Printer Friendly

Palynological characters and their phylogenetic signal in Rubiaceae.

V. Discussion


Pollen of the "average" Rubiaceae species is medium sized (30-40 [micro]m), 3(-4)-zonocolporate with an ecto- and endocolpus, subspheroidal, circular in outline, with a tectum perforatum mostly lacking supratectal elements. From the data gathered for the present article (see also Table I), it is clear that there are many variations on this theme.

The use of molecular data and cladistic procedures in systematics has resulted in well-supported hypotheses on the phylogeny of the major groups of Rubiaceae. We use a summary cladogram (Fig. 58) that combines recent insights into the phylogeny of Rubiaceae to discuss the pollen variation in the family. The systematic value of pollen characters at the subfamily to specific level is illustrated below by several case studies.


1. Subfamily Level

The summary phylogenetic tree of Rubiaceae (Fig. 58) shows three major clades, which correspond to a certain extent with three of the four subfamilies recognized by Robbrecht (1988a): Rubioideae, Cinchonoideae s.str., and Ixoroideae s.1. The taxa of Robbrecht's fourth subfamily, Antirheoideae, are scattered over the tree. The molecular support for the monophyly of these three subfamilies is fairly convincing (see Andersson & Rova, 1999; Andreasen & Bremer, 2000; Bremer & Manen, 2000), but morphological synapomorphies are scarce. For members of the Rubioideae, the presence of raphides is characteristic, but the other two subfamilies are mainly supported by molecular data.

Likewise, pollen morphological data by themselves are not useful in characterizing the subfamilies. No single pollen morphological character or complex of shared pollen features unambiguously identifies one of the subfamilies of Rubiaceae. The different character states of all pollen characters are scattered over the three subfamilies (Figs. 59-61; Table I) and hence originated several times in evolution. Verdcourt (1958: 226) already stated that "pollen grain characters are therefore not suitable as the main character for subdividing the family." It may be noted, however, that the pollen morphology of subfamily Cinchonoideae s.str, is more or less uniform, featuring small, 3(-4)-colp(or)ate and spheroidal grains often lacking supratectal elements (exceptions to this pollen type are found in the PECC-complex and Guettardeae); Rubioideae and Ixoroideae s.1. pollen grains show greater variation. Better sampling of the Cinchonoideae s.str., however, might prove that pollen is also more variable than currently estimated.

It is noteworthy that the more derived taxa of the Rubioideae often possess supratectal elements, whereas supratectal elements are lacking in the "basal" taxa. A similar pattern is observed among the Ixoroideae s.1., though supratectal elements are less common in this family (Fig. 59). In the subfamily Cinchonoideae s.str., suprateetal elements are present only in the PECC-complex and a few species of the Hillieae, Rondeletieae, and Guettardeae.


It is of special interest that Coptosapelta and Luculia Sweet, two genera taking an isolated and often conflicting position with respect to the rest of the family, feature very different pollen grains. Pollen of Coptosapelta is highly derived, characterized by pororate grains having a wall without columellae (Verellen et al., 2004). Similar pollen is found in some Guettarda species from Cuba (subfamily Cinchonoideae s.str.; Sotolongo Molina et al., 2002). Pollen of Luculia, on the contrary, matches what is thought to be the primitive pollen type for Rubiaceae; that is, small to medium-sized, more or less spheroidal, 3(-4)-colporate pollen with endocolpi and a reticulate tectum lacking supratectal elements. If the basal position of the genus Coptosapelta is confirmed in future phylogenetic analyses, its derived pollen illustrates the families' plasticity in pollen features from the beginning of its radiation.


2. Tribal Level

Although pollen is of little assistance on supporting the subfamilies defined by molecular studies, it often provides additional evidence for the circumscription of supertribes, tribes, and subtribes (Fig. 58). A few tribes are characterized by a pollen type that is not found in other members of the family. These pollen types are defined by a unique combination of pollen features rather than by the presence of a single diagnostic pollen character. A keen example is given by the pollen morphology of the PECC-complex. Molecular studies (e.g., Motley et al., 2005; Rova, 1999) have acknowledged the monophyly of this group, and Huysmans et al. (1999) demonstrated that pollen is remarkably stenopalynous within the complex. A unique pollen synapomorphy that characterizes all species is lacking, but the combination of relatively small grains, three ectocolpi, a perforated tectum beset with (micro)spines, and a complex endopattern comprising an endocolpus or endocingulum with numerous extensions is not observed in any other member of the family.

Other tribes that are remarkably stenopalynous and characterized by a peculiar pollen type are Danaideae (very small to small, 3-4(-5)-colporate pollen grains with a (micro)reticulate tectum), Rubieae (very small or small (rarely large) pluricolpate grains with a perforated tectum with (micro)spines), and Vanguerieae (po(ro)rate or short colporate grains, often with a costa). The tribe Naucleeae is also stenopalynous; at first sight, a special character that sets it apart from pollen grains of many other Rubiaceae, however, is lacking. Further research on the inside ornamentation of the grains may reveal that the H-shaped endoaperture that is frequently reported for the group is a useful synapomorphy (Verellen et al., submitted).

Sometimes, pollen morphological support for a clade is not immediately obvious. The clade formed by the Rubieae, Paederieae, and Theligoneae, for example, seems to be characterized by very different pollen grains: Rubieae have pluricolpate grains; Paederieae, 3-colpate grains; and Theligoneae, pantoporate grains. Pollen grain size, pollen shape, or peculiarities of the sexine are rather diverse and do not unite these three tribes. The synapomorphy is found at the inside of the pollen grains. All members of this clade seem to lack a distinct endoaperture, a feature otherwise almost omnipresent in the subfamily Rubioideae.


In some cases the pollen type observed is not unique to a single tribe. Pollen morphology may then be useful in rejecting or supporting molecular phylogenies. This is, for example, the case for the extended tribe Coccocypseleae (cf. Piesschaert et al., 2000b). A macromolecular study by Andersson and Rova (1999) proposed a close relationship among the genera Coccocypselum P. Br., Hindsia Benh. ex Lindl., and Declieuxia Kunth., three genera that were never considered be close allies. Piesschaert et al. (2000b) demonstrated that pollen of the three genera is remarkably similar. The pollen grains are medium to very large, are invariably 3-colporate, and feature a double reticulum. Although this pollen type is not unique to the tribe (it is also observed among genera from the tribe Spermacoceae s.1.), it does supply morphological support for a close relationship.

A few tribes are characterized not by a single pollen type but by a large number of pollen morphs. Pollen of the eurypalynous tribes is often not useful for delimiting the tribal boundaries but is rather useful at a lower taxonomic level (see below). Eurypalous tribes are Psychotrieae, Morindeae, Spermacoceae s.str., Coffeeae, Sabiceeae, and Gardenieae, as is the HedyotideaeHedyotis-group. In some of these tribes less-specialized pollen grains are found next to more derived types. Pollen morphology of tribes not cited above is less typical or often not adequately known to draw conclusions.

3. Generic and Infrageneric Level

In the eurypalynous tribes, different pollen types are observed. This pollen plasticity is not unique to Rubiaceae but is also recorded for genera of other angiosperm families (cf. Van Campo, 1976).

a. Generic Level

Persson (1993) studied in detail the pollen morphology of "Gardeniinae" (see Gardenieae in Table I). This subtribe comprises 64 genera, mainly trees, shrubs, and lianas. The eurypalynous character of the group is eye catching. Monads, tetrads, and polyads are observed; the pollen size ranges from very small to medium (rarely large); and the tectum is perforate, (micro)reticulate, foveolate, psilate, or rugulate. The pollen morphological variation within the genera turned out to be rather limited, however. Persson (1993: 581) wrote, "Contrary to the subtribe the majority of the genera are highly stenopalynous, the two neotropical genera Randia and Alibertia (A. Rich.) forming two striking exceptions." This situation agrees well with what was described by Dessein (2003) for the genera of Spermacoceae s.str., where Spermacoce forms the striking exception. Some other, higher taxa that contain stenopalynous genera are the tribe Morindeae (cf. Johansson, 1987a), the Hedyotideae-Pentas group (Dessein et al., 2000), and probably also the Hedyotideae-Hedyotis group.

b. Subgeneric and Specific Level

Pollen morphological data are sometimes useful in delimiting species within a genus; for example, in the pantropical genera Spermacoce and Psychotria. Johansson (1992) studied the pollen morphology of 168 species of Psychotria s.1. His study well illustrated the variation for almost each character, which resulted in the characterization of 22 pollen types. In conclusion he wrote, "Psychotria in a wide sense is very heterogeneous pollen morphologically and it is not possible to use pollen grain characteristics for the delimitation of the genus as now defined. ... Exine morphology presents powerful characters for distinguishing between species groups in Psychotria s.1. ... It is evident ... that pollen morphological analyses should form an integrated part of every future systematic investigation of the genus Psychotria s.1." (Johansson 1992: 29-30).

Pire (1996), Dessein et al. (2002a), and Dessein et al. (2005b) observed a similar wide variation in pollen characters in Spermacoce. In total almost 150 species (>50%) of this genus have been studied pollen morphologically, and 20 different pollen types have been recognized. In many cases, species can be identified on the basis of their pollen grains only, and pollen was used to solve taxonomic debates from the past (Dessein et al., 2003). In many other genera of the Rubiaceae, pollen is not so variable but is still useful for supporting groups of related species.

c. Infraspecific Variation

In Rubiaceae, infraspecific pollen variation is frequently observed. Sometimes, variation can be linked with heterostyly or coincides with infraspecific taxa. The number of apertures is probably the most variable character at the species level. Many species have predominantly 3 colpi, but 4 or, more rarely, 5 colpi are sometimes observed. The pollen grains with an aberrant number of apertures are often present in low frequencies, but in some cases 3- and 4-aperturate grains are observed in equal quantities (e.g., for some species of Pentas Benth.; Dessein et al., 2000). Pollen size is also variable at the species level, sometimes with pollen of different sizes in one anther, as in Ernodea (see the section on "Pollen Size," above).

The best-documented case of pollen polymorphy in Rubiaceae--i.e., plants having several kinds of palynomorphs--is observed in the genus Coffea. Chinnappa and Warner (1982) found that 13 species and varieties of Coffea s.1. are highly polymorphic. Coffea arabica L. beats the lot with 6 "distinct" pollen types, based on the number of colpi, aperture type (syncolporate or not), tectum peculiarities, and shape of the endoaperture. Some of these abnormal pollen types, however, were observed only in very low frequencies and may well represent malformed pollen grains. Stoffelen et al. (1997) considered the pollen polymorphism of Coffea an expression of a continuous variation within several characters. The occurrence of infraspecific plasticity certainly lowers the systematic value of pollen grains in some cases.


As illustrated above, certain pollen characters may be very plastic in one group and rather conserved in another. Hence, no overall systematic value can be assigned to these characters or character states, and their value should be assessed separately for each study group. This is very much in accordance with what is observed for most morphological characters within Rubiaceae. The "number and position of ovules in a locule" provides a good example. In former classification systems, much attention was paid to this character, and the subdivision into subfamilies by Hooker (1873) and Schumann (1891) was based on it. Even the much more polythetic system of Robbrecht (1988a) characterized one of the subfamilies (Antirheoideae) on the basis of the presence of a single pendulous ovule per locule, but molecular data completely reject the systematic value of this character at higher systematic levels.

This does not mean, however, that (pollen) morphological characters are useless in the systematics of the family. Morphological characters are still useful, but at levels different from (usually lower than) those that have been acknowledged (see below).


Our current knowledge of the presence and typology of orbicules within Rubiaceae is fragmentary. Only two subfamilies--Cinchonoideae s.str, and Ixoroideae s.1.--have been explored, by Huysmans et al. (1997) and by Vinckier et al. (2000), respectively. For Rubioideae, data on orbicules are available only for the tribe Rubieae and a few genera scattered all over the subfamily. Both Huysmans et al. (1997) and Vinckier et al. (2000) stated that tapetal orbicules can be useful taxonomic markers at the tribal level. In Ixoroideae s.1., for example, Vinckier et al. (2000) found that the tribes Coffeeae, Octotropideae, Ixoreae, and Pavetteae are characterized by specific orbicule types. The number of genera and species sampled in the studies available is very low, however, and more data are needed to prove that orbicule types really do coincide with tribal boundaries.

Another problem that needs to be addressed is the relationship among tapetum type, sexine ornamentation, and orbicule type. Pollen with supratectal spines often has echinate orbicules (type I)--e.g., in the PECC-complex (Huysmans et al., 1999)--whereas species with a smooth but perforate or microreticulate tectum often possess orbicules with perforations in their wall (Vinckier et al., 2000). According to Hesse (1986), these parallelisms are rooted in the homology of tapetum and sporogeneous tissue. The tapetum and the microspores contain homologous, nearly identical genetic information for the production, release, and accumulation of sporopollenin (Hesse, 1986; Pacini et al., 1985; Young et al., 1979). Studies of orbicule wall formation may clarify the underlying factors determining exine and orbicule wall patterning. Furthermore, orbicules are extracellular structures, independent of cytoplasmic control, as opposed to the pollen exine. These observations corroborate the idea proposed by Clement and Audran (1993a, 1993b) that orbicule wall development can represent a model for studying sporopollenin deposition.

The absence of orbicules in Rubieae, on the other hand, may merely indicate the presence of plasmodial tapeta in this tribe. This more evolved type of tapetum lost the potential to produce orbicules. At present, it is impossible to correctly interpret the absence of orbicules because so little is known about tapetum types in Rubiaceae (cf. Andronova, 1984).


Pollen characters are no doubt an important source of phylogenetically informative characters. As a result, they are frequently incorporated into morphologic cladistic analyses. Nevertheless, most of the analyses that include pollen lack (profound) discussions of character and character state delimitations. The lack of discussions on these issues may result in undetected, inappropriate coding that may bias the results of the phylogenetic analysis or produce high levels of homoplasy, which can be used as argument for excluding pollen from future analyses. Most pollen characters have been proved to and are expected to have homoplasy, but this should not discourage researchers from incorporating them into cladistic analyses. The existence of homoplasy does not necessarily mean that the character does not contain a phylogenetic signal. In the previous sections we showed that some character states are useful in circumscribing higher taxonomic levels, even though they may have more than a single, independent origin. In the process of phylogenetic analyses, homoplasy means ignorance that can be caused either by inappropriate character or character state definitions and/or coding or by our inability to differentiate homologous and nonhomologous conditions. Worrying about homoplasy is meaningful only if it is so high that it no longer provides useful grouping information. Otherwise, homoplasy should be considered necessary for exercising reciprocal illumination and as a tool with which to conduct targeted further investigations. In the case of pollen, this often means completing our knowledge by studying the fine ultrastructure and ontogeny of the grains.

In this section we will provide suggestions about how to code some of the commonly used pollen characters and other possible characters not commonly incorporated into phylogenetic analyses. The utility and the way to code a character frequently depends on the taxonomic level (hierarchy) at which the analysis is being conducted. Our discussion emphasizes higher taxonomic levels--i.e., tribes and above--but we hope that it will also be a useful guide for studies below these levels, which may include additional characters.

Distinguishing continuous and discrete characters may be difficult, but is important in phylogenetic analyses because it affects the primary homology assessments. With respect to pollen characters, obviously continuous characters are pollen size, length of polar axis, and equatorial diameter, as well as ectoaperture length. Less obvious are the shape of the grains, which is dependent on the ratio of P/E, and the type of tectum discontinuities (perforate, microreticulate, foveolate, or reticulate), which depend on the size of the discontinuities. Several alternatives for coding continuous characters have been proposed, such as Farris (1990), Strait et al. (1996), or Swiderski et al. (1998), or Thiele (1993). Of these proposals, the one of Thiele (1993) has been implemented in the computer program Morphocode, < sys/mc/>. In this method a continuous character is treated as an ordered multistate character in which raw measurements are transformed into character states by using a simple algorithm. The method in fact weights the differences between the measurements. Other methods, including simple statistical tests (e.g., Luna & Ochoterena, 2004), have been used. The application of a particular method to treat continuous characters depends, as in many other cases, on the case study and on the statistical properties of the data, and we recommend testing different methods by trial and error so that a reasonable amount of potential phylogenetic information is obtained by the groupings achieved.

Some of the characters that we discuss below can only be used in a few practical cases, because knowledge about them is generally lacking in many genera. Nevertheless, we include them in this section in the hope that the discussion below will stimulate more research on those topics.

Dispersal units (monads-tetrads-massulae) can be used as a potential source of phylogenetic information, but tetrad configuration (tetrahedral-uniplanar-decussate) does not seem to be reliable, because it may be variable even within a single individual. The characteristics of the apertures can be coded as several characters or as a few complex characters. Even though the latter option is widely used, we consider that it is not the most appropriate, because it results in a considerable reduction of potential phylogenetic information. One obvious condition to consider in phylogenetic analyses is the presence of apertures in different layers of the exine. Because most taxa have ectoaperturate pollen, this condition is implied when one codes the characteristics of that type of aperture. On the other hand, the presence of endo- and mesoapertures is not constant, so they should be coded as two independent characters. Regarding the ectoapertures, an obvious condition that appears to have phylogenetic content is their number. This is in fact a quantitative character, but within most of the taxa it is constant and at the same time variable among them. Therefore, we recommend using it as a simple numeric character, with as many states as the existing number of apertures. Nevertheless, in some cases, when the number of apertures exceeds 5, it may be sufficient to code them as "many" (e.g., 34-5-more than 5). Taxa with a variable number of apertures should be coded as (subset-) polymorphic, because coding these cases as different states will create groupings that do not necessarily reflect homology. The distribution of the ectoapertures can also be considered spread (pantoaperturate) or confined to the equator (zonoaperturate). The position of the ectoaperture with respect to the angles of the outline in polar view (angulaperturate-planaperturate) could also be incorporated in cladistic analyses. Nevertheless, this is a character that requires further attention, because it is seldom recognizable given the lack of angles in the common circular shape in polar view of Rubiaceae grains. It seems possible, however, that the relative position of both types of ectoapertures depends on the configuration of the tetrad. To explore this possibility, pollen ontogenetic studies should be undertaken to investigate whether the position of these two types of apertures is constant in the tetrad, thereby giving an indirect way to determine the character state even in pollen grains with a circular amb. One of the most obvious characters related to the ectoapertures is their shape (colpus-pore). Both conditions are normally easy to distinguish, but in Vanguerieae they seem to form a gradient. The presence-absence of a prominently protruding thickening of the exine around an ectoaperture, most commonly around a pore (aspis), can also be coded.

