella. The 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.

Figure 18.

Types of domatia in Conostegia. A, B examples of pocket domatia C example of vesicular pocket domatium D example of tuft domatium A C. ecuadorensis (J. Betancur 3202) B Clidemia hammelii (L. Acosta 133). C C. ombrophila (R. Kriebel 5730) D C. procera (W. Maxon 8949). Scale bar: 1 mm.

Indument

Trichomes in Conostegia are quite diverse and variable (Fig. 19). This 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. Difficulty 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 difficult 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 (Figs 2033). Apparently all taxa have minute glands on the abaxial leaf surface, and their shape is variable and may prove to be taxonomically informative. Their small size makes them difficult 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.

Figure 19.

Examples of trichome types in Conostegia. A C. dentata (J. Cuatrecasas 17668) B C. bracteata (M. Hopkins 22) C C. ortizae (D. Penneys 1857) D C. superba (W. Judd 6521) E C. extinctoria (H. David 1227). F C. brenesii (R. Kriebel 4907) G C. subcrustulata (L. Williams 27545) H C. rufescens (D. Penneys 1792) I C. consimilis (A. Jiménez 2326) J C. osaensis (R. Aguilar 12890).

Figure 20.

Trichomes in Conostegia. A–B C. allenii (G. de Nevers 7207) C–D C. ecuadorensis (J. Betancur 3202) E–F C. foreroi (F. Almeda 10336) G–H C. fraterna (R. Kriebel 5774). Scale bar: 100 µm.

Figure 21.

Trichomes in Conostegia A–B C. hammelii (L. Acosta 133) C–D C. ombrophila (R. Kriebel 5730) E–F C. henripittieri (R. Kriebel 5757) G–H C. subpeltata (R. Kriebel 5347). Scale bar: 100 µm.

Figure 22.

Trichomes in Conostegia. A C. trichosantha (F. Almeda 6491). B C. cinnamomea (T. Croat 6542) C–D C. brenesiana (J. Taylor 17646) E–F C. calocoma (R. Kriebel 5484) G–H C. centrosperma (R. Kriebel 5690). Scale bar: 100 µm.

Figure 23.

Trichomes in Conostegia A–B C. colliculosa (R. Kriebel 5721) C–D C. dissitiflora (R. Kriebel 5378) E–F C. dissitinervia (R. Kriebel 5377) G–H C. friedmaniorum (R. Kriebel 5721). Scale bar: 100 µm.

Figure 24.

Trichomes in Conostegia A–B C. galdamesiae (R. Kriebel 5836) C–D C. grayumii (T. McDowell 199) E–F C. incurva (W. Alverson 2747) G–H C. jefensis (R. Kriebel 5680). Scale bar: 100 µm.

Figure 25.

Trichomes in Conostegia A–B C. consimilis (A. Jimenez 2326) C–D C. oligocephala (R. Kriebel 8575) E–F C. osaensis (R. Aguilar 12890) G–H C. papillopetala (R. Kriebel 5718). Scale bar: 100 µm.

Figure 26.

Trichomes in Conostegia A–B C. peltata (R. Kriebel 5658) C–D C. povedae (F. Oviedo 1908) E–F C. schlimii (T. G. Yuncker 8780) G–H C. shattuckii (R. Kriebel 5688). Scale bar: 100 µm.

Figure 27.

Trichomes in Conostegia A–B C. bracteata (M. Hopkins 22) C–D C. brenesii (R. Kriebel 4907) E–F C. caelestis (A. Molina 8352) G–H C. cuatrecasii (R. Kriebel 5681). Scale bar: 100 µm.

Figure 28.

Trichomes in Conostegia A–B C. extinctoria (H. David 1227) C–D C. icosandra (R. Kriebel 5580) E–F C. jaliscana (F. Almeda 2450) G–H C. ortizae (D. Penneys 1857). Scale bar: 100 µm.

Figure 29.

Trichomes in Conostegia. A–B C. lasiopoda (J. Cuatrecasas 16953) C–D C. superba (W. Judd 6521) E–F C. macrantha (R. Kriebel 5406) G–H C. micrantha (R. Espinoza 1739). Scale bar: 100 µm.

Figure 30.

Trichomes in Conostegia A–B C. montana (R. Kriebel 5662) C–D C. monteleagreana (R. Kriebel 5747) E–F C. oerstediana (R. Kriebel 5408) G–H C. pittieri (R. Kriebel 5400). Scale bar: 100 µm.

Figure 31.

Trichomes in Conostegia A–B C. plumosa (D.W. Stevens 25247) C–D C. speciosa (S. B. Robbins 6173) E–F C. subcrustulata (L. O. Williams 27545) G–H C. xalapensis (D. Penneys 1758). Scale bar: 100 µm.

Figure 32.

Trichomes in Conostegia A–B C. polyandra (P. Acevedo 6905) C–D C. procera (W. Maxon 8949) E–F C. rhodopetala (R. Kriebel 5462) G C. rufescens (D. Penneys 1792) H C. rufescens (R. Kriebel 5687). Scale bar: 100 µm.

Figure 33.

Trichomes in Conostegia. A C. cuatrecasii (J. Cuatrecasas 17668). B C. setosa (M. Tirado 529) C–D C. superba (R. Kriebel 5582) E–F C. tenuifolia (R. Moran 7950) G–H C. volcanalis (R. Kriebel 5565). Scale bar: 100 µm.

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 inflorescence can be deflexed. 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 flowers 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 inflorescence, 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 difficult 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). These pedicels can further elongate as the flowers develop into fruits. Pedicels are absent (flowers 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. This 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 different 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 different types of indument on the hypanthium. The apex of the buds can also be helpful for identifying some species. An apiculate calyptra for example, is present in several distinct species from different 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).

Figure 34.

Longitudinal sections of floral buds in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. monteleagreana (R. Kriebel 5747) C C. ortizae (D. Penneys 1857) D C. tenuifolia (R. Kriebel 5773) E C. bernoulliana (R. Kriebel 5540). F C. bigibbosa (R. Kriebel 5522) G C. bracteata (R. Kriebel 5806) H C. brenesii (R. Kriebel 5631) I C. montana (R. Kriebel 5593) J C. oerstediana (R. Kriebel 5627) K C. pittieri (R. Kriebel 5400) L C. rhodopetala (R. Kriebel 5542). M C. setosa (R. Kriebel 5731) N C. volcanalis (R. Kriebel 5565) O C. peltata (R. Kriebel 5658) P C. consimilis (R. Kriebel 5726) Q C. subcrustulata (R. Kriebel s.n.) R C. xalapensis (R. Kriebel 5619). Scale bar: 1 mm.

Phylogenetic and anatomical analyses have revealed that the calyptra evolved at least three times within Conostegia (Kriebel et al. 2015). This 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. The adaptive value of the calyptra remains unknown.

The 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). The sclereids for the most part form a layer or two throughout the hypanthium and calyptra (Fig. 36). The main difference 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. The 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.

Figure 35.

Calyptra and calyx teeth anatomy in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. monteleagreana (R. Kriebel 5747) C C. tenuifolia (D. Santamaria 8863) D C. brenesii (R. Kriebel 5631) E C. icosandra (R. Kriebel 5578) F C. montana (R. Kriebel 5544) G C. oerstediana (R. Kriebel 5627) H C. rufescens (R. Kriebel 5635) I C. superba (R. Kriebel 5582) J C. cinnamomea (R. Kriebel 5330) K C. speciosa (R. Kriebel 5677) L C. subcrustulata (R. Kriebel 5653) M C. xalapensis (R. Kriebel 5629) N C. friedmaniorum (R. Kriebel 5641) O C. schlimii (R. Kriebel 5614). Scale bars: 100 µm.

Figure 36.

Ovary and hypanthium anatomy in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. monteleagreana (R. Kriebel 5747) C C. tenuifolia (D. Santamaria 8863) D C. brenesii (R. Kriebel 5631) E C. bracteata (R. Kriebel 5816) F C. cuatrecasii (R. Kriebel 5681) G C. icosandra (R. Kriebel 5578) H C. montana (R. Kriebel 5544) I C. oertsediana (R. Kriebel 5627) J C. setosa (R. Kriebel 5731) K C. superba (R. Kriebel 5582) L C. subcrustulata (R. Kriebel 5653). Scale bar: 500 µm.

Figure 37.

Examples of calyptra dehiscence modes in Conostegia. A The cleanly circumscissle calyptra of C. setosa B The calyptra of C. tenuifolia ruptured at one side.

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 floral 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 differences between the ovaries in section Geniculatae and the other two sections Australis and Conostegia, see below under the Gynoecium section.

Figure 38.

Ovary and hypanthium anatomy in Conostegia. A C. centrosperma (R. Kriebel 5690) B–D C. cinnamomea (R. Kriebel 5330) D under polarized light E C. dissitinervia (R. Kriebel 5377) F C. fraterna (R. Kriebel 5774) G–H C. grayumii (R. Kriebel 5807) H under polarized light I C. hammelii (R. Kriebel 5317) J C. ombrophila (R. Kriebel 3120) K C. schlimii (R. Kriebel 5614) L C. speciosa (R. Kriebel 5677) M C. trichosantha (R. Kriebel 5693). Scale bar: 500 µm except C and D: 100 µm.

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). This 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. The calyx teeth in calyptrate taxa are difficult 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 difficult. 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.

Figure 39.

Examples of irregularly rupturing calyces in Conostegia. A Flower bud from spirit collection of Conostegia subpeltata (R. Kriebel 3643) B Scanning electron micrograph of flower bud of C. papillopetala (R. Kriebel 5718). Scale bar: 1 mm.

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. There 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. This 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. The 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. The concept of Conostegia included in the present treatment includes many diplostemonous species, particularly those previously described as Clidemia and Miconia. These 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).

Figure 40.

Flowers in Conostegia section Australis (A–G), section Conostegia (H–T), and section Geniculatae (U–X). A C. centronioides (X. Cornejo 8160) B C. ortizae (D. Penneys 1857) C C. lasiopoda (R. Kriebel 5651) D C. monteleagreana (R. Kriebel 5354). E C. monteleagreana (P. Pedraza 1923) F C. polyandra (F. Almeda 10481) G C. tenuifolia (R. Kriebel 5773). H C. bracteata (R. Kriebel 5816). I C. brenesii (R. Kriebel 5631) J C. cuatrecasii (R. Kriebel 5673) K C. montana (R. Kriebel 5446) L C. rhodopetala (R. Kriebel 5542) M C. rufescens (R. Kriebel 5314) N C. setosa (R. Kriebel 5813) O C. superba (R. Kriebel 5582) P C. bigibbosa (R. Kriebel 5522) Q C. icosandra (R. Kriebel 5580) R C. macrantha (R. Kriebel 5406) S C. oerstediana (R. Kriebel 5408) T C. pittieri (R. Kriebel 5543) U C. cinnamomea (R. Kriebel 5330) V C. speciosa (R. Kriebel 5489) W C. subcrustulata (R. Kriebel 5333). C. xalapensis (R. Kriebel 5555). Flowers not to scale.

Figure 41.

Flowers in Conostegia section Geniculatae. A Conostegia xalapensis (R. Kriebel 5619) B C. osaensis (R. Aguilar 10200) C C. allenii (R. Aguilar 13243) D C. foreroi (F. Almeda 10336) E C. fraterna (R. Kriebel 5774) F C. hammelii (R. Kriebel 5737) G C. ombrophila (R. Kriebel 3120) H C. subpeltata (R. Kriebel 5347) I C. trichosantha (R. Kriebel 5693) J C. centrosperma (R. Kriebel 5690) K C. dissitiflora (R. Kriebel 5070) L C. dissitinervia (R. Kriebel 5046) M C. friedmaniorum (R. Kriebel 5641) N C. galdamesiae (R. Kriebel 5736) O C. grayumii (R. Kriebel 5807) P C. consimilis (R. Kriebel 5466) Q C. papillopetala (R. Kriebel 5718) R C. peltata (R. Kriebel 5658). S C. povedae (F. Oviedo 1215). T C. schlimii (R. Kriebel 5095). Flowers not to scale.

Corolla

The 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. The 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.

Figure 42.

Flowers of Conostegia from specimens collected in spirit with a longitudinal section at their side. A C. lasiopoda (R. Kriebel 5651) B C. monteleagreana (R. Kriebel 5747) C C. tenuifolia (R. Kriebel 5773) D C. bernoulliana (R. Kriebel 5540) E C. bracteata (R. Kriebel 5816) F C. brenesii (R. Kriebel 5631). G C. cuatrecasii (R. Kriebel 5673) H C. icosandra (R. Kriebel 5580) I C. montana (R. Kriebel 5751) J C. montana (R. Kriebel 5593) K C. oerstediana (R. Kriebel s.n.) L C. pittieri (R. Kriebel 5400) M C. rufescens (R. Kriebel 5314) N C. setosa (R. Kriebel 5731) O C. superba (R. Kriebel 5582). Scale bar: 1 mm.

Figure 43.

Flowers of Conostegia from specimens collected in spirit with a longitudinal section at their side. A C. brenesiana (R. Kriebel 3665) B C. centrosperma (R. Kriebel 5690) C C. cinnamomea (R. Kriebel 5330) D C. consimilis (R. Kriebel 5726) E C. friedmaniorum (R. Kriebel 5641) F C. galdamesiae (R. Kriebel 5736) G C. hammelii (R. Kriebel 5737) H C. papillopetala (R. Kriebel 5718) I C. peltata (R. Kriebel 5658) J C. schlimii (R. Kriebel 5614) K C. shattuckii (R. Kriebel 5681) L C. speciosa (R. Kriebel 5677) M C. subcrustulata (R. Kriebel s. n.) N C. trichosantha (R. Kriebel 5693) O C. xalapensis (R. Kriebel 5619). Scale bar: 1 mm.

Figure 44.

Petals of Conostegia to scale. a C. lasiopoda (R. Kriebel 5651) b C. monteleagreana (R. Kriebel 5747) c C. ortizae (D. Penneys 1857) d C. tenuifolia (R. Kriebel 5773) e C. bernoulliana (R. Kriebel 5540) f C. bigibbosa (R. Kriebel 5522) g C. bracteata (R. Kriebel 5816) h C. brenesii (R. Kriebel 5631) i C. cuatrecasii (R. Kriebel 5673) j C. fragrantissima (R. Kriebel 3174) k C. icosandra (R. Kriebel 5580) l C. macrantha (R. Kriebel 5406) m C. montana (R. Kriebel 5751) n C. montana (R. Kriebel 5593) o C. oerstediana (R. Kriebel 5627) p C. pittieri (R. Kriebel 5400) q C. rhodopetala (R. Kriebel 5542) r C. rufescens (R. Kriebel 5627) s C. setosa (R. Kriebel 5731) t C. superba (R. Kriebel 5582) u C. volcanalis (R. Kriebel 5565) v C. brenesiana (R. Kriebel 3665) w C. centrosperma (R. Kriebel 5690) x C. cinnamomea (R. Kriebel 5330) y C. consimilis (R. Kriebel 5726) z C. dissitinervia (R. Kriebel 5317) A C. fraterna (R. Kriebel 5774) B C. friedmaniorum (R. Kriebel 5641) C C. galdamesiae (R. Kriebel 5736) D C. grayumii (R. Kriebel 5807) E C. hammelii (R. Kriebel 5737) F C. ombrophila (R. Kriebel 3120). G C. papillopetala (R. Kriebel 5718) H C. peltata (R. Kriebel 5658) I C. povedae (F. Oviedo 231) J C. schlimii (R. Kriebel 5614) K C. shattuckii (R. Kriebel 5681) L C. speciosa (R. Kriebel 5677) M C. subcrustulata (R. Kriebel s.n.) N C. subpeltata (R. Kriebel 3643) O C. trichosantha (R. Kriebel 5693) P C. xalapensis (R. Kriebel 5619).

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 different 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 difference 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).

Figure 45.

Longitudinal sections of petals in Conostegia. A Representative section of a species with translucent petals, Conostegia friedmaniorum B Representative section of a species with white petals, C. pittieri. Scale bar: 100 µm.

Figure 46.

Scanning electron micrographs of Conostegia petal surfaces. A C. monteleagreana (R. Kriebel 5343) B C. brenesii (R. Kriebel 5631) C C. bernoulliana (R. Kriebel 5540) D C. montana (R. Kriebel 5496) E C. oerstediana (R. Kriebel 5338) F C. pittieri (R. Kriebel 5400) G C. rhodopetala (R. Kriebel 5542) H C. rufescens (R. Kriebel 5314) I C. hammelii (R. Kriebel 5317) J C. subpeltata (R. Kriebel 5347) K C. cinnamomea (R. Kriebel 5330) L C. dissitinervia (R. Kriebel 5377) M C. friedmaniorum (R. Kriebel 5641) N C. consimilis (R. Kriebel 5323) O C. schlimii (R. Kriebel 515). Scale bar: 100 µm.

Androecium

The androecium in Conostegia consists of 8–52 isomorphic stamens. The 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. These 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). There 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.

The 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. This 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.

Figure 47.

Example of stamens of species in Conostegia to scale. a C. lasiopoda (R. Kriebel 5651) b C. monteleagreana (R. Kriebel 5747) c C. ortizae (D. Penneys 1857) d C. tenuifolia (R. Kriebel 5773) e C. bernoulliana (R. Kriebel 5540) f C. bigibbosa (R. Kriebel 5522) g C. bracteata (R. Kriebel 5816) h C. brenesii (R. Kriebel 5631) i C. cuatrecasii (R. Kriebel 5673) j C. fragrantissima (R. Kriebel 3174) k C. icosandra (R. Kriebel 5580) l C. montana (R. Kriebel 5544) m C. macrantha (R. Kriebel 5406) n C. oerstediana (R. Kriebel 5338) o C. pittieri (R. Kriebel 5400) p C. rhodopetala (R. Kriebel 5542) q C. rufescens (R. Kriebel 5627). r C. setosa (R. Kriebel 5731) s C superba (R. Kriebel 5582). t C. volcanalis (R. Kriebel 5565) u C. brenesiana (R. Kriebel 3665). v C. centrosperma (R. Kriebel 5690) w C. cinnamomea (R. Kriebel 5330) x C. consimilis (R. Kriebel 5726) y C. dissitinervia (R. Kriebel 5317) z C. fraterna (R. Kriebel 5774) A C. friedmaniorum (R. Kriebel 5641) B C. galdamesiae (R. Kriebel 5736) C C. grayumii (R. Kriebel 5807) D C. hammelii (R. Kriebel 5737) E C. ombrophila (R. Kriebel 3120). F C. papillopetala (R. Kriebel 5718) G C. peltata (R. Kriebel 5658) H C. povedae (F. Oviedo 231) I C. schlimii (R. Kriebel 5614) J C. shattuckii (R. Kriebel 5681) K C. speciosa (R. Kriebel 5677) L C. subcrustulata (R. Kriebel s.n.) M C. subpeltata (R. Kriebel 3643) N C. trichosantha (R. Kriebel 5693) O C. xalapensis (R. Kriebel 5619).

The 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 (Figs 4749). 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. This 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 filament geniculations present them towards the apex of the filament, in one species, C. brenesiana, the geniculation is in the middle of the filament.

Figure 48.

