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Biodiv. Res. Conserv. 29: 1-23, 2013 BRC www.brc.amu.edu.pl DOI 10.2478/biorc-2013-0004 Submitted 27.01.2013, Accepted 31.03.2013

Reclassification of theAngraecum -alliance (, Vandoideae) based on molecular and morphological data

Dariusz L. Szlachetko*, Piotr Tukałło, Joanna Mytnik-Ejsmont & Elżbieta Grochocka

Department of and Nature Conservation, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland * corresponding author (e-mail: [email protected])

Abstract: Results of molecular analysis compared with morphological studies were used for reclassification of theAngraecum - alliance (Orchidaceae). For the purpose of this study we sequenced the ITS region (ITS1-5.8S-ITS2) of nrDNA representing nuclear genome and the region trnL-F (including intron of trnL gene and trnL-trnF intergenic spacer). The ITS matrix includes 97 samples representing 86 and the trnL-F matrix includes 94 samples representing 86 species. We focus mainly on the , however the other genera of are also included (, , , , , Lemurorchis, , , , , ). Additional 43 sequences, including an outgroup ( modesta) and other representatives of the subtribes () and (, , Erasanthe, ), were obtained from NCBI resources. Bayesian analysis using MrBayes 3.1.2 on the combined ITS/trnL-F matrix were performed. The monophyly of Angraecinae with an inclusion of Aerangidinae is highly supported by both methods (93 BP/100 PP). The Angraecoid taxa formed two well supported clades, namely clade I (89 BP/100 PP) and clade II (84 BP/100 PP). New classification based on both molecular and classical tax- onomy studies is presented including a key to the genera. The subtribe Angraecinae includes 36 genera, 18 of them, included within Angraecum by different authors so far, are treated here. Five new genera are described: Eichlerangraecum, Hermansia, Lesliegraecum, Pectianriella and Rudolfangraecum. Ten sections of Angraecum are raised to the generic status.

Key words: Angraecum, ITS, molecular phylogeny, new genus, taxonomy, trnL-F

1. Introduction probably evolved more than once in Asia and may have done so in . Orchids of the tribe Lindl. have usually­ Vandeae were first defined by Lindley (1835) who been divided into three subtribes: Aeridinae Pfitz. grouped the members of the group basing on the presence (=Sarcanthinae Benth.), Angraecinae Summerh. and of distinct “caudicles” of the pollinarium. In addition to Aerangidinae Summerh. The most important reason Vandeae sensu Dressler (1993; e.g. and Angraecum), for dividing Vandeae was the geographical criterion. a significant part (40%) of Lindley’s group comprises tropi- The great majority of Aeridinae occur in south-eastern cal American taxa that are now placed in Maxillarieae. Asia and Australasia, Angraecinae are found in Africa, Garay (1960, 1972) reinstated tribe Vandeae based and South and Central America, whereas on the presence of an incumbent anther, porrect rostel- Aerangidinae occur only in Africa and Madagascar. lum, and well-developed stipes. Vandeae sensu Garay Dressler (1993) suggested an independent of included tropical American subtribes: Cyrtopodiinae a leafless habit with photosynthetic in Asia, Africa Benth., Zygopetalinae Schltr. and Oncidiinae Benth. and tropical America. He also proposed that this habit as well as Vandinae Rchb.f. (Garay 1972). VARIABILITY, TAXONOMY AND PHYLOGENY TAXONOMY VARIABILITY,

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The subtribes Angraecinae and Aerangidinae were and Podochileae are closely related to described by Summerhayes in 1966. He officially reco­ Vandeae as Dressler (1993) proposed. gnized these two groups as the subtribes of Vandeae A morphological study of Orchidaceae based on which were earlier separated by Schlechter (1918), results of cladistic analysis was carried out by Freuden- according­ to the structure of the rostellum. In Angraeci- stein and Rasmussen (1999). According to their work, nae, the rostellum is deeply divided and in Aerangidinae Vandeae form a monophyletic tribe with several synapo­ is elongate and beak-like (Dressler 1993). Morphologi- morphies: isodiametric exodermal cell shape, mono- cally, the members of these two subtribes have similar podial growth habit, spherical stegmata and seeds with vegetative and floral features and are often referred to laterally compressed walls. Beacuse of the reduction collectively as “angraecoids”. Chromosome counts were of four pollinia to two, Aerangis and Angraecum were made by Jones (1967) and showed cytological support united and formed a sister clade to a paraphyletic grade for the subdivision of the angrecoid orchids (Angraeci- of Aeridinae ( and ). Freuden­stein nae and Aerangidinae) on the morphological basis and and Rasmussen (1999) also indicate a sister relationship revealed the chromosome number for Angraecinae n=19 with Schltr. and for Aerangidinae n=25. Arends et al. (1980) and In the classification of Orchidaceae presented by Arends and Van der Laan’s (1983) in their preliminary Chase et al. (2003), Vandeae form a monophyletic karyological studies questioned the previous division tribe within a large polytomy of advanced epidendroid of the African Vandeae. These authors established four groups (Cymbidieae and Agrostophyllinae Szlach.) and groups: the first one with a short rostellum and n=19 include the sympodial subtribe Polystachyinae. The (Aeranthes Lindl., Cryptopus Lindl., Jumella Schlecht. sister relationship of Vandeae sensu Dressler (1993) and some species of Angraecum Bory), the second and Polystachyinae is well-supported by the analyses of one with a short rostellum and n=21, 23, 24, 25 (some Cameron (2001) as well as van den Berg et al. (2005). species of Angraecum), the third one with a beak-like According to the results of molecular and morpho- rostellum and n=19 (Calyptrochilum Kraenzl.) and the logical studies conducted by Carlsward et al. (2006), the fourth one with a beak-like rostellum and n=23 to 27 monopodial Vandeae form a strongly supported clade (most of Aerangidinae). (>90 BP). Within this clade, Aeridinae also form a well- Dressler (1981) elevated units of the original tribe supported clade with >90 BP in all analyses. Carlsward et Epidendreae Lindl. delimited by Dressler and Dodson al. (2006) suggested that Aerangidinae and Angraecinae (1960) to form two subfamilies: Lindl. s. str. should be classified within Angraecinaesensu lato. ex Endl. and Vandoideae Endl.. Although Vandoideae­ The authors consider that individually these two subtribes have long been recognized (at various taxonomic are polyphyletic but together they form a well-supported levels)­ on the basis of floral morphology, there were monophyletic clade in all molecular analyses. Within few consistent characters to delimit Vandoideae from Angraecinae, a large clade of primarily Madagascan Epidendroideae. Dressler admitted that the only clear taxa (93 BP) is sister to an unresolved clade of the Old character delimiting these two subfamilies was anther and New World angraecoids (95 BP). The chromosome development and, several years later (1989), he rein- number common in many Aerangidinae n=25 suggests stated the members of Vandoideae into a broadly defined some affinities between Aerangidinae (primarily African) Epidendroideae, much like his original systematic treat- and the African species of Angraecum. ment with Dodson (Dressler & Dodson 1960). In his Angraecinae is a large and diverse group restricted classification system, Vandeae formed a well-defined mainly to Paleotropics. Garay (1973) distinguished 15 group of orchids splitted into three subtribes: Sarcan- African and four American genera within this subtribe. thinae, Angraecinae and Aerangidinae. Szlachetko (1995) transferred two genera (Angraecopsis Dressler (1993) united Vandeae with Dendrobieae Kraenzl. and Cribbia Senghas) to Angraecinae from and Podochileae in a dendrobioid subclade of Epiden- Aerangidinae, increasing the number of African genera droideae and replaced the illegitimate subtribal name within the subtribe to 19. Carlsward et al. (2003) included Sarcanthinae (Bentham 1881) with Aeridinae. Harrisella Fawc. & Rendle and Polyradicion Garay into Szlachetko in 1995 divided Vandeae into 14 subtibes the genus Campylocentrum Benth., reducing the number (Vandinae Rchb.f., Deceptorinae Szlach., Phalaenopsi­ of American genera from four to two. Stewart et al. (2006) dinae Szlach., Gastrochilinae Szlach., Aeridinae, accepted the Garay’s (1973) classification ofAngraecum , Diplocentrinae Szlach., Pelatantheriinae Szlach., Taeni- and Chase’s (2003) classification of the subtribes. ophyllinae Szlach., Bolusiellinae Szlach., Listrostach- Most of Angraecinae were initially included within yinae Szlach., Calyptrochilinae Szlach., Aerangidinae, the genus Angraecum until Schlechter (1918) reviewed Rhaesteriinae Szlach., Angraecinae) according to the the entire group and divided the genus into six sections. rostellum and pollinarium structure. Basing on the In 1925, he increased the number of the sections to gynostemium structure, he precluded possibility that thirteen, after acquaintance with the Perrier’s orchid