The presence-absence of mesoapertures seems easy to define in theory, but in practice it can be problematic. This is because it depends on the degree of thinning, which is sometimes difficult to observe. Therefore, this character should be explored with caution.

Regarding the endoaperture, it is necessary to consider whether it is congruent with the ectoaperture (relative size of the endoaperture with respect to the ectoaperture: congruent-noncongruent) and its shape (pore-colpus-cingulum). As for the ectoapertures, the presence-absence of a thickening of the nexine/endexine bordering an endoaperture, or following the outline of an ectoaperture (costa), can also be coded. In the latter case, sometimes it is possible to further specify the type of costa, depending on which aperture it is bordering (e.g., costa ectocolpi-costa endopri).

Several characters related to the exine can be included. The most obvious ones are the presence-absence of a tectum, columella layer, and free-standing bacula. In many cases these conditions are easy to recognize, but in some instances TEM observations may be required. The presence-absence of supratectal ornamentations is easy to recognize with SEM, but the definition of classes within this character (granules-microspines-spines) is problematic, because it corresponds to continuous variation. The reticulum of the exine can be simple or double, a character that is normally easy to code, but the nature of the double reticulum should be explored in more detail.

The ornamentation of the nexine can also be potentially informative, but it is a character that has been recorded only recently. To date, it seems possible to recognize different character states for the inner nexine (granular-endocracks-water drops-isolated patches), but it is possible that, as more information regarding this character is gathered, the distinction between the states will become fuzzy.

Orbicules also represent a potential source for several cladistically informative characters. Since they were first described, more researchers have paid attention to their presence-absence, but, as happens with several other morphological characters (e.g., raphides), it is not easy to use bibliographical references to code this character. This is mainly because absence is generally not reported. In order to have more confidence in the incorporation of this character in phylogenetic analyses, we encourage researchers to also report the absence of orbicules. If they are present, several characters can additionally be coded: the presentation of the orbicules (free-embedded); their shape (spiny-microruguate-smooth-folded-granular); the presence-absence of ornamentation on their walls; the presence-absence of perforations in their walls; and the characteristics of the core (electron dense-lucent). Evidently, taxa that lack orbicules need to be coded as inapplicable for the other characters.

Characters that would be interesting to include but are difficult to define and compare, given the current state of knowledge, are the presence-absence of protruding onci and pollen buds. Both structures are nonresistant to acetolysis and therefore in many cases have not been recorded, even though they may be frequent. In general appearance they may look similar, especially at the precursor states, but they can be unequivocally distinguished with the use of TEM.

Pollen dimorphism can be caused by several factors, so it requires special attention. In general, it is a difficult character to include in phylogenetic analyses, because it generally depends on sampling. For example, the most common cause for pollen dimorphism is heterostyly; the detection of two pollen-size modes depends on examination of both types of flowers (long and short style). If the sampling of pollen is specifically designed to cover this type of cases, it can be an interesting character to include in phylogenetic analyses. It is important to note that the cause of dimorphism (e.g., heterostyly) should not be included in the cladistic analysis at certain taxonomic levels, where it may be a completely dependent character.

Finally, we would like to suggest the exclusion of characters with intermediary conditions that make it difficult to define characters and character states. This is the case for: colpus ending and extensions of the endoaperture. Also, due to potential artifacts caused by acetolysis, we suggest excluding characters such as colpus membrane ornamentation and presence-absence of operculum, even though they may contain a phylogenetic signal.

VI. Future Research

Recently, enormous progress has been made in documenting the pollen morphology of Rubiaceae. Only now, by confronting the pollen data with the resulting phylogenies based on molecular data, can pollen evolution be inferred at all taxonomic levels. The present review of palynological literature on the family reveals the genera that were never studied and aims to stimulate further palynological exploration in Rubiaceae.

Apart from the need for pollen observations for the undocumented genera of Rubiaceae, many other pollen related topics deserve our attention. The relationship between pollen morphological traits and their ecological and/or functional role is in general and thus also in Rubiaceae largely underexplored. The possible link between pollinators and sexine ornamentation, habitat and apertural configuration, and flower morphology and pollen size also provide exciting research topics in several groups of Rubiaceae. A combined ontogenetic and evo-devo study in the eurypalynous genera such as Spermacoce may also provide new insights into the morphological diversity observed.

Another gap in our knowledge is the detailed development of the pollen grain wall and the orbicules. Only a few species have been investigated in this respect, and only two reports are based on ultrastructural observations (Mitriostigma: Hansson & El-Ghazaly, 2000; Rondeletia: El-Ghazaly et al., 2001). Regarding the orbicules, our understanding of their presence/absence pattern and their possible function(s) could improve considerably if more (ultrastructural) data on tapetum types in Rubiaceae become available.

We would greatly appreciate your feedback on this review. With your assistance we will make the pollen database of Rubiaceae as complete as possible and keep it updated.

IX. Appendix 1: Template for Pollen Descriptions

The pollen template below includes the essential characters for describing Rubiaceae pollen. Many other characters, such as the thickness of the different pollen wall layers or the size of the perforations and/or lumina, should be included whenever available. Also, special features, such as protruding onci, pollen buds, and margines, should be discussed when observed. If the species investigated are heterostylous, one should try to study pollen of all forms. Orbicules should also be described, at least naming the type, and, if there is certainty of negative reports (absence of orbicules), this should also be specified.

Pollen dispersed as (monads--tetrads--massulae)

Pollen grains (isopolar--heteropolar); (number of apertures) (zono--panto) aperturate; equatorial diameter (give size range); polar axis (give size range); equatorial shape (peroblate--oblate--suboblate--spheroidal--subprolate--prolate--perprolate, or give P/E range); polar outline or amb (triangular--quadrangular--circular--lobed-- ...)

Apertures (simple or compound); ectoapertures (colpus--porus)--mesoapertures (absent/present--colpus--porus)--aspis (absent/present)--endoapertures (absent/present--porus--colpus --endocingulum); costa (absent/present)

Tectum (perforate--microreticulate--reticulate-- ...); supratectal elements (absent/present--granules--microspines--spines) Inner nexine surface (granular--smooth); endocracks (absent/present--description)

X. Appendix 2: List of Genera in the Pollen Database, with Reference to the Literature

This appendix lists the genera for which "good" pollen morphological data are available, arranged in alphabetical order. In parentheses, the most relevant pollen sources are cited, with the number of species investigated for each contribution, the number of species investigated compared with the total number of species. If the number of species investigated cannot be deduced from the manuscript, a question mark is used. The total number of species given represents the best estimate we can make at present (a reference for each figure can be found in the database). For comparison, the number of species for each genus according to Mabberley (1997) is given in parentheses; if the genus is not listed or not recognized by Mabberley, a dash is used. More literature references can be found in the database at < bio/sys/pollen.htm>. Genera for which we found only poor and/or unreliable pollen data are listed in the "Results" section of the text.

Acranthera Am. ex Meisn. (Mathew & Philip, 1983: 1/(35)35).

Acunaeanthus Borhidi, Jarai-Koml. & Moncada (Borhidi et al., 1980: 1/(1)1)

Adenorandia Vermoesen (Malplanche, 1971: 1/(1)1, as Pseudogardenia; Keddam-Malplanche, 1985: 1/(1)1, as Pseudogardenia; Persson, 1993: 1/(1)1, as Pseudogardenia)

Adina Salisb. (Verellen et al., submitted: 2/(3)3)

Adinauclea Ridsdale (Verellen et al., submitted: 1/(1)1)

Agathisanthemum Klotzsch (Dessein, unpubl.: 3/(5-6)3-4)

Aidia Lour. (Puttock, 1992: 1/(18)60; Persson, 1993: 3/(18)60, 2 as Anomanthodia)

Aidiopsis Tirveng. (Persson, 1993: 1/(1)1)

Airosperma K. Schum. & Lauterb. (Darwin, 1980a: 6/(6)6)

Aitchisonia Hemsl. ex Aitch. (Robbrecht, 1982b: 1/(1)1)

Alberta E. Mey. (Puff et al., 1984: 4/(6)6)

Aleisanthia Ridl. (Tange, 1996a: 2/(2)2; Huysmans, 1998: 1/(2)2)

Aleisanthiopsis Tange (Tange, 1996b: 2/(-)2)

Alibertia A. Rich. (Persson, 1993: 13/(35)50)

Alleizettella Pit. (Robbrecht & Puff, 1986: 1/(1)2)

Alseis Schott (Andersson, 1993: 6/(20)16; Huysmans, 1998: 3/(20)16)

Amaioua Aubl. (Persson, 1993: 2/(25)9)

Amphiasma Bremek. (Dessein, unpubl. 3/(5-6)7-8)

Amphidasya Standl. (Huysmans et al., 1998a: 1/(7)8)

Ancyclanthos Desf. (Igersheim, 1989: 1+1/(5)1, as Ancylanthus; Lens et al., 2000: 1/(5)1)

Anthospermum L. (Robbrecht, 1982b: 2/(40)39)

Aoranthe Somers (Huysmans et al., 1998a: 2/(2)5; Keddam-Malplanche, 1980: 2/(2)5, as Porterandia spp.; Keddam-Malplanche, 1985: 4/(2)5, as Porterandia spp.)

Aphaenandra Miq. (Huysmans et al., 1998a: 1/(2)1)

Aphanocarpus Steyerm. (Piesschaert, 2001: 1/(1)1)

Appunia Hook. f. (Johansson, 1987a: 1/(-)ca. 15)

Arachnothryx Planch. (Borhidi, 1982: 1/(80)80)

Arcytophyllum Willd. (Huysmans, 1998: 1/(15)16)

Argocoffeopsis Lebrun (Robbrecht, 1981b: 6/(8)7; Chinnappa & Warner, 1981: 5/(8)7, as Coffea spp.)

Argostemma Wall. (Mathew & Philip, 1983: 1/(100)196)

Ariadne Urb. (Huysmans, 1998: 1/(2)1)

Asemnantha Hook. f. (Huysmans et al., 1999: 1/(1)1)

Asperula L. (Huysmans et al., 2003: 5/(90)150)

Atractocarpus Schltr. & K. Krause (Puttock, 1992: 3/(20)12, as Gardenia merikin, Randia hirta, R. stipularis; Persson, 1993: 1/(20)12)

Atractogyne Pierre (Persson, 1993: 1/(3)3)

Augusta Pohl (Huysmans, 1998: 1/(1)1)

Badusa A. Gray (Huysmans et al., 1999: 1/(3)3)

Balmea Martinez (Andersson, 1993: 1/(1)1; D'hondt et al., 2004: 1/(1)1)

Bathysa C. Presl (Andersson, 1993: 1/(2)14, as Schizocalyx; Huysmans, 1998: 1/(2)14, as Schizocalyx)

Batopedina Verdc. (Robbrecht, 1981a: 1/(3)3; Dessein et al., 2000: 3/(3)3)

Belonophora Hook. f. (Robbrecht, 1980: 1/(6)8)

Benkara Adans. (Persson, 1993: 1/4)

Bertiera Abul. (Robbrecht et al., 1993: 8/(55)50)

Bikkia Reinw. (Huysmans et al., 1999: 3/(20)20)

Blepharidium Standl. (D'hondt et al., 2004: 2/(2)2)

Borojoa Cuatrec. (Persson, 1993: 4/(8)11)

Brachytome Hook. f. (Persson, 1993: 1/(4)8)

Brenania Keay (Keddam-Malplanche, 1985: 1/(1)2; Persson, 1993: 1/(1)2)

Breonadia Ridsdale (Verellen et al., submitted: 1/(1)1)

Breonia A. Rich. (Verellen et al., submitted: 4/(5)20; Mathew & Philip, 1983: l/(5)20, as Anthocephalus)

Burchellia R. Br. (Malplanche, 1971: 1/(1)1; Persson, 1993: 1/(1)1)

Burttdavya Hoyle (Verellen et al., submitted: 1/(1)1)

Byrsophyllum Hook. f. (Persson, 1993: 2/(2)2)

Caelospermum Blume (Johansson, 1987a: 5/(7)7)

Calanda K. Schum. (Puff & Robbrecht, 1989: 1/(1)1)

Calochone Keay (Keddam-Malplanche, 1985: 2/(2)2; Persson, 1993: 1/(2)2)

Calycophyllum DC. (Andersson, 1993: 1/(3)6; Huysmans, 1998: 2/(3)6)

Calycosiphonia Pierre ex. Robbr. (Robbrecht, 1981b: 1/(2)2)

Canthium Lam. (Tilney, 1986: 6/(50)162; Verdcourt, 1987: 2/(50)162; Igersheim, 1989: 9/ (50)162; Tilney & Van Wyk, 1997: 9/(50)162; Lens et al., 2000: 6/(50)162; Mathew & Philip, 1983: 3/(50)162, as Plectronia neilgherrensis, P. rheedii, P. rheedii var. angustifolia)

Capirona Spruce (Andersson, 1993: 2/(5)2; Huysmans, 1998: 1/(5)2)

Captaincookia N. Halle (De Block & Robbrecht, 1998: 1/(1)1)

Carpacoce Sond. (Robbrecht, 1982b: 1/(7)7; Robbrecht, 1985: 7/(7)7)

Carphalea Juss. (Puff, 1988: 9/(10)10; Scheltens, 1998: 1/(10)10)

Casasia A. Rich. (Persson, 1993: 2/(11)10)

Catesbaea L. (Huysmans et al., 1999: 5/(20)ca. 16; Mathew & Philip, 1983: 1/(20)ca. 16)

Catunaregam Wolf (Verdcourt, 1981: 1/(5-6)6; Persson, 1993: 1/(5-6)6)

Cephalanthus L. (Huysmans, unpubl.: 5/(6)6-7)

Ceratopyxis Hook. f. (Huysmans et al., 1999: 1/(1)1)

Ceriscoides Hook. f. Tirveng. (Persson, 1993: 1/(7)6)

Chamaepentas Bremek. (Dessein et al., 2000: 1/(1)1)

Chassalia Comm. ex Juss. (Mathew & Philip, 1983: 1/(42)ca. 82, as Chasalia; Piesschaert et al., 1999c: 1/(42)ca. 80; Jansen et al., 1996a: 3/(42)ca. 80; Jansen, 1994: 4/(42)ca. 80)

Chazaliella E. M. A. Petit & Verdc. (Jansen, 1994: 3/(25)21; Jansen et al., 1996a: 3/(25)21)

Chimarrhis Jacq. (Huysmans, 1998: 1/(14)13; Delprete, 1999a: 10/(14)13)

Chiococca P. Br. (Huysmans et al., 1999: 2/(6)ca. 20)

Chione DC. (Huysmans, unpubl. 2/(15)15)

Chlorochorion Puff & Robbr. (Puff & Robbrecht, 1989; 1/(2)2)

Chomelia Jacq. (Jung-Mendacolli & Melhem, 1994: 1/(20)75)

Cinchona L. (Andersson, 1993: 7/(40)23; Mathew & Philip, 1983: 3/(40)23)

Cladoceras Bremek. (Robbrecht & Bridson, 1984: 1/(1)1; De Block & Robbrecht, 1998: 1/ (1)1)

Coccocypselum P. Br. (Piesschaert et al., 2000b: 4/(20)37; Rova & Andersson, 1995: 1/(20)37; Jung-Mendacolli & Melhem, 1995: 4/(20)37)

Coddia Verdc. (Verdcourt, 1981: 1/(1)1; Persson, 1993: 1/(1)1)

Coffea L. (Chinnappa & Warner, 1981, 1982: 17/(90)95-100; Stoffelen et al., 1997: 26/(90)95-100)

Coleactina N. Halle (De Block & Robbrecht, 1998: 1/(1)1)

Colletoecema E. M. A. Petit (Piesschaert et al., 2000a: 1/(1)1)

Condaminea DC. (Huysmans, 1998: 1/(3)1; Delprete, 1999a: 1/(3)1)

Conostomium (Stapf) Cufod. (Dessein, unpubl.: 4/(9)9)

Coprosma J. R. Forst. & G. Forst. (Robbrecht, 1982b: 2/(90)150; Mathew & Philip, 1983: 1/ (90)150)

Coptosapelta Korth. (Verellen, 2002: 11/(13)16; Verellen et al., 2004: 10/(13)16)

Coptosperma Hook. f. (De Block & Robbrecht, 1998:11/(-)50 10 spp. as Enterospermum, 1 as Tarenna nigrescens; De Block et al., 2001 [2002]: ?/(-)50)

Corynanthe Welw. (Huysmans, 1993: 3/(8)3)

Coryphothamnus Steyerm. (Piesschaert, 2001: 1/(1)1)

Cosmibuena Ruiz & Pav. (Andersson, 1993: 1/(12)4; D'hondt et al., 2004: 4/(12)4)

Coussarea Aubl. (Piesschaert, 2001: 3/(ca. 100)118; see also Jung-Mendacolli & Melhem, 1995)

Coutaportla Urb. (Huysmans et al., 1999: 1/(2)3-4)

Coutarea Aubl. (Huysmans et al., 1999: 1/(7)3-4; Rova & Andersson, 1995: 1/(7)3-4)

Crossopteryx Fenzl (Huysmans, unpubl.: 1/(1)1)

Crucianella L. (Huysmans et al., 2003: 3/(ca. 30)33)

Cruciata Mill. (Huysmans et al., 2003:2/(10)11)

Cruckshanksia Hook. & Am. (Dessein, unpubl.: 2/(7)7)

Crusea Chain. & Schltdl. (Anderson, 1972: 13/(13)14; Dessein, 2003: 2/(13)14)

Cubanola Aiello (Aiello, 1979: ?/(2)2)

Cuviera DC. (Verdcourt, 1987: 1/(20)23; Igersheim, 1989: 4/(20)23; Lens et al., 2000: 2/(20)23)

Cyaneuron Tange (Tange, 1998: 5/(-)5)

Damnacanthus C. F. Gaertn. (Johansson, 1987a: 2/(6) 10; Robbrecht et al., 1991: 1/(6) 10; Naiki & Nagamasu, 2003: 4/(6) 10)

Danais Comm. ex Vent. (Buchner & Puff, 1993: 15/(40)ca. 29; Scheltens, 1998: 1/(40)ca. 29; see also Huysmans, 1998)

Deccania Tirveng. (Persson, 1993: 1/(1)1)

Declieuxia Kunth (Piesschaert et al., 2000b: 15/(27)?40; Jung-Mendacolli & Melhem, 1995: 1/(27)?40)

Dentella J. R. Forst. & G. Forst. (Mathew & Philip, 1983: 1/(10)8)

Dialypetalanthus Kuhlm. (Piesschaert et al., 1997: 1/(1)1)

Dibrachionostylus Bremek. (Dessein, unpubl.: 1/(1)1)

Dichilanthe Thw. (Puff et al., 1996: 1/(2)2)

Dictyandra Hook. f. (Robbrecht, 1984: 2/(2)2; De Block & Robbrecht, 1998: 1/(2)2)

Didymosalpinx Keay (Keddam-Malplanche, 1985: 1/(4)4; Persson, 1993: 2/(4)4)

Diodella Small (Dessein, 2003: 1/(-)?)