Scanning electron micrographs of the side view of the stamens of Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. lasiopoda close up of anther filament junction (R. Kriebel 5651) C C. tenuifolia (D. Santamaría 8863) D C. monteleagreana (R. Kriebel 5343) E C. ortizae (D. Penneys 1857) F C. brenesii (R. Kriebel 5631) G C. icosandra (R. Kriebel 5540) H C. macrantha (R. Kriebel 5406) I C. montana (R. Kriebel 5496) J C. montana close up of anther filament junction (R. Kriebel 5496) K C. oerstediana (R. Kriebel 5338) L C. pittieri (R. Kriebel 5400) M C. rhodopetala (R. Kriebel 5542) N C. rufescens (R. Kriebel 5314) O C. setosa (R. Kriebel s.n.) P C. hammelii (R. Kriebel 5317) Q C. subpeltata (R. Kriebel 5347) R C. trichosantha (R. Kriebel 5693) S C. cinnamomea (R. Kriebel 5330) T Close up of anther filament junction in C. cinnamomea (R. Kriebel 5330). Scale bar: 100 µm.

Figure 49.

Scanning Electron Micrographs of the side view of the stamens of Conostegia. A C. speciosa (R. Kriebel 5489) B C. subcrustulata (R. Kriebel s.n.) C C. xalapensis (R. Kriebel 5555) D C. brenesiana (R. Kriebel 3665) E C. centrosperma (R. Kriebel 5690) F C. dissitinervia (R. Kriebel 5377) G C. friedmaniorum (R. Kriebel 5641) H C. consimilis (R. Kriebel 5323) I C. peltata (R. Kriebel 5658) J C. schlimii (R. Kriebel 515). Scale bar: 100 µm.

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. The 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). This character was recently documented in some species of Tococa (Michelangeli 2005) and Pachyanthus (Bécquer 2008). The 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.

Figure 50.

Scanning electron micrographs of the ventral and dorsal view of the stamens in Conostegia. A C. ortizae (D. Penneys 1857) B C. lasiopoda (R. Kriebel 5651) C C. monteleagreana (R. Kriebel 5343) D C. tenuifolia (D. Santamaria 8863) E C. brenesii (R. Kriebel 5631) F C. bernoulliana (R. Kriebel 5540) G C. montana (R. Kriebel 5496) H C. oerstediana (R. Kriebel 5338) I C. pittieri (R. Kriebel 5400) J C. rhodopetala (R. Kriebel 5542) K C. rufescens (R. Kriebel 5314) L C. hammelii (R. Kriebel 5317) M C. subpeltata (R. Kriebel 5347) N C. trichosantha (R. Kriebel 5693) O C. cinnamomea (R. Kriebel 5330) P C. subcrustulata (R. Kriebel s.n.). Scale bar: 100 µm.

Anther shape in Conostegia varies a lot even when considering only the species treated traditionally in the genus. This 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. These 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). They are definitely not all ovoid. Also, they can be recurved or arcuate making them diffciult 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. The shape of the anther apices and anther pores is quite variable in Conostegia and merits further research. This 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).

Figure 51.

Scanning Electron Micrographs of the ventral and dorsal view of the stamens in Conostegia. A C. xalapensis (R. Kriebel 5555) B C. brenesiana (R. Kriebel 3665) C C. dissitinervia (R. Kriebel 5377) D C. friedmaniorum (R. Kriebel 5641) E C. consimilis (R. Kriebel 5323) F C. papillopetala (R. Kriebel 5718) G C. peltata (R. Kriebel 5658) H C. schlimii (R. Kriebel 515). Scale bar: 100 µm.

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).

Figure 52.

Anther anatomy in Conostegia. Each image is a transversal cut of an anther with a photograph under polarized light next to it. A–H are examples of anthers with sporangia positioned side by side. K–P are examples of anthers with sporangia superposed A–B C. lasiopoda (R. Kriebel 5651) C–D C. tenuifolia (D. Santamaría 8863) E–F C. oerstediana (R. Kriebel 5627) G–H C superba (R. Kriebel 5582) I–J C. subcrustulata (R. Kriebel 5653) K–L C. fraterna (R. Kriebel 5774) M–N C. friedmaniorum (R. Kriebel 5641) O–P C. ombrophila (R. Kriebel 3120). Scale bar: 100 µm.

Pollen of species in the Melastomataceae has been rarely documented in taxonomic treatments. This 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. The 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).

Figure 53.

Example of anther pores and pollen in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. ortizae (D. Penneys 1857) C C. tenuifolia (D. Santamaría 8863) D C. bernoulliana (R. Kriebel 5540) E C. brenesii (R. Kriebel 5631) F C. oerstediana (R. Kriebel 5338) G C. rhodopetala (R. Kriebel 5542) H C. rufescens (R. Kriebel 5314) I C. brenesiana (R. Kriebel 3665) J C. cinnamomea (R. Kriebel 5330) K C. consimilis (R. Kriebel 5323) L C. friedmaniorum (R. Kriebel 5641) M C. schlimii (R. Kriebel 515) N C. cinnamomea (R. Kriebel 5330) O C. hammelii (R. Kriebel 5317) P C. schlimii (R. Kriebel 515) Q C. subpeltata (R. Kriebel 5347) R C. macrantha (R. Kriebel 5406) S C. montana (R. Kriebel 5496) T C. oerstediana (R. Kriebel 5338). Scale bar: 100 µm (A–M); 10 µm (N–T).

Gynoecium

The gynoecium can have from 4 to 25 carpels (Schnell 1996). Placentation is axillary as in most Melastomataceae (Renner 1993). The 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.

Figure 54.

Example of placenta shapes in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. macrantha (R. Kriebel 5406) C C. schlimii (R. Kriebel 5614). Scale bar: 1 mm.

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). These 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. The 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. The 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 flowering plants display the natural posture of the flowers.

Figure 55.

Example of styles of species in Conostegia to scale. a C. lasiopoda (R. Kriebel 5651) b C. monteleagreana (R. Kriebel 5747) c C. ortizae (D. Penneys 1857) d C. tenuifolia (R. Kriebel 5773) e C. balbisiana (G. Proctor 10285) f C. bernoulliana (R. Kriebel 5540) g C. bigibbosa (R. Kriebel 5522) h C. bracteata (R. Kriebel 5816) i C. brenesii (R. Kriebel 5631) j C. cuatrecasii (R. Kriebel 5673) k C. fragrantissima (R. Kriebel 3174) l C. icosandra (R. Kriebel 5580) m C. macrantha (R. Kriebel 5406) n C. montana (R. Kriebel 5544) o C. oerstediana (R. Kriebel 5338) p C. pittieri (R. Kriebel 5400) q C. rhodopetala (R. Kriebel 5542) r C. rufescens (R. Kriebel 5627) s C. setosa (R. Kriebel 5731) t C. superba (R. Kriebel 5582) u C. volcanalis (R. Kriebel 5565) v C. brenesiana (R. Kriebel 3665) w C. centrosperma (R. Kriebel 5690) x C. cinnamomea (R. Kriebel 5330) y C. consimilis (R. Kriebel 5726) z C. dissitinervia (R. Kriebel 5317) A C. fraterna (R. Kriebel 5774) B C. galdamesiae (R. Kriebel 5736) C C. hammelii (R. Kriebel 5737) D C. ombrophila (R. Kriebel 3120) E C. papillopetala (R. Kriebel 5718) F C. peltata (R. Kriebel 5658) G C. povedae (F. Oviedo 231) H C. schlimii (R. Kriebel 5614) I C. shattuckii (R. Kriebel 5681) J C. speciosa (R. Kriebel 5677) K C. subcrustulata (R. Kriebel s.n.) L C. subpeltata (R. Kriebel 3643) M C. trichosantha (R. Kriebel 5693) N C. xalapensis (R. Kriebel 5619).

Figure 56.

Scanning electron micrographs of stigmas in Conostegia. A C. hammelii (R. Kriebel 5317) B C. subpeltata (R. Kriebel 5347) C C. trichosantha (R. Kriebel 5693) D C. cinnamomea (R. Kriebel 5330) E C. montana (R. Kriebel 5496) F C. oerstediana (R. Kriebel 5338) G Close up of a stigma lobe of C. oerstediana (R. Kriebel 5338) H C. pittieri (R. Kriebel 5400) I C. rufescens (R. Kriebel 5314) J C. speciosa (R. Kriebel 5489) K C. tenuifolia (D. Santamaría 8863) L C. xalapensis (R. Kriebel 5555) M C. brenesiana (R. Kriebel 3665) N C. dissitinervia (R. Kriebel 5377) O C. consimilis (R. Kriebel 5323) P C. schlimii (R. Kriebel 515). Scale: 100 µm.

There are two main clades where the style is not exserted beyond the stamens (not herkogamous). The 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. The 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 Conostegia and convergently Conostegia ortizae in section Australis present this phenomenon. This 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. This character is helpful to confirm 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. These results also coincide with Schnell’s (1996) observations of a lack of a cylinder in C. subcrustulata. The stele found in the style of C. ortizae, in general, differs from those in species of section Conostegia because it has fewer vascular traces.

Figure 57.

Two examples of longitudinal sections of flowers of Conostegia. Note the presence of a stele within the style in A Conostegia superba (R. Kriebel 5582) and the lack of a stele in B C. tenuifolia (D. Santamaría 8863). Scale bar: 1 mm.

Figure 58.

Style transversal anatomy in Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. tenuifolia (D. Santamaría 8863) C C. brenesii (R. Kriebel 5631) D C. pittieri (R. Kriebel 5543) E C. setosa (R. Kriebel 5731) F C. superba (R. Kriebel 5582) G C. friedmaniorum (R. Kriebel 5641) H C. hammelii (R. Kriebel 5317) I C. schlimii (R. Kriebel 515) J C. subcrustulata (R. Kriebel 5653) K C. trichosantha (R. Kriebel 5693) L C. xalapensis (R. Kriebel 5629). Scale bar: 100 µm.

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.

Figure 59.

Berries of some species of Conostegia. A C. lasiopoda (R. Kriebel 5651) B C. monteleagreana (R. Kriebel 5354) C C. polyandra (X. Cornejo 8126) D C. caelestis (R. Kriebel 5588) E C. cuatrecasii (R. Kriebel 5673) F C. icosandra (R. Kriebel 5580) G C. montana (R. Kriebel 5446) H C. oerstediana (R. Kriebel 5338) I C. rufescens (E. Saliceti s.n.) Photograph by E. Saliceti. J C. superba (R. Aguilar 12103) K C. subcrustulata (R. Kriebel 5333) L C. xalapensis (R. Kriebel s. n.) M C. ombrophila (R. Kriebel 5396) N C. pittieri (R. Kriebel 5757) O C. calocoma (R. Kriebel s. n.) P C. dissitiflora (R. Kriebel 5378) Q C. friedmaniorum (R. Kriebel 5497) R C. osaensis (R. Aguilar s. n.) S C. shattuckii (R. Kriebel 5688) T C. schlimii (R. Kriebel 5329). All photographs by the author except where specified.

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. The 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 6062) (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. The 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.

The conclusion of this study is that despite the difficulty 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.

Figure 60.

Seeds in Conostegia. A C. allenii (M. Nepokroeff 722) B C. hammelii (R. Kriebel 1420) C C. ombrophila (R. Kriebel 4887) D C. pittieri (D. Quiroz 800) E C. trichosantha (F. Almeda 6491) F C. cinnamomea (R. Hartman 12257) G C. brenesiana (R. Kriebel 4614) H C. dissitiflora (R. Kriebel 5378) I C. dissitinervia (R. Kriebel 5377) J C. peltata (R. Kriebel 5658) K C. grayumii (M. grayum 1834) L C. consimilis (A. Jiménez 2326) M C. oligocephala (R. Thorne 41087) N C. pendula (D. Santamaría 6672) O C. calocoma (R. Kriebel s.n.). Scale bar: 100 µm.

Figure 61.

Seeds in Conostegia. A C. brenesii (A. Tonduz 12580) B C. caelestis (R. Kriebel 5617) C C. chiriquiensis (E. Alfaro 2365) D C. hirtella (A. Molia 3189) E C. icosandra (F. Ventura 20608) F C. macrantha (C. Schnell 1077) G C. micrantha (R. Kriebel 694) H C. montana (W. Stevens 25434) I C. centronioides (E. Little 6215) J C. extinctoria (H. David 1227) K C. ortizae (D. Penneys 1857) L C. lasiopoda (L. Fournier 303) M C. monteleagreana (F. Almeda 6075) N C. pittieri (C. Todzia 2045) O C. plumosa (P. Gentle 9258) P C. polyandra (P. Azevedo 6905). Scale bar: 100 µm.

Figure 62.

Seeds in Conostegia. A C. procera (N. Britton 3607) B C. rhodopetala (C. Schnell 1081) C C. rufescens (O. Valverde 13) D C. setosa (D. Solano 1448) E C. speciosa (F. Almeda 2863) F C. subcrustulata (R. Kriebel 5333) G C. superba (J. Steyermark 47891) H C. tenuifolia (A. Rodríguez 11693) I C. volcanalis (E. Matuda 2644) J C. xalapensis (A. Soto 1772) K C. osaensis (R. Aguilar 12228) L C. papillopetala (R. Kriebel 5718) M C. ecuadorensis (S. Stern 328) N C. balbisiana (N. Britton 558) O C. bracteata (E. Killip 12158) P C. shattuckii (R. Kriebel 5688). Scale bar: 100 µm.

Taxonomic treatment

Conostegia D. Don

Conostegia D. Don, Mem. Wern. Soc. 4: 316. 1823. Lectotype: C. procera (Sw.) D. Don ex DC. (= Melastoma procera Sw.), designated here.

Synodon Raf., Sylva Tellur. 95. 1838. Lectotype:—Conostegia montana (Sw.) D. Don ex DC.(= Melastoma montana Sw.), designated here.

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 defined 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 defining “distinct” is subjective and varies across organisms. Nonetheless, it is the most common species concept applied to plants (McDade 1995), and an effort was made to recognize only species that appear distinct. This author views species as hypotheses that are subject to testing. Thus 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 defining 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 confidence, 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.