Unauthenticated Download Date | 1/3/17 2:10 PM Biodiv. Res. Conserv. 29: 1-23, 2013 3 collection from Madagascar. The taxonomic revision 2. Materials and methods of the genus was presented by Garay (1973), who dis- tinguished nineteen sections within Angraecum. 2.1. Molecular markers and taxon sampling The representatives of Angraecinae are monopodial, We collected 56 samples from living specimens rep- epiphytic or lithophytic with a wide variety of resenting Angraecum sensu lato and minor angraecoid habits, from specimens with elongate stems and well- genera. Leaf samples were obtained from the botanical developed leaves to those with reduced stems and small gardens of Hamburg, Heidelberg, Munich and Wien, scale leaves. They are pollinated mostly by moths in our filed trips to New World, Africa and Madagascar accordance with the structure of . The subtribe as well as from the living collection of the University Angraecinae is well characterized by a structure of the of Gdańsk. Basing on an availability of the sequence rostellum which is deeply notched, dome-like, wide data in GenBank from previous studies (Carlsward et al. and short. Tegula is single or double, usually small, 2006), we chose two molecular markers. For the purpose sometimes papillate near the attachment of the pollinia. of this study we sequenced the ITS region (ITS1-5.8S- Viscidium is single or double, oblong to elliptic, as large ITS2) of nrDNA representing nuclear genome and the as or smaller than the tegula (Szlachetko 2003). plastid region trnL-F (including intron of trnL gene and Angraecum, the most species rich genus of the sub- trnL-trnF intergenic spacer). Additional 43 sequences tribe, includes about 200 species, distributed in tropical including an outgroup and other representaives of the Africa and Madagascar with one species occurring in subtribes Aeridinae and Aerangidinae were obtained and the (Angraecum zeylanicum from NCBI resources. Lindl.). Most of the taxa occur in Madagascar, about 50 species have been recorded from the mainland of 2.2. DNA extraction Africa (Stewart et al. 2006). The representatives of Total genomic DNA was extracted from 15-20 mg of Angraecum are monopodial herbs of various size. Stems silica-dried leaves using Genomic Mini AX Plant (A&A are short or elongate, branching or unbranched. Leaves Biotechnology, Gdynia, Poland), following manufac- are dorsiventrally flattened or laterally compressed, turer protocols. Precooled in -45ºC Lysing Matrix A imbricating basally or well-spaced, thin-textured or tubes and FastPrep instrument (MP Biomedicals, USA) fleshy, coriaceous. is erect or pendent, were used for samples homogenization. Pelleted DNA single- to many-flowered. are tiny to large, was then dried overnight and resuspended in 50 ul of resupinate or not. and are similar (one dd H2O and stored in 4ºC for further usage. another). Lip is simple, usually entire, furnished with a 2.3. PCR and sequencing callus or not, spurred. Gynostemium is short and mas- sive, erect. Stigma is large, deeply concave, elliptic. Amplification and sequencing were carried in Bi- Anther is incumbent, operculate, dorsiventrally flat- ometra TGradient and Eppendorf Mastercycler thermal- tened, thin-walled, notched in front. Two pollinia are cyclers. Polymerase chain reaction (PCR) was carried in ellipsoid or obovate, cleft, dorsiventrally flattened. The a final volume of 25 µl using Color Perpetual Taq DNA rostellum is deeply notched, short, dome-like. Double Polymerase kit (Eurx, Gdansk, Poland) with addition of viscidia are oblong, thin, delicate. Double tegulae is dimethyl sulfoxide (DMSO) to the final concentration linear, delicate, lamellate. In many species single tegula of 5% per sample. or tegula variously splitted or divided is attached to the The ITS region (ITS1-5.8S-ITS2) was amplified single viscidium. The rostellum is deeply notched after using two sets of primers: 17SE and 26SE of Sun et removal of pollinaria (Szlachetko 2003). al. (1994). In a few cases, the ITS4 and ITS5 (White Phylogenetic relationships of the closely related et al. 1990) primers were used for the nested PCR genera and species within the Angraecum subtribe were technique. The PCR conditions for ITS were adopted presented in Carlsward et al. (2006). The authors ana- from Carlsward et al. (2006): initial denaturation for 6 lyzed the ITS nrDNA, trnL-F plastid DNA, and matK min in 98°C, pause at 80ºC, addition of Taq, followed plastid DNA. The results of these analyses revealed that by 33 cycles with 45 sec denaturation in 95°C, 45 sec Angraecum is clearly polyphyletic. The genera Bonniera annealing in 57°C (for AB101/AB102 primer set) or Cordem., Oeoniella Schltr. and SobennikoffiaSchltr. are 52°C (for ITS4/ITS5 primer set) and 1 min elongation embedded within the clade composed primarily of the in 72°C, with a final extension for 7 min in 72°C. species of Angraecum. For the trnL-F region, primers from Taberlet et al. Because Angraecum is a polyphyletic group and (1991) were used with PCR condition adopted from the results of detailed morphological studies support Shaw et al. (2007, ‘slow and cold’): intial denaturation reclassification of theAngraecum alliance, we propose for 6 min in 98ºC, pause at 80ºC – added Taq, followed a new classification of Angraecinae and the results of by 33 cycles with 1 min denaturation in 95ºC, 1 min our studies are presented below. of annealing in 50ºC and 4 min of primer extension in