Diodia L. (Pire, 1997a: 1/(30)5; Dessein, 2003: 2/(30)5)

Dioecrescis Tirveng. (Persson, 1993: 1/(1)1)

Dioicodendron J. A. Steyermark (Delprete, 1999a: 1/(2)1)

Diplospora DC. (Ali & Robbrecht, 1991: 3/(10)19; Mathew & Philip, 1983: 1/(10)19)

Discospermum Dalzell (Ali & Robbrecht, 1991: 1/(6)6)

Dolichodelphys K. Schum. (Delprete, 1999a: 1/(1)1)

Dolicholobium A. Gray (Andersson, 1993: 1/(28)28; Huysmans, 1998: 1/(28)28)

Doricera Verdc. (Verdcourt, 1983: 1/(1)1)

Dunnia Tutcher (Ying et al., 1993: 1(2)2)

Duperrea Pierre ex Pit. (De Block, 1997: 1/(2)2)

Duroia L. f. (Persson, 1993: 5/(20)34)

Durringtonia R. J. F. Hend. & Guymer (Puff & Robbrecht, 1988: 1/(1)1)

Ecpoma K. Schum. (Huysmans et al., 1998a: 1/(1)5+1)

Elaeagia Wedd. (Huysmans, 1998: 2/(10)22)

Emmenopterys Oliv. (Ying et al., 1993: 1/(2)2)

Emmeorhiza Pohl. ex Endl. (Pire & Cabral, 1992: 1/(1)1; Dessein, 2003: 1/(1)1)

Erithalis P. Br. (Huysmans et al., 1999: 2/(10)9)

Ernodea Sw. (Negron-Ortiz, 1996: 4/(9)4; Dessein, 2003: 1/(9)4)

Euclinia Salisb. (Persson, 1993: 1/(3)3)

Exallage Bremek. (Mathew & Philip, 1983: 1/(-)24)

Exostema (Pers.) Rich. ex Humb. & Bonpl. (Huysmans et al., 1999: 2/(45)47)

Fadogia Schweinf. (Verdcourt, 1987: 2/(45)44; Igersheim, 1989: 5+1/(45)44, 1 as Ancylanthus rogersii; Lens et al., 2000: 1/(45)44)

Fadogiella Robyns (Igersheim, 1989: 1/(2)3; Lens et al., 2000: 1/(2)3)

Fagerlindia Tirveng. (Persson, 1993: 3/(6)9)

Faramea Aubl. (Jung-Mendacolli & Melhem, 1995: 1/(125) 130; Piesschaert, 2001: 2/(125) 130)

Ferdinandusa Pohl (Andersson, 1993: 10/(24)20; Huysmans, 1998: 3/(24)20)

Gaertnera Lam. (Jansen, 1994: 4/(30)60; Jansen et al., 1996b: 7/(30)60; Pailler & Thompson, 1997: 8/(30)60)

Gaillonia A. Rich. (Robbrecht, 1982b: 3/(17)27, 1 as Crocyllis anthospermoides, 2 as Neogaillonia spp.; Leonard, 1984: 1/(17)27)

Galianthe Griseb. (Pire & Cabral, 1992: 20/(-)50; Pire, 1997a: 8/(-)50; Dessein, 2003 (2/(-)50)

Galium L. (Huysmans et al., 2003: 16/(ca. 300)400; Mathew & Philip, 1983: 1/(ca. 300)400)

Galopina Thunb. (Robbrecht, 1982b: 1/(-)1)

Ganguelia Robbr. (Robbrecht et al., 1996: 1/(-)1)

Gardenia Ellis (Persson, 1993: 2/(60)154; Mathew & Philip, 1983: 2/(60)154; Puttock, 1992: 18/(60)154)

Genipa L. (Persson, 1993: 2/(7)4)

Gentingia J. T. Johanss. & K. M. Wong (Johansson & Wong, 1988: 1/(1)1)

Geophila D. Don (Piesschaert et al., 1999b: 6/(ca. 20) 19; Es, 1999: 5/(ca. 20) 19; Vanthournout, 2002: 10/(ca. 20)19; Mathew & Philip, 1983: 1/(ca. 20)19)

Gleasonia Standl. (Rogers, 1984: 2/(5)4)

Glionnetia Tirveng. (Tirvengadum, 1984: 1/(1)1)

Glossostipula Lorence (Lorence, 1986: 1/(2)3; Persson, 1993: 1/(2)3)

Gomphocalyx Baker (Dessein et al., 2005a: 1/(1)1)

Gonzalagunia Ruiz & Pav. (Huysmans et al., 1998a: 3/(15)43)

Gouldia A. Gray (Huysmans et al., 1998a: 1(3x)/(-)3, three different species given, but two placed in synonymy

Greeniopsis Merr. (Huysmans, unpubl.: 4/(6)6)

Guettarda L. (Sotolongo Molina et al., 2002: 19/(80)ca. 85; Mathew & Philip, 1983: 1/(80)ca. 85)

Gynochthodes Blume (Johansson, 1987a: 3+2/(ca. 20)14)

Gyrostipula J.-F. Leroy (Verellen et al., submitted: 1/(2)1-2)

Habroneuron Standl. (Darwin, 1980b: 1/(1)1)

Haldina Ridsdale (Verellen et al., submitted: 1/(1)1)

Hallea J.-F. Leroy (Huysmans et al., 1994: 3/(3)3)

Hamelia Jacq. (Huysmans, unpubl.: 6/(16)17; Mathew & Philip, 1983: 1/(16)17)

Hedyotis L. (Mathew & Philip, 1983: 1+2(250)65, as Oldenlandia pruinosa, O. ramarowii)

Hedythyrsus Bremek. (Dessein, unpubl.: 2/(2)2)

Heinsia DC. (Huysmans et al., 1998a: 3/(4-5)6)

Hekistocarpa Hook. f. (Dessein et al., 2001b: 1/(1)1)

Henriquezia Spruce ex Benth. (Rogers, 1984: 3/(7)3)

Hillia Jacq. (Andersson, 1993: 1/(24)24; D'hondt et al., 2004: 19/(24)24); see also D'hondt, 2002)

Himalrandia T. Yamazaki (Lobreau-Callen, 1978: 1/(2)3; Verdcourt, 1981: 1/(2)3)

Hindsia Benth. ex Lindl. (Piesschaert et al., 2000b: 2/(8)11; Huysmans, 1998: 1/(8)11)

Hintonia Bullock (Huysmans et al., 1999: 3/(4)3)

Hippotis Ruiz. & Pav. (Rova & Andersson, 1995: 2/(12)11)

Hoffmannia Sw. (Huysmans, unpubl.: 5/(ca. 45)119)

Homollea Atenes (De Block & Robbrecht, 1998: 1/(3)3)

Homolliella Arenes (De Block & Robbrecht, 1998: 3/(1)5)

Houstonia L. (Dessein, unpubl.: 15/(-)20; Terrell et al., 1986: ?/(-)20, as Hedyotis)

Hutchinsonia Robyns (Igersheim, 1989: 1/(2)2)

Hydrophylax L. f. (Puff, 1986b: 1/(1)1; Dessein, 2003:1/(1)1

Hymenocoleus Robbr. (Robbrecht, 1977:11/(12) 12)

Hymenodictyon Wall. (Huysmans, 1993: 5/(20)26; Mathew & Philip, 1983: 1/(20)26)

Hyperacanthus E. Mey. ex Bridson (Bridson & Robbrecht, 1985a: 1/(2)2; Persson, 1993: 1/(2)2)

Isertia Schreb. (Bosser & Lobreau-Callen, 1998: 1/(13)14; Huysmans et al., 1998a: 5/(13)14)

Isidorea A. Rich. (Huysmans et al., 1999: 3/(20)17)

Ixora L. (De Block & Robbrecht, 1998: 29/(300)400, 5 unidentified; see also De Block, 1998; Mathew & Philip, 1983: 4+1/(300)400, 1 as Pavetta calycina)

Jackiopsis Ridsdale (Robbrecht, 1980: 1/(1)1)

Janotia J.-F. Leroy (Verellen et al., submitted: 1/(1)1)

Jaubertia Guill. (Robbrecht, 1982b: 1/(16)1, as Neogaillonia aucheri)

Javorkaea Borhidi et J. Komlodi (Borhidi & Jarai-Komlodi, 1983: 1/(1)1)

Joosia H. Karst. (Andersson, 1993: 6/(7) 11; Huysmans, 1998:1/(7) 11)

Jovetia Guides (Robbrecht, 1980: 1/(2)1)

Kailarsenia Tirveng. (Persson, 1993: 1/(6)6; Yirvengadum, 1993: 1/(6)6)

Kajewskiella Merrill et Perry (Delprete, 1999a: 1/(2)2)

Keetia E. Phillips (Tilney, 1986: 1/(40)31, as Canthium gueinzii; Verdcourt, 1987: 1/(40)31; Igersheim, 1989:14/(40)31; Tilney & Van Wyk, 1997:1/(40)31: Lens et al., 2000:3/(40)31)

Kelloggia Torr. ex Benth. (Robbrecht, 1982b: 1/(2)2)

Kerianthera J. H. Kirkbr. (Delprete, 1999a: 1/(1)1)

Knoxia L. (Mathew & Philip, 1983: 2/(9)9)

Kochummenia K. M. Wong (Persson, 1993: 1/(2)2)

Kohautia Cham. & Schltdl. (Dessein, unpubl., 20/(-)31)

Kutchubaea Fisch. ex DC. (Robbrecht, 1980:1/(11) 11; Persson, 1993: 2/(11) 11)

Ladenbergia Klotzsch (Andersson, 1993: 6/(55)34; Huysmans, 1998: 1/(55)34)

Lagynias E. Mey ex Robyns (Igersheim, 1989: 1/(4)5; Lens et al., 2000: 1/(4)5)

Lamprothamnus Hiem (Robbrecht, 1980:1/(1)1)

Landiopsis Capuron ex Bosser (Bosser & Lobreau-Callen, 1998: 1/(-)1)

Larsenaikia Tirveng. (Puttock, 1992: 3(-)3, as Gardenia ochreata, G. suffruticosa, G. jardinei; Tirvengadum, 1993: 1/(-)3)

Lasianthus W. Jack. (Adams et al., 1987: 1/(170)>180; Huang, 1972: 8/(170)>180; Piesschaert et al., 2000a: ?/(170)>180)

Lathraeocarpa Bremek. (Dessein et al., 2005a: 1/(2)2)

Lecananthus Jack (Puff & Buchner, 1998: 2/(-)3)

Lelya Bremek. (Dessein, unpubl.: 1/(1)1)

Leptactina Hook. f. (Robbrecht, 1984: ?5/(25)<20; De Block & Robbrecht, 1998: 6/(25)<20)

Leptodermis Wall. (Robbrecht, 1982b: 1/(30)41)

Leptostigma Am. (Robbrecht, 1982b: 2/(6)6; 1 as Coynula pilosa, 1 as Nertera arnottianum [sic])

Leucocodon Gardner (Puff & Buchner, 1998: 1/(-)1)

Limnosipanea Hook. f. (Huysmans, 1998: 1/(7)5)

Lindenia Benth. (Darwin, 1976: 3/(3)3)

Ludekia Ridsdale (Verellen et al., submitted: 2/(2)2)

Luculia Sweet (Huysmans, 1993: 2/(5)5; Mathew & Philip, 1983: 2/(5)5)

Macrocnemum P. Br. (Andersson, 1993: 3/(20)8; Huysmans, 1998: 1/(20)8)

Macrosphyra Hook. f. (Persson, 1993: 1/(3)3; Robbrecht, 1980: 2/(3)3)

Manettia Mutis ex L. (Huysmans, 1998: l/(80)ca. 130; Rova & Andersson, 1995: 1/(80)ca. 130; Jung-Mendagolli & Melhem, 1994: 2/(80)ca. 130; Jung-Mendacolli & Melhem, 1995: 1/(80)ca. 130)

Manostachya Bremek. (Dessein, unpubl.: 2/(3)3)

Mantalania Capuron ex J.-F. Leroy (Persson, 1993: 1/(2-3)2; Leroy, 1974: 2/(2-3)2)

Maschalocorymbus Bremek. (Huysmans, 1998:1 + 1/(-)4)

Massularia (K. Schum.) Hoyle (Keddam-Malplanche, 1985: 1/(1)1; Persson, 1993: 1/(1)1)

Mastixiodendron Melch. (Huysmans et al., 1999: 2/(7)7; Darwin, 1977: 7/(7)7)

Melanopsidium Colla (Persson, 1993: 1/(?)1)

Metabolos Blume (Puff & Igersheim, 1994a: 1/(-)1)

Metadina Bakh. f. (Verellen et al., submitted: 1/(1)1)

Meyna Roxb. ex Link (Igersheim, 1989: 1/(11)10; Lens et al., 2000: 1/(11)10)

Mitchella L. (Robbrecht, 1982b: 1/(3)2); Robbrecht et al., 1991: 1/(3)2)

Mitracarpus Zucc. ex Schult. & Schult. f. (Dessein, 2003: 2/(30)45)

Mitragyna Korth. (Huysmans et al., 1994: 7/(10)7; Mathew & Philip, 1983: 1/(10)7, as Stephegyne)

Mitrasacmopsis Jovet (Groeninckx, 2005: 1/(1)1)

Mitriostigma Hochst. (Persson, 1993: 1/(5)4; Hansson & E1-Ghazaly, 2000: 1/(5)4)

Molopanthera Turcz. (Huysmans et al., 1999: 1/(1)1)

Monosalpinx N. Halle (Persson, 1993: 1/(1)1)

Morelia A. Rich. (Persson, 1993: 1/(1)1; Malplanche, 1971: 1/(1)1; Robbrecht, 1980: 1/(1)1)

Morierina Vieill. (Huysmans et al., 1999: 1/(2)2)

Morinda L. (Mathew & Philips, 1983: 3/(80) 115; Johansson, 1987a: 8/(80) 115)

Motley(i)a J. T. Johanss. (Johansson, 1987a: 1/(1)1)

Mouretia Pit. (Tange, 1997: 4/(1)4)

Multidentia Gilli (Verdcourt, 1987:1/(11)11; Igersheim, 1989:1/(11)11)

Mussaenda L. (Huysmans et al., 1998a: 4/(100)216; Bosser & Lobreau-Callen, 1998: 2/ (100)216; Rova & Andersson, 1995: 1/(100)216; Mathew & Philip, 1983: 3/(100)216)

Mussaendopsis Baill. (Huysmans, 1993: 2/(2)2; Puff & Igersheim, 1994b: ?2/(2)2)

Mycetia Reinw. (Huysmans et al., 1998a: 3/(25)44)

Myonima Comm. ex Juss. (De Block & Robbrecht, 1998: 2/(4)4)

Myrioneuron R. Br. ex Hook. f. (Huysmans et al., 1998a: 1/(-)14)

Myrmecodia Jack. (Robbrecht, 1988a: 1/(26)26; see also Huxley & Jebb, 1993)

Myrmeconauclea Merr. (Verellen et al., submitted 3/(3)3)

Nauclea L. (Verellen et al., submitted: 9/(10)10; Mathew & Philip, 1983: 2/(10)10)

Neanotis W. H. Lewis (Lewis, 1966: ?/(28)33; Mathew & Philip, 1983: 2/(28)33)

Nematostylis Hook. f. (Puffet al., 1984: 1/(1)1)

Nenax Gaertn. (Robbrecht, 1982b: 1/(9)9+2)

Neohymenopogon Bennet (Huysmans, 1998: 1/(3)3)

Neolamarckia Bosser (Verellen et al., submitted: 1/(2)2)

Neomussaenda Yange (Tange, 1994: 2/(-)2)

Neonauelea Merr. (Verellen et al., submitted: 9/(65)66)

Neopentanisia Verde. (Puff & Robbrecht, 1989: 1/(2)2)

Nernstia Urb. (Aiello, 1979: 1/(1)1, as Cigarilla; Huysmans et al., 1999: 1/(1)1)

Nertera Banks & Sol. ex Gaertn. (Robbrecht, 1982b: 1+?/(9)15)

Neurocalyx Hook. (Mathew & Philip, 1983: 1/(5)4)

Nichallea Bridson (De Block & Robbrecht, 1998: 1/(1)1)

Normandia Hook. f. (Robbrecht, 1982b: 1/(l)1)

Ochreinauclea Ridsdale & Bakh. f. (Verellen et al., submitted: 1/(2)2)

Octotropis Bedd. (Yissot et al., 1994: (1/(2)1)

Oldenlandia L. (Dessein, unpubl.: 7/(300)ca. 100; Mathew & Philip, 1983: 7/(300)ca. 100)

Oligocodon Keay (Persson, 1993: 1/(1)1)

Opercularia Gaertn. (Robbrecht, 1982b: 2/(18) 16)

Ophiorrhiza L. (Piesschaert et al., 2000a: ?/(150)313; Huang, 1972: 4/(150)313; Mathew & Philip, 1983: 4/(150)313)

Osa Aiello (Aiello, 1979: 1/(1)1)

Otiophora Zuec. (Robbrecht & Puff, 1981: 3/(20) 18); Scheltens, 1998: 3/(20) 18)

Otomeria Benth. (Dessein et al., 2000: 5/(8)8)

Oxyanthus DC. (Persson, 1993: 1/(40)40)

Oxyceros Lour. (Persson, 1993: 3/(?)14)

Pachystigma Hochst. (Igersheim, 1989: 3/(10)17; Lens et al., 2000: 1/(10)17)

Pachystylus K. Schum. (De Block & Robbrecht, 1998: 1/(2)2)

Paederia L. (Robbrecht, 1982b: 1/(30)29; Igersheim, 1991: 28/(30)29)