Key to the sections and species of Conostegia

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 C. centrosperma
4' Sepals hyaline and almost not visible during flowering; anther apex acute C. dissitinervia
3' Sepals not evidently fused at anthesis and rupturing into irregular lobes
5 Calyx teeth adnate to the calyx lobes, barely discernable and not projecting beyond them; petals papillose C. fulvostellata
5' Calyx teeth oblong or linear, usually evident and exceeding the calyx lobes; petals glabrous
6 Abaxial indument rusty colored; calyx teeth usually longer than 2 mm and hooked C. incurva
6' Abaxial indument white; calyx teeth up to 1.5 mm long and not hooked C. oligocephala
2' Leaves not discolorous, the abaxial surface visible
7 Inflorescence axillary or appearing so
8 Leaves peltate; indument on abaxial leaf surface and hypanthia stellate; flowers 4-merous C. subpeltata
8' Leaves not peltate; indument on abaxial leaf surface and hypanthia glabrous or stellulate lepidote; flowers 4–5 merous
9 Leaves and hypanthium glabrous or appearing so
10 Leaf base decurrent on the petiole and lacking domatia at the base abaxially; inflorescence of pedunculate glomerules with sessile flowers and ovate bracteoles C. fraterna
10' Leaf base acute and usually with domatia at the base abaxially; inflorescence a laxly branched dichasium with pedicellate flowers and linear to lanceolate bracteoles
11 Leaves with four domatia located abaxially at the base of the leaf C. ecuadorensis
11' Leaves lacking, or usually with two domatia located abaxially at the base of the leaf C. ombrophila
9' Leaves and hypanthium densely pubescent with long simple or dendritic hairs (some populations of C. allenii with glabrous leaves)
12 Petals acuminate at the apex C. trichosantha
12' Petals rounded at the apex
13 Leaves with domatia at the base abaxially C. hammelii
13' Leaves lacking domatia at the base abaxially
14 Leaves sessile or subsessile C. allenii
14' Leaves distinctly petiolate C. foreroi
7' Inflorescence terminal
15 Leaves peltate C. peltata
15' Leaves petiolate from the lamina base or sessile, not peltate
16 Hypanthium densely covered with simple hairs sometimes intermixed with stellate hairs; the leaves with simple hairs on both surfaces or just on the abaxial surface; calyx teeth linear to subulate and 2–4 mm long C. allenii
16' Hypanthium glabrous to densely covered with stellate and or dendritic hairs; the leaves adaxially glabrous and abaxially glabrescent to densely covered with stellate or dendritic trichomes mostly on the veins; calyx teeth tuberculate to triangular or, if linear, usually less, but up to, 2 mm long
17 Inflorescences with sessile flowers in bracteate clusters
18 Leaves sessile and the base decurrent C. povedae
18' Leaves petiolate, the base obtuse to acute or rounded
19 Calyx lobes not evidently fused at anthesis and rupturing, ovate to orbicular and mucronate, the same consistency as the hypanthium C. colliculosa
19' Calyx lobes fused at anthesis, hyaline, and rupturing into irregular lobes C. galdamesiae
17' Inflorescences with pedicellate flowers not in clusters
20 Calyx fused in bud and rupturing into irregular hyaline lobes
21 Flower 4 merous C. calocoma
21' Flower 5 merous
22 Leaves subsessile with rounded to cordate bases; plants overall appearing glabrous but bearing minute inconspicuous trichomes almost invisible to the naked eye; stamens alternately two sizes C. dissitiflora
22' Leaves clearly petiolate or if short petiolate (sometimes in C. papillopetala) with acute to decurrent bases; plants evidently pubsecent, especially on the apical nodes and leaf abaxial surface; all stamens mostly of one size
23 Petals linear oblong; Costa Rican endemics
24 Leaves 5 plinerved; indument of stem apices of asperous headed hairs only; inflorescence branched at the base C. friedmaniorum
24' Leaves 7 plinerved; indument of stem apices lanate at least in part; inflorescence branched above the base C. pendula
23' Petals ovate; Panamanian endemics
25 Flower regularly arranged at the end of the inflorescence branches and with pink papillose petals C. papillopetala
25' Flowers usually clustered at the end of the inflorescence branches and with white glabrous petals C. galdamesiae
20' Calyx not fused in bud and the lobes not rupturing into irregular hyaline lobes, the lobes mostly low and undulate
26 Flowers large, the petals 9 mm long or more; stigma capitate; ovary totally inferior C. schlimii
26' Flowers small, the petals usually no more than 5 mm long; stigma truncate to punctiform; ovary at least partly superior
27 Leaves 5–7 nerved with a broadly rounded to cordate base C. shattuckii
27' Leaves 5–7 plinerved with rounded to mostly acute to attenuate base
28 Petals magenta C. jefensis
28' Petals white
29 Petals ovate; anthers with broad pores C. brenesiana
29' Petals linear-oblong; anthers with small pores
30 Leaves 3 plinerved, the leaf blade 1.7–9.5 × 0.4–1.7 cm C. iteophylla
30' Leaves 5 plinerved, the leaf blade 4.2–28.4 × 1.4–9.2 cm C. consimilis
1' Sepals fused into a calyptra, calyx teeth absent, or if present, restricted to the apex of the calyptra; flowers usually pleiostemonous
31 Calyptra often rupturing at one side or breaking into pieces; style exserted (section Australis and C. cinnamomea)
32 Leaf bases with pouch like formicaria; indument of the stems with long simple hairs C. dentata
32' Leaf bases without pouch like formicaria; indument of stems absent or of stellate, dendritic or asperous hairs
33 Stem apices and leaf abaxial surface glabrous or nearly so
34 Flowers in glomerules at the end of inflorescence branches and subtended by evident bracts and bracteoles; stamens usually less, but up to 15 C. monteleagreana
34' Flowers regularly arranged at the end of inflorescence branches and without evident bracts; stamens 16 or more in number
35 Inflorescence a reflexed and pseudoaxillary panicle; flowers (4-) 5-merous C. cinnamomea
35' Inflorescence a terminal panicle; flowers usually 6-merous
36 Leaves with an entire margin and caudate apex C. tenuifolia
36' Leaves with a serrulate to serrate margin and acuminate apex
37 Leaves linear to linear elliptic, up to 3.5 cm wide, attenuate at the base; stamens 16–21 C. attenuata
37' Leaves ovate, 2–7.3 cm wide, obtuse to rounded at the base; stamens 26–36 C. polyandra
33' Stem apices and leaf abaxial surfaces evidently pubescent at least on the veins
38 Leaf base strongly decurrent on the petiole; leaves sessile or nearly so C. ortizae
38' Leaf base acute to rounded; leaves petiolate
39 Leaves linear to linear elliptic, up to 3.5 cm wide C. attenuata
39' Leaves elliptic to ovate, more than 5 cm wide
40 Petioles setose adaxially; flower buds covered by foliaceous bracteoles C. lasiopoda
40' Petioles glabrescent or variously pubescent, but not setose adaxially; flower buds not covered by foliaceous bracteoles
41 Floral buds up to 4(-5.5) mm long; petals up to 5 mm long; style lacking a collar around the base C. extinctoria
41' Floral buds more than (4.7-)5.5 mm long; petals 7 mm long or more; base of the style enveloped by a collar
42 Indument on stems evidently stipitate stellate, not rusty colored; stamens more than 30 C. lancifolia
42' Indument on stems sometimes stipitate stellate but not evidently so, rusty colored; stamens up to 24
43 Flower buds (7.5-) 10–18 mm long, the apex long apiculate C. apiculata
43' Flower buds 4.7–11 mm long, the apex rounded to short apiculate
44 Flower buds with the surface sparsely pubescent resulting in a visible surface, constricted below the torus; petioles 1–5 cm long C. centronioides
44' Flower buds with the surface densely pubescent resulting in a hidden surface, not to slightly constricted below the torus; petioles 0.4–2.5 cm long C. rubiginosa
31' Calyptra circumscissle and falling as a unit; style shorter or of equal length than the stamens (section Conostegia & section Geniculatae, pro minore parte)
45 Stigma lobed and/or with an evident hole in the middle
46 Flower buds pyriform to ellipsoid, with an attenuate to apiculate apex, longer than wide
47 Stem apices and leaf abaxial surface slightly pubescent to evidently pubescent; plants from Mexico
48 Stems and leaf abaxial surfaces furfuraceous C. jaliscana
48' Stems and leaf abaxial surfaces densely beset with stellate and stalked stellate trichomes C. arborea
47' Stem apices and leaves glabrous or with scattered stellate trichomes in C. chiriquensis; plants from Costa Rica and Panama
49 Flower buds 5–7 mm long; petals to 5 mm long, white with a red band at the base; anthers to 2.3 mm long C. fragrantissima
49' Flower buds 7–14 mm long; petals at least 8 mm long, white to less commonly lavender; anthers at least 2.5 mm long
50 Stigma peltate and with a hole in the middle C. pitteri
50' Stigma barely expanded and lacking a hole in the middle C. chiriquensis
46' Flower buds mostly ovoid to spherical with a rounded, mucronate, or broadly acute apex, as long as wide or slightly longer than wide
51 Stigma capitate, but lacking a large hole in the middle; stigma lobes not conspicuous C. icosandra
51' Stigma with large hole in the middle; stigma lobes conspicuous
52 Leaves with two tubercles at the apex of the petiole abaxially; petals pink C. bigibbosa
52' Leaves lacking tubercles at the apex of the petiole abaxially; petals white
53' Floral buds obovoid, constricted near the torus, acute at the apex, the surface smooth; leaves narrowly elliptic C. bernoulliana
53 Floral buds spherical, not constricted near the torus, abruptly mucronate apically, the surface inconspicuously to usually evidently tuberculate; leaves elliptic to usually ovate
54 Bracteoles replaced by persistent clusters of setae; pedicels up to 3 mm long C. setifera
54' Bracteoles not consisting of setae, usually early deciduous; pedicels usually more than 3 mm
55 Style straight; petals apically retuse; leaves stellate pubsecent abaxially C. macrantha
55' Style curved; petal apex rounded, truncate to emarginate; leaves stellate sometimes stellate pubescent abaxially when young, usually glabrous with age
56 Leaf margin entire to denticulate; floral buds usually conspicuously tuberculate; Nicaragua to Panama C. oerstediana
56' Leaf margin undulate dentate; floral buds inconspicuously tuberculate; Guatemala and Mexico C. volcanalis
45' Stigma not lobed or with a hole in the middle
57 Leaf bases with pouch like formicaria; indument of the stems with long simple hairs C. setosa
57' Leaf bases lacking formicaria; indument of the stems various but not of long simple hairs
58' Flowers subtended by persistent foliaceous bracts
59 Stems and leaves glabrous or with inconspicuous trichomes C. monteleagreana
59' Stems and leaves densely pubescent with simple and somewhat branched trichomes C. bracteata
58' Flowers not subtended by persistent foliaceous bracts
60 Leaves with the abaxial surface covered with white to tan trichomes, making the actual surface hidden or almost so
61 Abaxial leaf surface and hypanthium covered with lepidote trichomes; tree reaching about 25 meters tall; calyptra translucent, less pubescent than the hypanthium; seeds about 1.5 mm long; Endemic to the Osa Peninsula, Costa Rica C. osaensis
61' Abaxial leaf surface and hypanthium covered with stellate trichomes; tree reaching about 15 meters tall; calyptra tan to whitish colored, not translucent, and as pubsecent as the hypanthium; seeds to about 1 mm long
62 Calyptra with linear appendages at the apex about 2.5 mm long C. plumosa
62' Calyptra lacking linear appendages at the apex, at most minute bumps present on the apex C. xalapensis
60' Leaves with the abaxial surface glabrous or pubescent, if pubescent, never with the surface hidden
63 Stem apices and leaf abaxial surfaces pubescent
64 Leaf adaxial surface and floral buds densely hirsute; inflorescence and bud pubescence purple C. speciosa
64' Leaf adaxial surface glabrous and floral buds glabrous or beset with stellate or dendritic trichomes; inflorescence and bud pubescence rusty or whitish
65 Leaves 5–9 plinerved; flowers with small appendages on the calyptra apex; floral buds wider above the torus; petals apically slightly mucronate to acute, especially when seen at anthesis; filaments with an evident geniculation; seeds about 1 mm long C. subcrustulata
65' Leaves 3–5 nerved or plinerved; flowers without small appendages on the calyptra apex; floral buds usually not wider above the torus; petals rounded, emarginate or truncate; filaments without an evident geniculation; seeds about 0.5 mm long
66 Floral buds, veins on leaf abaxial surface and stem apices rusty with small brown dendritic hairs; petals 7 or more mm long; ovary apex elevated into a conspicuous collar around the style base C. rufescens
66' Leaf abaxial surface and stem apices sparsely to densely covered with stellate or stipitiate stellate trichomes, floral buds glabrous to sparsely or densely covered with stellate or stipitiate stellate trichomes; petals 6.5 mm long or less; ovary apex usually not elevated into a conspicuous collar around the style base (somewhat evident in C. superba)
67 Stem apices, leaf abaxial surfaces and floral buds densely covered with sessile stellate and/or stipitate stellate trichomes
68 Leaf and bud indument combining sessile stellate and stipitate stellate trichomes; floral buds with the lobes slightly differentiated at the apex; plants from Cuba and the Dominican Republic C. lindenii
68' Leaf and bud indument of stipitate stellate trichomes; floral buds with the lobes not differentiated at the apex; plants from Central America
69 Trees reaching 12 m high; Belize, Guatemala and Honduras C. caelestis
69' Shrubs rarely reaching 4 m high; Costa Rica C. brenesii
67' Stem apices and leaf abaxial surfaces sparsely to densely covered with stellate trichomes or sparsely beset with stipitate stellate trichomes, floral buds glabrous or sparsely beset with sessile or less commonly some stipitate stellate trichomes
70 Leaves 7.1–36 × 2.3–16 cm; inflorescence 7–27.5 cm long, with flowers disposed in umbels terminating the inflorescence branches C. superba
70' Leaves 3.8–21.5 × 1.2–10.5 cm; inflorescence 2.7–18.1 cm, long with flowers not disposed in umbels terminating the inflorescence branches
71 Stem apices and abaxial leaf surfaces with sessile and stipitate stellate trichomes C. hirtella
71' Stem apices and abaxial leaf surfaces with sessile stellate trichomes
72 Leaves densely stellate pubescent on abaxial leaf surface; lowlands of the Caribbean slope of Nicaragua, Costa Rica and Panama C. micrantha
72' Leaves with trichomes mostly restricted to the veins; widely distributed C. montana
63' Stem apices and leaf abaxial surfaces glabrous or with very inconspicuous trichomes
73 Inflorescence pendant; flowers with purple petals; seeds muriculate C. muriculata
73' Inflorescence erect; flowers with white or pink petals; seeds smooth
74 Flowers sessile or on pedicels up to about 1 mm long C. montana
74' Flowers on pedicels 1.5 mm long or usually longer than 2 mm
75 Flowers clustered at the end of the inflorescence branches; flower buds up to and usually less than 11 mm long; Southern Mexico (Oaxaca and Veracruz) through Central, and South America, only C. superba reaching the Caribbean including Jamaica
76 Petals 7–12 mm; anthers 4–4.5 mm long; stigma capitate; bracteoles up to 6 mm long; flowers 6–8 merous C. cuatrecasii
76' Petals to 6 mm, less commonly to up to 8.3 mm; anthers to 3.25 mm long; stigma cylindrical to slightly expanded; bracteoles absent or to about 1 mm long; flowers (4-)5–7 merous
77 Flower buds acute to slightly apiculate at the apex, not contricted in the middle, usually white; inflorescence rachis usually white C. superba
77' Flower buds apiculate at the apex, constricted in the middle, usually pink; inflorescence rachis pink C. rhodopetala
75' Flowers not clustered at the end of the inflorescence branches; flower buds 10 mm long or usually longer; Southwest Mexico (Guerrero and Jalisco) or Jamaica
78 Leaf margin serrulate and cilate; pedicels 2–5 mm long; plants from Southwest Mexico (Guerrero and Jalisco) C. jaliscana
78' Leaf margin not serrulate and cilate; pedicels 5–13 mm long; plants from Jamaica
79 Hypanthium ribbed, at least towards the base; abaxial leaf surfaces with tuft domatia at the base C. procera
79' Hypanthium not ribbed at the base; abaxial leaf surface lacking tuft domatia at the base
80 Petals pink C. balbisiana
80' Petals white C. pyxidata

Conostegia sect. Conostegia

Diagnosis

A mostly Central American and Caribbean group, section Conostegia is distinguished by the following combination of characters: calyx calyptrate with the calyptra falling as a unit, lacking calyx teeth altogether, and with conspicuous sclereids internally. Flowers generally pleiostemonous, staminal filaments not evidently geniculate and transitioning smoothly to the anther thecae, style mostly the same length or shorter than the stamens, with a stele within, mucilage inside the ovary, seeds ovoid and smooth.

Conostegia arborea (Schltdl.) Steud.

Conostegia arborea (Schltdl.) Steud., Nomencl. ed. II. 1: 405. 1841. Melastoma arboreum Schltdl., Linnaea 13: 424. 1839. Type: Mexico. “Inter Tioselo et Jicochimalco”, August 1829, Schiede s.n. (lectotype: GOET!, designated here; isolectotypes: BM!, K!, LE).

Conostegia galeottii Naudin, Ann. Sci. Nat. Bot. ser. 3 16: 107. 1850. Type: Mexico. Veracruz: June-October 1940, H. Galeotti 2917 (holotype: P, isotypes: BR!, K!, NY!).

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, abaxially 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.

Figure 63.

Distribution of Conostegia arborea.

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. The amount of indument on floral buds is variable. The flowers of C. arborea have been reported to have a good fragrance (Ventura 1141-NY).

Specimens examined

MEXICO. Puebla: Texcaco, Gold 7 (NY); El Reparo, municipio de Hueytamalco, Ventura 415 (IEB, NY). Veracruz: 1 km al norte de Rancho Viejo sobre el río Pixquiac, Marquez et al. 849 (NY); along very winding road from Naolinco to Misantla 13 km by road S of turn off to Yecuatla and 6 km by road N of Paz de Enríquez, Mun. Yecuatla, Nee et al. 26343 (NY); near Jalapa, Pringle 8170 (NY); Tatzayanala, Municipio de Atzalan, Ventura 1141 (NY).

Conostegia balbisiana Ser. ex DC.

Conostegia balbisiana Ser. ex DC., Prodr. 3: 174. 1828. Type: Jamaica. 1822, C. Bertero s.n. (holotype G!).

Conostegia grisebachii Cogn., DC. Monog. Phan. 7: 700. 1891. Type: Jamaica. 1857, W. Marsh 598 (holotype: GOET!; isotype: BR-fide Schnell (1996), K!, TCD!).

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.

Figure 64.

Distribution of Conostegia balbisiana.

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. The short style characteristic of section Conostegia is evident in specimens Proctor 10432 and 10285 both at NY.

Specimens examined

JAMAICA. Hanover: East Slope of Dolphin Head, Proctor 10432 (NY). Manchester: Holmwood P.T.C., near Christiana, Proctor 10285 (NY). Portland: Woodlands eastern slopes of south end of John Crow Mountains, Harris and Britton 10730 (NY). St. Andrew: Near Diamond (Redhills District), Robertson and Wynter s.n. (NY). St. Ann: Soho, Harris 12024 (NY). St. Catherine: Holly Mount, Mt. Diablo, Harris 8984 (NY); west of Hollymount, Mt. Diablo, Hespenheide 1301 (NY). Ocho Rios and vicinity, Britton and Hollick 2690 (NY). Mandeville, Maxon 2590 (NY).

Conostegia bernoulliana Cogn.

Fig. 65

Conostegia bernoulliana Cogn., DC. Monog. Phan. 7: 698. 1891. Type: Guatemala. Sarnayara: April 1877, K. Bernoulli & O. Cairo 2884 (lectotype: GOET! (image seen), designated here; isolectotype: K!).

Conostegia sphaerica Triana, Trans. Linn. Soc. London 28: 98. 1872. Type: Mexico. Teotalcingo (probably in Oaxaca): June 1842, Liebmann 2842 (holotype: P, isotypes: BM!, BR, C, F!, K!). Also numbered as Liebmann 18 and Liebmann s.n.

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 differentiated, 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.

Figure 65.

Conostegia bernoulliana. A Habit and inflorescence B Leaf abaxial surface C Frontal view of flower D Lateral view of the flower E Longitudinal section of flower bud F Lateral view of the flower from pickled material G Dissection of lateral view of the flower from pickled material H Petal I Stamen J Style. Photos of specimen vouchered R. Kriebel 5578.

Distribution

(Fig. 66). Mexico, Guatemala and on the pacific slope of the Costa Rica cordilleras, reaching the lowlands of the Osa Península, 50–1500 m in elevation.

Figure 66.

Distribution of Conostegia bernoulliana.

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. The main differences 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) noted that the other syntype, Wendland 545 (GOET), pertains to C. oerstediana Berg ex Triana.

Specimens examined

MEXICO. Chiapas: Finca Mexiquito, Purpus 6785 (MO, NY).

GUATEMALA. Huehuetenango: vicinity of Maxbal about 17 miles north of Barillas, Sierra de los Cuchumatanes, between Maxbal and lake to the southeast, Steyermark 48726 (NY). Sololá: south-facing slopes of Volcán Atitlán above Finca Mocá, Steyermark 47935 (NY). Suchitepéquez: Finca Mocá, Hunnewell 14756 (NY); Finca Moca, Skutch 2068 (NY); southern lower slopes of Volcán Zunil, vicinity of Finca Las Nubes, along Quebrada Chita, east of Pueblo Nuevo, Steyermark 35407 (NY).

COSTA RICA. Guanacaste: Liberia, P.N. Guanacaste, cuenca del Tempisque, Sector Cacao, Acosta et al. 1166 (CR, INB, MO, NY). Puntarenas: Distrito Sierpe, Reserva Forestal Golfo Dulce, Rincón, cerca de Banegas, Los Charcos, 1 km. al Este del centro del pueblo Banegas Estación Biológica Los Charcos de Osa, Aguilar 10855 (NY); camino a Rancho Quemado, Kriebel et al. 5540 (INB). San José: Río San Isidro, Jiménez 3840 (NY); Perez Zeledón, Vicinity of El General, Skutch 2604, 4159 (MO, NY).

Conostegia bigibbosa Cogn.

Fig. 67

Conostegia bigibbosa Cogn., Bull. Soc. Roy. Bot. Belg. 30: 252. 1892. Type: Costa Rica. San José: Dans la forest a General, 800 m, February 1891, A. Tonduz 3793 (holotype BR!; isotypes BR! (2), US!). Cited as Pittier 3793 in the original description.

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 undifferentiated, 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.

Figure 67.

Conostegia bigibbosa. A Lenticellate internode B Tubercles on the base of the abaxial leaf surface C Inflorescence D Flowers at anthesis E Longitudinal section of a flower bud F Ventral view of a pickled flower with the petals removed and showing a stamen stuck under the stigma G Lateral view of a pickled flower with the petals removed H Longitudinal section of a hypanthium I Stamen J Style. Photos of specimen vouchered R. Kriebel 5522.

Distribution

(Fig. 68). Endemic to the pacific side of the Cordillera de Talamanca in Costa Rica, 800–1800 m elevation.

Figure 68.

Distribution of Conostegia bigibbosa.

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. The two species are not sympatric however and maintain the petal color difference as well as the petiole tubercle difference in natural populations.

Specimens examined

COSTA RICA. Puntarenas: Finca Las Alturas NW of lechería, Almeda et al. 6697 (CAS, CR, NY); E.B. Las Cruces Trailhead to río Java, Boyle et al. 6265 (CR, NY). San José: Cantón de Pérez Zeledón, Fila Costeña, Fila Tinamastes, en la Fila del Farallón, Hammel et al. 20145 (INB, CR, NY); Aserrí, Quebrada Lajas, ca. 2.5 km al noroeste de Altos el Aguacate, Kriebel et al. 3975 (INB); Pérez Zeledon, Bajo Bonitas, Kriebel et al. 5522 (INB); Vicinity of El General, Skutch 2623 (NY).

Conostegia bracteata Triana

Fig. 69

Conostegia bracteata Triana, Jour. Bot. 4: 209. 1867. Type: Nicaragua. Chontales: 12 September 1867, B. Seemann 36 (holotype: K!; isotypes: BM!, EM, LE; BR).

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 differentiated, 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.

Figure 69.

Conostegia bracteata. A Habit B Leaf abaxial surface C Flowers at anthesis D Infuctescence E Pickled flowers at anthesis F Lomgitudinal section of hypanthia of flowers at anthesis G Bract H Petal I Stamen J Style. Photos of specimen vouchered R. Kriebel 5806.

Distribution

(Fig. 70). Nicaragua to Colombia, 0–1150 m in elevation.

Figure 70.

Distribution of Conostegia bracteata.

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). Specimens 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”. These species look a lot alike in their leaf shape and pubescence but not in their flowers. Conostegia bracteata occurs sympatrically with M. barbinervis in some localities like at La Selva Biological Statation. They can easily be distinguished on the basis of the stipitate stellate trichomes present in M. barbinervis and simple trichomes in C. bracteata.

Specimens examined

NICARAGUA. Chontales: Jinotega (fide Schnell): Cerro San Pedro, Comarca Kilambe, Sandino 183 (MO).