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65ºC, with a final extension step for 7 min in 65ºC. Three We also tested data congruence using the partition samples failed to amplify (Angraecum drouhardii, A. homogeneity test in PAUP* version 4.0b10, implemen- crassum, A. montanum). tation of ILD (Incongruence length difference) test by The PCR products were purified using the High Pure Farris et al. (1994). Heuristic searches for the test were PCR Product Purification Kit (Roche Diagnostic GmbH, performed using 1.000 replicates and an TBR algorithm Germany) following the manufacturer’s protocol. but saving no more than 10 trees per replicate. Proba­ Sequencing ­reaction was carried using Big Dye termina- bility values (p) greater than 0.05 were used to identify tor v 3.1 chemistry (Applied Biosystems, Inc.) with the whether the data sets were not significantly different same primers used for PCR amplification in a total of 10 from each other and could therefore be combined. µl reaction volume. Cycle sequencing conditions were as 2.5. Bayesian analysis follow: 25 cycles each with 15 sec denaturation (94°C), 5 sec annealing (50°C) and 4 min elongation (60°C). We also performed a bayesian analysis using Mr- The sequencing reaction products were then purified and Bayes 3.1.2 (Ronquist & Huelsenbeck 2003) on the sequenced on ABI 3720 automated capillary DNA se- combined ITS/trnL-F matrix. The generalized model quencer in the Institute of Biochemistry and Bio­physics, of DNA substitution was calculated using MrModeltest Polish Academy of Sciences (Warsaw). Sequences were 2.2 (Nylander 2004). The posterior probabilities (PP) inspected/edited in Chromas (Technelysium Pty Ltd.) of clades were estimated by sampling trees from the PP and assembled in AutoAssembler (Applied Biosystems, distribution using Markov chain Monte Carlo simula- Inc). All sequen­ces were checked against BLAST (Basic tions. Two parallel runs with four simultaneous chains Local Alignment Search Tool) from NCBI for possible were executed for 5.000.000 generations with trees fungal or other non-orchid contamination. For both ITS sampled every 100 generation. A plot of generations and trnL-F regions, final alignments were done manually against likelihood scores of the sampled trees was then in Seaview (Galtier et al. 1996) following the guidelines examined in order to establish “burn-in” required for from Kelchner (2000). Resulting indels were coded both runs to converge on a stationary probability value. manually as additional binary characters following the Burn-in trees were then discarded from the study. The simple coding method of Simmons and Ochoterena remaining trees were used to calculate a majority-rule (2000). consensus tree using “sumt” command in MrBayes. 2.4. Parsimony analysis 3. Results and discussion We performed a cladistic analysis using Fitch par- simony (Fitch 1971) with PAUP* 4.0b10 (Swofford 3.1. ITS 2003) and Polystachya modesta designated as a single species outgroup. The ITS matrix includes 97 samples representing 86 Heuristic searches were performed using tree bi- species. Considering 867 analyzed aligned sequence po- section reconnection (TBR) branch swapping, simple sitions (799 after excluding indels), 442 were potentially­ sequence addition and maxtrees limit set to 10.000. parsimony informative (374 after excluding indels). All characters were equally weighted and gaps were Heuristic analysis produced +10.000 equally parsimo- treated as missing data. Internal support for clades was nious trees (trees not shown) with length (L) of 1517 estimated using bootstrap (BP) percentages (Felsenstein steps, consistency index (CI) – 0.404, and retention 1985) with 1.000 bootstrap replicates, simple addition index (RI) – 0.747. and TBR branch swapping but saving no more than According to the ITS results, monophyly of An- 10 trees per replicate (Salamin et al. 2003). We define graecine with Aerangidinae against the outgroup boostrap support as weak for values from 50-74, moder- and Aeridinae is weakly supported with 50 BP. The ate for 75-89 and high for 90-100. remaining ingroup taxa form two weakly supported Initially, the ITS and trnL-F matrices were analyzed clades. The large clade (56 BP) consists almost entirely separately. We used bootstrap trees generated for each of the Malagasy-Mascarene angraecoid species (with region to manually compared them for congruence, representatives on Seychelles and ), with the following guidelines from Wiens (1998). If there is no exception of continental A. conchiferum, A. dives and conflict, the well-supported clades (bootstrap percent- partially A. eburneum. The smaller clade (65 BP) is age higher than 74) between the regions, the datasets formed by continental species of Angraecum, N eotropi- for each region can be combined into a single matrix. cal Angraecinae, minor Malagasy angraecoid genera The samples that failed to amplify in one of the regions and representatives of the genus Aerangis (Aerangidi- or were unavailable from NCBI were coded as missing. nae). However, the resolution within each of these two The combined analysis was performed with the same nodes was weak with rather low to moderate bootstrap heuristic search strategies as described above. support.