Pagamea Aubl. (Jansen et al., 1996b: 3/(24)28)

Pagameopsis Steyerm. (Piesschaert et al., 2001: 2/(2)2)

Palicourea Aubl. (Jung-Mendacolli, 1984: 4/(200+)ca. 200; Piesschaert, unpubl.: 9/(200+)ca. 200)

Paracephaelis Baill. (De Block & Robbrecht, 1998: 4/(1)ca. 10; Bridson & Robbrecht, 1985b: 1/(1)ca. 10, as Tarenna trichantha)

Parachimarrhis Dueke (Delprete, 1999a: 1/(1)1)

Paracorynanthe Capuron (Huysmans, 1993: 2/(2)2)

Paragenipa Baill. (Tirvengadum & Robbrecht, 1985: 1/(1)1)

Parapentas Bremek. (Dessein et al., 2000: 3/(3-4)3+1)

Pauridiantha Hook. f. (Bangoura, 1992:11/(25)39; Ntore et al., 2003: 2/(25)39; Ntore, 2004: ?/(25)39)

Pausinystalia Pierre ex Beille (Huysmans, 1993: 4/(13)5; Verellen et al., submitted: 3/(13)5)

Pavetta L. (De Block & Robbrecht, 1998: 14/(400)400; Mathew & Philip, 1983: 1/(400)400)

Payera Baill. (Buchner & Puff, 1993: 6/(1)9)

Pelagodendron Seem. (Persson, 1993: 1/(4)1)

Pentagonia Benth. (Rova & Andersson, 1995: 2/(20)25-30)

Pentanisia Harv. (Puff & Robbrecht, 1989: 1/(15)14)

Pentas Benth. (Dessein et al., 2000: 24/(34)39; Mathew & Philip, 1983: 2/(34)39)

Pentodon Hochst. (Dessein, unpubl.: 1/(2)2)

Peponidium (Igersheim, 1989: 1/(20)20)

Perakanthus Robyns (Verdcourt, 1987: 1/(1)2)

Perama Aubl. (Erdtman, 1971: 1/(9)13; !LM only)

Pertusadina Ridsdale (Verellen et al., submitted: 1/(4)4)

Petitiocodon Robbr. (Robbrecht, 1988b: 1/(1)1)

Phellocalyx Bridson (Bridson et al., 1980: 1/(1)1; Persson, 1993: 1/(1)1)

Phyllis L. (Robbrecht, 1982b: 1/(2)2)

Phylohydrax Puff (Puff, 1986b: 2/(2)2; Dessein, 2003: 2/(2)2)

Picardaea Urb. (Delprete, 1999a: 1/(2)1)

Pinckneya Rich. (Aiello, 1979: 1/(1)1; Delprete, 1999a: 1/(1)1)

Placocarpa Hook. f. (Huysmans et al., 1999: 1/(1)1)

Platycarpum Bonpl. (Rogers, 1984:7/(10) 12)

Pleiocoryne Rauschert (Persson, 1993: 1/(1)1)

Plocama Aiton (Robbrecht, 1982b: 1/(1)1; Lecuona Neumann et al., 1987: 1/(1)1)

Poecilocalyx Bremek. (Bangoura, 1992: 1/(2)4)

Pogonolobus F. Muell. (Johansson, 1987a: 1/(1)1)

Pogonopus Klotzsch (Delprete, 1999a: 3/(2-3)3)

Polysphaeria Hook. f. (Verdcourt, 1980: 5/(20)21)

Pomax Sol. ex DC. (Robbrecht, 1982b: 1/(-)1)

Porterandia Ridl. (Persson, 1993: 3/(9-10)10)

Portlandia P. Br. (Huysmans et al., 1999: 1/(6)5; Mathew & Philip, 1983: 1/(6)5)

Posoqueria Aubl. (Persson, 1993: 2/(12)16)

Praravinia Korth. (Huysmans, 1998: 2+1/(50)49)

Preussiodora Keay (Robbrecht, 1978b: 1/(1)1; Persson, 1993: 1/(1)1)

Prismatomeris Thwaites (Johansson, 1987a: ?/(15) 17; Johansson, 1987b: 13/(15) 17)

Pseudomantalania J.-F. Leroy (Persson, 1993: 1/(1-2)1; Leroy, 1974 1/(1-2)1)

Pseudomussaenda Wernham (Huysmans et al., 1998a: 2/(4-5)6; see also Puff et al., 1993b)

Pseudopyxis Miq. (Robbrecht, 1982b: 1/(2)2; Puff, 1989: 2/(2)2)

Pseudosabicea N. Halle (Huysmans et al., 1998a: 2/(12)13)

Psilanthus Hook. f. (Stoffelen et al., 1997: 6/(ca. 20)20; Chinnappa & Warner, 1981 & 1982: 3/ (ca. 20)20, as Coffea spp.)

Psychotria L. (Mathew & Philip, 1983: 6/(800-1500)>1200; Johansson, 1992: 168/(8001500)>1200;

Jansen, 1994: 8/(800-1500)>1200; Jansen et al., 1996b: 6/(800-1500)>1200; Piesschaert, unpubl.: 7/(800-1500)>1200)

Psydrax Gaertn. (Mathew & Philip, 1983: 3/(100)51, as Plectronia didyma, P pergracilis, P. umbellatum; Tilney, 1986: 2/(100)51, as Canthium locuples, C. obovatum; Igersheim, 1989: 15/(100)51; Tilney & Van Wyk, 1997: 4/(100)51; Lens et al., 2000: 2/(100)51)

Psyllocarpus Mart. & Zucc. (Kirkbride, 1979: 8/(8)8; Dessein, 2003: 2/(8)8)

Putoria Pers. (Robbrecht, 1982b: 1/(-)3)

Pygmaeothamnus Robyns (Robbrecht, 1980: 1/(4)2; Lens et al., 2000: 1/(4)2)

Pyrostria Comm. ex Juss. (Verdcourt, 1987: 2+1/(45)37, 1 as Dinocanthium; Igersheim, 1989: 2/(45)37; Lens et al., 2000: 1+1/(45)37)

Ramosmania Yirveng. (Tirvengadum, 1982: 1/(2)2)

Randia L. (Persson, 1993: 5/(100)ca. 90; Mathew & Philip, 1983: 4/(100)ca. 90)

Raritebe Wernh. (Huysmans et al., 1998a: 1/(1)2)

Remijia DC. (Andersson, 1993: 8/(25)45)

Rennellia Korth. (Johansson, 1987a: 1+4/(4)4, 4 unidentified species)

Retiniphyllum Bonpl. (Cortes-B. & Huysmans, submitted: 23/(20)23)

Rhipidantha Bremek. (Bangoura, 1992: 1/(1)1; Huysmans, unpubl.: 1/(1)1)

Richardia L. (Pire, 1997b: 7/(15)15; Dessein, 2003: 2/(15)15; Mathew & Philip, 1983: l/ (15)15)

Riodocea Delprete (Delprete, 1999b: 1/(-) 1)

Robbrechtia De Block (De Block, 2003: 2/(-)2)

Rogiera Planch. (Borhidi, 1982: 1/(18)?)

Roigella Borhidi & M. Fernandez Zeq. (Borhidi & Fernandez Zequeira, 1981a: l/(1)1)

Rondeletia L. (Igersheim, 1993b: 20/(130)286; Bosser & Lobreau-Callen, 1998: 1/(130)286; El-Ghazaly et al., 2001: 1/(130)286; Mathew & Philip, 1983: 1/(130)286)

Rosenbergiodendron Fagerlind (Gustafsson, 1998: 4/(-)4)

Rothmannia Thunb. (Keddam-Malplanche, 1985: 12/(40)ca. 40; Persson, 1993: 5/(40)ca. 40; Robbrecht, 1980: 1/(40)ca. 40)

Rubia L. (Huysmans et al., 2003: 2/(60)72; Mathew & Philip, 1983: 1/(ca. 60)72)

Rubovietnamia Tirveng. (Tirvengadum, 1998: 1/(-)1)

Rudgea Salisb. (Jung-Mendacolli, 1984: 3/(150)>120; Piesschaert, unpubl.: 6/(150)>120)

Rustia Klotzsch (Aiello, 1979: ?/(15)14; Huysmans, 1998: 1/(15)14; Delprete, 1999a: 12/(15)14)

Rutidea DC. (De Block & Robbrecht, 1998: 15/(22)22)

Rytigynia Blume (Igersheim, 1989: 5/(60-70)83; Lens et al., 2000:3/(60-70)83; Verdcourt, 1980: 6/(60-70)83)

Sabicea Aubl. (Huysmans et al., 1998a: 4/(120)ca. 130)

Sacosperma G. Taylor (Dessein, unpubl.: 1/(2)2)

Saldinia A. Rich. ex DC. (Piesschaert et al., 2000a: ?/(2)22)

Salzmannia DC. (Huysmans et al., 1999: 1/(1)1)

Saprosma Blume (Mathew & Philip, 1983: 1/(1)1)

Sarcocephalus Afzel. ex Sabine (Verellen et al., submitted: 2/(2)2; Mathew & Philip, 1983: 1/(2)2)

Scandentia E. L. Cabral & Bacigalupo (Cabral & Bacigalupo, 2001:4/(-)4; Dessein, 2003: 1/(-)4)

Schismatoclada Baker (Buchner & Puff, 1993: 3/(20)18)

Schizenterospermum Homolle ex Arenes (De Block & Robbrecht, 1998: 2/(4)4)

Schizomussaenda Li (Huysmans et al., 1998a: 1/(-)1; see also Puff et al., 1993b)

Schizostigma Am. ex Meisn. (Huysmans et al., 1998a: 1/(1)1)

Schmidtottia Urb. (Aiello, 1979: ?/(15)14)

Schradera Vahl (Puff et al., 1993a & Puff & Buchner, 1998: ?/(25)64)

Schumanniophyton Hams. (Keddam-Malplanche, 1985: 1/(5)6; Persson, 1993: 1/(5)6)

Schwendenera K. Schum. (Dessein, 2003: 1/(1) 1)

Scleromitrion (Wight & Am.) Meisn. (Mathew & Philip, 1983:1/(-) 12, as Oldenlandia nitida)

Scolosanthus Vahl (Huysmans et al., 1999: 1/(27)27)

Scyphiphora C. f. Gaertn. (Puff & Rohrhofer, 1993: 1/(1) 1)

Scyphochlamys Ball. f. (Verdcourt, 1983: 1/(1)1)

Semaphyllanthe L. Andersson (Andersson, 1993: 3/(/)6, as Calycophyllum spp.)

Sericanthe Robbr. (Robbrecht, 1978a: 1/(15)17)

Serissa Comm. (Robbrecht, 1982b: 1/(2)2; Mathew & Philip, 1983: 1/(2)2)

Sherardia L. (Huysmans et al., 2003: 1/(1)1)

Sherbournia G. Don (Persson, 1993: 1/(10)13; Keddam-Malplanche, 1980: 7/(10)13)

Simira Aubl. (Huysmans, 1998: 1/(35)41)

Sinoadina Ridsdale (Verellen et al., submitted: 1/(1)1)

Sipanea Aubl. (Huysmans, 1998: 2/(17)19; Dessein, unpubl.: 2/(17)19)

Sipaneopsis Steyerm. (Huysmans, 1998: 1/(6)8)

Sommera Schltdl. (Rova & Andersson, 1995: 2/(12)13)

Spermacoce L. (Dessein et al., 2002a: 48/(150)275; Dessein et al., 2002b: 1/(150)275; Dessein et al., 2005b: 47/(150)275; Pire, 1996: 50/(150)275, most as Borreria; Mathew & Philip, 1983: 4/(150)275, 3 as Borreria spp., 1 as Mitracarpus verticillatus)

Spermadictyon Roxb. (Robbrecht, 1982b: 1/(1)1)

Sphinctanthus Benth. (Persson, 1993: 2/(3)7)

Squamellaria Becc (Robbrecht, 1988a: 1/(3)3)

Stachyarrhena Hook. f. (Persson, 1993: 4/(10)11)

Staelia Cham. & Schltdl. (Dessein, 2003: 4/(12)15)

Stelechantha Bremek. (Bangoura, 1992: 2/(1)4; Huysmans, unpubl.: 1/(1)4)

Stephanococcus Bremek. (Dessein, unpubl.: 1/(1)1)

Stevensia Poit. (Huysmans, 1998: 1/(8)11)

Stilpnophyllum Hook. f. (Andersson, 1993: 2/(1)4)

Stipularia P. Beauv. (Huysmans et al., 1998a: 2/(-)2)

Strumpfia Jacq. (Igersheim, 1993a: 1/(1)1; Bridson & Robbrecht, 1985b: 1/(1)1)

Suberanthus Borhidi & M. Fernandez Zeq. (Borhidi & Fernandez-Zequeira, 1981b: ?/(7)5; Borhidi & Fernandez-Zequeira, 1983: ?/(7)5)

Sukunia A. C. Sm. (Persson, 1993: 1/(2)2)

Sulitia Merr. (Persson, 1993: 1/(-)1)

Syringantha Standl. (McDowell, 1996: 1/(1)1)

Tamilnadia Tirveng. (Persson, 1993: 1/(-)1)

Tammsia H. Karst. (Rova & Andersson, 1995: 1/(1)1)

Tapiphyllum Robyns (Havard & Verdcourt, 1987: 7/(20) 18; Igersheim, 1989: 2/(20) 18; Lens et al., 2000: 1/(20)18)

Tarenna Gaertn. (De Block & Robbrecht, 1998: 26/(180)219; De Block et al., 2001 [2002]: ?/ (180)219)

Tarennoidea Tirveng. (Persson, 1993: 1/(?)2)

Temnopteryx Hook. f. (Huysmans et al., 1998a: 1/(1)1)

Tennantia Verdc. (Verdcourt, 1981: 1/(-)1; De Block & Robbrecht, 1998: 1/(1)1)

Theligonum L. (Rutishauser et al., 1998: 1/(3)3; Behnke, 1975: 1/(3)3)

Timonius DC. (Darwin, 1993: 6/(150)180; Darwin, 1994: 18/(150)180)

Tobagoa Urb. (Dessein, 2003: 1/(1)1)

Tocoyena Aubl. (Persson, 1993: 3/(20)28)

Trailliaedoxa W. W. Smith & Forrest (Ying et al., 1993: 1/(1)1)

Tresanthera H. Karst. (Aiello, 1979: 1/(2)1; Delprete, 1999a: 1/(2)1)

Triainolepis Hook. f. (Dessein, unpubl.: 1/(8)11)

Tricalysia A. Rich. (Robbrecht, 1979, 1982a, 1983, 1987 give short comments on pollen)

Trichostachys Hook. f. (Piesschaert et al., 2000a: ?/(10)15)

Trukia Kaneh. (Puttock, 1992: 1/(5)5, as Randia fitzalanii; Persson, 1993: 1/(5)5)

Uncaria Schreb. (Huysmans, unpubl.: 3/(34)42; Verellen et al., submitted: 4/(34)42; Mathew & Philip, 1983: 1/(34)42)

Valantia L. (Huysmans, unpubl.: 1/(3-4)7)

Vangueria Juss. (Verdcourt, 1987: 2/(15)19; Igersheim, 1989: 4/(15)19; Lens et al., 2000: 2/ (15)19)

Vangueriella Verdc. (Tilney, 1986:1/(21) 18, as Canthium spinosum; Verdcourt, 1987:4/(21) 18; Igersheim, 1989: 2/(21)18)

Vangueriopsis Robyns (Verdcourt, 1987: 3/(4)4; Igerhseim, 1989: 2/(4)4; Lens et al., 2000: 1/ (4)4)

Versteegia Val. (De Block & Robbrecht, 1998: 2/(2)4; Bridson & Robbrecht, 1985b: 2/(2)4)

Vidalasia Tirveng. (Yirvengadum, 1998: 1/(-)5)

Virectaria Bremek. (Huysmans et al., 1998a: 2/(7)8; Dessein et al., 2001a: 6/(7)8)

Wendlandia Bartl. ex DC. (Huysmans, 1998: 2/(70)80; Mathew & Philip, 1983: 1/(70)80)

Wittmackanthus Kuntze (Andersson, 1993: 1/(1)1)

Xanthophytum Reinw. ex Blume (Tange, 1995: ?/(30)33).

XI. Appendix 3: List of Genera according to Tribes

This appendix gives the list of genera considered in this study, by tribe. The tribes are ordered by subfamily and therein as they appear on the summary cladogram starting with the basal taxa (Fig. 58). The delimitation of some tribes/groups as used here is still controversial and often based on relatively poor molecular evidence. These tribes are marked with an asterisk. Genera with an uncertain position are added at the end; these are not included in Table I, but their data will be found on the forthcoming pollen Internet site of the Laboratory of Plant Systematics, <>.


Coptosapelta; Luculia


Ophiorrhizeae--Colletoecema: Colletoecema; Neurocalyx; Ophiorrhiza; Xanthophytum

Pauridiantheae: Pauridiantha; Poecilocalyx; Rhipidantha; Stelechantha

Urophylleae: Amphidasya; Maschalocorymbus; Praravinia

Lasiantheae:* Lasianthus; Metabolos; Perama; Saldinia; Trichostachys

Cruckshanksieae: Cruckshanksia

Coussareeae:* Coussarea; Faramea

Coccocypseleae:* Coccocypselum; Declieuxia; Hindsia

Craterispermeae: /

Psychotrieae: Chassalia; Chazaliella; Geophila; Hymenocoleus; Myrmecodia; Palicourea; Rudgea; Psychotria; Squamellaria.

Note: The delimitation of several genera of the Psychotrieae is still in state of flux. Because it is remarkably variable in the tribe, pollen should be investigated to help determine generic boundaries.

Morindeae: Appunia; Caelospermum; Damnacanthus; Gentingia; Gynochthodes; Morinda; Motley(i)a; Pogonolobus; Prismatomeris; Rennellia

Schradereae: Schradera; Lecananthus; Leucocodon

Gaertnereae: Gaertnera; Pagamea

Danaideae: Danais; Payera; Schismatoclada

Hedyotideae--Pentas group:* Batopedina; Carphalea; Chamaepentas; Otiophora; Otomeria; Parapentas; Pentas

Knoxieae:* Calanda; Chlorochorion; Knoxia; Neopentanisia; Pentanisia

Triainolepideae:* Triainolepis

Hedyotideae--Hedyotis group:* Agathisanthemum; Amphiasma; Arcytophyllum; Conostomium; Dentella; Dibrachionostylus; Exallage; Gomphocalyx; Gouldia; Hedyotis; Hedythyrsus; Houstonia; Kohautia; Lelya; Manettia; Manostachya; Mitrasacmopsis; Neohymenopogon; Neanotis; Oldenlandia; Pentodon; Phylohydrax; Scleromitrion

Spermacoceae s.str.: Crusea; Diodella; Diodia; Emmeorhiza; Ernodea; Galianthe; Hydrophylax; Mitracarpus; Psyllocarpus; Richardia; Scandentia; Schwendenera; Spermacoce; Staelia; Tobagoa

Note." The Hedyotideae--Hedyotis group and the Spermacoceae s.str, form a monophyletic group, which is here called "Spermacoceae s.1."