COSTA RICA. Limón: R.B. Hitoy Cerere, siguiendo el sendero el Espavel hasta la cima de un cerro innominado, González et al. 3309 (INB, NY). Heredia: La Selva, OTS field station, La Sura trail at 1070 m, Hopkins 22 (NY); Sarapiquí, sendero Tres ríos y lindero el Peje, Kriebel et al. 3606 (INB, NY). Puntarenas: Osa, Sierpe, Reserva Forestal Golfo Dulce, Rancho Quemado, camino a Chiqueron en la parte mas alta de la fila al Sur de Rancho Quemado, Aguilar 6704 (NY); Golfito, P. N. Corcovado, Estación Agujas, Cerro Rincón, Mora et al. 721 (INB, MO, NY).

PANAMA. Bocas del Toro: Duwebdulup Peak No. of Río Teribe across from Quebrada Huron, Kirkbride and Duke 585 (MO, NY); Above Chiriqui Grande 10 road-miles from continental divide and c. 2 miles along road to east, McPherson 12837 (MO, NY). Canal Zone: Zetek trail 550, Croat 6628 (MO, NY); Madden Forest, Las Cruces Trail, Gentry 1380 (NY). Darién: 0–2 mi. E. of Tres Bocas along shortest headwater of Río Coasi, Kirkbride and Duke 1194 (MO, NY); Coasi-Cana Trail on Cerro Campamiento E. of Tres Bocas headwater of Río Coasi, Kirkbride and Duke 1252 (MO, NY); Vicinity of Cana, Stern et al. 514 (NY). Panamá: 2.5 m N of Goofy Lake on road to Cerro Azul, Croat 11533 (MO, NY); Along highway near top of Cerro Campana, Croat 12089 (MO, NY); 10 miles from highway on road to Cerro Jefe, Croat 15186 (MO, NY); Cerro Azul, Dwyer 2083 (MO, NY); Trail into forest 4.8 miles north of highway just west of El Llano, Gentry 5077 (MO, NY); Between Frijoles and Monte Lirio, Killip 12158 (NY); Parque Nacional Chagres, Sendero hacia Cerro Jefe que sale de la urbanización Altos de Cerro Azul, Kriebel and Burke 5683 (PMA, NY).

COLOMBIA. Antioquia: vicinity of Planta Providencia, 26 km S & 23 km W of Zaragoza, in valley of Río Anorí between Anorí and Dos Bocas, Denslow 2312 (US-not seen).

Conostegia brenesii Standl.

Fig. 71

Conostegia brenesii Standl., Field Mus. Publ. Bot. 18: 801. 1938. Type: Costa Rica. Alajuela: La Palma de San Ramón, 1275–1300 m, 7 August 1927, A. Brenes 5577 (holotype: F!, isotypes: CR!, NY!). Note: at least one sheet bearing this number (also at NY) is of C. montana (Sw) Don ex DC. Schnell (1996).

Description

Shrubs to small trees 1.5–4 m tall with tetragonal young stems that soon become terete and which are densely covered with stipitate-stellate trichomes; the nodal line inconspicuous to absent. Leaves of a pair equal to somewhat unequal in length. Petioles 0.6–4.9 cm. Leaf blades 5–13.2 × 2.5–6.5 cm, 5 nerved or more commonly slightly plinerved, with the innermost pair of veins arising up to 1.5 cm above the base in opposite to sub opposite fashion, elliptic to elliptic-ovate, the base acute or obtuse, the apex acute and acuminate, the margin denticulate or entire, densely hirsute with rigid hairs on both surfaces. Inflorescence a terminal panicle 4.8–9.6 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, the rachis greenish-purple to purple, densely covered with stipitate stellate trichomes, the bracts absent or very early deciduous, bracteoles 1–4 mm, deciduous. Flowers 5–6 merous, calyptrate, flower buds 5.2–7.51 × 2.4–4 mm, rounded at the base, acute apically, the calycine and hypanthium portions weekly differentiated, slightly constricted at the torus; hypanthium 3–3.5 × 2.75–3.25 mm, covered with stipitate-stellate trichomes. Petals 5–6.5 × 6–6.25 mm, pink, light violet or white, obovate, spreading, rounded and emarginate, glabrous. Stamens 11–15, 6.25–7.25 mm, slightly zygomorphic, forming a 45 degree angle, the filament 3.75–4.25 mm, not evidently geniculate, white, anthers 2.5–3 × 0.5–1 mm, linear-oblong and somewhat recurved, laterally compressed, the base sagittate, yellow, the pore 0.1–0.2 mm, terminal. Ovary 6 (-7)-locular, inferior, glabrous and elevated into a collar around the base of the style. Style 5.5–6.5 mm, bending below the stigma, distance between the anther and the stigma -0.1 – -0.3 mm, horizontal distance absent; stigma subcapitate, 1.4–1.6 mm wide. Almost mature berry 5–6 × 5–6, probably purple at maturity like its close relatives. Seeds 0.4–0.6 mm, pyramidal, smooth.

Figure 71.

Conostegia brenesii. A Branch with inflorescence B Leaf abaxial surface C Inflorescence D Flower at anthesis E Longitudinal section of flower bud F Pickled flower at anthesis G Pickled flower at anthesis with half of the petals and stamens removed H Petal I Stamen J Style K Longitudinal section of the hypanthium. Photos of specimen vouchered R. Kriebel 5631.

Distribution

(Fig. 72). Endemic to cloud forests on the Caribbean slope of the Central and Tilaran cordilleras in Costa Rica, 1100–1750 m in elevation.

Figure 72.

Distribution of Conostegia brenesii.

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 stellate 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. The flowering time differs 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. The molecular phylogeny does not place these species as sister taxa, which suggests convergent evolution in the dense stipitate stellate indument (Fig. 1). Conostegia brenesii falls sister to the C. montana-C. setosa complex in the molecular phylogeny.

Specimens examined

COSTA RICA. Alajuela: Zapote, San Carlos, Caribe watershed, Smith 1102 (NY); La Palma de San Ramón, Brenes 5633 (CR, NY); San Carlos, P. N. Juan Castro Blanco, entrando por San Vicente faldas del Cerro Platanar, Rodríguez et al. 6050 (INB, NY); Vara Blanca de Sarapiquí, north slope of the Central Cordillera, Skutch 3161 (NY); Forests of La Palma, Tonduz 12580 (NY). Puntarenas: R. B. Monteverde, Haber 352 (CR). San José: Cuenca del Sarapiquí, Braulio Carrillo, cerca de el túnel, Kriebel 4907, 5631 (INB).

Conostegia caelestis Standl.

Fig. 73

Conostegia caelestis Standl., Field Mus. Nat. Hist. Publ. Bot. Series. 4: 318. 1929. Type: British Honduras (= Belize). Big Creek: Mullins river road, 15 m, 8 March 1929, W. Schipp 63 (holotype: F!, isotypes: A, BM, CAS!, GH, MICH, MO!, NY!, S, UC, US!).

Conostegia hondurensis Standl. ex Yuncker, Field Mus. Nat. Hist. Publ. Bot. Series. 9: 322. 1940. Type: Honduras. Atlántida: bank of Danto river, slopes of Mt. Cangrejal, vicinity of La Ceiba, 300 m, 6 August 1938, T. Yuncker, J. Koepper, & K. Wagner 8818 (holotype: F!, isotypes: BM, GH, K!, MO!, NY!, US!).

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.

Figure 73.

Conostegia caelestis. A Leaf abaxial surface. B Infructescence. Photos of specimen vouchered R. Kriebel 5588.

Distribution

(Fig. 74). Known from Mexico through Guatemala and Belize to Honduras from sea level to 1000 m in elevation.

Figure 74.

Distribution of Conostegia caelestis.

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 differences under C. brenesii.

Specimens examined

MEXICO. Chiapas: west end of Laguna Ocotal Grande, Municipio de Ocosingo, Breedlove 15700 (NY).

BELIZE. El Cayo District: Humming Bird Highway, Gentle 8633 (MO, NY).

GUATEMALA. Alta Verapaz: Sebol in high forest about 2 km east, Contreras 4587 (NY). Izabal: Vicinity of Lago Izabal 1–2 km south of village of Izabal, Jones et al. 3014 (NY); Sierra Caral, Quebrada atravesada por el sendero al noreste de la casa de investigadores, hacia la Finca Bonanza, Kriebel et al. 5588, 5617 (NY, USCG). HONDURAS. Atlantida: along Río Danta road to La Presa vicinity of La Ceiba, Molina 20822 (NY). Olancho: Quebrada Catacamas cerca de la presa en Montaña Peña Blanca, Molina 8352 (NY); Lancetilla mountain, Molina and Molina 25611 (NY).

Conostegia chiriquensis Gleason

Conostegia chiriquensis Gleason in R. E. Woodson, Jr. and R. W. Schery, (Eds), Flora of Panama. Ann. Missouri Bot. Gard. 45: 203–304. 1941. Type: Panama. Chiriquí: Vicinity of Finca Lérida, 1750 m. elev., 7–11 July 1940, R. Woodson & R. Schery 376 (holotype: NY!, isotype: GH!).

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 glabrous 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.

Figure 75.

Distribution of 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 differs 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 differs 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 (Almeda et al. 6599-CAS, NY). Schnell (1996) reports the infestation of anthers in unopened flower buds by gall wasps on the specimen Woodson and Schery 479 (NY).

Specimens examined

COSTA RICA. Puntarenas: Lumber road along Fila Tigre S and E of Las Alturas between Río Cotón and Río Quebrada Nochebuena, Almeda et al. 6597, 6599 (CAS, NY); Cantón de Coto Brus, Z. P. Las Tablas, Est. Biológica Las Alturas, Alfaro 2365 (INB, NY); Cantón de Coto Brus, Z. P. Las Tablas, Hacienda La Amistad, Zona La Neblina, Solano et al. 752 (INB, NY).

PANAMA. Chiriquí: Along the rock road to Lago del Volcán Barú and due SW of El Hato del Volcán, Almeda et al. 6205 (NY); Trail from Paso Ancho to Monte Liro upper valley of Río Chririquí Viejo, Allen 1480 (NY); R. Chiriquí Viejo valley, in Bambita Woods, White 47 (NY); Valley of the upper Río Chiriquí Viejo, White 324 (NY); west of Aserradero Cerro de Punta, Stern and Chambers 64 (NY); Vicinity of Callejón Seco, Volcán de Chiriquí, Woodson and Schery 479 (NY).

Conostegia cuatrecasii Gleason

Fig. 76

Conostegia cuatrecasii Gleason, Bull. Torrey Bot. Club 72: 473. 1945. Type: Colombia. Depto. del Valle: Silva, Río Cajambre, Costa del Pacífico, 5–80 m, 5–15 May 1944, J. Cuatrecasas 17612 (holotype: NY!, isotype: F!).

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.

Figure 76.

Conostegia cuatrecasii. A Leaf abaxial surface B Frontal view of flower at anthesis C Inflorescence with side view of flower at anthesis D Infructescence E Pickled flower at anthesis F Petal G Stamen H Longitudinal section of flower at anthesis with petals removed. Photos of specimen vouchered R. Kriebel and Burke 5673.

Distribution

(Fig. 77). Ranging from Panama to Colombia, Venezuela and Ecuador, and the coastal range of Venezuela, 0–1000 m in elevation.

Figure 77.

Distribution of Conostegia cuatrecasii.

This 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 (Kriebel and Burke 5681-NY).

Specimens examined

PANAMA. Panamá: P. N. Chagres, sendero hacia Cerro Jefe, Kriebel and Burke 5681 (NY, PMA).

COLOMBIA. Antioquia: Rio Chigorodo, Forest on Quebrada Congo 11 km east of Chigorodo 40 km south of Turbo, Haught 4713 (NY); In damp forest on ridge west of Quebrada Mercedes east of Turbo, Haught 4973 (NY). Chocó: Río San Juan margen derecha, Quebrada del Taparal, Cuatrecasas 21468 (NY); Banks of Quebrada Togoromá, Killip and Cuatrecasas 39089 (NY); Rio Mutata tributary of Río El Valle between base of Alto de Buey and mouth of river, Gentry and Fallen 17283 (MO, NY). El Valle: Cordillera Occidental, vertiente occidental, hoya del río Anchicayá ado derecho bosques entre Pavas y Miramar, Cuatrecasas 14386 (NY); Cordillera Occidental, vertiente occidental, Hoya del Río Digua lado izquierdo, Piedra de Moler, Cuatrecasas 15142 (NY); Costa del Pacífico, río Cajambre, Barco, Cuatrecasas 17001 (NY); Costa del Pacífico, río Cajambre, Quebrada de Ordonez, Cuatrecasas 17273 (NY); Río Digua Valley between La Elsa and Río Blanco, Killip 34712 (NY).

ECUADOR. Bolívar: Trip to Bucay 87 km e. of Eloy Alfaro in walk of 5 km up trail n.e. in foothills along pipeline to intake of Guayaquil water supply, Little 6736 (NY). Esmeraldas: Quininde Cantón, Bilsa Biological Station, Mache mountains 35 km W of Quinindé 5 km W of Santa Isabel, Clark and Troya 680 (NY); Carretera Lita-Alto Tambo-La Punta, Gudiño and Moran 1286 (MO, NY); Río San Miguel one hour upstream from San Miguel de Cayapas, Holm-Nielsen et al. 25494 (MO, NY); Rio Onzole on estero Chontaduro, Játiva and Epling 1103 (NY); near Borbon, Játiva and Epling 2203 (NY). Los Rios: Hacienda Clementina, Samama, Harling 284 (NY); Hacienda Clementina, Cerro Samama, Cornejo and Yoza 8188 (NY). Imbabura: Lita, Palacios 12231 (NY). Manabí: Pedernales, Reserva Ecológica Mache-Chindul, Comunidad Ambache (vía marginal de la costa-Chindul), Clark et al. 4229 (MO, NY). Pichincha: Distrito Metropolitano de Quito, carretera Mashpi-Los Bancos, Ulloa et al. 1935 (MO, NY).

VENEZUELA (fide Schnell). Carabobo: Mpio. Autónomo Mora, cuenca del río Moron, Díaz 153 (US).

Conostegia fragrantissima Almeda

Fig. 78

Conostegia fragrantissima Almeda, Proc. Calif. Acad. Sci. 46: 327. 1990. Type: Panama. Bocas del Toro: Fortuna Dam area, along continental divide bordering Chiriqui province, 1200-1300 m, 10 March 1988, F. Almeda, T. Daniel, & G. McPherson 6064 (holotype: CAS!, isotypes: MO!, US, AAU, BM, BR, CR!, DUKE, F, MEXU, PMA, TEX, NY!).

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.

Figure 78.

Conostegia fragrantissima. A Inflorescence B Close up of flowers at anthesis C Flowers bud with longitudinal section to the side D Pickled flower at anthesis E Style F Longitudinal section of the hypanthium G Stamen H Petal. Photos A-B of specimen vouchered F. Almeda 6040 (CAS) and taken by Frank Almeda C–H of specimen vouchered R. Kriebel and D. Solano 3174.

Distribution

(Fig. 79). Known from south eastern Costa Rica and west Panama, 1200–1700 m elevation.

Figure 79.

Distribution of Conostegia fragrantissima.

Schnell (1996) and Almeda (2009) synonimized Conostegia fragrantissima under C. montana. Despite the heterogeneity present in C. montana, there was no stigma lobes 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 (Almeda et al. 6040-CAS, MO, NY). The 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.

Specimens examined

COSTARICA. Puntarenas: Coto Brus, Z. P. Las Tablas, sendero Las Tablas camino a Cotoncito, Kriebel and Solano 3174 (INB, NY).

PANAMA. Chiriquí: SE slopes and summit of Cerro Pate Macho, trail between Rio Palo Alto, 4 km NE of Boquete, Sytsma et al. 4884 (MO, NY).

Conostegia hirtella Cogn

Conostegia hirtella Cogn. in J. D. Smith, Bot. Gaz 16: 4. 1891. Type: Guatemala. Alta Verapaz: Pansamala, 1170 m, May 1887, H. von Tuerckheim 1233 (holotype: BR; isotypes: BR, GH, K!, NY!, P (fide Almeda in Schnell (1996)), PH!, US!).

Conostegia gleasoniana Standl. & Steyerm., Field Mus. Publ. Bot. 22: 361. 1940. Type: Guatemala. Alta Verapaz: Damp forest, region of Cocola, NE of Carcha,1200 m, 2 April 1938, P. Standley 70317 (F!).

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.

Figure 80.

Distribution of Conostegia hirtella.

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. The much sparser pubescence in C. hirtella compared to the dense indument in all the plant in C. caelestis makes it diffcult to confuse them. Unfortunately C. hirtella was not included in the molecular phylogeny to further help elucidate its close relatives.

Specimens examined

GUATEMALA. Alta Verapaz: Bosque mixto de Chamal, margenes del Río Cobán, Molina 12142 (NY); Mountains east of Tactic on road to Tamahú, Standley 71305 (NY); Wet forest near Tactic, above the bridge across Río Frío, Standley 90285 (NY).

HONDURAS. Comayagua: Bosque nuboso de Cordillera de Misoco o Volcán de Guaimaca entre los Departamentos de Olancho y Morazán, Molina 3189 (NY); Bosque Pino-Liquidambar de Montana, la Choca, Cordillera Comayagua, cerca de Coyocutena, Molina 7120 (NY). Morazán: Bosque de nubes del Volcán de Guaimaca, Cordillera de Misoco, Molina 6094 (NY).

NICARAGUA. Estelí: Dept. of Estelí on the border with Madriz, Cerro Pataste, Neill 121 (NY); on border with Madriz, Cerro Pataste, Neill 128 (NY). Matagalpa: Cut over cloud forest area El Porvenir, Cordillera Central de Nicaragua, Molina 20522 (MO, NY); Cut over cloud forest of El Picacho east of Sant María de Ostuma, Cordillera Dariense, Molina and Molina 30522 (NY); Cut over cloud forest area road to Aranjuez, Cordillera Central de Nicaragua, Williams 20143 (NY); road to La Fundadora, cloud forest area north of Sta. María de Ostuma, Cordillera Central de Nicaragua, Williams 24858, 24926 (MO, NY).

Conostegia icosandra (Swartz in Wikstr.) Urban

Fig. 81

Conostegia icosandra (Swartz in Wikstr.) Urban, Rep. Sp. Nov. 17: 404. 1921. Melastoma icosandrum Swartz in Wikstrom, Kongl. Vetensk. Akad. Handl. 64. 1827. Type: O. Swartz s.n. (not seen; application of name based on Urban’s treatment). Lectotype: Guadeloupe: J. Forsstrom s.n. (designated by Howard and Kellogg, J. Arnold Arb. 67: 244. 1986): S!).

Conostegia subhirsuta DC, Prodr. 3: 174. 1828. Lectotype (designated here): Cuba. Havana: 1825, J. de la Ossa s. n. (G!). Additional syntypes: Guadeloupe. Richard s.n. (G!, photograph of P specimen at IJ (Schnell 1996)).

Conostegia mexicana Cogn., DC. Monog. Phan. 7: 707. 1891. Lectotype (designated here): Mexico. Monte Pelado, July 1840–1849, H. Galeotti 2963 (BR!); Mexico. Huatusco, 1335 m, August 1888, Comisión Geografico-Exploradora 401 (BR). Later homonym of C. mexicana (Bonpl.) Ser. ex DC, Prodr. 3: 175. 1828 (see “Excluded taxa or uncertain names” section for further discussion of this name).

Conostegia icosandra var. crenata Urban, Rep. Sp. Nov. 22: 222. 1926. Type: Cuba. Oriente: Arroyo Jimenez, Sierra Maestra, 600–700 m, Ekman 14783 (holotype: S!; isotype: NY!). Conostegia icosandra subsp. crenata (Urban) Borhidi and Muniz, Bot. Kozelem. 58: 176. 1971.