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3.2. trnL-F we did not observe substantial changes in trees topology­ after the exclusion of indels, the bootstrap support for The trnL-F matrix includes 94 samples representing­ major clades was significantly lower than when the 86 species with 1905 aligned positions (1010 after indels were included. exluding ­indels) of which 649 positions were potentially The combined analysis resulted in much better resol­ parsimony informative (588 after exluding indels). Heu- ved trees compared to the separate analysis of each ristic search produced +10.000 equally parsimonious dataset. The Bayesian analysis produced approximately trees with length (L) of 1127 steps, consistency index the same tree topology as parsimony. One of the most (CI) – 0.684, and retention index (RI) of 0.787 (trees not parsimonious trees annoted with BP and PP values is shown). The monophyly of Angraecinae with Aerangis shown in the Fig 1. The monophyly of Angraecinae with (Aerangidinae) against the outgroup and Aeridinae is an inclusion of Aerangidinae is highly supported by both moderately suppported with 81 BP. Similar to the ITS methods (93 BP/100 PP). The Angraecoid taxa fall into results, the ingroup taxa form two clades, although, only two well supported clades, namely clade I (89 BP/100 the smaller clade, consisting of the continental Angrae- PP) and clade II (84 BP/100 PP). We cannot indicate, cum species, neotropical and Malagasy Angraecine and however, any synapomorphy for either of clades. three accessions of Aerangis, is moderately supported Clade I contains nearly exclusively the Mascarenian, with 81 BP. The resolution and overall bootstrap support Malagasy and Malagasy-Mascarenian Angraecum spe- was much higher for the trnL-F matrix than ITS. cies (with three exceptions described above) and some 3.3. Combined analysis angraecoid genera (Aeranthes, Bonniera, Jumellea, Lemurorchis, Oeniella and Sobennikoffia) with similar The results of partition homogenity test for the nrITS distribution range. Species included in this clade pos- and trnL-F datasets indicate that the partitions were sess both resupinate () and signi­ficantly different from random partitions (p = 0.01), nonresupinate () flowers, with thus should not be combined. However, the ILD test can variously developed flower’s segments and reproduc- sometimes reveal unreliable results (Dolphin et al. 2000, tive structures (single pollinarium in Angraecum sect. Reeves et al. 2001) and the observed incongruence can Arachnangraecum, double pollinaria in Angraecum be more likely a result of an insufficient taxon sampling sect. Gomphocentrum and intermediate state in Angrae- or weak phylogenetic signal and homoplasy (Wendel cum magdalenae). & Doyle 1998). Visual inspection of separate bootstrap The genus Jumellea embraces species clearly trees revealed no major conflicting clades with higher separated from all other Angraecinae, taken into this bootstrap support (BP >75). Therefore, we decided to analysis due to the peculiar position of the lip versus combine both datasets into a single matrix and proceed gynostemium. The lip of Jumellea is narrow and often with a combined analysis. The tree statistics for the clawed at the base and never enfolds the gynostemium, combined and separate analysis is summarized in the what results in the location ofthe lip below the gyno­ Table 1. stemium. In our opinion, this significant feature is a The combined ITS/trnL-F matrix, representing 98 good background for retaining Jumellea as a separate acessions, consisted of 2770 aligned positions (2639 genus. That is obvious that an outgroup to Jumellea, after excluding indels) with 621 parsimony informa- i.e. Angraecum section Perrierangraecum, and part of tive characters (548 excluding indels). Heuristic search Angraecum represented by Angraecum sesquipedale resulted in 652 equally parsimonious trees with length and A. sororium deserve a generic status. (L) of 2736 steps, consistency index (CI) – 0.502 and Clade A includes such distinct groups of angrae- retention index (RI) of 0.740. An exclusion of the coids as Angraecum sect. Humblotiangraecum, sect. coded indels decreased the overall trees length but also Arachnangraecum, sect. Gomphocentrum, Lemurochis, increased the number of resultant trees to 777. Although Oeoniella, Aeranthes and Bonniera, what means that

Table 1. Tree statistics for maximum parsimony analyses

ITS trnL-F ITS/trnL-F Number of aligned position 867 1905 2770 Number of informative characters 297 326 621 Number of trees saved +10 000 +10 000 652 Lenght of MP trees 1517 1127 2736 Consistency index CI 0.404 0.684 0.502 Retention index RI 0.747 0.787 0.740

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Fig. 1. One of the most parismonious trees from the combined analysis using maximum parsimony. The branch length is shown above and bootstrap support versus bayesian posterior probabilities below the branch (bp/pp); gb denotes sequences obtained from genbank resources. Single arrowheads on the right designate taxa with problematic placement and are discussed further in the text.