Anthospermeae: Anthospermum; Carpacoce; Coprosma; Durringtonia; Galopina; Leptostigma; Nenax; Nertera; Normandia; Opercularia; Phyllis; Pomax

Argostemmateae:* Argostemma; Cyaneuron; Mouretia; Mycetia

Paederieae:* Aitchisonia; Gaillonia; Jaubertia; Kelloggia; Leptodermis; Paederia; Plocama; Pseudopyxis; Putoria; Saprosma; Serissa; Spermadictyon

Theligoneae: Theligonum

Rubieae: Asperula; Crucianella; Cruciata; Galium; Rubia; Sherardia; Valantia


Condamineeae-Calycophylleae-Hippotideae-complex:* Alseis; Bathysa; Calycophyllum; Capirona; Chimarrhis; Condaminea; Dialypetalanthus; Dioicodendron; Dolichodelphys; Elaeagia; Emmenopterys; Ferdinandusa; Hippotis; Macrocnemum; Mastixiodendron; Parachimarrhis; Pentagonia; Picardaea; Pinckneya; Pogonopus; Rustia; Semaphyllanthe; Simira; Sommera; Tammsia; Tresanthera; Wittmackanthus

Note: The delimitation of the Condamineeae, Calycophylleae, and Hippotideae is highly problematic. Rova (1999) suggests that the Hippotideae and Calycophylleae might be monophyletic, but their relation with the other genera of the complex is unknown. Until more data become available, we prefer not to split the complex into smaller entities.

Mussaendeae: Aphaenandra; Heinsia; Landiopsis; Mussaenda; Neomussaenda; Pseudomussaenda; Schizomussaenda

Sabiceeae:* Ecpoma; Pseudosabicea; Sabicea; Stipularia

Virectarieae:* Hekistocarpa; Virectaria

Sipaneeae: Limnosipanea; Sipanea; Sipaneopsis

Henriquezieae:* Gleasonia; Henriquezia; Platycarpum. Tentatively included: Molopanthera; Posoqueria.

Retiniphylleae: Retiniphyllum

Ixoreae: Captaincookia; Doricera; Ixora; Myonima; Versteegia. Tentatively included: Scyphiphora.

Vanguerieae: Ancyclanthos; Canthium; Cuviera; Fadogia; Fadogiella; Hutchinsonia; Keetia; Lagynias; Meyna; Multidentia; Pachystigma; Peponidium; Perakanthus; Psydrax; Pygmaeothamnus; Pyrostria; Rytigynia; Scyphochlamya; Tapiphyllum; Vangueria; Vangueriella; Vangueriopsis

Alberteae: Alberta; Nematostylis

Coffeeae: Bertiera; Coffea; Diplospora; Discospermum; Psilanthus; Sericanthe; Tricalysia

Octotropideae: Jovetia; Lamprothamnus; Octotropis; Paragenipa; Polysphaeria; Ramosmania

Pavetteae: Cladoceras; Coleactina; Coptosperma; Dictyandra; Homollea; Homolliella; Leptactina; Nichallea; Pachystylus; Paraeephaelis; Pavetta; Robbreehtia; Rutidea; Schizenterospermum; Tarenna; Tennantia

Gardenieae: Adenorandia; Aidia; Aidiopsis; Alibertia; Alleizettella; Amaioua; Atractoearpus; Atractogyne; Benkara; Borojoa; Brachytome; Brenania; Burchellia; Byrsophyllum; Caloehone; Casasia; Catunaregam; Ceriscoides; Coddia; Deccania; Didymosalpinx; Dioecreseis; Duperrea; Duroia; Euclinia; Fagerlindia; Ganguelia; Gardenia; Genipa; Glossostipula; Himalrandia; Hyperacanthus; Kailarsenia; Kochummenia; Kutchubaea; Larsenaikia; Maerosphyra; Mantalania; Massularia; Melanopsidium; Mitriostigma; Monosalpinx; Morelia; Oligoeodon; Oxyanthus; Oxyceros; Pelagodendron; Phellocalyx; Pleiocoryne; Porterandia; Preussiodora; Pseudomantalania; Randia; Riodocea; Rosenbergiodendron, Rothmannia; Rubovietnamia; Schumanniophyton; Sherbournia; Sphinctanthus; Stachyarrhena; Sukunia; Sulitia; Tamilnadia; Tarennoidea; Tocoyena; Trukia; Vidalasia. Tentatively included: Argocoffeopsis.

Note: Some general pollen comments on the tribe Gardenieae s.l. are given by Robbrecht and Puff (1986). The delimitation of several genera of the Gardenieae is still in state of flux. Because it is remarkably variable in the tribe, pollen should be investigated to help determine generic boundaries.


Cinchoneae: Cinchona; Dolieholobium; Joosia; Ladenbergia; Remijia; Stilpnophyllum

Isertieae: Isertia; Kerianthera; Raritebe; Schizostigma; Temnopteryx

PECC complex: Asernnantha; Badusa; Bikkia; Catesbaea; Ceratopyxis; Chiococca; Coutaportla; Coutarea; Erithalis; Exostema; Hintonia; Isidorea; Morierina; Nernstia; Osa; Portlandia; Salzmannia; Schmidtottia; Scolosanthus; Syringantha. Sister to PECC: Strumpfia. Pollen of this species differs from pollen of the PECC complex in the absence of supratectal elements.

Nancleeae (including Hymenodictyoneae): Adina; Adinauclea; Breonadia; Breonia; Burttdavya; Cephalanthus; Corynanthe; Cubanola; Gyrostipula; Haldina; Hallea; Hymenodictyon; Janotia; Ludekia; Metadina; Mitragyna; Myrmeconauclea; Nauclea; Neolamarekia; Neonauclea; Ochreinauclea; Paracorynanthe; Pausinystalia; Pertusadina; Sarcocephalus; Sinoadina; Unearia

Hillieae/Hamelieae: Cosmibuena; Hamelia; Hillia; Hoffmannia. Tentatively included: Balmea; Chione.

Rondeletieae:* Blepharidium; Roigella; Rondeletia; Suberanthus. Tentatively included: Aeunaeanthus; Glionnetia; Habroneuron; Rogiera (pro parte); Stevensia.

Guettardeae:* Arachnothryx; Chomelia; Gonzalagunia; Guettarda; Javorkaea; Timonius. Tentatively included: Diehilanthe.


Genera incertae sedis: Aeranthera; Airosperma; Aleisanthia; Aleisanthiopsis; Aoranthe; Aphanocarpus; Ariadne; Augusta; Calycosiphonia; Coryphothamnus; Crossopteryx; Dunnia; Greeniopsis; Jaekiopsis; Kajewskiella; Lathraeocarpa; Lindenia; Mitchella; Mussaendopsis; Myrioneuron; Pagameopsis; Petitiocoden; Plaeoearpa; Sacosperma; Stephanoeoccus; Trailliaedoxa; Wendlandia

VII. Acknowledgments

The authors wish to thank all former and present students at the Laboratory of Plant Systematics who studied Rubiaceae pollen and orbicules for their individual contributions to this review. Anja Vandeperre and Marcel Verhaegen are acknowledged for technical assistance.

This study was supported by grants from the Research Council of the Catholic University of Leuven (OT/01/25) and F.W.O.-Vlaanderen (G.0268.04). Steven Dessein and Stefan Vinckier are postdoctoral fellows of the F.W.O.-Vlaanderen. Suzy Huysmans gratefully acknowledges I.W.T. and F.W.O.-Vlaanderen for the personal grants that made it possible to focus on Rubiaceae pollen and orbicules for eight years. Visits by Helga Ochoterena to the Laboratory of Plant Systematics were financially supported by the F.W.O.-Vlaanderen and the Coimbra Group.

VIII. Literature Cited

Adams, L. G., D. M. Bridson & E. Robbrecht. 1987. The identity of Lasianthus graciliflorus Bailey (Rubiaceae). Kew Bull. 42: 209-214.

Aiello, A. 1979. A reexamination of Portlandia (Rubiaceae) and associated taxa. J. Arnold Arbor. 60: 38-126.

Ali, S. J. & E. Robbrecht. 1991. Remarks on the tropical Asian and Australian taxa included in Diplospora or Tricalysia (Rubiaceae-Ixoroideae-Gardenieae). Blumea 35: 279-305.

Anderson, W. R. 1972. A monograph of the genus Crusea (Rubiaceae). Mem. New York Bot. Gard. 22: 1-128.

--. 1973. A morphological hypothesis for the origin of heterostyly in the Rubiaceae. Taxon 22: 537-542.

Andersson, L. 1993. Pollen characteristics of the tribes Calycophylleae, Cinchoneae, and Hillieae (Rubiaceae). Nordic J. Bot. 13: 405-417.

-- & J. H. E. Rova. 1999. The rps 16 intron and the phylogeny of the Rubioideae (Rubiaceae). Pl. Syst. Evol. 214: 161-186.

Andreasen, K. & B. Bremer. 2000. Combined phylogenetic analysis in the Rubiaceae-Ixoroideae: Morphology, nuclear and chloroplast DNA data. Amer. J. Bot. 87: 1731-1748.

--, B. G. Baldwin & B. Bremer. 1999. Phylogenetic utility of the nuclear rDNA ITS region in subfamily Ixoroideae (Rubiaceae): Comparisons with cpDNA rbcL sequence data. Pl. Syst. Evol. 217: 119-135.

Andronova, N. N. 1984. The structure of the anther and pollen development in the Rubiaceae. Bot. Zhurn. 69: 43-54 (in Russian).

Baker, H. G. 1956. Pollen dimorphism in the Rubiaceae. Evolution 10: 23-31.

Bangoura, D. 1992. Beitrage zur Klarung der Gattungsabgrenzungsprobleme und Verwandtschafts-verhaltnisse in der Tribus Pauridiantheae (Rubiaceae). Ph.D. diss., Univ. Wien, Vienna.

Barrett, S. C. H. 1992. Heterostylous genetic polymorphisms: Model systems for evolutionary analysis. Pp. 1-29 in S. C. H. Barrett (ed.), Evolution and function of heterostyly. Monographs on Theoretical and Applied Genetics, 15. Springer-Verlag, Berlin.

Behnke, H. D. 1975. Electronenmikroskopische Untersuchungen zur Frage der verwandtschaftlichen Beziehungen zwischen Theligonum und Rubiaceae: Feinbau der Siebellement-Plastiden und Anmerkungen zur Struktur der Pollenexine. Plant. Syst. Evol. 123: 317-326.

Bir Bahadur. 1963. Heterostylism in Oldenlandia umbellata L. J. Genet. 58: 429-439.

--. 1966. Heterostyly in Oldenlandia scopulorum. Bull. J. Genet. 59: 267-272.

--. 1968a. Pollen dimorphism in three heterostyled Rubiaceae. Rev. Palaeobot. Palynol. 7: 233-239.

--. 1968b. Heterostyly in Rubiaceae: A review. Osmania Univ. J. Sci. [Golden Jubilee Vol.] 4: 207-238.

--. 1970. Heterostyly in Hedyotis nigricans (Lam.) Fosb. J. Genet. 60: 175-177.

--- & N. Rama Swamy. 1993. Pollen exine dimorphism in ten heterostylous species. J. Palynol. 29: 77-88.

Borhidi, A. 1982. Studies in Rondeletieae (Rubiaceae), III. The genera Rogiera and Arachnothryx. Acta Bot. Acad. Sci. Hung. 28: 65-71.

-- & M. Fernandez Zequeira. 1981a. Studies in Rondeletieae (Rubiaceae), I. A new genus: Roigella. Acta Bot. Acad. Sci. Hung. 27: 309-312.

-- & --. 1981b. Studies in Rondeletieae (Rubiaceae), II. A new genus: Suberanthus. Acta Bot. Acad. Sci. Hung. 27: 313-316.

-- & --. 1983. Studies in Rondeletieae (Rubiaceae), V. Los limites del genero Suberanthus. Acta Bot. Hung. 29: 29-34.

-- & M. Jarai-Komlodi. 1983. Studies in Rondeletieae (Rubiaceae), IV. A new genus: Javorkaea. Acta Bot. Hung. 29: 13-27.

-- & M. Moncada. 1980. Acunaeanthus, a new genus of Rubiaceae. Acta Bot. Acad. Sci. Hung. 26: 277-287.

Bosser, J. & D. Lobreau-Callen. 1998. Landiopsis Capuron ex Bosser, genre nouveau de Rubiaceae de Madagascar. Adansonia, ser. 3, 20: 131-137.

Bremekamp, C. E. B. 1952. The African species of Oldenlandia L. sensu Hiern et K. Schumann. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect., 48: 1-297.

--. 1963. On pollen dimorphism in heterostylous Psychotrieae, especially in the genus Mapouria Aubl. Grana Palynol. 4: 53-63.

--. 1966. Remarks on the position, the delimitation and the subdivision of the Rubiaceae. Acta Bot. Neerl. 15: 1-33.

Bremer, B. & M. Thulin. 1998. Collapse of Isertieae, re-establishment of Mussaendeae, and a new genus of Sabiceeae (Rubiaceae): Phylogenetic relationships based on rbcL data. Pl. Syst. Evol. 211: 71-92.

-- & J. F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Pl. Syst. Evol. 225: 43-72.

--, R. K. Jansen, B. Oxelman, M. Backlund, H. Lantz & K. J. Kim. 1999. More characters or more taxa for a robust phylogeny: Case study from the coffee family (Rubiaceae). Syst. Biol. 48: 413-435.

Bridson, D. M. & E. Robbrecht. 1985a. Validation of the African genus Hyperacanthus E. Mey. (Rubiaceae tribe Gardenieae). Kew Bull. 40: 273-286.

-- & --. 1985b. Further notes on the tribe Pavetteae (Rubiaceae). Bull. Jard. Bot. Belg. 55: 83-115.

--, P. Gasson & E. Robbrecht. 1980. Phellocalyx, a new tropical African genus in the Rubiaceae (Gardenieae). Kew Bull. 35: 315-321.

Buchner, R. & C. Puff. 1993. The genus complex Danais-Schismatoclada-Payera (Rubiaceae): Character states, generic delimitation and taxonomic position. Bull. Mus. Natl. Hist. Nat., B, Adansonia 15: 23-74.

Cabral, E. L. & N. M. Bacigalupo. 2001. Scandentia, nuevo genero de Rubiaceae-Spermacoceae. Darwiniana 39: 29-41.

Chaw, S. M. & S. P. Darwin. 1992. A systematic study of the paleotropical genus Antirhea (Rubiaceae: Guettardeae). Tulane Stud. Zool. Bot. 28: 25-118.

Chennaveeraiah, M. S. & P. M. Shivakumar. 1983. Pollen bud formation and its role in Ophiorrhiza spp. Ann. Bot. 5l: 449-452.

Chinnappa, C. C. & B. G. Warner. 1981. Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance. Bot. J. Linn. Soc. 83: 221-236.

-- & --. 1982. Pollen morphology in the genus Coffea (Rubiaceae), II. Pollen polymorphism. Grana 21: 29-37.

Clement, C. & J.-C. Audran. 1993a. Orbicule wall surface characteristics in Lilium (Liliaceae). Grana 32: 348-353.

-- & --. 1993b. Electron microscope evidence for a membrane around the core of the Ubisch body in Lilium (Liliaceae). Grana 32: 311-314.

Cortes-B., R. & S. Huysmans. Submitted. A survey of pollen and orbicules morphology of Retiniphyllum (Rubiaceae-Ixorideae). Rev. Palaeobot. Palynol.

Darwin, S. P. 1976. The genus Lindenia (Rubiaceae). J. Arnold. Arbor. 57: 426-449.

--. 1977. The genus Mastixiodendron (Rubiaceae). J. Arnold. Arbor. 58: 349-381.

--. 1979. A synopsis of the indigenous genera of Pacific Rubiaceae. Allertonia 2: 1-44.

--. 1980a. Notes on Airosperma (Rubiaceae), with a new species from Fiji. J. Arnold. Arbor. 61: 95-105.

--. 1980b. Habroneuron Standley, a little-known genus of Mexican Rubiaceae. Brittonia 32: 343-347.

--. 1993. A revision of Timonius subgenus Timonius (Rubiaceae: Guettardeae). Allertonia 7: 1-39.

--. 1994. Systematics of Timonius subgenus Abbottia (Rubiaceae: Guettardeae). Syst. Bot. Monogr. 42: 1-86.

De Block, P. 1997. Biosystematic studies in the tribe Pavetteae (Rubiaceae-Ixoroideae). Ph.D. diss., Univ. Instelling Antwerpen, Antwerp.

--. 1998. The African species of Ixora (Rubiaceae: Pavetteae). Opera Bot. Belg. 9: 1-218.

--. 2003. Robbrechtia, a new Rubiaceae genus from Madagascar. Syst. Bot. 28: 145-156.

-- & E. Robbrecht. 1998. Pollen morphology of the Pavetteae (Rubiaceae, Ixoroideae) and its taxonomic significance. Grana 37: 260-275.

--., J. Degreef & E. Robbrecht. 2001 [2002]. Reinstatement of the Afro-Malagasy genus Coptosperma (Rubiaceae, Ixoroideae, Pavetteae). Syst. Geogr. Pl. 71: 455-492.

Delprete, P. G. 1999a. Rondeletieae (Rubiaceae). Part I. Fl. Neotrop. Monogr. 77: 1-226.

--. 1999b. Riodocea (Rubiaceae, Gardenieae), a new genus from the Brazilian Atlantic forest. Brittonia 51: 15-23.

Dessein, S. 2003. Systematic studies in the Spermacoceae (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.

--, A. Scheltens, S. Huysmans, E. Robbrecht & E. Smets. 2000. Pollen morphological survey of Pentas (Rubiaceae-Rubioideae) and its closest allies. Rev. Palaeobot. Palynol. 112: 189-205.