Conostegia lundellii Gleason, Publ. Carnegie lnst. Wash. 522: 348. 1940. Type: British Honduras (= Belize). El Cayo District: San Augustín, Mountain Pine Ridge, July–August 1936, C. Lundell 6587 (holotype NY!, isotypes MICH!, NY!).

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 differentiated; 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 difficult 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.

Figure 81.

Conostegia icosandra. A Leaf abaxial folia surface B Side view of flower at anthesis C Frontal view of flower at anthesis D Infructescence E Flower bud with detail of bracteoles at the base F Pickled flower at anthesis G Longitudinal section of flower at anthesis H Frontal view of flower at anthesis I Petal J Stamen. Photos of specimen vouchered R. Kriebel 5578 (flowers) and R. Kriebel 5580 (fruits).

Distribution

(Fig. 82). Ranging from Mexico throughout Central America and the Caribbean, reaching the coast of Venezuela, sea level to 1500 (-2000) m.

Figure 82.

Distribution of Conostegia icosandra.

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). These 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. These 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. The 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. The specimen Davidse, González and León 18607 (NY) from Zulia, Venezuela, unlike any other specimens studied has extremely narrow leaves.

Specimens examined

CUBA. Oriente: Sierra Maestra, Pinar de Papagayo, Ekman 9273 (NY); Loma del Gato, Cobre Range, Clemente 1823, León 10054 (NY). Santa Clara: Trinidad Mountains, Aguacate, Britton and Wilson 5377 (NY); Southern Oriente and Pico Turquino, High Maestra, Ekman 10944 (NY); Pinar de la Caridad, Southeast of Yara, Ekman 14689 (NY); Buenos Aires, Trinidad Hills, Jack 6844 (NY); Loma Las Divisiones, Banao Mts., León 7864 (NY); Sierra de Gavilanes, Sanoti-Spiritus Mts., León and Clement 6616 (NY); Lomas de Banao, Luna 172 (NY); Sevilla Estate near Santiago, Jiquarito, Sierra Maestra, Taylor 501 (NY); Vicinity of La Sabana, Buenos Aires, Trinidad Mountains, Smith et al. 3384 (NY). Las Villas: Buenos Aires, Hills East of Cienfuegos, León and Jack 13934 (NY).

GUADALOUPE. Basse Terre: Mosciu district, south of La Citerne, Proctor 20129 (NY). Gran Savane, Duss 3472 (NY); along road from Bains Jaunes to Soufriere, Howard 19773 (NY); Dr. Madiana s. n. (NY); Grand Etang, Martin and Hus 488 (NY).

HAITI. Massif de la Hotte: Depts. Sud-Grand Anse límite, zona rural “Gerard”, 18 km Norte de Camp Perrin, en la carretera a Beaumont y Jérémie, Zanoni et al. 25696 (NY).

JAMAICA. Portland: East slope of the John Crow Mts. 1.5–2 miles southwest of Ecclesdown, Proctor 9975 (NY); about 1.5–2 miles southwest of Ecclesdown, Yuncker 18537 (NY); Vicinity of Moody’s Gap, Britton 3388 (NY); John Crow Mountains, Britton 4181 (NY); Woodland eastern slopes of south end of John Crow Mountains, Harris and Britton 10721 (NY). St. Andrew: Coopers Hill, Red Hills, Proctor 8483 (NY). St. Thomas: Cuna Cuna Gap, Britton 4050 (NY).

MARTINIQUE. Sta. Marie, Duss 917 (NY); Hauteux de Case-Pelote, Petons du Cartel, Duss 4626 (NY); along road from Morne Rouge to Mt. Pelée, Holdridge 465 (NY); Sieber 119, 466 (NY).

MONTSERRAT. Pond Mountain, Shafer 683 (NY).

SAINT LUCIA. Forest reserve between Qilesse and Morne Troumasse, Howard 11666 (NY); 1.5 miles south west of Millet on the Millet River, Slane 137 (NY).

SAINT VINCENT. Upper Richmond Valley, Smith 489 (NY).

TOBAGO. Roxborough-Parlatuvier road, 8th-9th mileports, Sandwith 1924 (NY).

MEXICO. Chiapas: 18–20 km north of Ocozocoautla along road to Mal Paso, Municipio of Ocozocoautla de Espinoza, Breedlove and Thorne 21011 (CAS, MO, NY); between Colonia Francisco I. Madero and Colonia A. Lopez Mateos, Breedlove 50594 (CAS, NY); Loc. a 3 km al este de Tziscao en el parque natural lagos de Montebello, Cabrera, Mendez and Cabrera 2957 (NY); Avalinero, Palenque, Matuda 3601 (NY); Finca Mexiquito, Purpus 6785 (MO, NY). Oaxaca: Juchitan to the east of Sarabia, MacDougall s.n. (NY); Palomares, Juchitán, MacDougall 88 (NY); Distrito Choapam, Yaveo, Mexia 9141; Sierra de Juárez, Paray 127 (NY); San Juan Teotalcingo and Santiago Choapam District of Choapam, Schultes 568 (NY); Between Monte Negro de Lalana and San Juan Lalana District of Choapam, Schultes and Ro 796 (NY). Puebla: Moist roadside soil near Ocostoc below Teziutlan, Sharp 45823 (NY); Atecocomo, municipio de Cuetzalan, Ventura 1115 (NY). Tabasco: Km. 4 de la Est. Chontalpa hacia Malpaso, carretera Huimanguillo Malpaso, Cowan et al. 2545 (NY); Km 31.9 de la desviación de Huimanguillo hacia Fco. Rueda, Cowan 3329 (MO, NY); Achotal, Balancan, Matuda 3068 (NY); Carretera Huimanguillo-Francisco Rueda, 8.5 km, Ventura 20608 (MO, NY). Veracruz: Minatlitlán, Mell s.n. (NY); About 6 km by air S of Tlapacoyan on road to Altotonga, Mun. Tlapacoyan, Nee and Diggs 24874 (NY); Vicinity of La Calavera 10 km N of Altotonga (13 km by road) on road to Tlapacoyan, Nee and Diggs 24891 (NY); Jaltipan, Orcutt 6515 (NY); Zacuapan, Purpus 7510 (NY).

BELIZE. Cayo: Kinlocks Camp Road, Balick 3345 (MO, NY); Along Hummingbird Highway south of Belmopan between 30–38 mi., Croat 24837 (MO, NY); Mountain Pine Ridge, San Agustin, Lundell 6587 (NY); Near Río On along road to Augustine, Nee et al. 46782 (MO, NY); Mountain Pine Ridge on main road to Augustine, Ratter 5168 (NY). Stann Creek: in broken cohune ridge near Carasow Hill, Gentle 8212 (NY); MI 32 Hummingbird HWY, Margaret Creek Village, Holmes 4613 (NY); Mullins River Road, Schipp 145 (MO, NY). Toledo: 1.4 km on S turnooff ”Farmer’s Road” 10km W of Punta Gorda Town, Arvigo 969 (NY); in edge in wooded island Condemn Branch Pine Ridge, Gentle 5283 (NY); ca. 3 km W of coast of Punta Gorda, Nee et al. 46934 (MO, NY).

GUATEMALA. Alta Verapaz: Tucuru, Finca de la Concepción, Boeke and Utzschneider 2932 (NY); Chacirociha, Finca Seaway, Hatch 193 (NY); entre San Pedro Carchá y Sacoyoú, Molina and Molina 12124 (NY); large swamp just east of Tactic, Steyermark 43988, 92629 (NY); Saquija 43 km. northeast of Cobán, Standley 70110 (NY); von Turckheim 2236 (NY). Izabal: 1–2 km south of Izabal, Jones 3015 (F, NY); South shore of Lake Izabal east of village of Izabal at sea level, Jones 3143 (NY); San Tomas de Castilla on coastline road to Las Pavas, Marshall et al. 332 (MO, NY); Vicinity of Quiriguá, Standley 23924 (NY). Quiché: Finca Chaila, “Zona Reyna”, Skutch 1809 (NY).

HONDURAS. Comayagua: 14 km SE of Taulabé, Davidse and Pohl 2228 (NY); Matorrales en colinas rocosas de los alrededores de La Libertad, Molina 7074 (NY); Bosque de Montania La Choca en Cordillera de Comayagua cerca de Coyocutena, Molina 8129 (NY). Cortés: Río Amapa on road to Lake Yojoa, Howard 587 (NY); Matorrales húmedos de Peña Blanca, nacimiento del Río Lindo, Molina 6777 (NY); bosque de pino-roble entre Cofradia y Cusuco, Molina 7294 (NY); Bosque lluvioso entre Agua Azul y Pito Solo Lago de Yojoa, Molina 7331 (NY); Montaña La Cumbre caserío las Piñitas, Molina 10522 (NY); Matorrales y bosque mixto de Cascada El Chorrito 16 kms. al S.O. de Siguatepeque, Molina 10901 (NY); Montaña San Idalfonso entre Cusuco y San Isirdo, Molina 11544 (NY); above la Misión, Molina 12341 (NY). Yoró: Ocotales en sabanas pedrejosas de Piedra Colorada, Molina 6886 (NY). El Paraíso: Montaña Teupasenti entre El Junquillo y Teupasenti, Molina 11866 (NY); Matorrales Quebrada El Coyol, Sierra El Chile entre El Junquillo y El Robledal, Molina 14158 (NY); Bosque mixto Quebrada Tapahuasca, Molina 14658 (NY). Gracias a Dios: Mocorón 60 km SO de Puerto Lempira, Rivas 131 (NY); Alrededores de Mocorón 60 km al SO de Puerto Lempira, Torres 136 (NY). Intibucá: Cordillera Opalaca, between Calaveras and Pela Naríz road to La Esperanza, Molina 22600 (NY). Morazán: entre la Pirámide y Zambrano, Molina 11030 (MO, NY); carretera a Maraita, Molina 13781 (NY); along Agua Amarilla creek 5 kms. north of El Zamorano, Molina 26269 (NY); Region of Agua Amarilla above El Zamorano, Standley et al. 5074 (NY); Agua amarilla, Williams and Molina 10142, 12156 (MO, NY). Siguatepeque: near El Achote, hills above the plains of Siguatepeque, Yuncker et al. 5822, 5825, 5909 (MO, NY).

NICARAGUA. Nueva Segovia: in gorge on Cerro Mogoton, Atwood and Neill AN15 (MO, NY); San Mateo, 16 millas al sur de Tronquera cerca de Río Wawa, Molina 15075 (MO, NY); frecuente a orillas de Río Leicus cerca del campo de aviación de Tronquera 35 kms S. O. de Waspan, Molina 15174 (MO, NY).

BRAZIL: Bahia: Camaça, km la 2 da estrada do lado L de Camaça, dos Santos 1340 (NY).

COLOMBIA. Guajira: Municipio Riohacha, Sierra Nevada de Santa Marta, sector de Cenuá, Dueñas 1779 (COL). Magdalena: Cincinati, Giacometto 138 (NY); along quebrada in S portion of finca Reflejo, Kirkbride 2146 (NY); Around San Andrés de la Sierra, western slope of Cordillera de Santa Marta, Pittier 1684 (NY); Plants of Santa Marta above Agua Dulce, Smith 765 (NY).

TRINIDAD. Arima: Cumaca Roadover Morn Croix, Adams 14331 (NY); Arima-Blanchisseuse Road, Kalloo B.1056 (NY); St. Pat’s, Arima Valley, Snow and Fleming 310 (NY); Arima Valley, Snow DS-H (NY); Blanchiseusses River, Snow DS-D (NY).

VENEZUELA. Falcón: Cerro Santa Ana, ascensión del lado sur desde el pueblo de Santa Ana, Steyermark and Braun 94654 (NY). Miranda: Los Guayabitos (arriba de Baruta), Aristeguieta 2880 (NY); Bernardi s. n. (NY). Monagas: Cuenca del Río Caripe al E de Caripe vía Las Margaritas, Quebrada Pajaral 3 km al E de Escuela Rural El Aguacate, 11.2 Km al E del puente sobre el Río Colorado, Michelangeli and Alford 612 (NY); P. G. Pacanilla, Parque Nacional Guatopo, Quebrada al inmediatamente al S de P.G.P Macanilla, 9 km S de Los Alpes, Michelangeli and Michelangeli 803 (NY); Montaña de Aguacate, along Quebrada de Pajarral, tributary to Río Caripe, northeast of Alto de Aguacate, between Caripe and Caripito, Steyermark 62200 (NY). Zulia: Distrito Colón, 3 km E of the Río de Oro settlement on the Río de Oro, Davidse et al. 18607 (NY).

Conostegia jaliscana Standl.

Conostegia jaliscana Standl., Field Mus. Publ. Bot. 4: 245. 1929. Type: Mexico. Jalisco: Streamside, Arroyo de los Hornos, Hacienda del Ototal, E of San Sebastian, Sierra Madre, 1500 m, 5 March 1927, Y. Mexía 1819 (holotype: F!, isotypes: A!, BM!, C, CAS!, GH!, LA, MICH, MO!, NY!, S!, TEX!, UC, US!).

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 differentiated; 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.

Figure 83.

Distribution of Conostegia jaliscana.

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. This latter hypothesis is the one supported in the molecular phylogeny.

Specimens examined

MEXICO. Guerrero (fide Schnell): in Sierra Madre del Sur, 20 mi S of Chilpancingo on the Acapulco Highway, Smith M73 (TEX). Jalisco: ca. 27.8 mi sse of Puerto Vallarta on a new dirt road to silver mine, left hand turnoff hwy 200 toward Manzanillo, Almeda 2540 (CAS).

Conostegia lindenii Cogn.

Fig. 84

Conostegia lindenii Cogn, DC. Monog. Phan. 7: 705. 1891. Type: Cuba. La Guinea, 600 m, no date, J. Linden 2204 (holotype: LE, isotype: BR!).

Conostegia lomensis Urban, Fedde Rep. Sp. Nov. 17: 161. 1921. Type. Dominican Republic. Santo Domingo: Barahona, La Loma, 1000 m, September 1911, M. Fuertes 1028 (holotype: A!).

Conostegia furfuracea Urban & Ekman, Arkiv. Bot. 23A (11): 15. 1931. Type: DOMINICAN REPUBLIC. Santo Domingo: Duarte, Cordillera Septentrional, Loma Quita-Espuela, c. 700 m, 25 April 1929, E. Ekman H12273 (holotype: S!, isotypes: A!, GH!, NY!, US!).

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.

Figure 84.

Conostegia lindenii. A Inflorescence B Close up of flower. Photos by Eldis Bécquer.

Distribution

(Fig. 85). Cuba and the Dominican Republic, 750–1300 m elevation.

Figure 85.

Distribution of Conostegia lindenii.

Note that in Dominican Republic a pubescent morphotype of C. superba has been collected in the same place. They 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.

Specimens examined

CUBA. Oriente: Loma del Gato and vicinity Cobre Range of Sierra Maestra, Edmond 84 (NY); Sierra Maestra, El Gigante on the high ridge, on Río Guisa, Ekman 16084 (NY).

DOMINICAN REPUBLIC. Monseñor Nouel: Road up to Alto Casabito ca. 8 km W of jct. with Highway Duarte on road from Bonao to Constanza, Judd et al. 6521 (NY); Firme de Banilejo Piedra Blanca, Liogier and Liogier 19940 (NY).

Conostegia macrantha O. Berg ex Triana

Fig. 86

Conostegia macrantha O. Berg ex Triana, Trans. Linn. Soc. London 28: 97. 1872. Type: Costa Rica. San José: Candelaria, no date, A. Oersted 12 (lectotype: BR!, designated here; isolectotypes (fide 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 differentiated; 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.

Figure 86.

Conostegia macrantha. A Leaf abaxial foliar surface B Inflorescence C Lateral view of flower at anthesis D Frontal view of flower at anthesis E Close up of the surface of the hypanthium F Longitudinal section of a hypanthium of a flower at anthesis with its parts removed G Petal H Stamen I Style J Stigma. Photos of specimen vouchered R. Kriebel 5406.

Distribution

(Fig. 87). Endemic to Costa Rica, 1300–3000 m elevation.

Figure 87.

Distribution of Conostegia macrantha.

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). This species is also reported from northern Panama by Schnell (1996) but I have not seen any Panamanian specimens. The 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) noted some specimens from Vara Blanca showed evidence of introgression from C. oerstediana in their smaller flowers.

Specimens examined

COSTA RICA. Alajuela: Near Continental Divide, vicinity of Vara Blanca, Chrysler 5348 (NY); Alfaro Ruiz hills above laguna, Smith 10075 (NY). Cartago: Volcán Turrialba on road between Lechería La Central and La Trinidad, Schnell 1077 (MO, NY); south slope of Turrialba Volcano above Santa Cruz, Williams 19677 (NY); near La Sierra, about 25 km south of Cartago, Cordillera de Talamanca, Williams et al. 28022 (NY). Heredia: Sacramento, Finca Ingrid Steinvorth, Kriebel 5406 (INB, NY); near Río Las Vueltas N.E. of Volcán Barba, Lent 2644 (CR, NY). San José: San Cristóbal Norte, Desamparados, Antonio 706 (CR, NY); Property of Hacienda Forestales above Cascajal, Lumer 1301 (NY); NE of Coronado, Chrysler 5348 (NY); Riviera de los Arcángeles, Escazú, Pittier 13053 (NY).

Conostegia micrantha Standl.

Fig. 88

Conostegia micrantha Standl., Field Mus. Nat. Hist, Bot. Series 4: 246. 1929. Type: Panama. Bocas del Toro: Buena Vista Camp on Chiriquí Trail, Almirante, 400 m, January–March 1928, G. Cooper 578 (holotype F!, isotype NY!).

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.

Figure 88.

Conostegia micrantha. A Branch apex B Abaxial view of leaf base. Photos of specimen vouchered R. Kriebel 5553.

Distribution

(Fig. 89). Nicaragua to Panama and in Ecuador, 100–1200 m elevation.

Figure 89.

Distribution of Conostegia micrantha.

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. This is indeed a possibility supported also by the molecular phylogeny where C. micrantha falls in a clade of several specimens of C. montana. The question does remain as to whether C. montana should be considered a single species.

Specimens examined

NICARAGUA. Río San Juan: in tall forest near Río San Juan at ‘‘El Relos” ca. midpoint between El Castillo and Delta de San Juan, Bunting and Licht 771 (NY). Jinotega: Cordillera Isabelia near Río Bote Comarca de Bocaycito 117 km from Matagalpa, Neill 7170 (NY).

COSTA RICA. Alajuela: Cataratas de San Ramón, Brenes 13654 (CR, NY); Forested stream edge and cleared slopes below the Methodist Rural Center Quebrada Marín, About 7 km east of Ciudad Quesada, Burger and Stolze 4990 (CR, F, NY); Upala, San Cristobal, Sendero toma de agua, Espinoza 1739 (INB, NY); R. V. S. Bosque Alegre, Laguna Hule, Kriebel and Larraguivel 694 (INB, NY); Woods N.E. base of Arenal Volcano 5 km W of Fortuna San Carlos, Lent 2524 (NY); San Carlos, Fortuna, R. B. Arenal Mundo Aventura, Rodríguez 8715 (INB, NY); Bijagua, Parque Nacional Volcán Tenorio, estación Pilón, Santamaría and Azofeifa 1104 (INB, NY); Villa Quesada, San Carlos, Smith 2508 (CR, NY); P. N. Tenorio, Sector Pilón, Río Celeste, Desviación hacia la catarata, Vargas and Villalobos 1243 (INB, NY); Near Artezalea and Methodist Rural Center about 8 km N.E. of Villa Quesada, Williams 17200 (NY). Cartago: entre Pavones y Chitaria, Turrialba, Jiménez 734 (CR, NY). Heredia: Finca La Selva, the OTS Field Station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, Southern boundary, SW corner, Folsom 9525 (INB, NY); Cantón de Sarapiquí, entre Bijagual y Magsasay, Bosque ripario sobre el río Mojón, Rodríguez et al. 3110 (INB, NY). Limón: Pococí, P.N. Braulio Carrillo, Estación Quebrada González, entre sendero El Ceibo y sendero Botarrama, Rodríguez et al. 5076 (INB, NY); Talamanca, Bratsi ca. 1 km NW de Laguna Dabagri, Rodríguez et al. 11296 (INB, NY).