Unauthenticated Download Date | 1/3/17 2:10 PM Biodiv. Res. Conserv. 29: 1-23, 2013 7 they have no mutual evolutionary advanced feature. of Aerangis represent completely different level of or- Moving down within clade A we can divide it into 3 ganization – its rostellum and single tegula are greatly subclades, neither of which is monomorphic enough elongate, occasionally longer than column part, and to be defined by a set of features: viscidium is relatively small. subclade 1 – Bonniera, Angraecum sect. Arachnan- There is a group of Angraecum species that are nested graecum, Aeranthes, Gomphocentrum, Sobennikoffia; in puzzling places on the tree presented in this article. subclade 2 – Oeoniella, Lemurorchis, Angraecum Angraecum multinominatum Rendle is a member of the s.str. & A. sect. Boryangraecum; section Afrangraecum Summerh., typified by Angrae- subclade 3 – Angraecum sect. Humblotiangraecum cum reygaertii De Wild. and it does not appear to be & A. sect. Rudolfangraecum. drastically different from other member of the section. Clade II consists of two well supported clades of the We can not explain its close relation with Angraecop- continental African Angraecum species. One of these sis parviflora (Thouars) Schltr. Analogical situation is clades is sister to some genera included in Aerangidinae observed in the case of Angraecum dives-Lemurorchis (Aerangis, Soelnangis) and the other is sister to the well and Angraecum zeylanicum-Oeonia. Angraecum cras- supported (99 BP/100 PP) Neotropical Angraecinae sum Thouars was included in the nominal section of the (Campylocentrum, Dendrophylax). The monotypic genus Angraecum Garay (1973), but in our analysis it is Malagasy genus Erasanthe (formerly Aeranthes henrici sister to sublcade 2 including, amongst others, Oeoniella sensu Cribb et al. 2007) remains sister to the groups and Lemurorchis. described above. Another well supported clade (100 BP/100 PP) within clade II consists of some Malagasy 3.4. The role of pollination in evolution angraecoid genera (Cryptopus, Oeonia, Neobathiea) of Angraecoid orchids and an interesting species of Angraecum zeylanicum Most of Angraecinae produce long-spurred white known from Sri Lanka and Seychelles. Another­ interest- flowers emitting crepuscular scent, what is an adapta- ing exception stands for Calyptrochilum christyanum. tion to pollination by hawkmoths; these floral traits are This species remains sister to the rest of the taxa in the consistent with the general syndrome of hawkmoth clade II although its separated position is weakly sup- pollination (Grant 1985; Haber & Frankie 1989). The ported by bootstrap (62 BP). Clade II, similarly like angraecoids share similarities in floral morphology clade I, comprises so strongly diversified species that and one can deduce that play an important it is impossible to indicate any synapo­morphy, even if role in their evolution and speciation (Dressler 1981). we follow the procedure accepted ­above, i.e., to step In many species infundibular base of the spur provides down towards the species level. We were able to find enough space for insertion of the base of the pollina- out characteristic combination of features at the level tor’s head into the flower. On contrary, there is a large corresponding roughly to the sections of Angraecum. group of angraecoids with narrow spur base, which can Therefore, we propose the generic status for most of be penetrated by the narrow and long proboscis only. the sections proposed by Schlechter (1918) and Garay In many species a keel thickening is observed on the (1973), but with modified species contents. basal part of the lip and it can play a role of a guidepost, Most of the sectional arrangements within the genus directing the proboscis to the spur exactly below pol- seems to be unnatural (Fig. 1), with few exceptions for linarium. In this case, the flower produces two separate the highly supported (100 BP/100 PP) African sections – pollinaria, each consisting of single viscidium, tegula Dolabroflia (erected to the generic status by Szlachetko and pollinium. These features, along with the various and Romowicz in 2007), Conchoglossum and Pecti- lengths of the spur, size of the flowers, their number naria. The latter section is not clearly monophyletic due and arrangement in inflorescence, as well as the large to outstanding position of Angraecum dasycarpum in variability of vegetative parts responsible for adaptation the clade I, as one of the three members of this section to different environmental conditions, make possible occurs outside continental Africa. the exploitation of a large spectrum of animals acting The position of Solenangis Schltr., Oeoniella Schltr. as pollinators. and particularly Aerangis Rchb.f. on the nrITS and An interesting pattern of spur morphology and trnL-F combined tree is enigmatic. Solenangis has been production was recently observed by Martins classified to Bolusiellinae and Oeoniella to Listros- and Johnson (2007) in the Aerangis species examined tachyinae by Szlachetko (1995) and it shares the similar in the eastern Africa. The authors revealed that some type of rostellum, rostellum remnant and pollinarium long-spurred species (Aerangis brachycarpa and A. with such genera as Nephrangis (Schltr.) Summerh. confusa) have straight spurs, full of nectar, whereas in A. and Schltr. This type of gynostemium can be thomsonii and A. kotschyana, the nectar is located only compared with Angraecopsis, Cribbia or some species in the basal third/quarter of the spur and, what is more of Campylocentrum. However, reproductive structures interesting, the spurs of the latter species are spirally