--, S. Jansen, S. Huysmans, E. Robbrecht & E. Smets. 2001a. A morphological and anatomical survey of Virectaria (African Rubiaceae), with a discussion of its taxonomic position. Bot. J. Linn. Soc. 137: 1-29.

--, L. Andersson, E. Robbrecht & E. Smets. 2001b. Hekistocarpa (Rubiaceae): A member of an emended tribe Virectarieae. Pl. Syst. Evol. 229: 59-78.

--, S. Huysmans, E. Robbrecht & E. Smets. 2002a. Pollen of African Spermacoce species (Rubiaceae). Grana 41: 69-89.

--, S. Jansen, E. Robbrecht & E. Smets. 2002b [2004]. A new species of Spermacoce (Rubiaceae) from the Manika high plateau (Katanga; R. D. Congo). Nordic J. Bot. 22: 513-523.

--, S. Ntore, E. Robbrecht & E. Smets. 2003. Pollen and seeds reveal that Spermacoce thymoidea s. lat. (African Rubiaceae, Spermacoceae) represents three endemic or disjunct species from the Zambezian high plateaus. Syst. Bot. 28: 130-144.

--, L. Andersson, K. Geuten, E. Smets & E. Robbrecht. 2005a. Gomphocalyx and Phylohydrax (Rubiaceae): Sister taxa excluded from the Spermacoceae s.s., featuring a remarkable case of convergent evolution. Taxon 54: 91-107.

--, R. Harwood, S. Smets & E. Robbrecht. 2005b. Pollen of the Spermacoce species from the Northern Territory of Australia: Morphology and taxonomic significance. Austral. Syst. Bot. 18: 367-382.

D'hondt, C. 2002. Pollen--en orbiculemorfologie van Hillieae (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.

--, P. Schols, S. Huysmans & E. Smets. 2004. Systematic relevance of pollen and orbicule characters in the tribe Hillieae (Rubiaceae). Bot. J. Linn. Soc. 146: 303-321.

Di Maio, F. R. 1996 [1998]. Revisao taxonomica de genero Hindsia Bentham (Rubiaceae, Hedyotideae). Arch. Jar. Bot. Rio de Janeiro 34: 51-92.

El-Ghazaly, G. & S. Huysmans. 2001. Re-evaluation of a neglected layer in pollen wall development with comments on its evolution. Grana 40: 3-16.

-- & E. Smets. 2001. Pollen wall development of Rondeletia odorata (Rubiaceae). Amer. J. Bot. 88: 14-30.

Erdtman, G. 1971. Pollen morphology and plant taxonomy--Angiosperms. An Introduction to Palynology, I. Hafner Publishing, New York.

Es, K. 1999. Geophila (Psychotrieae, Rubiaceae): Revisie van de neotropische taxa. Licentiate thesis, Catholic Univ. of Leuven.

Farooq, M. & M. Inamuddin. 1969. The embryology of Oldenlandia nudicaulis Roth. J. Indian Bot. Soc. 48: 166-173.

Farris, S. S. 1990. Phenetics in camouflage. Cladistics 6: 91-100.

Groeninckx, L 2005. Zoektocht naar de taxonomische positie van Mitrasacmopsis (Rubiaceae) op basis van moleculaire en morfologische data. Licentiate thesis, Catholic Univ. of Leuven.

Gustafsson, C. G. R. 1998. The neotropical Rosenbergiodendron (Rubiaceae, Gardenieae). Brittonia 50: 452-466.

-- & C. Persson. 2002. Phylogenetic relationships among species of the neotropical genus Randia (Rubiaceae, Gardenieae) inferred from molecular and morphological data. Taxon 51: 661-674.

Hansson, T. & G. El-Ghazaly. 2000. Development and cytochemistry of pollen and tapetum in Mitriostigma axillare (Rubiaceae). Grana 39: 65-89.

Havard, A. & B. Verdcourt. 1987. A pollen survey of Tapiphyllum (Rubiaceae-Vanguerieae). Kew Bull. 42: 605-609.

Hesse, M. 1986. Orbicules and the ektexine are homologous sporopollenin concretions in Spermatophyta. P1. Syst. Evol. 153: 37-48.

Hooker, J. D. 1873. Ordo LXXXIV. Rubiaceae. Pp. 7-151 in G. Bentham & J. D. Hooker (eds.), Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata defirmata. London.

Huang, T. C. 1972. Pollen flora of Taiwan, VI. National Taiwan Univ., Botany Dept. Press, Taipei.

Huxley, C. R. & M. H. P. Jebb. 1993. The tuberous epiphytes of the Rubiaceae, 5. A revision of Myrmecodia. Blumea 37: 271-334.

Huysmans, S. 1993. De pollenmorfologie van de Coptosapelteae (Rubiaceae-Cinchonoideae). Licentiate thesis, Catholic Univ. of Leuven.

--. 1998. Palynology of the Cinchonoideae (Rubiaceae): Morphology and development of pollen and orbicules. Ph.D. diss., Catholic Univ. of Leuven.

--, E. Robbrecht & E. Smets. 1994. Are the genera Hallea and Mitragyna (Rubiaceae-Coptosapelteae) pollen morphologically distinct? Blumea 39: 321-340.

--, G. El-Ghazaly, S. Nilsson & E. Smets. 1997. Systematic value of tapetal orbicules: A preliminary survey of the Cinchonoideae (Rubiaceae). Canad. J. Bot. 75: 815-826.

--, E. Robbrecht & E. Smets. 1998a. A collapsed tribe revisited: Pollen morphology of the Isertieae (Cinchonoideae-Rubiaceae). Rev. Palaeobot. Palynol. 104: 85-113.

--, G. El-Ghazaly & E. Smets. 1998b. Orbicules in Angiosperms: Morphology, function, distribution, and relation with tapetum types. Bot. Rev. (Lancaster) 64: 240-272.

--, E. Robbrecht, P. Delprete & E. Smets. 1999. Pollen morphological support for the Catesbaeeae-Chiococceae-Exostema-complex (Rubiaceae). Grana 38: 325-338.

--, G. El-Ghazaly & E. Smets. 2000. Orbicules: Still a well hidden secret of the anther. Pp. 201-212 in B. Nordenstam, G. El-Ghazaly & M. Kassas (eds.), Plant systematics for the 21st century. Wenner-Gren International Series, 77. Portland Press, London.

--, S. Dessein, E. Smets & E. Robbrecht. 2003. Pollen morphology of NW European representatives confirms monophyly of Rubieae (Rubiaceae). Rev. Palaeobot. Palynol. 127: 219-240.

Igersheim, A. 1989. Beitrage zur Klarung der Gattungsabgrenzung-probleme innerhalb der Rubiaceae--Vanguerieae. Ph.D. diss., Botanical Institute, Vienna.

--. 1991. Palynological investigations of Paederia L. (Rubiaceae-Paederieae). Pp. 135-149 in C. Puff (ed.), The genus Paederia L. (Rubiaceae-Paederieae): A multidisciplinary study. Opera Bot. Belg., 3. National Botanic Garden of Belgium, Meise.

--. 1993a. The character states of the Caribbean monotypic endemic Strumpfia (Rubiaceae). Nordic J. Bot. 13: 545-559.

--. 1993b. The palynology of the genus Rondeletia L. (Rubiaceae-Cinchonoideae-Rondeletieae). Grana 32: 321-326.

-- & M. Weber. 1993 [1994]. "Pollen bud" formation in Ophiorrhiza (Rubiaceae). Pp. 51-59 in E. Robbrecht (ed.), Advances in Rubiaceae Macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.

Jansen, S. 1994. Een houtanatomische en palynologische studie van Psychotrieae (Rubiaceae-Rubioideae). Licentiate thesis, Catholic Univ. of Leuven.

--, E. Robbrecht & E. Smets. 1996a. The systematic value of endexine ornamentation in some Psychotrieae pollen (Rubiaceae-Rubioideae). Grana 35: 129-137.

--, H. Beeckman & E. Smets. 1996b. Gaertnera and Pagamea: Genera within the Psychotrieae or constituting the tribe Gaertnereae? A wood anatomical and palynological approach. Bot. Acta 109: 466-476.

Johansson, J. T. 1987a. Pollen morphology of the tribe Morindeae (Rubiaceae). Grana 26: 134-150.

--. 1987b. Revision of the genus Prismatomeris Thw. (Rubiaceae, Morindeae). Opera Bot. 94: 1-62.

--. 1992. Pollen morphology in Psychotria (Rubiaceae, Rubioideae, Psychotrieae) and its taxonomic significance: A preliminary survey. Opera Bot. 115: 1-71.

-- & K. M. Wong. 1988. The identity of Prismatomeris subsessilis King & Gamble (Rubiaceae, Rubioideae). Blumea 33: 351-356.

Jung-Mendacolli, S. L. 1984. Contribuicao ao estudo palinologico das Rubiaceae. Ph.D. diss., Univ. of Sao Paulo.

-- & T. S. Melhem. 1994. Flora polinica da reserva do parque estadual das fontes do Ipiranga (Sao Paulo, Brasil). Hoehnea 21: 131-155.

-- & --. 1995. Graos de polen de especies heterostilicas de Rubiaceae (Pollen grains of heterostylous Rubiaceae). Revista Brasil. Bot. 18: 61-93.

Keddam-Malplanche, M. 1980. Etude palynologique comparative des especes lianescentes dans les genres Sherbournia et Porterandia (Rubiacees-Gardeniees). Adansonia, ser. 2, 19: 429-434.

--. 1985. Le pollen et les stomates des Gardeniees (Rubiacee) du Gabon. Morphologie et tendances evolutives. Mem. Mus. Nation. Hist. Nat. Nouv. Ser., Ser. B, 29. Paris.

Kirkbride, J. H., Jr. 1979. Revision of the genus Psyllocarpus (Rubiaceae). Smithsonian Contr. Bot. 41: 1-32.

Lecuona Neumann, R. M., I. La Serna Ramos, B. Mendes Perez & W. Wildpret de la Torre. 1987. Contribucion al estudio palinologico de la flora endemica macronesica. Pollen & Spores 29: 359-390.

Lens, F., S. Jansen, S. Huysmans, E. Robbrecht & E. Smets. 2000. Pollen morphological variation in Vanguerieae (Ixoroideae-Rubiaceae). Grana 39: 1-13.

Leonard, J. 1984. Contribution a la connaissance de la flore de l'Iran, VI. Le "complexe Gaillionia A. Rich. ex DC." (Rubiaceae). Bull. Jard. Bot. Belg. 54: 493-497.

Leroy, J. F. 1974. Recherches sur les Rubiacees de Madagascar: Les genres Mantalania et Pseudo-mantalania (Gardeniees). Adansonia 14: 29-52.

Lewis, W. H. 1965a. Cytopalynological study of African Hedyotideae (Rubiaceae). Ann. Missouri Bot. Gard. 52: 182-211.

--. 1965b. Pollen morphology and evolution in Hedyotis subgenus Edrisia (Rubiaceae). Amer. J. Bot. 52: 257-264.

--. 1966. The Asian genus Neanotis nomen novum (Anotis) and allied taxa in the Americas (Rubiaceae). Ann. Missouri Bot. Gard. 53: 32-46.

Lobreau-Callen, D. 1978. L'aperture composee des Rubiaceae. Annales Mines Belg. 2: 167-173.

Lorence, D. H. 1986. Glossostipula (Rubiaceae), a new genus from Mexico and Guatemala. Candollea 41: 453-461.

Luna, I. & H. Ochoterena. 2004. Phylogenetic relationships of the genera of Theaceae based on morphology. Cladistics 20: 223-270.

Mabberley, D. J. 1997. The plant-book: A portable dictionary of the vascular plants. Ed. 2. Cambridge Univ. Press, Cambridge.

Malplanche, M. 1971. Etude palynologique de trois genres de Rubiacres--Gardeniees d'Afrique. Adansonia, ser. 2, 11: 343-355.

Mathew, P. M. & O. Philip. 1983. Studies in the pollen morphology of South Indian Rubiaceae. Vol. 10 of P. K. K. Nair (ed.), Advances in pollen-spore research. New Delhi: Today and Tomorrow's Printers.

-- & --. 1987. Developmental and systematic significance of pollen bud formation in Ophiorrhiza Linn. New Botanist 14: 47-54.

McDowell, T. 1996. Syringantha coulteri (Hooker f.) T. McDowell, a new combination, and remarks on the relationships of the monotypic Mexican genus Syringantha Standley (Rubiaceae). Novon 6: 273-279.

Motley T. J., K. J. Wurdack & P. G. Delprete. 2005. Molecular systematics of the Catesbaeeae-Chiococceae complex (Rubiaceae): Flower and fruit evolution and biogeographic implications. Amer. J. Bot. 92: 316-329.

Muller, J. 1981. Fossil pollen records of extant angiosperms. Bot. Rev. (Lancaster) 47: 1-142.

Naiki, A. & H. Nagamasu. 2003. Distyly and pollen dimorphism in Damnacanthus (Rubiaceae). J. Pl. Res. 105-113.

Negron-Ortiz, V. 1996. Reproductive biology of Ernodea (Rubiaceae-Spermacoceae) in the Bahamas and Puerto Rico. Pp. 403-412 in E. Robbrecht, C. Puff & E. Smets (eds.), Second Int. Rubiaceae Conf., Proceedings. Opera Bot. Belg., 7. National Botanic Garden Belgium, Meise.

Nilsson, S. & J. Praglowski. 1992. Erdtman's handbook of palynology. Ed. 2. Munksgaard, Copenhagen.

Ntore, S. 2004. Contribution a la connaissance systematique du genre afrotropical Pauridiantha (Rubiaceae). Ph.D. diss., Catholic Univ. of Leuven.

--, P. De Block, S. Huysmans, E. Robbrecht & S. Dessein. 2003. Two new species from Gabon show the need to reduce Commitheca to the synonymy of Pauridiantha (Rubiaceae, Pauridiantheae). Bot. J. Linn. Soc. 141: 105-117.

Ochoterena, H. 2000. Systematics of Hintonia Bullock and the Portlandia complex (Rubiaceae). Ph.D. diss., Cornell Univ.

Pacini, E. & G. G. Franchi. 1993. Role of the tapetum in pollen and spore dispersal. Pl. Syst. Evol., Suppl. 7: 1-11.

--, G. G. Franchi & M. Hesse. 1985. The tapetum: Its form, function and possible phylogeny in embryophyta. Pl. Syst. Evol. 149: 155-185.

Pailler, T. & J. D. Thompson. 1997. Distyly and variation in heteromorphic incompatibility in Gaertnera vaginata (Rubiaceae) endemic to La Reunion Island. Amer. J. Bot. 84: 315-327.

Persson, C. 1993. Pollen morphology of the Gardenieae-Gardeniinae (Rubiaceae). Nordic J. Bot. 13: 561-582.

Philip, O. & P. M. Mathew. 1975. Cytology of exceptional development of the male gametophyte in Ophiorrhiza mungos. Canad. J. Bot. 53: 2032-2037.

Piesschaert, F. 2001. Carpology and pollen morphology of the Psychotrieae (Rubiaceae-Rubioideae): Towards a new tribal and generic delimitation. Ph.D. diss., Catholic Univ. of Leuven.

--, S. Jansen, S. Huysmans, E. Smets & E. Robbrecht. 1999a. Chassalia petitiana (Rubiaceae-Psychotrieae), an overlooked epiphytic species hidden in the African canopy. Syst. Bot. 24: 315-322.

--, E. Robbrecht, A. D. Poulsen & E. Smets. 1999b. Pyrene and pollen observations in the pantropical genus Geophila (Rubiaceae-Psychotrieae). Nordic J. Bot. 19: 93-100.

--, -- & E. Smets. 1999c. Chassalia subcordatifolia, a new combination in African Rubiaceae (Rubioideae, Psychotrieae). Syst. Geogr. Pl. 69: 189-194.

--, L. Andersson, S. Jansen, S. Dessein, E. Robbrecht & E. Smets. 2000a. Searching for the taxonomic position of the African genus Colletoecema (Rubiaceae): Morphology and anatomy compared to an rps 16-intron analysis of the Rubioideae. Canad. J. Bot. 78: 288-304.

--, S. Huysmans, I. Jaimes, E. Robbrecht & E. Smets. 2000b. Morphological evidence for an extended tribe Coccocypseleae (Rubiaceae-Rubioideae). Pl. Biol. 2: 536-546.

--, S. Jansen, I. Jaimes, E. Robbrecht & E. Smets. 2001. Morphology, anatomy, and taxonomic position of Pagameopsis (Rubiaceae-Rubioideae). Brittonia 53: 490-504.

Pire, S. M. 1996. Palynological study of American species of Borreria (Rubiaceae-Spermacoceae). Pp. 416-423 in E. Robbrecht, C. Puff & E. Smets (eds.), Second Int. Rubiaceae Conf., Proceedings. Opera Bot. Belg., 7. National Botanic Garden Belgium, Meise.

--. 1997a. Genero Galianthe subg. Ebelia (Rubiaceae: Spermacoceae): Estudio palinologico. Ann. Missouri Bot. Gard. 84: 878-887.

--. 1997b. The pollen of Brazilian species of Richardia L. (Rubiaceae-Spermacoceae). Rev. Univ. Guarulhos, Geosiencias II Sp. Vol.: 184-191.

-- & E. L. Cabral. 1992. El valor del polen en la revalidacion de Galianthe (Spermacoceae-Rubiaceae). Darwiniana 31: 1-10.

Puff, C. 1986a. A biosystematic study of the African and Madagascan Rubiaceae. Anthospermae. Springer-Verlag, Vienna

--. 1986b. Phylohydrax (Rubiaceae-Spermacoceae)--A new genus to accommodate the African and Madagascan "Hydrophylax" species. Pl. Syst. Evol. 154, 343-366.

--. 1988. Observations on Carphalea Juss. (Rubiaceae, Hedyotideae), with particular reference to the Madagascan taxa and its taxonomic position. Bull. Jard. Bot. Belg. 58: 271-323.

--. 1989. The affinities and relationships of the Japanese endemic Pseudopyxis (Rubiaceae: Paederieae). Pl. Spec. Biol. 4: 145-155.

-- & R. Buchner. 1998. Lecananthus and Leucocodon, two genera to be added to the tribe Schradereae (Rubiaceae). Blumea 43: 265-286.