PANAMA. Bocas del Toro: Región of Almirante, Buena Vista camp on Chiriquí trail, Cooper 619 (NY). Coclé: n. side Gaital above El Valle, Darwin 2780 (NY); El Valle de Antón, Trailside woods near La Mesa about 5 miles north of El Valle, Wilbur and Luteyn 11721 (NY).

ECUADOR (fide Schnell). Esmeraldas: near Río Palavi Awá encampment, Hoover et al. 3741 (MO). Los Ríos or Pichincha: Montañas de Ila, on road from Patricia Pilar to 24 de Mayo, Dodson et al. 8673 (MO).

Conostegia montana (Swartz) D. Don ex DC

Figs 90, 91

Conostegia montana (Swartz) D. Don ex DC, Prodr. 3: 175. 1828. Melastoma montana Swartz, Prodr. Veg. Ind. Occ. 69. 1788. Type: Jamaica. no date, O. Swartz s.n. (holotype: S!)

Melastoma calyptrata Desr. in Lam. Encycl. Meth. Bot. 4: 51. 1797. Type: Antilles, 177 in herb. Surian (v.s. Apud. D. de Jussieu) (not seen: Cogniaux’s monograph cites a Richard specimen at P, apparently the type, of which there is a photograph in IJ; the latter has served to authenticate the name).

Conostegia alpina Macfad., Fl. Jamaica 2: 72. 1850; nom. inval.

Conostegia calyptrata (Desr.) DC, Prodr. 3: 174. 1828.

Conostegia cooperi Cogn, DC. Mon. Phan. 7: 705. 1891. Type: Costa Rica: Cartago: 1500 m, 1888, J. Cooper 290 (distributed by Donnell Smith as 5740) (holotype: BR; isotypes: A, CAS!, F, GH!, K!, M!, US!).

Conostegia petiolata Gleason, Brittonia 1: 184. 1932. Type: British Guiana (= Guyana). Demerara, no date, Parker s.n. (holotype: K!).

Conostegia multiflora Gleason, Bull. Torrey Bot. Club 66: 416. 1939. Type: Ecuador. Esmeraldas: Playa Rica, Parroquia de Concepción, 105 m, 7 December 1936, Y. Mexía 8409 (holotype: NY!; isotypes: BM, F!, GH, K!, MO, NY!, S!, UC!, US!).

Description

Shrubs to small trees 2–11 m tall with apically tetragonal stems that are glabrous or pubescent with stellate hairs or occasionally furfuraceous with sessile and stipitate stellate hairs; the nodal line present yet slight. Leaves of a pair equal to unequal in length. Petioles 0.3–6.7 cm long, sometimes bearing to small tubercles abaxially at the lamina/petiole junction. Leaf blades 3.8–21.3 × 1.2–9.6 cm, 3–5 nerved or slightly plinerved, narrowly ovate to ovate, elliptic or obovate, glabrous adaxially, glabrous or with some scattered stellulate hairs abaxially. The base acute to rounded, the apex acute to acuminate, the margin entire to serrate. Inflorescence a terminal panicle 2.7–18.1 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, rarely a spike, accessory branches infrequent, the rachis glabrous or with some stellulate hairs, sometimes strongly flattened, subulate, lanceolate or slightly ovate, bracteoles 0.5–1 × 0.25–0.5 mm, early deciduous or persistent. Flowers sessile or more commonly with pedicels to 1mm long. Flowers (4-)5(-7) merous, calyptrate. Flower buds 4–7(-10) × 2.25–4.3 mm, variable in shape, mostly obovoid to ellipsoid, the base rounded, the apex acute to apiculate, slightly constricted below the calyptra; the hypanthium 2.85–3.25 × 4–4.5 mm, glabrous or with few scattered stellate trichomes. Petals 4.5–5.25 × 4–4.25 mm, white to pink or lilac, oblong or obovate, spreading to somewhat reflexed, the apex rounded to emarginate, glabrous. Stamens (9-)12–16(-19), slightly zygomorphic, the filament 2.5–3.5 mm, white, 1.5–2.5 × 0.5–0.75 mm, yellow, the pore 0.1–0.25 mm wide. Ovary (4-)5–7(-8) locular, inferior, the apex glabrous and elevated into a collar around the style. Style 3.25–3.75 mm, straight and curving towards the apex, vertical distance between the anther and the stigma -2 – -0.5 mm, horizontal distance absent, stigma truncate to capitellate, 1.3–1.6 mm wide. Berry 4.5–6 × 4.5–6 mm, purple black. Seeds 0.6 mm, pyramidal, smooth.

Figure 90.

Morphological variation in Conostegia montana. A Fertile branch B Abaxial leaf surface C Flower and floral buds D Fruit E Flower and floral buds F Abaxial leaf surface G Flower and floral buds H Branch showing leaf bases (note tubercles towards the apex of the petioles) I Inflorescence J Abaxial leaf surface (note serrate margin) K Inflorescence L Flower and floral buds M Inflorescence N Leaf abaxial surface (note coriaceous leaves) O Infructescence A–D from specimen vouchered R. Kriebel 5446 E from specimen vouchered R. Kriebel 5544 F–G from specimen vouchered R. Kriebel 5446 H–I from specimen vouchered R. Kriebel 5593 J–K from specimen vouchered R. Kriebel 5751 L–M from specimen vouchered R. Kriebel 4895 N–O from specimen vouchered R. Kriebel 5662.

Figure 91.

Conostegia montana. A Longitudinal cut of a flower bud B Petal C Stamen, lateral view D Style E Flower bud F Flower G Longitudinal cut of a flower H Petal I Stamen, lateral view J Style K Flower bud L Flower M Longitudinal cut of a flower N Petal O Stamen, lateral view P Style Q Longitudinal cut of a flower bud R Flower S Longitudinal cut of a flower T Petal U Stamen, lateral view V Style. A–D from specimen vouchered R. Kriebel 5544 E–J from specimen vouchered R. Kriebel 5568. K–P from specimen vouchered R Kriebel 5751 from specimen vouchered R. Kriebel 5544 Q–V from specimen vouchered R. Kriebel 5593.

Distribution

(Fig. 92). Lesser Antilles, Jamaica, in Mesoamerica from Mexico (Chiapas) to Panama (except Belize and El Salvador), western Colombia, northern Ecuador, and coastal Venezuela, at 0–2100 m in elevation.

Figure 92.

Distribution of Conostegia montana.

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. The molecular phylogeny (Kriebel et al. 2015) including several specimens of different 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. This 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 differences between both, see the discussion under the latter species. Schnell (1996) clarified the precedence of the Swartz name Melastoma montana over M. calyptrata.

Specimens examined

DOMINICA. Roseau: Laudat, Beard 1467 (NY); Near the Fresh Water Lake on southeast side of Morne Micotrin and along old road on side of the mountain 1–1/2 miles east of Laudat, Chambers 2563 (MO NY); Castle Bruce Road and Trace, Cowan 1622 (NY); Lloyd 188 (NY); St. John. Morne Diablotin on NW ridge, Webster 13325 (MO, NY); Sylvania, Rainforest bordering Imperial Road, Hodge 3972 (NY); Proctor 95 (NY); Lower slopes of Morne Plat Pays above Bellevue along trail to Grand Bay, Wilbur et al. 7879 (NY).

GUADALOUPE. Basse Terre: Grand Etang, Barrier 2404 (NY); loc. cit., Martin and Gus 489 (NY); Forest above Bains Jaunes, Howard 19386 (NY); Forest near les Mamelles, Howard 19778 (NY).

JAMAICA. Portland: Hillside Cura Cura Gap, Britton 3527 (NY); North side of Cuna Cuna Pass, Harris 10652 (NY); 0.5 mile north of Hardwar Gap, Port Royal Mts, Proctor 9346 (NY). St. Andrew: Blue Mountains, along the road from Newcastle to Catherine’s Peak, Skean and Slantis 1872 (MO, NY); Blue Mountains, Hollywell Gardens, along the ”Waterfall Trail” originating at the rental cabins, Skean and Slantis 1903 (NY). St. George: Vicinity of Cinchona, between Moodee’s Gap and Vinegar Hill, Britton 163, 221 (NY); Below Vinegar Hill, Harris 6336 (NY). St. Thomas: Saint Thomas and Portland Parish boundary, Blue Mountains, Trail from just below Blue Mountain Peak (Middle Peak) to Portland Gap, Judd 5288 (NY); Portland Gap, Proctor 8204, 9597 (NY); West slope of Blue Mt. Peak, Proctor 9443 (NY); West slopes of Blue Mountain Peak between Portland Gap and the top, Skean and Slantis 1919 (NY).

MARTINIQUE. Saint-Pierre: Near L’Alma, Bailey and Bailey 284 (NY); Valle du Carbet St. Denis, Duss 108924 (NY).

ST. VINCENT. Valley of north fork of Cumberland River, Morton 5519 (MO, NY); Smith and Smith 994 (NY).

MEXICO (fide Schnell). Chiapas: km 18 Col. Cuahutemoc trinitaria Chiapas, Shilom Ton 8196 (MO).

GUATEMALA. Alta Verapaz: Wet forest near Tactic above the bridge across Río Frío, Standley 90312 (NY); Large swamp east of Tactic, Standley 92640 (NY). Baja Verapaz: Entrada al Biotopo Universitario para la Conservación del Quetzal, Kriebel et al. 5568 (NY, USCG); Biotopo del Quetzal WNW of Purulhá, Stevens et al. 25434 (MO, NY). Izabal: Sierra Caral, sendero al sur de la casa de investigadores hacia la cima de la Sierra Caral, mitad del camino, Kriebel et al. 5593 (NY, USCG).

HONDURAS. Comayagua: Bosque de pino-liquidambar de Montaña El Cedral Cordillera Montecillos, Molina 7202 (NY). Cortés: frecuente en el bosque nebuloso de Cusuco, Montaña Idalfonso, norte de Cofradía, Molina 7256, 8252 (NY). Intibucá: Bosque abierto de El Cedral 20 kms al suroeste de Siguatepeque camino a Jesus de Otoro, Molina 6150 (NY). La Paz: Cut over cloud forest of Montaña Verde on Cordillera Guajiquiro, Molina 24374 (NY). Ocotepeque: a 35 km al NE de Nvo Ocotepeque camino a San Pedro Sula, Martínez and Téllez 12986 (MO, NY).

COSTA RICA. Alajuela: La Palma de San Ramón, Brenes 5577, 5647, 6282 (CR, NY); Guatuso, P.N. Volcán Tenorio, Cuenca del Río Frío, Alto Masís, Chaves and Muñoz 335 (INB, NY); Finca La Paz, San Ramón, Kriebel 1476 (INB, NY); Parque Nacional Volcán Tenorio, Estación El Pilón, Sendero hacia Cerro Montezuma, Kriebel et al. 5496 (INB, NY); Vara Blanca de Sarapiquí, north slope of Central Cordillera, Skutch 3314 (NY). Cartago: approximately 2 km E of Peñas Blancas bordering the Río Naranjo, Almeda and Nakai 3958 (CR, NY); P.N. Tapantí, alrededores de la estación, Kriebel 5352 (INB, NY); Kiri Lodge sendero a la catarata, Kriebel et al. 5466 (INB, NY); south slope of Turrialba Volcano above Santa Cruz, Williams 19662 (NY). Heredia: Z. P. La Selva, camino al lindero sur, Kriebel et al. 3984 (INB, NY). Limón: Parque Internacional La Amistad, Camp 2, ridge above camp, Monro and Santamaria 5540 (INB, NY); Talamanca, P.I. La Amistad, colectando en transecto 2 y alrededores, Solano 4165 (INB, NY); Valle de la Estrella, Fila Matama, Cerca de 11 km SW del pueblo de Aguas Zarcas, Sitio Helechal, Solano et al. 4759 (INB, NY). Puntarenas: Monteverde cloud forest reserve, Lumer 1025, 1101 (CR, NY); Buenos Aires Sabanas Esperanza y Bosques Aledaños, Santamaría et al. 4407 (INB, NY). San José: Pérez Zeledón, Río Nuevo, Savegre Abajo, Finca de Julio Mena, Estrada et al. 2194 (CR, NY); Río Quebradas a orillas de la carretera Interamericana camino a Pérez Zeledón, Kriebel and Hammel 3338 (INB, NY).

PANAMA. Bocas del Toro: Robalo Trail Northern slopes of Cerro Horqueta, Allen 4942, 5006 (NY); along road to Chiriquí Grande c. 10 road miles from continental divide and about 2 miles along road east of highway, McPherson 10834 (MO, NY). Canal Zone: Militar Reserva Fuerta Sherman and adjacent canal zone, road S-1 between Gatun and Pina, Liesner 1360 (MO, NY); Premontane wet forest along road S1 5.5 km W of Gatun Dam, Nee and Hale 9692 (NY). Chiriquí: Quebrada Velo Vcty. Finca Lerida, Allen 4684 (NY); heavily forested slope above the Río Caldera beyond Bajo Mono in the vecinity of Boquete, Wilbur et al. 11075 (MO, NY). Colón: Carretera hacia el poblado de Piñas, Galdames and Guillén 3317, 3320 (MO, NY); Santa Rita Ridge 2-3 miles from Transisthmian Hwy, Gentry 1860 (NY). Darién: Coasi-Cana trail on Cerro Campamento E. of Tres Bocas headwater of Río Coasi, Kirkbride and Duke 1253 (MO, NY). Panamá: between peaks of Cerro Trinidad saddle on SE slope, Kirkbride and Duke 1643 (MO, NY); El Llano-Cartí road about 4.6 miles north of junction with Pan-American Highway, trail to east, McPherson 12514 (MO, NY). San Blas: El Llano-Carti road near Nusagandi along Sendero Nusagandi, c. 11 road-miles from Pan-American Highway, McPherson 12731 (MO, NY). Veraguas: Serranía de Tute, Aranda et al. 2672 (NY); above Santa Fé on slopes of Cerro Tute below Agricultural School, Gentry 6217 (MO, NY).

COLOMBIA. Antioquia: Municipio Campamento, 6 km NO del pueblo en la vía a Mina Las Brisas, Callejas et al. (MO, NY). Chocó: cerca del río Atrato en los alrededores de Quibdó, Araque and Barkley 129 (NY); Rio Munguido afluente del rio Atrato, alrededores de Altagracia, Forero et al. 1504 (MO, NY); Top of Serranía del Darien ca. due east of Unguia, Gentry et al. 16790 (MO, NY); Top of Serranía del Darien exactly on the frontier with Panama, N.E. of Cerro Mali, Gentry et al. 16982 (MO, NY); North ridge of Alto de Buey east-southwest of El Valle, Gentry and Fallen 17381 (NY); 7 km W. of Tutenendo on road to Quibdo, Gentry and Fallen 17591 (MO, NY). El Valle: Cordillera Occidental, vertiente occidental, Hoya del río Digua lado izquierdo, Piedra de Moler, Cuatrecasas 14983, 15100 (NY); Costa del Pacífico, río Yurumanguí, El Aguacate, Cuatrecasas 16146 (NY); Río Calima (región del Chocó), entre Herradura de Ordónez y Peña de Campotriste, Cuatrecasas 16633, 16689 (NY); Costa del Pacífico, río Cajambre, Barco, Cuatrecasas 16950, 17247 (NY); Punta Magdalena, Haught 5589 (NY). Gorgona Island: “Sr. George” Expedition 705 (NY). Nariño: Barbacoas, Alston 8490 (NY).

ECUADOR. Esmeraldas: San José km 321 along railroad from Ibarra to San Lorenzo, Boom 1325 (MO, NY); The Mache-Chindul Ecological Reserve, Bilsa Biological Station, Mache mountains 35 km W of Quinindé, collected on Ramon Loor’s property, Clark 3088 (NY); Bilsa Biological Station, Reserva Ecologica Mache-Chindul 40 km NW of Quinindé, Loma de los Guerrilleros, Permanent plot 2, Clark 4054 (MO, NY); Territorio Indígena Awá, Mataje village, Neil et al. 12498 (MO, NY); Reserva Biologica Bilsa, sendero Amarillo, Stern and Tepe 392 (NY). Pichincha: Carretera Quito-Puerto Quito km 113, Betancourt 139 (NY); Reserva de ENDESA km 113 along Quito-Pto. Quito rd, Luteyn and Borchsenius 13343 (NY).

VENEZUELA (fide Schnell). Zulia: San José de los Altos, Sierra Perijá, Delascio and Benkowsky 2952 (US).

Conostegia muriculata Almeda

Fig. 93

Conostegia muriculata Almeda, Proc. Calif. Acad. Sci. 46: 330. 1990. Type: Panama. Bocas del Toro: above Chiriqui Grande 10 road-miles from continental divide and 2 mi along road to E (0855'N 8210'W, 300 m), 6 August 1988, G. McPherson 12836 (holotype: CAS!, isotypes: CR!, DUKE, MO!, PMA, US!).

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 differentiated; 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.

Figure 93.

Conostegia muriculata. A Inflorescence B Lateral view of flower C Frontal view of flower. Photographs by Laurencio Martínez.

Distribution

(Fig. 94). Endemic to the Caribbean slope of south eastern Costa Rica and northern Panama, 40–1200 m in elevation.

Figure 94.

Distribution of Conostegia muriculata.

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) mentions the fact that several specimen labels point to an understory habitat and suggests it is probably a shade tolerant species.

Specimens examined

COSTA RICA. Limón: Sixaola, San Miguel, Finca Albergue ASACODE, Quesada 356 (CR, NY).

PANAMA. Bocas del Toro: above Chiriquí Grande on a side road about 10 road miles below the Continental Divide about 2.5 miles east on that road, Almeda et al. 6328 (CAS, CR, MO, NY). Veraguas: Distrito de Santa Fé, Serranía de Tute, Galdames et al. 3122 (MO, NY).

Conostegia oerstediana O. Berg ex Triana

Fig. 95

Conostegia oerstediana O. Berg ex Triana, Trans. Linn. Soc. 28: 98. 1872. Type: Costa Rica. Naranjo, no date, A. Oersted 11 (holotype: C; isotypes: BR!, K!, GH).

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 undifferentiated; 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 purple-black. Seeds 0.6–0.8 mm, pyramidal, the testa smooth.

Figure 95.

Conostegia oerstediana. A Leaf abaxial surface B Flower at anthesis showing many stamens stuck below the stigma C Lateral view of a flower showing bent style D Infructescence E Longitudinal section of flower bud F View of pickled flower from above G Lateral view of pickled flower with petals removed H Longitudinal section of a flower at anthesis I Petal J Stamen K Style A and D from specimen vouchered R. Kriebel 5633 B, C, E–K from specimen vouchered R. Kriebel 5627.

Distribution

(Fig. 96). Nicaragua to west Panama where it occurs in the mountains from (550)1000–2400 m elevation.

Figure 96.

Distribution of Conostegia oerstediana.

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. The 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. The latter name was considered a synonym of C. oerstediana. See the discussion under C. bigibbosa for the differences 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, 5845-INB, MO). These 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).