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twisted (Martins & Johnson 2007). Similar observa- nism are still scanty and give no authorization to tions were made by Nilsson et al. (1985) in Angraecum formulate any hypothesis regarding significance of arachnites in Madagascar. behavior and morphology in the speciation Despite the fact that most of Angraecinae is pol- and evolution of this orchid group. linated by the representatives of , there are three Mascarenian species classified within the section A key to the genera of Angraecinae: Hadrangis Schltr. pollinated by birds and crickets, what 1. Plants aphyllous ...... 2 was recently discovered (Micheneau et al. 2006, 2010, 1. Plants leafy ...... 4 respectively). Angraecum striatum Thou., A. bracteo- 2. Flowers more than 1 cm in diameter. Spur filiform or sum Balf.f. & S.Moore and A. cadetii Bosser, endemic to cylindrical, much longer than lip, at least three times as , are pollinated by small song-birds, long as the lip ...... Dendrophylax Zosterops olivaceus and Z. borbonicus (Zosteropidae), 2. Flowers smaller than 1 cm in diameter. Spur usually the bird species endemic also to Rèunion (Micheneau et very short, up to twice as long as the lip ...... 3 al. 2006). What is surprising, the orchids produce white 3. Spur globose at the apex, with callus in the centre of flowers. Except for the colour, the flower morphology the lip median lobe ...... Harrisella of A. striatum and A. bracteosum matches orchid-bird 3. Spur blunt or saccate, never globose at the apex, lip (Micheneau et al. 2006). The bill without any callus ...... Campylocentrum of Z. borbonicus matches the flower entrance diameter 4. Leaves equitant, basally imbricating ...... 5 perfectly and is suitable for both extracting nectar and 4. Leaves conduplicate ...... 6 performing pollination (Micheneau et al. 2006). Many 5. Plants stemless, leaves oblong falcate, inflorescence examples show that flower colour is not necessarily few-flowered, flowers large, spur much longer than correlated with pollination type (i.e. Momose et al. pedi­cellate ovary, orifice spacious ...... 1998; Johnson & Steiner 2000) and flower colour within ...... Humblotiangraecum Angraecinae may be a conservative character. The third 5. Plants with elongate stem, leaves dolabriform to species, A. cadetii occurring on Rèunion and Mauri- elongate, inflorescence single-flowered, flowers small, tius, has a highly surprising pollinator, a raspy cricket spur almost equal in length to pedicellate ovary, orifice (Gryllacrididae, the order Orthoptera) (Micheneau et narrow ...... Dolabrifolia al. 2010). 6. Spur clavate to saccate, apically blunt, predominantly The section Hadrangis, entirely endemic to the subequal in length to the pedicellate ovary ...... 7 Mascarenes, represents a case of an Angraecum intra- 6. Spur filiform, occasionally somewhat swollen at the archipelago radiation (Micheneau et al. 2008a), the acute apex, orifice spacious, mostly much longer than atypical flower morphology resulted from specific pedicellate ovary ...... 11 adaptation to the local pollinators, linked to the oceanic 7. Leaf petiole twisted, hence, blades lying in one plane context of the Mascarene Archipelago (Micheneau et or so ...... Pectinariella al. 2006). 7. Leaves not as above ...... 8 There are also some long-spurred angraecoids 8. Flowers resupinate ...... 9 (more than 9 cm) on Rèunion, however all these spe- 8. Flowers nonresupinate ...... 10 cies endemic to the island became totally independent 9. Floral bracts oblong-ovate, much exceeding pedicel- of pollinators and are capable of autonomous self- late ovary ...... Hadrangis pollination (Micheneau et al. 2006), what is probably 9. Floral bracts inconspicuous, shorter than pedicellate caused by loosing an orchid-hawkmoth interaction ovary ...... Lepervenchea (T. Pailler and C. Micheneau, unpublished). Therefore, 10. Stem elongate, inflorescence shorter than leaves ... an auto-pollination is linked to the absence of specific ...... Lemurangis pollinator during the island colonization and species 10. Stem abbreviated, inflorescence longer than leaves establishment, oceanic islands are known for the paucity ...... Lesliegraecum of their insect fauna and often whole groups of insects 11. Viscidium single ...... 12 are missing (Micheneau et al. 2008b). 11. Viscidia 2 ...... 18 These wide range of pollination syndrome within 12. Rostellum slightly bent forwards, short and wide, the angraecoid orchids on Rèunion is probably related obscurely 3-lobulate ...... Coenadenium to their recent dispersal to the Mascarene Islands, where 12. Rostellum dome-like, with deep and acute or shal- specific long-tongue pollinators of ancestral orchid colo- low and gentle sinus in front ...... 13 nists were absent (Micheneau et al. 2008b [Jumellea]). 13. Basal part of the lip below the gynostemium, not The floral morphology of Angraecinae is diverse surrounding it ...... Jumellea suggesting some variation in pollination systems. 13. Basal part of the lip cochleate, more or less sur- Unfortunately,­ the observations of pollination mecha- rounding the gynostemium ...... 14

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14. Flowers resupinate, spur infundibuliform at the base 30. Lip fleshy, transversely elliptic, spur relatively short, ...... 15 hardly exceeding pedicellate ovary, straight, swollen 14. Flowers nonresupinate, spur orifice rather narrow .. and blunt at the apex ...... Boryangraecum ...... 16 31. Lip 3-lobed, the middle lobe often bilobulate ...... 15. Viscidia double, but connate marginally to one an- ...... Oeonia other, forming a single structure. Lip large, ecallose ... 31. Lip not as above ...... 32 ...... Rudolfangraecum 32. Inflorescence usually densely and many-flowered 15. Viscidium single, lanceolate to ovate, oblong. Lip ...... Campylocentrum very large, transversely elliptic to obreniform, with 32. Inflorescence usually 1- to few-flowered ...... 33 prominent keel along midvein in the basal part ...... 33. Stem with elongate internodes, inflorescence always ...... Eichlerangraecum 1-flowered ...... Pseudojumellea 16. Tegula more or less plit apically, hence V-shaped .. 33. Stem with abbreviated internodes, inflorescence ...... Angraecoides usually few-flowered ...... 34 16. Tegula entire, more or less oblong ...... 17 34. Flowers small, inconspicuous, resupinate, greenish 17. Inflorescence single-flowered, lip ecallose ...... to yellowish, thin in texture, rather diaphanous ...... Arachnangraecum ...... Gomphocentrum 17. Inflorescence multiflowered, elongate, lip with basal 34. Flowers large, showy, ivory-coloured, not diapha- keel ...... Angraecum nous ...... 35 18. Flowers spurless or spur greatly reduced to shallow 35. Spur base very narrow, filiform ...... sac ...... Bonniera ...... Perrierangraecum 18. Spur with prominent spur ...... 19 35. Spur base narrowly conical ...... Macroplectrum 19. Rostellum median lobe much longer than the column part ...... Ambrella 3.5. Taxonomic treatment 19. Rostellum median lobe obscure, much shorter than 1. Angraecoides (Cordem.) Szlach., Mytnik & Gro- the column part, digitate or reduced ...... 20 chocka, stat. et gen. nov. 20. Viscidia 2, tegula single ...... Sobennikoffia Basionym: Lindl. sect. Angraecoides 20. Viscidia 2, tegulae 2 ...... 21 Cordem., Rev. Gen. Bot. 11: 421. 1900; G e n e r i - 21. Column foot prominent, longer than the column part type: Angraecum pingue Frapp. [≡ Angraecoides ...... 22 pingue (Frapp.) Szlach., Mytnik & Grochocka]. = An- 21. Column foot usually absent or inconspicuous .....23 graecum Bory sect. Conchoglossum Schltr., Beih. Bot. 22. Lip margins entire ...... Aeranthes Centralbl. 36(2): 157. 1918; Ty p e s p e c i e s (Garay 22. Lip margins fimbriate ...... Erasanthe 1973: 499): Angraecum viride Kraenzl. 23. Rostellum lateral lobes ribbon-like, prominent, curved Plants medium-sized. Stem elongate, internodes down at the base and then abruptly upcurved ...Distylodon prominent. Leaves in two rows, ligulate to lanceola­ 23. Rostellum not as above ...... 24 te-ovate, unequally bilobed at the apex, sheaths com- 24. Gynostemium elongate, slender ...... 25 pressed, loosely enclothing stem. Inflorescence usually 24. Gynostemium short, massive ...... 26 1-flowered. Peduncle usually shorter than internodes, 25. Petals strongly asymmetric, partially connate with enclothed basally by 1-2 adpressed bracts. Flowers lateral sepals ...... Angraecopsis small to medium-sized, greenish, yellowish, pinkish- 25. Petals more or less symmetric, free from lateral brown to white, nonresupinate. Floral bracts incon- sepals ...... Cribbia spicuous. Pedicellate ovary terete. Sepals and petals 26. Petals clawed, with apical more or less transversely el- subsimilar, narrowly lanceolate or linear-lanceolate, liptic plate. Tegulae densely and stiffly hairy ...Cryptopus acute to acuminate. Lip longer than wide, often peta- 26. Petals not clawed, widest at the base. Tegulae gla- loid, concave in the center, with more or less oblong brous ...... 27 or linear callus in the middle, acuminate. Spur more or 27. Spur entrance very large, its base broadly conical ... less as long as the lip, sometimes slightly exceeding ...... Neobathiea it or shorter, slender, narrowly cylindrical, somewhat 27. Spur base cylindrical ...... 28 swollen at the apex, orifice narrow. Tegula more or less 28. Flowers nonresupinate ...... 29 split at the apex, hence V-formed, viscidium single. 28. Flowers resupinate ...... 30 The genus, as proposed here, combines two 29. Lip prominently 3-lobed ...... Garay’s (1973) sections of Angraecum, i.e. Angrae- 29. Lip unlobed or obscurely 3-lobed ...... 31 coides (Cordem.) Garay and Conchoglossum Schltr. 30. Lip thin-textured, ovate-lanceolate, spur longer than In our opinion, the distinguishing character of these pedicellate ovary, filiform, straight, somewhat swollen sections (peduncle prominent vs usually short) is at the apex, with narrow entrance ...... Hermansia neither constant nor important enough to substantiate