-- & A. Igersheim. 1994a. The character states and taxonomic position of Metabolos B1. (syn. Allaeophania Thw.) (Rubiaceae). Bull. Jard. Bot. Belg. 63: 241-262.

-- & --. 1994b. The character states of Mussaendopsis Baill. (Rubiaceae-Coptosapelteae). Flora 189: 161-178.

-- & E. Robbrecht. 1988. The taxonomic position of the Australian endemic Durringtonia (Rubiaceae). Austral. Syst. Bot. 1: 191-197.

-- & --. 1989. A survey of the Knoxieae (Rubiaceae-Antirheoideae). Bot. Jahrb. Syst. 110: 511-558.

-- & U. Rohrhofer. 1993 [1994]. The character states and taxonomic position of the monotypic mangrove genus Scyphiphora (Rubiaceae). Pp. 143-172 in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.

--, E. Robbrecht & V. Randrianasolo. 1984. Observations on the SE African-Madagscan genus Alberta and its ally Nematostylis (Rubiaceae, Alberteae), with a survey of the species and a discussion of the taxonomic position. Bull. Jard. Bot. Belg. 54: 293-366.

--, L. Andersson, U. Rohrhofer & A. Igersheim. 1993a. The tribe Schradereae (Rubiaceae) reexamined. Bot. Jahrb. Syst. 114: 449-479.

--, A. Igersheim & U. Rohrhofer. 1993b. Pseudomussaenda and Schizomussaenda (Rubiaceae): Close allies of Mussaenda. Bull. Jard. Bot. Belg. 62: 35-68.

--, R. Buchner & A. Igersheim. 1996. Dichilanthe, an unusual Asiatic Rubiaceae with "Lonicera flowers" and "dipterocarp fruits." Nordic J. Bot. 16:145-164.

Punt, W., S. Blackmore, S. Nilsson & A. Le Thomas. 1994. Glossary of pollen and spore terminology. LPP Foundation, Utrecht. (Updated online version: <>.)

Puttock, C. F. 1992. Systematics of the Australian Gardenieae (Rubiaceae). Ph.D. diss., Univ. of New South Wales.

Ramam, S. S. 1954. Gametogenesis and fertilization of Stephegyne parviflora Korth. Agra Univ. J. Res., Sci. 3: 243-348.

Razafimandimbison, S. G. & B. Bremer. 2001 [2002]. Tribal delimitation of Naucleeae (Cinchonoideae, Rubiaceae): Inference from molecular and morphological data. Syst. Geogr. Pl. 71: 515-538.

Reitsma, T. 1969. Size modifications of recent pollen grains under different treatments. Rev. Palaeobot. Palynol. 9: 175-202.

Robbrecht, E. 1977. The tropical African genus Hymenocoleus (Rubiaceae-Psychotrieae): Additions. Bull. Jard. Bot. Belg. 47: 3-29.

--. 1978a. Sericanthe, a new African genus of Rubiaceae (Coffeeae). Bull. Jard. Bot. Belgique 48: 3-78.

--. 1978b. Some observations in Preussiodora Keay (African Rubiaceae, Gardenieae). Bull Soc. Roy. Bot. Belgique 111: 3-9.

--. 1979. The African genus Tricalysia A. Rich. (Rubiaceae-Coffeeae), 1. A revision of the species of subgenus Empogona. Bull. Jard. Bot. Belg. 49: 239-360.

--. 1980. Bijdragen tot de classificatie van de Ixoroideae en tot de revisie van Tricalysia s.1. (Rubiaceae). Ph.D. diss., Univ. Gent.

--. 1981a. Studies in tropical African Rubiaceae, I. 1. A new species of Tricalysia subgenus Empogona from Gabon. 2. The position of "Tricalysia auriculata" and "Neorosea adamii." 3. A new species of Batopedina (Hedyotideae) from Shaba. 4. The distribution of Batopedina and Parapentas. Bull. Jard. Bot. Belg. 51: 165-189.

--. 1981b. Studies in tropical African Rubiaceae, II. 5. A survey of Argocoffeopsis. 6. A revision of Calycosiphonia. Bull. Jard. Bot. Belg. 51: 359-378.

--. 1982a. The African genus Tricalysia A. Rich. (Rubiaceae-Coffeeae), 2. Ephedranthera, a new section of subgenus Tricalysia. Bull. Jard. Bot. Belg. 52: 311-339.

--. 1982b. Pollen morphology of the tribes Anthospermeae and Paederieae (Rubiaceae) in relation to taxonomy. Bull. Jard. Bot. Belg. 52: 349-366.

--. 1983. The African genus Tricalysia A. Rich. (Rubiaceae), 3. Probletostemon revived as a section for subgenus Tricalysia. Bull. Jard. Bot. Belg. 53: 299-320.

--. 1984. The delimitation and taxonomic position of the tropical African genera Leptactina and Dictyandra (Rubiaceae). Pl. Syst. Evol. 145: 105-118.

--. 1985. Further observations on the pollen morphology of the South African genus Carpacoce (Rubiaceae-Anthospermeae). Rev. Palaeobot. Palynol. 45: 361-371.

--. 1987. The African genus Tricalysia A. Rich. (Rubiaceae), 4. A revision of the species of section Tricalysia. Bull. Jard. Bot. Belg. 57: 39-208.

--. 1988a. Tropical woody Rubiaceae: Characteristic features and progressions: Contributions to a new subfamilial classification. Opera Bot. Belg., 1. National Botanic Garden of Belgium, Meise.

--. 1988b. Studies in tropical African Rubiaceae, 13. Petitiocodon, a new genus to accommodate Didymosalpinx parviflora (Gardenieae-Diplosporinae). Bull. Jard. Bot. Belg. 58: 109-120.

--. 1993 [1994]. Supplement to the 1988 outline of the classification of the Rubiaceae. Index to genera. Pp. 173-196 in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.

-- & D. M. Bridson. 1984. The taxonomic position of the East African genus Cladoceras (Rubiaceae). Bull. Soc. Roy. Bot. Belgique 117: 247-251.

-- & C. Puff. 1981. Mericocalyx Bamps, synonymous with Otiophora Zucc. (Rubiaceae). Bull. Jard. Bot. Nat. Belg. 51: 143-151.

-- & --. 1986. A survey of the Gardenieae and related tribes. Bot. Jahrb. Syst. 108: 63-138.

-- & A. Igersheim. 1991. Evidence for the close alliance between Mitchella and Damnacanthus, with comments on the campylotropy in the Rubiaceae and the circumscription of the Morindeae. Blumea 35: 307-345.

--, U. Rohrhofer & C. Puff. 1993 [1994]. A survey of Bertiera (Rubiaceae), including a discussion of its taxonomic position. Pp. 101-141 in E. Robbrecht (ed.), Advances in Rubiaceae macrosystematics. Opera Bot. Belg., 6. National Botanic Garden of Belgium, Meise.

--, S. Huysmans & E. Figueiredo. 1996. The generic status of Oxyanthus gossweileri (Rubiaceae) in Angola. S. African J. Bot. 62: 17-22.

Rogers, G. K. 1984. Gleasonia, Henriquezia, and Platycarpum (Rubiaceae). Fl. Neotrop. Monogr. 39: 1-135.

Rova, J. 1999. The Condamineeae Rondeletieae Sipaneeae complex (Rubiaceae). Ph.D. diss., Univ. Goteborg.

-- & L. Andersson. 1995. A reevaluation of the tribes Hippotideae and Tammsieae (Rubiaceae). Nordic J. Bot. 15: 269-284.

--, P. G. Delprete, L. Andersson & V. A. Albert. 2002. A trnL-F cpDNA sequence study of the Condamineeae-Rondeletieae-Sipaneeae complex with implications on the phylogeny of the Rubiaceae. Am. J. Bot. 89: 145-159.

Rutishauser, R., L. P. Ronse Decraene, E. Smets & I. Mendoza-Heuer. 1998. Theligonum cynocrambe: Developmental morphology of a peculiar rubiaceous herb. Pl. Syst. Evol. 210: 1-24.

Scheltens, A. 1998. Pollenmorfologische studie van de Afrikaanse Hedyotideae (Rubiaceae). Licentiate thesis, Catholic Univ. of Leuven.

Schumann, K. 1891. Rubiaceae. Pp. 1-156 in A. Engler & K. Prantl (eds.), Die naturlichen Pflanzenfamilien. Engelmann, Leipzig.

Sotolongo Molina, L., M. Fernandez Zequeira & P. Herrera Oliver. 2002. Pollen morphology of some Cuban Guettarda species (Rubiaceae: Guettardeae). Grana 41: 142-148.

Stoffelen, P., E. Robbrecht & E. Smets. 1997. Pollen morphology of Coffea and Psilanthus (Rubiaceae-Coffeeae), mainly from Africa. Grana 36: 313-327.

Strait, D. S., M. A. Moniz & P. T. Strait. 1996. Finite mixture coding: A new approach to coding continuous characters. Syst. Biol. 45: 67-78.

Swiderski, D. L., M. L. Zelditch & W. L. Fink. 1998. Why morphometrics is not special: Coding quantitative data for phylogenetic analyses. Syst. Biol. 47: 508-519.

Tange, C. 1994. Neomussaenda (Rubiaceae), a new genus from Borneo. Nordic J. Bot. 14: 495-500.

--. 1995. The identity of Siderobombyx and a new species of Xanthophytum (Rubiaceae). Nordic J. Bot. 575-581.

--. 1996a. Studies in SE Asiatic Rondeletieae, I: The West Malaysian endemic genus Aleisanthia (Rubiaceae). Nordic J. Bot. 16: 571-578.

--. 1996b. Studies in SE Asiatic Rondeletieae, II: Aleisanthiopsis (Rubiaceae), a new genus from Borneo. Nordic J. Bot. 16: 563-570.

--. 1997. A revision of the genus Mouretia (Rubiaceae). Nordic J. Bot. 123-132.

--. 1998. Cyaneuron (Rubiaceae), a new genus from Borneo and Sulawesi. Nordic J. Bot. 147-158.

Terrell, E. E., W. H. Lewis, H. Robinson & J. W. Nowicke. 1986. Phylogenetic implications of diverse seed types, chromosome numbers and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot., 73: 103-115.

Thiele, K. 1993. The holy grail of the perfect character: The cladistic treatment of morphometric data. Cladistics 9: 275-304.

Tilney, P. M. 1986. The taxonomic significance of anatomical and morphological characters in the southern African species of Canthium Lam. (Rubiaceae). Ph.D. diss., Univ. of Pretoria.

-- & A. E. Van Wyk. 1997. Pollen morphology of Canthium, Keetia and Psydrax (Rubiaceae: Vanguerieae) in southern Africa. Grana 36: 249-260.

Tirvengadum, D. D. 1982. Ramosmania, a new monotypic genus of Mascarene Rubiaceae. Nordic J. Bot. 2: 323-327.

--. 1984. Glionnetia, nouveau genre de Rubiacees (Rondeletiees) des Seychelles. Bull. Mus. Natl. Hist. Nat., B, Adansonia 6: 197-205.

--. 1993. Larsenaikia, a new genus of the Rubiaceae from Australia. Nordic J. Bot. 13: 175-184.

--. 1998. Novelties in Rubiaceae from the limestone flora of southeast Asia. Biogeographica (The Hague) 74: 163-175.

-- & E. Robbrecht. 1985. Remarks on three Hypobathreae from Rodrigues, Seychelles and Sri Lanka. Nordic J. Bot. 5: 455-461.

Tissot, C. & R. W. J. M. van der Ham. 1994. Seventh bibliographic index to the pollen morphology of Angiosperms. Trav. Sect. Sci. Techn. Inst. Franc. Pondicherie 27: 1-345.

--, H. Chikhi & T.S. Nayar. 1994. Pollen of wet evergreen forests of the western Ghats, India. Trav. Sect. Sci. Techn. Inst. Franc. Pondicherie 35: 1-133.

Van Campo, M. 1976. Patterns of pollen morphological variation within taxa. Pp. 125-137 in I. K. Ferguson & J. Muller (eds.), The evolutionary significance of the exine. Linnean Soc. Syrup. ser. 1. Acad. Press, London.

--. 1978. La face interne de l'exine. Rev. Palaeobot. Palynol. 26: 301-311.

Vanthournout, S. 2002. Geophila. Revisie van de paleotropische taxa. Licentiate thesis, Catholic Univ. of Leuven.

Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. Etat. 28: 209-290.

--. 1980. A conspectus of Polysphaeria (Rubiaceae). Kew Bull. 35: 97-130.

--. 1981. Notes on African Rubiaceae. Kew. Bull. 36: 493-557.

--. 1983. Notes on Mascarene Rubiaceae. Kew Bull. 37: 521-574.

--. 1987. Notes on African Rubiaceae-Vanguerieae. Kew Bull. 42: 123-199.

Verellen, J. 2002. Palynologische studie en revisie van Coptosapelta (Rubiaceae). Licentiate thesis, Catholic Univ. of Leuven.

--, E. Smets & S. Huysmans. 2004. The remarkable genus Coptosapelta (Rubiaceae): Pollen and orbicule morphology and systematic implications. J. Pl. Res. 117: 57-68.

--, S. Huysmans, S. G. Razafimandimbison & E. Smets. Submitted. Pollen morphology of the tribe Naucleeae s. lat. (Rubiaceae-Cinchonoideae) and its taxonomic and phylogenetic significance. Bot. J. Linn. Soc.

Vinckier, S., S. Huysmans & E. Smets. 2000. Morphology and ultrastructure of orbicules in the subfamily Ixoroideae (Rubiaceae). Rev. Palaeobot. Palynol. 108: 151-174.

Weber, M. & A. Igersheim. 1994. "Pollen buds" in Ophiorrhiza (Rubiaceae) and their role in pollenkitt release. Bot. Acta 107: 257-262.

Wodehouse, R. P. 1935. Pollen grains. McGraw-Hill, New York.

Ying, T.-S., Y.-L Zhang & D. E. Boufford. 1993. The endemic genera of seed plants of China. Science Press, Beijing.

Young, B. A., J. Schulz-Schaeffer & T. W. Caroll. 1979. Anther and pollen development in male-sterile intermediate wheatgrass plants derived from wheat x wheatgrass hybrids. Canad. J. of Bot. 57: 602-618.
Table I
Some variable pollen morphological characters for the tribes
of the family Rubiaceae

Key. D. U. = dispersal unit; M = monad; T = tetrad; Ma = massulae.
Size: very small <20 [micro]m; small 20-30 um; medium
30-40 [micro]m; large 40-65 [micro]m; very large >65 [micro]m.
Shape: O = oblate; SO = suboblate; S = spheroidal; SP = subprolate;
P = prolate. N. A. = number of apertures. Aperture position:
Z = zonoaperturate; P = pantoaperturate; S = spiraperturate.
Aperture type: PR = pororate; CPR = colporate; CP = colpate;
PO = porate. Endoaperture: Ep = endoporus; Eco = endocolpus;
Eci = endocingulum. Sexine: ar = areolate; cla = clavate;
dre = double reticulum; fo = foveolate; ha = hamulate;
[micro]re = microreticulate; pe = perforate; psi = psilate;
re = reticulate; ru = rugulate; si = sinuate; str = striate.
/ = no data. - = absent; + = present. + * = present on
infrareticulum. Doubtful data are preceded by a question
mark and are further discussed in the database.

Genus or tribe D. U. Size Shape

Unresolved or basal position

 Coptosapelta M Very small O to S
 to medium

 Luculia M Small SO to S
 to medium

 Ophiorrhizeae M Very small to SO to SP

 Pauridiantheae M Very small SO to SP,
 O & P

 Urophylleae M Very small to SO-S

 Lasiantheae M Small to very S

 Cruckshanksieae M Medium to large S

 Coussareeae M Small to medium O-SO

 Coccocypseleae M Medium to very S

 Craterispermeae / / /

 Psychotrieae M Small to very O to S,
 large rarely SP

 Morindeae M Small to (very) O to S,
 large rarely P

 Schradereae M Small to SO to SP
 rarely very

 Gaertnereae M Small to large O to SO

 Danaideae M Very small to S

 Hedyotideae- M Small to large, S to SP
 Pentas rarely very

 Knoxieae M Small to S

 Triainolepideae M Medium S

 Hedyotideae- M Very small to (SO)S to SP
 Hedyotis medium,

 Spermacoceae M Very small O to SP
 s.str. to very

 Anthospermeae M Small to O to S

 Argostemmateae M Very small to S

 Paederieae M Medium to SO to P

 Theligoneae M Small SO

 Rubieae M Very small S to SP
 to small,

Ixoroideae s. lat.

 Condamineeae- M Very small to SO to S,
 Hippotideae- medium, rarely SP
 Simireae- rarely or SO
 Calycophylleae large

 Mussaendeae M Very small to SO to S

 Sabiceeae M Very small to O to S

 Virectarieae M Medium to S to P

 Sipaneeae M Very small to SP to S

 Henriquezieae M/T Very small O to S
 to very

 Retiniphylleae M Small to S to P

 Ixoreae M Very small to S, rarely
 large SP or SO

 Vanguerieae M, Small to SO to S,
 [sup.?] large, rarely SP
 Ma rarely
 very large

 Alberteae M Medium S-SP
 to large

 Coffeeae M Very small to SO to SP

 Octotropideae M Very small to SO-S

 Pavetteae M Very small to SO-SP

 Gardenieae M/T/ Very small SO to S,
 Ma to medium, rarely
 rarely O or SP
s. str.

 Cinchoneae M Very small SO to SP
 to small,

 Isertieae M Small SO to S
 to large

 PECC M Very small SO to SP,
 to medium, rarely O

 Naucleeae M Very small to SO to S,
 small rarely SP

 Hilheae/ M Very small to S to SP
 Hamelieae large

 Rondeletieae M Very small to S, rarely SP

 Guettardeae M Very small to SO-S

Genus or tribe N. A. Position type

Unresolved or basal position

 Coptosapelta 3-10 Z & P PR

 Luculia 3 Z CPR


 Ophiorrhizeae 3-4 Z CPR

 Pauridiantheae 3 Z CPR or
 Urophylleae 3 Z CPR

 Lasiantheae (2-)3- Z CPR,
 4(-5) [sup.?]CP

 Cruckshanksieae 3 Z CPR

 Coussareeae 2-4 Z PO

 Coccocypseleae 3 Z CPR

 Craterispermeae / / /

 Psychotrieae 0-5(-150) Z & P PO, CPR,
 Morindeae 3(-4-6) Z CPR

 Schradereae 2-3(-4) Z Porate or

 Gaertnereae (2-)3(-4) Z CPR,
 rarely PO

 Danaideae 3-4(-5) Z CPR

 Hedyotideae- 3-4(-5) Z CPR

 Knoxieae [3.sup.?] Z CPR

 Triainolepideae 3 Z CPR

 Hedyotideae- (2-)3-4 Z CPR or
 Hedyotis (-5-12) [sup.?]CP

 Spermacoceae 3-30 Z & P CPR or
 s.str. PR

 Anthospermeae [sup.?]0/ Z CPR

 Argostemmateae 3(-4) Z CPR or
 Paederieae (2-)3 Z CP,
 (-4-5) [sup.?]CPR

 Theligoneae 4-8 Z Porate

 Rubieae 5-13 Z CP

Ixoroideae s. lat.