Specimens examined

NICARAGUA. Granada: in forest on Mombacho Volcano, Williams 20009, 20039 (MO, NY).

COSTA RICA. Alajuela: Los Angeles de Heredia, Brenes 14646 (CR, NY); Los Angeles de San Ramón, Brenes 168, 3820, 6093, 6711, 13630 (CR, NY); El Socorro de San Ramón (Haut de la Palma), Brenes 5242 (CR, NY); San Miguel Arriba de Grecia, Finca Arnold Haehner, Kriebel 5408 (INB, NY); Zarcero, Upper edge tropical zone, Pacific watershed edge of woodland, Smith 7 (NY); near Tapezco River, Cordillera Central, about 10 km north of Zarcero, Williams et al. 28927 (NY). Cartago: Paraíso, Parque Nacional Tapantí Macizo de La Muerte, a orilla de la calle el cruce de Río Humo al tunnel, Acosta and Ramírez 572 (INB, MO, NY); a 20 metros del río Grande de Orosi, Tapantí, Jiménez 1596 (CR, NY); Paraíso, Parque Nacional Tapantí Macizo de La Muerte, Alrededores de la estación, Kriebel 5338 (INB, NY); Woods beside Río Sombrero, 1.5 km S. of Muneco, Lent 1904 (NY); P. N. Tapantí, 500 meters after Rangers Station, Vargas and Villalobos 3827 (INB, MO, NY). Heredia: Collected along the slopes of Cerro Zurquí above the Río Para Blanco, Taylor 17631 (NY). Puntarenas: Cordillera de Tilarán, Road to San Luís about 3 km downslope from the Monteverde Cloud Forest Reserve Station, Almeda and Daniel 7179 (CAS, NY); Monteverde San Luis Cerro Chomogo, Kriebel et al. 5515 (INB, NY); Monteverde property of John Campbell, Lumer 1013 (NY). San José: Vicinity of Altos Tablazo about 10 km. W of Tablon and SE of Higuito, Almeda et al. 2841 (CR, NY); Old cart road to Limon, 5.6 km N of San Jeronimo, near La Palma, Hill et al. 17791 (CR); Acosta, Salvaje, Kriebel 5416 (INB, NY); Alto La Palma, Lumer 1341 (NY); La Palma, Tonduz 7427 (NY); Heavily pastured slopes between Volcán Barba and Irazú about 5 miles northeast of San Vicente, Wilbur and Stone 9666 (CR, NY); in mountains near Tarbaca about 15 kms south of San José, Williams 19466 (NY).

PANAMA. Chiriquí: Vicinity of Gualaca ca 8 mi from Planes de Hornito, La Fortuna on road to damsite, Antonio 5157 (MO, NY); Fortuna Dam region, along Quebrada Arena near continental divide, McPherson 8723 (NY); above Boquete near Parque Nacional Volcan Barú, along Río Caldera in Alto Chiquero, Penneys and Olmos 1738 (NY); Distrito de Bugaba, Santa Clara, Hartmann’s finca, van der Werff and Herrera 7091 (MO, NY).

Conostegia pittieri Cogn. Ex T. Durand

Fig. 97

Conostegia pittieri Cogn. Ex T. Durand, Bull. Soc. Roy. Bot. Belg. 27: 176. 1888. Type: Costa Rica. Alto del Roble, Massif du Barba, 1800–2000 m, 6 July 1888, H. Pittier 212 (holotype: BR; isotypes: F!, US!).

Conostegia donnell-smithii Cogn., DC. Monog. Phan. 7: 700. 1891. Type: Costa Rica. Cartago, 4000 ft., 1888, J. Cooper 327 (holotype BR; isotypes BR, F, GH!, NY!, US!). Schnell (1996) notes: Some sheets of this number have been distributed by John Donnell Smith under his number 5471, but since the latter was also used for Cooper 334 (US) care must be exercised in determining isotypes. The Donnell Smith number at least on the GH specimen is 5741 and not 5471.

Conostegia pittieri Cogn. ex Durand var. brevifolia Cogn., DC. Monog. Phan. 7: 704. 1891. Type: Costa Rica. Rio Segundo, Massif du Barba, 2000 m, Tonduz 1732 (lectotype BM!, designated here; isolectotypes BR, M!, US!). Other syntypes: Costa Rica. Rancho Flores, versant S du Massif du Barba, 2050 m, Costa Rica, Pittier 290 (BR, US!). The 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).

Description

Trees 2–18 m tall with tetragonal stems when young which become terete with age and glabrous; the nodal line present. Leaves of a pair equal to somewhat unequal in length. Petioles 0.5–4.3 cm. Leaf blades 3.5–18.5 × 1.6–6.1 cm, 3–5 nerved or generally 3–5 plinerved, with the innermost diverging from the mid vein just above the blade base in opposite or alternate fashion, elliptic to ovate, the base acute to decurrent on the petiole, the apex abruptly acute to acuminate, the margin entire to undulate-denticulate. Inflorescence a terminal panicle 4–12.1 cm long branching above the base, bracts and bracteoles to 3 mm, linear, deciduous. Pedicels 4–11 mm long. Flowers (5-) 6–8 (-10) merous, calyptrate, floral buds 7–14 × 2.75–6.75 mm, pyriform or ellipsoid, the base rounded, the apex apiculate, slightly constricted in the middle; the hypanthium 3.25–3.75 × 3.75–4.25 mm, glabrous and smooth. Petals 8–14 × 5–9 mm, white, obovate, rotate, rounded-truncate to emarginate, glabrous on both surfaces, rotate, entire, persistent after the stamens and style have fallen. Stamens 14–23, 7–9 mm, opposing the style resulting in zygomorphy, the filament 5.5–6.5 mm, white, anthers 2.5–3.5 × 0.75–1.25 mm, linear and slightly recurved, the base sagittate, yellow, not conspicuously compressed, the pore 0.2–0.26 mm, terminal. Ovary 7–12 locular, inferior. Style 6–6.5 mm, bending to one side of the flower, opposite the stamens, lacking a basal collar, vertical distance between the anthers and the stigma absent, horizontal distance 0.5–3 mm; stigma peltate, consisting of 7–11(-12) laterally compressed lobes, 3–3.5 mm wide. Berry ca. 1 × 1 cm, purple black. Seeds 0.5–0.7 mm, ovoid, the testa smooth.

Figure 97.

Conostegia pittieri. A Leaf abaxial B Flower buds C Flower at anthesis D Immature fruit E Longitudinal section of a floral bud F Pickled flower at anthesis G Longitudinal section of a flower at anthesis H Petal I Stamen J Style. Photos of specimen vouchered R. Kriebel 5400.

Distribution

(Fig. 98). In cloud forests from southern Nicaragua through central and northern Costa Rica to northwestern Panama, (500-)1000–2400 m.

Figure 98.

Distribution of Conostegia pittieri.

Conostegia pittieri is one of the few species of Conostegia that retains its petals once the stamens and style have fallen. The functional significance of petal retention, if any, might have to do with continuing to make inflorescences attractive (Bertin 1982).The similar species Conostegia chiriquensis also retains the petals and the style after the stamens have fallen. See the discussion under the latter species for differences 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.).

Specimens examined

NICARAGUA (fide Schnell). Rivas: Volcán Maderas, Isla Ometepe, Moreno 19760 (MO).

COSTA RICA. Alajuela: Upala, Bijagua, P. N. Tenorio, Aguilar 6452 (NY); Palmira de Naranjo, Brenes 3519 (CR, NY); Cerros de San Antonio de San Ramón, Brenes 5664 (NY); La Palma de San Ramón, Brenes 5528, 6787, 6807 (NY); Posada Volcán Póas, Viento Fresco, Vara Blanca, Kriebel and Solano 3266 (INB, NY); Alto Palomo 9 km W of top of Poás Volcano, Lent 1655 (CR, NY); Remnant cloud forest on mountains of Cordillera Central about 2 kms east of Zarcero, Molina et al. 17060 (NY); Cordillera Central near Palmira about 5 kms east of Zarcero, Molina et al. 17769 (NY); Fraijanes Volcán Poás, on road from San Pedro de Poás to Vara Blanca, Schnell 1084 (CR, NY); Palmira (Canton de Alfaro Ruiz), Smith 40 (NY). Heredia: Wet secondary forest about 15 m high with many epiphytes with frequent wind and rain from the Caribbean, Rio Vueltas (Upper Río Patria), eastern slope of Volcan Barba near Continental Divide, near Finca Montecristo, Barringer and Christenson 3412 (F, NY); Sacramento, Finca Ingrid Steinvorth, Kriebel 5400 (INB, NY); 5 km down a jackknife turn to the east off of Highway 9, 3.3 km north of Vara Blanca, Luteyn 685 (MO, NY); Vara Blanca de Sarapiquí north slope of Central Cordillera between Póas and Barba volcanoes, Skutch 3415, 3563 (MO, NY); entre Porosatí y Sacramento, Solano et al. 2696 (INB, NY); Collected along the slopes of Cerro Zurquí above Río Para Blanco, Taylor 17481 (NY); Monte de la Cruz, above Finca Monte Cristo, Todzia and Moran 2045 (CR, NY); Forets du Barba, Tonduz 1947 (NY); woods on steep bank of stream about 7 miles northeast of Barba on Route 114, on flank of Volcán Barba, Wilbur and Teeri 13681 (MO, NY). Puntarenas: near the Continental Divide about 2 to 5 km east and southeast of Monteverde, Burger and Gentry 8728 (F, NY). San José: Vázquez de Coronado, P. N. Braulio Carrillo, Cuenca del Sarapiquí, Santa Elena, Acosta and Ramírez 442 (INB, MO, NY). PANAMA. Chiriquí: Gualaca-Chiriqui Grande road, 4.5 mi N of middle of bridge over Fortuna Lake, just S of Bocas del Toro border, Croat 66710 (MO); Fortuna Dam, trail from highway near forestry nursery down to Rio Hornito, McPherson 12498 (MO); Cerro Colorado 50 km N of San Felix on continental divide, Mori and Dressler 7803 (MO).

Conostegia procera (Swartz) D. Don ex DC

Conostegia procera (Swartz) D. Don ex DC., Prodr. 3: 174. 1828. Melastoma procera Swartz, Prodr. Veg. Ind. Occ. 68. 1788. Type Jamaica. no date, O. Swartz s.n. (lectotype: S!, designated here; isolectotype: BM!).

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.

Figure 99.

Distribution of Conostegia procera.

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. The 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.

Specimens examined

JAMAICA. St. Andrew: Southwest slope of Mt. Horeb above Hardwar Gap, Proctor 10321 (NY). St. Thomas: Hillside, Mansfield, Britton 3607 (NY); Mountain trail between House Hill and Cuna Cuna Gap, Maxon 8949 (NY).

Conostegia pyxidata Proctor

Conostegia pyxidata Proctor, Bull. Inst. Jam. Sci. Series 16: 38. Pl. 15, p. 39. 1967. Type: Jamaica. Portland Parish: East slope of John Crow Mts, 1.5–2.5 miles southwest of Ecclesdown, 1500–2500 ft, 11 August 1956, G. R. Proctor 10468 (lectotype: NY!, designated here; isolectotypes: IJ, LIL!).

Conostegia subprocera Proctor, Bull. Inst. Jam. Sci. ser. 16: 38. Pl. 16, p.40. 1967. Type: Jamaica. Portland Parish: east slope of John Crow Mts, c. 1-1.5 miles southwest of Ecclesdown, 1500 ft., 6 August 1954, G. R. Proctor 9229 (holotype: IJ).

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.

Figure 100.

Distribution of Conostegia pyxidata.

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 differences in a small area; 2) the disjunct range of C. balbisiana; and 3) differences 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 different stamen morphology. Schnell (1996) argued that these morphological and phenological differences 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. The type of Conostegia subprocera was not studied but Schnell (1996) cited this name as a synonym of C. pyxidata.

Specimens examined

JAMAICA. Portland: east slope of the John Crow Mts 1.5–2 miles southwest of Ecclesdown, Proctor 9814 (NY); east foothills of John Crow Mts Along Ecclesdown road ca. 2 miles N of Ecclesdown, Skean and Slantis 1864 (MO, NY).

Conostegia rhodopetala Donn. Smith

Fig. 101

Conostegia rhodopetala Donn. Smith, Bot. Gaz. 42: 295. 1906. Type: Costa Rica. San José: La Palma de San José, 1500–1700 m, 22 May 1898, Tonduz 12347 (lectotype: NY!, designated here; isolectotypes: BR!, F!, US!). Other syntypes: COSTA RICA. San José: La Palma de San José, 1500–1700 m, A. Tonduz 9702 (BR!(2), NY!, US!), H. Pittier 10169 (BR!(2), M!, US!); Heredia: San Isidro, Rio de Las Lajas,1500 m, H. Pittier 14022 (F!, NY!, US!).

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, campanulate 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. -1 – -0.5 mm, horizontal distance ca. 0–1mm, stigma slightly expanded, 0.75–1 mm wide. Berry 6–7 × 6–7 mm, purple-black. Seeds 0.3–0.55 mm, ovoid to pyramidal, the testa smooth.

Figure 101.

Conostegia rhodopetala. A Inflorescence B Infrutescence C Longitudinal section of a flower bud D Pickled flower collected at late anthesis E Longitudinal section of a flower at late anthesis with petals removed F Petal G Stamen H Style. Photos A, C–H of specimen vouchered R. Kriebel 5542 B from specimen vouchered R. Kriebel 5462.

Distribution

(Fig. 102). Cloud forests in Costa Rica reaching western Panama, 700–1900 m in elevation.

Figure 102.

Distribution of Conostegia rhodopetala.

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.

Specimens examined

COSTA RICA. Alajuela: San Ramón, Reserva Monteverde, Sendero El Camino, Haber and Zuchowski 12398 (INB, NY); San Carlos, La Tigra, entre la Fila divisoria de agua (Rincón de Cedral) y cerro Bekom, Herrera et al. 8999 (CR, MO, NY); vicinity of La Palma, Maxon 445 (NY); Zapote de San Carlos, Smith 381 (NY); Guadalupe de Zarcero, cantón de Alfaro Ruiz, Smith 702 (NY); P. N. Juan Castro Blanco. Cuenca alta del río La Vieja. Colectando a orillas del río y en la fila, Solano et al. 2667 (INB, NY). Cartago: Jiménez, Pejibaye, Turrialba, Refugio El Copal, Sendero Galbula, Kriebel et al. 5462 (INB, NY); Volcan Turrialba on road from La Trinidad to Lechería La Central, Schnell 1076 (NY). Heredia: Vara Blanca on road to Cariblanco, Schnell 1081 (MO, NY); Vara Blanca de Sarapiquí north slope of Central Cordillera, Skutch 3224 (NY). Puntarenas: Coto Brus, Sendero Las Tablas camino a Cotoncito, Kriebel and Solano 3176 (INB, NY). San José: Moravia, P.N. Braulio Carrillo, Cuenca del Sarapiquí, Camino al Bajo de la Hondura, Acosta and Ramírez 389 (INB, NY); Woods high above Río Cascajal 3 km N.E. of Cascajal, Lent 2177 (NY); Vicinity of El General, Skutch 2989 (MO, NY).

PANAMA. Chiriquí: Sendero Rio Hornito, Kriebel and Burke 5760 (PMA, NY).

Conostegia rufescens Naudin

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).

Conostegia formosa Macfad., Fl. Jamaica 2: 70. 1850; nom. inval.

Conostegia puberula Cogn., DC Monog. Phan. 7: 703. 1891. Type: NICARAGUA. Chontales: no date, B. Seemann 30 (lectotype: BM!, designated here; isolectotypes: BR, K!, LE, W). Additional syntype: Costa Rica. Naranjo, Wendland s.n. (GOET).

Conostegia hotteana Urban & Ekman, Ark. Bot. 22a 17: 29. 1929. Type: Haiti. Massif de la Hotte, western group, Dame-Marie, Montagniac, 500 m, Ekman 10324 (holotype: S!; isotypes: A!, C, IJ, NY!, US!).

Conostegia affinis Urban, Arkiv. Bot. 22a 17: 29. 1929. Type: Haiti. Massif de la Hotte, western group, Jeremie, near La Source Chaude, E. Ekman 10264 (holotype: S!).

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.

Figure 103.

Conostegia rufescens. A Habit and morphotype with revolute leaf base B Leaf abaxial surface C Flower at anthesis D Berry E Longitudinal section of a flower bud F Pickled flower at anthesis G Longitudinal section of a flower at anthesis H Petal I Stamen J Style. Photograph A from voucher R. Kriebel 5635 B, C from R. Kriebel 5314 D by E. Salicetti E–J from R. Kriebel 5687.

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.

Figure 104.

Distribution of Conostegia rufescens.

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. This 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. The latter two species differ from C. rufescens in their exserted styles. Schnell (1996) noted limited local flowering seasons and differentiation 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).

JAMAICA. Portland: Vicinity of Moody’s Gap, Britton 3392 (NY). Westmoreland: Copse Mt. Woods c. 1 mile southwest of Rat Trap, Proctor 21476 (NY).

PUERTO RICO. Luquillo: Northeastern Luquillo Mts. Rd. 191, Woodbury s.n. (NY).

NICARAGUA (fide Schnell). Jinotega: Cordillera Isabelia, Macizos de Peñas Blancas, Neill 7180 (GH). Matagalpa: N & NW sides of Cerro Musun above Salto Grande de Q. Negra, Río Bilampi, Neill 1797 (MO). Zelaya: 20 km O de Awas Tingi, S. de Rio Wawa, Little and Delvis 23354 (US).

COSTA RICA. Alajuela: La Palma de San Ramón, Brenes 4401, 5688, 6283, 16203 (CR, NY); Cerro de “La Muralla” de San Ramón, Brenes 5688 (NY); San Carlos, Fortuna, R. B. Arenal, Mundo Aventura, Rodríguez 8606, 8830 (INB, NY); Villa Quesada, Cantón de San Carlos, Smith 2556 (NY); 3.5 km west of Fortuna, 2.5 km northeast of New Volcan Arenal along sloping base, Taylor and Taylor 11625 (NY); La Palma, Tonduz 12434 (NY). Cartago: P.N. Tapantí, sendero Oropéndula, ca. 10 mfrom river, Penneys and Blanco 1792 (FLAS, NY). Limón: Quebrada González, sendero Las Palmas, Kriebel 1422, 5314 (INB, NY); 6 miles inland from mouth of Estrella River, Stork 4614 (NY); Talamanca, Fila Carbón, Buena Vista, Finca Corredor Biológico, Valverde 13 (CR, NY). Puntarenas: Forested slopes east of Las Cruces and 5 to 6 km south of San Vito on and around the property of Mr. Robt. Wilson, Burger and Matta 4470 (F, NY); R.B. Monteverde, Cordillera de Tilarán, Pacific slope wet forest road to Continental divide, Haber 11514 (INB, MO, NY). San José: Tarrazú, San Lorenzo, Estribaciones sureste de cerro Toro, Estrada et al. 684 (CR, NY); Límite del P. N. Braulio Carrillo, 2 Km después del peaje hacia Guápiles, Kriebel 1938 (INB, NY); Pérez Zeledón, Río Nuevo, El Brujo, 0.5 km NO del Andaribel del Brujo, Rodríguez et al. 6974 (INB, CR, NY); San Isidro de Dota, albergue Tinamú, Santamaría and Morales 813 (INB, NY).