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such proposal. Angraecoides ­appears to be related to Angraecoides erecta (Summerh.) Szlach., Mytnik Eichlerangraecum from which it is easily separable & Grochocka, comb. nov. by the flower and tegula morphology. The species Basionym: Angraecum erectum Summerh., Kew of Angraecoides produce smaller flowers, which are Bull. 11: 232. 1956. inconspicuous, with narrow spur orifice and petaloid Angraecoides keniae (Kraenzl.) Szlach., Mytnik & lip, what is probably an adaptation to pollination by Grochocka, comb. nov. small, long-proboscis insects. Large flowers with a Basionym: Angraecum keniae Kraenzl., Bot. Jahrb. broad and spacious, infundibular spur entrance, seen Syst. 17: 59. 1893. in Eichlerangraecum, can easily accomodate head and Angraecoides modica (Summerh.) Szlach., Mytnik thorax of larger insects. Tegula of the Angraecoides & Grochocka, comb. nov. species is V-shaped, whereas in Eichlerangraecum is Basionym: Angraecum modicum Summerh., Kew single, undivided. Bull. 13: 84. 1958. The genus includes 25 species. Angraecoides nasuta (Schltr.) Szlach., Mytnik & Angraecoides angustipetala (Rendle) Szlach., Myt- Grochocka, comb. nov. nik & Grochocka, comb. nov. Basionym: Angraecum nasutum Schltr., Repert. Basionym: Angraecum angustipetalum Rendle, Spec. Nov. Regni Veg. Beih. 33: 315. 1925. Cat. Pl. Oban: 106. 1913. Angraecoides obversifolia (Frapp. ex Cordem.) Angraecoides brevicornu (Summerh.) Szlach., Szlach., Mytnik & Grochocka, comb. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum obversifolium Frapp. ex Basionym: Angraecum brevicornu Summerh., Kew Cordem., Fl. Réunion: 212. 1895. Bull. 16: 311. 1962. Angraecoides panicifolia (H. Perrier) Szlach., Myt- Angraecoides chermezonii (H. Perrier) Szlach., nik & Grochocka, comb. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum panicifolium H.Perrier, Basionym: Angraecum chermezonii H. Perrier, Notul. Syst. (Paris) 7: 105. 1938. Notul. Syst. (Paris) 7: 107. 1938. Angraecoides pingue (Frapp. ex Cordem.) Szlach., Angraecoides cilaosiana (Cordem.) Szlach., Mytnik Mytnik & Grochocka, comb. nov. & Grochocka, comb. nov. Basionym: Angraecum pingue Frapp. ex Cordem., Basionym: Mystacidium cilaosianum Cordem., Fl. Réunion: 214. 1895. Rev. Gén. Bot. 11: 424. 1899. ≡ Angraecum cilao- Angraecoides rhizomaniaca (Schltr.) Szlach., Myt- sianum (Cordem.) Schltr., Beih. Bot. Centralbl. 33(2): nik & Grochocka, comb. nov. 432. 1915. Basionym: Angraecum rhizomaniacum Schltr., Angraecoides clavigera (Ridl.) Szlach., Mytnik & Repert. Spec. Nov. Regni Veg. Beih. 33: 315. 1925. Grochocka, comb. nov. Angraecoides rostrata (Ridl.) Szlach., Mytnik & Basionym: Angraecum clavigerum Ridl., J. Linn. Grochocka, comb. nov. Soc., Bot. 21: 485. 1885. Basionym: Angraecum rostratum Ridl., J. Linn. Angraecoides cultriforme (Summerh.) Szlach., Soc., Bot. 21: 485. 1885. Mytnik & Grochocka, comb. nov. Angraecoides scalariforme (H. Perrier) Szlach., Basionym: Angraecum cultriforme Summerh., Kew Mytnik & Grochocka, comb. nov. Bull. 13: 272. 1958. Basionym: Angraecum scalariforme H.Perrier, Angraecoides curvicaule (Schltr.) Szlach., Mytnik Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 265. & Grochocka, comb. nov. 1955. Basionym: Angraecum curvicaule Schltr., Repert. Angraecoides sedifolia (Schltr.) Szlach., Mytnik & Spec. Nov. Regni Veg. Beih. 33: 346. 1925. Grochocka, comb. nov. Angraecoides curvipes (Schltr.) Szlach., Mytnik & Basionym: Angraecum sedifolium Schltr., Repert. Grochocka, comb. nov. Spec. Nov. Regni Veg. Beih. 33: 316. 1925. Basionym: Angraecum curvipes Schltr., Bot. Jahrb. Angraecoides stolzii (Schltr.) Szlach., Mytnik & Syst. 38: 21. 1905. Grochocka, comb. nov. Angraecoides egertonii (Rendle) Szlach., Mytnik Basionym: Angraecum stolzii Schltr., Bot. Jahrb. & Grochocka, comb. nov. Syst. 53: 603. 1915. Basionym: Angraecum egertonii Rendle, Cat. Pl. Angraecoides triangulifolia (Senegas) Szlach., Oban: 107. 1913. Mytnik & Grochocka, comb. nov. Angraecoides elliotii (Rolfe) Szlach., Mytnik & Basionym: Angraecum triangulifolium Senghas, Grochocka, comb. nov. Adansonia, n.s., 4: 310. 1964. Basionym: Angraecum elliotii Rolfe, J. Linn. Soc., Angraecoides viride (Kraenzl.) Szlach., Mytnik & Bot. 29: 54. 1891. Grochocka, comb. nov.