 Condamineeae- (2-)3(-4) Z CPR,
 Hippotideae- rarely PO
 Simireae- or PR

 Mussaendeae 3-4(-5) Z CPR,

 Sabiceeae (2-)3- Z CPR, [sup.?]
 4(-5) CP, PR

 Virectarieae 3 Z CPR

 Sipaneeae 3-4 Z CPR

 Henriquezieae 3-5(-6) Z CPR or PR

 Retiniphylleae 3 Z CPR

 Ixoreae 3 Z CPR

 Vanguerieae (2-)3(-4) Z Po(ro)rate,

 Alberteae 3 Z CPR

 Coffeeae (2-)3-5 Z(S) CPR

 Octotropideae 3 Z CPR

 Pavetteae 3(-4) Z CPR

 Gardenieae 3(-4-5), Z(P) Po(ro)rate,
 rarely up also CPR
 to >10

s. str.

 Cinchoneae 3-4(-5) Z CPR

 Isertieae 3-4(-5) Z CPR

 PECC 3((-4)) Z CPR or

 Naucleeae 3((-4)) Z CPR

 Hilheae/ 3 Z CPR

 Rondeletieae 3-4(-5); Z CPR
 [sup.?] 1

 Guettardeae [sup.?] Z CPR or
 (0-2-) PR

Genus or tribe aperture Annulus Costa

Unresolved or basal position

 Coptosapelta Ep + -/+

 Luculia Eco + +


 Ophiorrhizeae / - -

 Pauridiantheae Eco or ?- - -

 Urophylleae Eco or / /
 Lasiantheae Eco or ?- / /

 Cruckshanksieae Eco / /
 Coussareeae / + -

 Coccocypseleae Eco or - /
 Craterispermeae / / /
 Psychotrieae / - +/-

 Morindeae / - /

 Schradereae / +/- /

 Gaertnereae / - +

 Danaideae / - -

 Hedyotideae- Eco or -/+ -
 Pentas Eci

 Knoxieae Eco - /

 Triainolepideae Eci / /

 Hedyotideae- Eco or
 Hedyotis Eci

 Spermacoceae Eco, EP - -
 s.str. or Eci

 Anthospermeae Eco or - -

 Argostemmateae Eco -/+ /

 Paederieae - - -

 Theligoneae / - -

 Rubieae - - -

Ixoroideae s. lat.

 Condamineeae- Eco or -/+ -/+
 Hippotideae- Eci

 Mussaendeae Eco +(-) +(-)

 Sabiceeae Eco or - - -

 Virectarieae Eco - -

 Sipaneeae Eci / -

 Henriquezieae [sup.?]Eco

 Retiniphylleae Eco -/+ -

 Ixoreae Eco or +(-) -

 Vanguerieae EP or -, -/+ +(-)

 Alberteae / +/- /

 Coffeeae / +/- /

 Octotropideae Eco -/+ -/+

 Pavetteae Eco, -/+ -/+

 Gardenieae Eco or -/+ -/+

s. str.

 Cinchoneae Eco or - -

 Isertieae Eco - -

 PECC Eco or - -
 with ex-

 Naucleeae Eco, -/+ -/+

 Hilheae/ Eco or- -/+ +

 Rondeletieae Eci - -

 Guettardeae Eci or -/+ +/-

 Endo- Supra
Genus or tribe pattern Tectum elem.

Unresolved or basal position

 Coptosapelta Droplet fo-([micro]) -

 Luculia Granular re -


 Ophiorrhizeae / pe-([micro])re -

 Pauridiantheae Granular pe-([micro])re -

 Urophylleae / pe- [micro]re -

 Lasiantheae / psi-pe-[micro] -/+

 Cruckshanksieae / re -/+

 Coussareeae / pe-[micro]re -

 Coccocypseleae Granular dre +*

 Craterispermeae / / /

 Psychotrieae / pe-([micro]) -/+

 Morindeae / ([micro])re -/+

 Schradereae / pe-([micro])re -/+

 Gaertnereae Granular, ([micro])re -/+
 with en-
 Danaideae Granular ([micro])re -/+

 Hedyotideae- Granular, pe-([micro])re -/+
 Pentas often with

 Knoxieae Granular ([micro])re -/+
 with en-

 Triainolepideae Granular pe +

 Hedyotideae- Granular, pe-([micro])re, -/+
 Hedyotis often with often dr,
 endocracks rarely si
 to ar

 Spermacoceae Granular, pe-([micro])re +/(-)
 s.str. often with -psi, dre,
 endocracks rarely to,

 Anthospermeae Granular, ru, cla +
 often with pe-ru, also
 endocracks pe

 Argostemmateae Granular (u)re, -/+
 sometimes dre

 Paederieae / pe-(u)re +(-)
 rarely ru
 or fo

 Theligoneae / [micro] re +

 Rubieae Granular, pe +
 with en-

Ixoroideae s. lat.

 Condamineeae- Granular pe-([micro])re [-.sup.?] (+)
 Hippotideae- nexine
 Simireae- islands

 Mussaendeae Granular psi-pe- -

 Sabiceeae Granular, pe-([micro])re -

 Virectarieae Granular pe -

 Sipaneeae / pe-[micro]re -

 Henriquezieae Smooth ([micro])re -

 Retiniphylleae Granular pe-([micro])re -

 Ixoreae Granular, pe-([micro])re, -
 sometimes rarely
 endocracks atectate

 Vanguerieae Smooth or pe-([micro])re, -(+)
 granular rarely fo
 or psi

 Alberteae / pe-([micro])re -

 Coffeeae / pe-([micro])re +/-

 Octotropideae / pe-(u)re -

 Pavetteae Granular, pe-(p)re -/+
 sometimes -str

 Gardenieae Granular; pe-(u)re -(+)
 rarely -fo-psi
 with -ru

s. str.

 Cinchoneae Granular pe, rarely -
 with en- [micro] re

 Isertieae Granular, pe-([micro])re -

 PECC Granular pe +
 with en-

 Naucleeae Granular pe-([micro]) -

 Hilheae/ Granular psi-pe- -/+
 Hamelieae ([micro])re

 Rondeletieae Granular pe-([micro])re -(+)

 Guettardeae Granular, pe-([micro])re -(+)
 sometimes -fo-psi
 with en-

Table 11
Summary of orbicule data for Rubiaceae. The order of
tribes follows the topology of the cladogram in Figure 58.

Key: # Sp. = number of species investigated/(number of
species in Mabberley 1997) total number of species most
recent reference; + = present; - = absent;
SE = supratectal elements; IR = infrareticulum;
dtc = difficult to classify in existing types;
? = unknown; PECC = Portlandia-Exostema-Catesbaeeae-Chiococceae
clade. For an explanation of types, see the text.

Tribe Genus

Unresolved Luculia Sweet
or basal Coptosapelta Korth.
Pauridiantheae Pauridiantha Hook. f.
Coccocypseleae Coccocaypselum P. Br.
Psychotrieae Palicourea Aubl.
 (Pentas group) Pentas Beath.
 (Hedvotis group) Gouldia A. Gray
Paederieae Paederia L.
 Serissa Comm. ex Juss.
Rubieac Asperula L.
 Crucianella L.
 Cruciata Mill.
 Galium L.
 Rubia L.
 Sherardia L.
 Valantia L.
Condamineeae Rustia Klotzsch
 Simira Aubl.
Mussaendeae Mussaenda L.
Retiniphylleae Retiniphyllum Bonpl.
Gardenieae Aidia Lour.
 Atractogyne Pierre
 Colochone Keay
 Ganguelia Robbr.
 Gardenia Ellis
 Macrosphyra Hook. f.
 Mitriostigma Hochst.
 Oxvanthus DC.
 Oxyceros Lour.
 Pleiocoryne Rauschert
 Rothmannia Thunb.
Gardenieae Alibertia A. Rich.
 (Alibertia Amaioua Aubl.
Gardenieae Aulacocalvx Hook. f.
 (Aulacocalyceae) Belonophora Hook. f.
 Himalrandia T. Yamaz.
?Gardenieae Argocoffeopsis Lebrun
?Gardenieae/ Augusta Pohl
Pavetteae Leptactina Hook. f.
 Pavetta L.
 Rutidea DC.
 Tarenna Gaertn.
Octotropideae Feretia Delile
 Paragenipa Baill.
 Galiniera Delile
Coffeeae Bertiera Aubl.
 Coffea L.
 Diplospora DC.
 Psilanthus Hook. f
 Sericanthe Robbr.
 Tricalysia A. Rich. ex DC.
?Coffeeae Calycosiphonia Pierre ex Robbr.
Ixoreae Ixora L.
 Scvphiphora C. F. Gaertn.
Cinchoneae Cinchona L.
Isertieae Iserlia Schreb.
PECC Bodusa A. Gray
 Bikkia Reinw.
 Catesbaea L.
 Coutaportla Urb.
 Coutarea Aubl.
 Exostema (Pers.)
 Rich. ex Humb.
 & Bonpl.
 Hintonia Bullock
 Isidorea A. Rich.
 Morierina Vieill.
 Osa Aiello
 Phialanthus Griseb.
 Portlandia P. Br.
 Salzmannia DC.
 Schmidtottia Urb.
 Scolosanthus Vahl
 Siemensia Urb.
 Thogsennia Aiello
sister Strumpfia Jacq.
 of PECC
Naucleeae Adina Salisb.
 Adinauclea Ridsdale
 Breonia A. Rich.
 Burttdavya Hoyle
 Cephalanthus L.
 Gyrostipula J.-F. Leroy
 Janotia J.-F. Leroy
 Ludekia Ridsdale
 Metadina Bakh. f.
 Nauclea L.

Hillieae & Hillia Jacq.
 Hamelieae Cosmibuena Ruiz & Pav.
 Hamelia Jacq.
 Hoffmannia Sw.
?Hillieae/ Balmea Martinez

Rondeletieae Blepharidium Standl.
 Rondeletia L.

Guettardeae Gonzalagunia Ruiz & Pav.

Tribe # Sp. Orbicules

Unresolved 2/(5)5 + IIIb
 or basal 11/(13)16 -/+ IIIa,b
Pauridiantheae 11/(25)39 + III
Coccocypseleae 1/(20)37 + ?
Psychotrieae 1/(200)200 -
 (Pentas group) 1/(34)39 + ?
 (Hedvotis group) 1/(-)3 -
Paederieae 1/(30)29 - ?
 1/(2)2 - ?
Rubieac 5/(90)150 -
 3/(30)33 -
 21/(10)11 -
 16/(300)400 -
 2/(60)72 -
 1/(1)1 -
 1/(3-4)7 -
Condamineeae 1/(15)14 + IIIa
 1/(35)41 -
Mussaendeae 1/(100)216 + IIIa
Retiniphylleae 23/(20)23 + III-VI
Gardenieae 1/(18)60 -
 1/(3)3 T dtc
 1/(2)2 +
 l/(/)1 -
 2/(60)154 -
 1/(3)3 -
 2/(5)4 -
 1/(40)40 -
 1/(?)14 + V
 1/(1)1 -
 1/(40)40 + dtc
Gardenieae 1/(35)50 + dtc
 (Alibertia 1/(25)9 + dtc
Gardenieae 1/(8)9 + dtc
 (Aulacocalyceae) 2/(6)8 -
 1/(2)3 + V
?Gardenieae 2/(8)7 -
?Gardenieae/ 1/(1)1 - II
Pavetteae 1/(25)<20 IIIb
 1/(400)400 + IIIb
 1/(22)22 + IIIa
 2/(180)219 + IIIb
Octotropideae 1/(2)2-3 + dtc
 1/(1)1 + V
 1/(2)2 + V
Coffeeae 1/(55)50 + VI
 3/(90)95-100 + VI
 1/(10)19 + dtc
 2/(20)20 + VI
 1/(15)17 -
 1/(95)95 + VI
?Coffeeae 1/(2)2 -
Ixoreae 2/(300)400 + IIIa
 1/(1)1 + dtc
Cinchoneae 1/(40)23 + IIIa
Isertieae 1/(13)14 + IIIa
PECC 1/(3)3 + I
 8/(20)20 + I
 5/(20)16 + I
 3/(2)3-4 + I
 1/(7)3-4 + I
 2/(45)47 + I
 3/(4)3 + I
 3/(20)17 + I
 1/(2)2 + I
 1/(1)1 + I
 2/(18)19 + I
 3/(6)5 + I
 1/(1)1 + I
 3/(15)14 + 1
 5/(27)21 + I
 1/(1)1 + I
 1/(1)1 + I
sister 1/(1)1 + ?
 of PECC
Naucleeae 2/(3)3 + III
 1/(1)1 + III
 3/(5)20 + III
 1/(1)1 + IV
 3/(6)6-7 + III
 1/(2)1-2 + III
 1/(1)1 + III
 1/(2)2 + III
 1/(1)1 + III
 1/(10)10 + inter III
 & IV
Hillieae & 19/(24)24 + III
 Hamelieae 4/(12)4 + III
 1/(16)17 + ?
 1/(45)119 + ?
?Hillieae/ 1/(1)1 + III

Rondeletieae 2/(2)2 + III
 1/(130)286 + IIIa

Guettardeae 1/(15)43 + ?

Tribe References SE

Unresolved Huysmans et al., 1997 -
 or basal Verellen, 2002; -
 Verellen et al., 2004
Pauridiantheae Ntore, unpubl.
Coccocypseleae Huysmans et al., 1998b + on IR
Psychotrieae Huysmans et al., 2000 -/+
 (Pentas group) Huysmans et al., 1998b -/+
 (Hedvotis group) Huysmans et al., 1997 +
Paederieae Huysmans et al., 1998b -/+
 Huysmans et al., 1998b +
Rubieac Huysmans et al., 2003 +
 Huysmans et al., 2003 +
 Huysmans et al., 2003 +
 Huysmans et al., 2003 +
 Huysmans et al., 2003 +
 Huysmans et al., 2003 +
 Huysmans, unpubl. +
Condamineeae Huysmans et al., 1997
 Huysmans et al., 1997 -
Mussaendeae Huysmans et al., 1997 -
Retiniphylleae Cortes and Huysmans, subm.
Gardenieae Vinckier et al., 2000 -
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -
 Robbrecht et al., 1996 -
 Vinckier et al., 2000 -/+
 Vinckier et al., 2000 -
 Hansson and El-Ghazaly, 2000; -
 Vinckier et al., 2000
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -
Gardenieae Vinckier et al., 2000 -
 (Alibertia Vinckier et al., 2000 -
Gardenieae Vinckier et al., 2000 -
 (Aulacocalyceae) Vinckier et al., 2000 ?
 Vinckier et al., 2000 -
?Gardenieae Vinckier et al., 2000 -
?Gardenieae/ Huysmans et al., 1997 -
Pavetteae Vinckier et al., 2000 -
 Vinckier et al., 2000 +/-
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -(+)
Octotropideae Vinckier et al., 2000 ?
 Vinckier et al., 2000 -
 Vinckier et al., 2000 ?
Coffeeae Vinckier et al., 2000 -
 Vinckier et al., 2000 -/+
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -/+
 Vinckier et al., 2000 -
 Vinckier et al., 2000 -
?Coffeeae Vinckier et al., 2000 -
Ixoreae Vinckier et al., 2000 -
 Vinckier et al., 2000 -
Cinchoneae Huysmans et al., 1997 -
Isertieae Huysmans et al., 1997 -
PECC Huysmans et al., 2000 +
 Huysmans et al., 2000, +
 1997, unpubl.
 Huysmans et al., 2000, 1997 +
 Huysmans et al., 2000, +
 Huysmans et al., 2000 +
 Huysmans et al., 2000 +
 Huysmans et al., 2000, 1997 +
 Huysmans et al., 2000 +
 Huysmans et al., 2000 +
 Huysmans, unpubl. +
 Huysmans, unpubl. +
 Huysmans, unpubl. +
 Huysmans et al., 2000 +
 Huysmans, unpubl. +
 Huysmans, unpubl. +
 Huysmans, unpubl. +
 Huysmans, unpubl. +
sister Igersheim, 1993a; -
 of PECC Huysmans, unpubl.
Naucleeae Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans et al., 1997; -
 Huysmans, unpubl.
 Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans, unpubl. -
 Huysmans et al. 1997; -
 Huysmans unpubl.
Hillieae & D'hondt, 2002 -
 Hamelieae D'hondt, 2002 -
 IT uysmans et al., 1998b -/+
 Huysmans et al., 1998b -
?Hillieae/ D'hondt, 2002 -

Rondeletieae D'hondt, 2002 -
 Huysmans et al., 1997; -
 El-Ghazaly et al., 2000
Guettardeae Huysmans et al., 1998b -
COPYRIGHT 2005 New York Botanical Garden
No portion of this article can be reproduced without the express written permission from the copyright holder.
Copyright 2005 Gale, Cengage Learning. All rights reserved.

Article Details
Printer friendly Cite/link Email Feedback
Publication:The Botanical Review
Geographic Code:1USA
Date:Jul 1, 2005
Previous Article:Palynological characters and their phylogenetic signal in Rubiaceae.
Next Article:Drowning by numbers: rereading Nelson's "Nullius in Verba".

Related Articles
A consensus classification for the order Gentianales with additional details on the suborder Apocynineae.
Apomorphy-based definition also pinpoints a node, and PhyloCode names prevent effective communication.
Palynological characters and their phylogenetic signal in Rubiaceae.
Interpreting the Past: Essays on Human, Primate, and Mammal Evolution in Honor of David Pilbeam.
Insights on using morphologic data for phylogenetic analysis in sedges (Cyperaceae).

Terms of use | Copyright © 2018 Farlex, Inc. | Feedback | For webmasters