PANAMA. Bocas del Toro: Cricamola Valley, Cooper 486 (NY); Buena Vista Camp on Chiriquí trail, Cooper 581 (NY); Along road towards Chiriquí Grande along trail leaving road, McPherson 12553 (MO, NY). Coclé: Vicinity of La Mesa, Croat 13325 (F, MO, NY); La Mesa, Gentry 5636 (MO, NY); Vicinity of La Mesa above El Valle, Gentry 7427 (MO, NY). Veraguas: Along road on Pacific slope 1-3 km above Escuela Agricola Alto Piedra, Croat 25996 (NY); Area between La Junta and Limón, 5 hours walk north of Alto Calvario, Folsom 5877 (MO, NY). Colón: Around Dos Bocas, Río Fató valley, Pittier 4218 (NY).

COLOMBIA. Nariño: Espriella, Tumaco, Romero-Castañeda 2801 (NY). Pichincha: 20 km W of Santo Domingo de los Colorados, Cazalet 5110 (NY).

ECUADOR. Esmeraldas: Parroquia Mataje, Reserva Etnica Awá, Centro Mataje, Aulestia et al. 442 (MO, NY, QCNE); Parroquia de Concepción, bank of Río Santiago, above Playa Rica, Mexia 8471 (NY); Eloy Alfaro, Reserva Ecológica Cotacachi-Cayapas, Parroquia Luis Vargas Torres, Río Santiago, Estero Angostura, Tirado et al. 647 (MO, NY, QCNE).

Conostegia setifera Standl.

Fig. 105

Conostegia setifera Standl., Field Mus. Nat. Hist., Bot. ser. 18: 805. 1938. Type: Costa Rica. Alajuela: Camino de la Finca Johanson, Los Angeles de San Ramón, 15 March 1928, A. Brenes 6041 (holotype: F!, isotypes: CR!, NY!).

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. 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.

Figure 105.

Conostegia setifera. A Inflorescence B Lenticellate node C Flower at anthesis. Photos by E. Salicetti.

Distribution

(Fig. 106). Ranging from Nicaragua to Costa Rica, 0–1350 m elevation.

Figure 106.

Distribution of Conostegia setifera.

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 differentiate 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. This 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. This species is usually encountered in the forest understory and Schnell (1996) noted that within this habitat, only well lit branches produce flowers.

Specimens examined

NICARAGUA (fide Schnell). Río San Juan: near Caño Chontaleño, 20 km NE of El Castillo, Neil and Vincelli 3519 (MO); 1 km E de Río Sábalos, Moreno 23190 (MO); boca del Sábalo, camino a Buena Vista, Moreno 25625 (MO).

COSTA RICA. Alajuela: Los Angeles de San Ramón, Brenes 13582 (CR, NY); Cataratas de San Ramón, Brenes 13652 (CR, NY); Eastern slopes of Volcán Miravalles west of Bijagua near the Río Zapote, Burger et al. 11709 (CR, NY); Guatuso, P. N. Volcán Tenorio, Cuenca del Río Frío, El Pilón, Chaves and Muñoz 436 (CR, INB, NY); San Ramón, Villa Blanca, González 680 (INB, NY); N side of Volcan Arenal, Lent et al. 3311 (NY); along trail from macadamia village SE around base of Cerro Chato to Catarata de Fortuna, Smith 10900 (CR, NY). Heredia: Finca La Selva, the OTS Field Station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, Holdridge trail 1800 m line, Folsom 9139 (MO, NY); along Río Peje about .5 km SW of back end of Vargas property, aprox. in the area where an imaginary line drawn between Magsasay (colonia penal) and Puerto Viejo de Sarapiquí would cross the Río Peje, Hammel 11216 (MO, NY). Limón: Reserva Biológica Hitoy Cerere, rainforest along the sloping banks of Río Cerere from the Reserve Station to the big waterfall, Almeda et al. 6841 (CR, MO, NY); Hacienda Tapezco-Hda. La Suerte 29 air km W of Tortuguero, Davidson 6968 (NY). San José: Sendero La Montura, entre Estación Quebrada González y Estación Zurquí, Rodríguez et al., 5799 (INB, NY).

Conostegia setosa Triana

Fig. 107

Conostegia setosa Triana, Trans. Linn. Soc. London 28: 99. 1872. Type: Colombia. Chocó: Cordillera Occidental, between Tuquerres and Barbacoas, 1851–1857, J. Triana 3940 (holotype: BM!, isotypes: BR!, K!, P (fide Almeda in Schnell 1996), W).

Cryptophysa setosa Standl. & J. F. Macbr., Field Mus. Publ. Bot. 4: 244. 1929. Type: Panama. Bocas del Toro: Buena Vista, Almirante, January-March 1928, G. Cooper 219 (holotype: F!, isotype: NY!). Conostegia hirsuta Gleason, Phytologia 3: 359. 1959. Non Conostegia setosa Triana.

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 surfaces. 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.

Figure 107.

Conostegia setosa. A Habit B Inflorescence and leaf base showing formicaria C Close up of flowers D Close up of morphotype with red indument E Flower buds and maturing fruit in the middle F Longitudinal section of a flower bud G Pickled flowers at anthesis H Longitudinal section of a flower at anthesis I Petal J Stamen K Style. Photos A, D–K of specimen voucher R. Kriebel 5731 B–C from R. Kriebel s. n.

Distribution

(Fig. 108). From Nicaragua through Costa Rica and Panama to Colombia and Ecuador, 0–1400 m elevation.

Figure 108.

Distribution of Conostegia setosa.

This 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. The latter taxon differs 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. The 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. The 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) found a lot of variation in the ant inhabitants of C. setosa.

Specimens examined

COSTA RICA. Alajuela: San Carlos, Boca Tapada, Laguna de Lagarto Lodge, Solano 894, 1448 (INB, NY). Cartago: Jiménez, Pejibaye, Reserva El Copal, Kriebel 2474 (INB); Jiménez, Pejibaye, Refugio de Vida Silvestre La Marta, Kriebel 4484 (INB); Heredia: La Selva, Sendero Holdridge a Parcelas, Kriebel 3565 (INB). Limón: North end of Tortuguero National Park and near the Boca de las Lagunas de Tortuguero, Burger and Antonio 11274 (CR, F, NY); Between Cerro Jacrón and Cerro Bitárkara, Trail between “Sitio Rangalle and Cuen”, Hazlett 5125 (NY); Pococí, R. B. Bosque Lluvioso, Sendero derecho, Vargas et al. 3539 (INB, NY).

PANAMA. Coclé: 7 km from Llano Grande on road to Coclesito near Continental Divide, Antonio 1365 (NY); Vicinity of La Mesa, N of El Valle de Antón, along steep slopes above water reservoirs, ca. 1 km W of road between Finca Mandarinas and Finca Furlong, Croat 67169 (MO, NY); Parque Nacional Omar Torrijos, Sendero Cuerpo de Paz, Penneys and Blanco 1760 (FLAS, NY). Darién: 0-2 mi. E of Tres Bocas along shortest headwater of Río Coasi, Kirkbride and Duke 1170 (MO, NY); Mannene to the mouth of Río Coasi, Kirkbride and Bristan 1492 (MO, NY); Ensenada Guayabo, between Punta Guayabo Grande and Punta Guayabo Chiquita, Stern and Chambers 178 (NY). Panamá: On Atlantic side Llano-Carti Road, 12 miles from Pan-American Highway, Antonio 3308 (MO, NY); Road to Carti (San Blas), 19 km north of El Llano, Busey 894 (MO, NY); Near summit of Cerro Camapan, Croat 22814 (MO, NY); Cerro Azul, D’Arcy and D’Arcy 6233 (MO, NY); Cerro Jefe, Dwyer and Gentry 10257 (MO, NY); On trails radiating from end of road which passes Campana water tank near Cerro Campana, Kirkbride and Hayden 311 (MO, NY); along El Llano Carti-Tupile road, 12 mi above Pan-Am Hwy, Liesner 1132 (MO, NY); Forest 10 miles north of Highway 1 towards Cerro Jefe, Luteyn 1323 (MO, NY); Cerro Campana, Sendero La Cruz, near the summit, Penneys and Blanco 1678 (FLAS, NY).

COLOMBIA. Chocó: North ridge of Alto de Buey, above Dos Bocas del Río Mutatá, tributary of Río El Valle, ESE of El Valle, Gentry and Fallen 17413 (MO, NY); Nuquí, Alto de Buey, von Sneidern 7 (NY). El Valle: Cordillera Occidental, vertiente occidental, hoya del río Anchicayá, lado derecho, bosques entre Pavas y Miramar, Cuatrecasas 14380 (NY); Costa del Pacífico, río Cajambre, Silva, Cuatrecasas 17555 (NY).

ECUADOR. Esmeraldas: Eloy Alfaro, Reserva Ecológica Cotacachi-Cayapas, Parroquia Luis Vargas Torres, Río Santiago, estero Pote, Tirado et al. 529 (MO, NY). Pichincha: Carretera Quito-Puerto Quito km 113, Betancourt 82, 111, 166, 169, 218 (NY); Carretera Quito-Puerto Quito km 113, 10 km al Norte de la carretera principal, Freire 1060 (NY); Reserva Forestal ENDESA, Río Silanche: “Corporación Forestal Juan Manuel Durini”, km 113 de la carretera Quito-Pto. Quito, faldas occidentales a 10 km al Norte de la carretera principal, Jaramillo 5202, 6412 (NY).

VENEZUELA (fide Schnell). Zulia: Caño Helena, Sierra Perijá, Delascio and Benkowsky 3191 (US).

Conostegia superba Naudin

Fig. 109

Conostegia superba Naudin, Ann. Sci. Nat. Bot. ser. 3. 16: 108. 1850. Type: Jamaica. Wilson s.n. (holotype: P, isotype: K!; Schnell (1996) cites a fragment in F).

Conostegia macrophylla Naudin, Ann. Sc. Nat. ser 3 16: 112. 1850. Type: Mexico. “In montibus Mex. prope Oaxaca and Chinantla”, 700 m, April–November 1840, H. Galeotti 2941 (isotype: BR!).

Conostegia alternifolia Macfad., Fl. Jamaica 2: 71. 1850; nom. inval.

Conostegia clidemioides Wright ex Grisebach, Cat. PI. Cuba 98. 1866. Type: Cuba. La Perla, eastern Cuba, 1861, C. Wright 2503 (holotype: GOET!; isotypes: BM!, BR, GH!, K!, LE, MO!, S!).

Conostegia poeppigii Cogn., Mart. Fl. Bras. 14(4): 211. 1886. Type: Peru. Provo Maynas, Poeppig s. n. (lectotype: LE, designated here; isolectotype: ?BR). Other syntype: Colares, Provo Para, Brazil, June 1832, Poeppig s. n. (W). “The Brazilian syntype is sterile and cannot be identified with certainty” Schnell (1996).

Conostegia purpusii Brandegee, Univ. Calif. Publ. Bot. 6: 57. 1914. Type: Mexico. Chiapas: Finca Mexiquita, July 1913, C. Purpus 6784 (holotype: UC!; isotypes: A, BM!, CAS!, F!, GH, MO!, NY!, US). “This collection is mixed with a species of Miconia in many herbaria.” Schnell (1996)

Conostegia pentaneura Standl., Field Mus. Publ. Bot. 8: 146. 1930. Type: Honduras. Lancetilla valley near Tela, 100 m, 8 August 1929, F. Salvoza 875 (holotype: A!, photograph: GH).

Miconia bailloni Gomez, Anal. Hist. Nat. Madrid 23: 69. 1894. Nom. inval. (no specimen cited)

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, sul-cate, 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 constricted 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, purple-black. Seeds 0.4–0.6 mm long, narrowly pyramidal, the testa smooth.

Figure 109.

Conostegia superba. A Leaf abaxial surface B Inflorescence C Close up of the flower D Infructescence E Longitudinal section of a flower bud F Pickled flower at anthesis G Longitudinal section of a flower at anthesis wit petals and most stamens removed H Petal I Stamen J Style. Photos of A–C and E–J from specimen vouchered R. Kriebel 5582 D taken by Reinaldo Aguilar.

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.

Figure 110.

Distribution of Conostegia superba.

The 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. The 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) noted that some isotypes of Conostegia purpusii are mixed with a species of Miconia in many herbaria.

Specimens examined

CUBA. Oriente: El Yunque, Ekman 3970 (NY); La Prenda, Hioram and Maurel 4776 (NY).

DOMINICAN REPUBLIC. Barahona: Sierra del Bahoruco, Municipio Paraíso La Víbora, Clase, Montilla and Schuber 4383 (NY); Fuertes 969 (NY). Bonao: subida al Casabito, Liogier 30328 (NY). Monseñor Nouel: Road up to Alto Casabito, ca. 8 km W of jct. with Highway Duarte on road from Bonao to Constanza, Judd and McDowell 6523 (NY); Cordillera Central on road to Alto Casabito ca. 2.8 km W on highway from Autopista Duarte N of Bonao to El Rio and Constanza, Judd et al. 8218 (NY); Cordillera Central, “Zumbador”, 1.5 km al sudeste de Juan Aldian, en la confluencia de los rios Zumbador and Calle Estrecha, Zanoni et al. 31550 (NY).

PUERTO RICO. Cordillera Central: Toro Negro Forest, in Vereda del Bolo, Vives 138 (NY).

JAMAICA. Portland: Upper Swift River Study Site of Ecological Survey, Blue Mt. Multipurpose Project near Mossman’s Peak, Bretting 61, 254 (NY); North side of Cuna Cuna Pass, Harris and Britton 10560 (NY); Foothills of the John Crow Mts. Along road between Fair Prospect and Hartford and Ecclesdown, ca. 3.9-4 mi SE of jct. with Rt. A-4 (coastal road), Judd 5327 (NY). St. Thomas: Banks Devil’s River, Britton 3582 (NY); near Milepost 18 southeast of Bowden Pen. upper valley of the Río Grande, Proctor 26603 (NY).

MEXICO (fide Schnell). Chiapas: crest of ridge 3 km E of Francisco Madero, NE of Cintalapa, Breedlove 38701 (MO). Oaxaca: entre Puerto Eligio y Comaltepec, km 149 entre Tuxtepec a Oaxaca S. Juarez, Martinez Calderon 388 (GH, MICH, UC, US). Veracruz: Lado SE de Laguna Catemaco, arriba de Rio Cuetzalapan, Beaman 5165 (GH).

GUATEMALA. Izabal: Sierra Caral, camino de terracería cerca de la entrada a la Finca La Firmeza, desde Morales, Kriebel et al. 5582 (USCG, NY). Sololá: south-facing slopes of Volcán Atitlán above Finca Mocá, Steyermark 47891 (NY).

HONDURAS. Cortés: entre Agua Azul y Pito Solo Lago de Yojoa, Molina 7326 (NY).

NICARAGUA (fide Schnell). Rivas: Isla Ometepe, Volcán Maderas, Hacienda La Argentina, Robleto 845 (MO). Zelaya: near Bil Tingnia, 6 km NW of Bonanza, 150 m, Neill 4000 (M).

COSTA RICA. Puntarenas: about 5 km. west of Rincón de Osa, Osa Península, Burger and Gentry 9006 (CR, NY); Buenos Aires, Potrero Grande, Sabanas Helechales, bosques y potreros aledaños al camino, Santamaría et al. 4194 (NY). San José: about 1.8 km north of Platanillo on the road to Dominical, Almeda and Nakai 4123 (CR, MO, NY); Basin of El General, Skutch 4842 (CR, NY).

PANAMA. (fide Schnell). Chiriquí: Burica Peninsula, 4-9 mi S of Puerto Armuelles, Croat 22101 (MO).

COLOMBIA. Putumayo: Municipio Mocoa, corregimiento de San Antonio, vereda Alto Campucana, finca La Mariposa, Vertiente Amazónica de Colombia, Betancourt et al. 4957 (NY); Umbría, Klug 1909 (NY).

ECUADOR. Morona-Santiago: Centro Shuar Yukutais 8 km SW of Sucua, Andrade 571 (NY); small ravine ca. 7 km N of Limón, Moran et al. 7613 (NY). Napo: Puyo, in rastrojo, Asplund 18907 (NY); Mera in forest on shore of Río Pastaza, Asplund 19128 (NY); Hacienda San Antonio del Barón von Humboldt 2 km al NE de Mera, Baker et al. 5370 (MO, NY); Hacienda San Antonia von Humboldt 2 km al NE de Mera, Baker et al. 5501 (NY); Near El Topo along trail to La Gloria, Valley of the río Pastaza and adjacent uplands, Camp 2403 (NY); Loreto, Faldas del Volcán Sumaco, al oeste de Avila Viejo, Bloque 19 línea sísmica 8, Compania Triton, Freire and Cerda 136 (MO, NY); Chaco rastrojal, Harling 3888 (NY); Río Napo between Coca (Puerto Francisco de Orellana) and Armenia Vieja, Harling and Andersson 11978 (NY); Santa Rosa at Rio Napo, Lugo 173 (MO, NY); between Banios and Mera, Mexia 6967 (NY); ca. 50 km NE of Baeza Cascada de San Rafael along Río Quijos, Moran et al. 7561 (NY); Hacienda San Antonio del Baron von Humboldt, 2 km al NE de Mera, Neill et al. 5869 (MO, NY); Hda. San Francisco below Banos, Penland and Summers 279 (NY); Vicinity of Puyo, Eastern foothills of the Andes, Skutch 4518 (NY); carretera Hollin-Loreto-Coca, en las orillas del Río Hollin, Zak and Jaramillo 3140 (NY). Tungurahua: Rio Negro, Asplund 18366 (NY); Colonia Mexico 4 km de Topo, Lugo 648 (NY). Zamora-Chinchipe: Road La Saquea-Yacuambi 1 km N Chapintza, Harling and Anderson 23885 (NY).

PERU (fide Schnell). Cuzco: Mapitunari valley, 5-7 km from Hda. Luisiana and the Apurimac river, Cordillera Villacabamba, Madison 10066-7 (NA, US). Huanaco: Tingo Maria, Asplund 12992 (US).

VENEZUELA. Amazonas: Trail S from Cerro Neblina camp 5, Gentry and Stein 46530 (NY); Cerro Neblina Campamento 5 N base of Pico Cardenas, Gentry and Stein 46650 (NY); Departamento Río Negro, Cerro de La Neblina Camp V Valley north base of Pico Cardona, Liesner and Stannard 16893 (MO, NY).

Conostegia volcanalis Standl. & Steyermark

Fig. 111

Conostegia volcanalis Standl. & Steyermark, Field Mus. Nat. Hist., Bot. sere 23: 136. 1944. Type: Guatemala. Quetzaltenango: Damp forest, Chiquihuite, 1410 m, 8 March 1939, P. Standley 68152 (holotype: F!, isotypes: A!, NY!).

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 stalked-stellate 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.

Figure 111.

Conostegia volcanalis. A Leaf abaxial surface B Inflorescence with flower buds C Longitudinal section of a flower bud D Lateral view of a flower at late anthesis E Petal F Stamen. Photos from specimen vouchered R. Kriebel 5565.

Distribution

(Fig. 112). From Mexico through Guatemala to Honduras, 500–2200 m in elevation.

Figure 112.

Distribution of Conostegia volcanalis.

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”. These morphotypes were considered on the “borderline” of deserving species status (Schnell 1996). The three morphotypes differed in their distribution, habitat preference, indument density, and floral part size. The first morphotype is found in Guerrero and Jalisco and (Schnell 1996) noted that as C. jaliscana. These plants prefer streamsides in pine forests. The leaves are is more pubescent and have larger floral parts. These trees flower January through April. The second morphotype recognized by Schnell (1996) occurs in cloud forests of Chiapas, San Marcos and Quetzaltenango in Mexico. The plants from this morphotype are larger trees with less pubescent leaves and flowering November through March. This morphotype is reminiscent of C. icosandra and had been described as C. sphaerica Triana. The third race occurs in cloud forests of central Guatemala, Honduras, El Salvador and Nic