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Basionym: Angraecum viride Kraenzl., Bot. Jahrb. Arachnangraecum ampullaceum (Bosser) Szlach., Syst. 51: 395. 1914. Mytnik & Grochocka, comb. nov. Angraecoides zaratananae (Schltr.) Szlach., Mytnik Basionym: Angraecum ampullaceum Bosser, Adan- & Grochocka, comb. nov. sonia, n.s., 10: 109. 1970. Basionym: Angraecum zaratananae Schltr., Repert. Arachnangraecum conchiferum (Lindl.) Szlach., Spec. Nov. Regni Veg. Beih. 33: 317. 1925. Mytnik & Grochocka, comb. nov. Basionym: Angraecum conchiferum Lindl., Com- 2. Angraecum Bory panion Bot. Mag. 2: 205. 1836. Voy. 1: 359, t. 19. 1804; Arachnangraecum danguyanum (H. Perrier) Generitype: Angraecum eburneum Bory. Szlach., Mytnik & Grochocka, comb. nov. Plants usually very large, often caespitose. Stem Basionym: Angraecum danguyanum H. Perrier, elongate, stout, leafy. Leaves long, linear, thick, leathery, Notul. Syst. (Paris) 7: 107. 1938. stiff, unequally bilobed at the apex. Inflorescence elon- Arachnangraecum expansum (Thouars) Szlach., gate, multiflowered. Flowers large, fleshy, conspicuous, Mytnik & Grochocka, comb. nov. nonresupinate. Floral bracts prominent, amplexicaul. Basionym: Angraecum expansum Thouars, Hist. Pedicellate ovary ribbed. Sepals and petals subsimilar, Orchid.: 57. 1822. lanceolate, acute. Lip more or less transversely elliptic, Arachnangraecum germinyanum (Hook. f.) concave, cochleate, with basal keel, apically obscurely Szlach., Mytnik & Grochocka, comb. nov. 3-lobed, long acuminate. Spur narrowly cylindrical to Basionym: Hook. f. Bot. filiform, acute, base narrow, pendent. Viscidium single, Mag. 115: t. 7061. 1889. elliptic-ovate, tegula elongate, oblanceolate, longer than Arachnangraecum humbertii (H. Perrier) Szlach., tegula. Mytnik & Grochocka, comb. nov. The genus as proposed here is limited to the species Basionym: Angraecum humbertii H. Perrier., Notul. of Angraecum eburnem-complex. In their habit they are Syst. (Paris) 8: 46. 1939. similar to both Lemurorchis and Hadrangis, but have Arachnangraecum linearifolium (Garay) Szlach., completely different, nonresupinate flowers. Mytnik & Grochocka, comb. nov. Basionym: Angraecum linearifolium Garay, Bot. 3. Arachnangraecum (Schltr.) Szlach., Mytnik & Gro- Mus. Leafl. 23: 160. 1972. chocka, stat. et. gen. nov. = Angraecum palmiforme H. Perrier, Mém. Inst. Sci. Basionym: Angraecum Bory sect. Arachnangrae- Madagascar, Sér. B, Biol. Vég. 6: 268. 1955, nom. cum Schltr., Repert. Sp. Nov. Regni Veg., Beih. 33: 309. illeg. 1925; Lectotype (Garay 1973: 501): Angraecum Arachnangraecum mirabile (Schltr.) Szlach., Myt- ramosum Thouars [≡ Arachnangraecum ramosum nik & Grochocka, comb. nov. (Thouars) Szlach., Mytnik & Grochocka]. Basionym: Angraecum mirabile Schltr., Repert. Stem elongate, erect to pendent. Leaves numerous, Spec. Nov. Regni Veg. 15: 338. 1918. ligulate to linear, unequally bilobed. Inflorescence Arachnangraecum pseudofilicornu (H. Perrier) 1-flowered. Flowers showy, relatively large, nonresupi- Szlach., Mytnik & Grochocka, comb. nov. nate. Peduncle slender, longer than internodes, with 1-2 Basionym: Angraecum pseudofilicornu H. Perrier, sheaths at the base. Floral bracts inconspicuous. Sepals Notul. Syst. (Paris) 7: 108. 1937. and petals subsimilar, usually long-acuminate to cau- Arachnangraecum ramosum (Thouars) Szlach., date, acute. Lip large, suborbicluar, concave at the base, Mytnik & Grochocka, comb. nov. usually long-acuminate at the apex, ecallose. Spur long, Basionym: Angraecum ramosum Thouars, Hist. filiform, acute, somewhat wider at entrance. Viscidium Orchid.: 59. 1822. single, elliptic, small. Tegula oblong-oblanceolate to Arachnangraecum scottianum (Rchb. f.) Szlach., ligulate, longer than viscidium, single. Mytnik & Grochocka, comb. nov. The genus includes the species producing flow- Basionym: Angraecum scottianum, Rchb. f., Gard. ers which resemble those of Angraecum, especially Chron., n.s., 10(2): 556. 1878. in the position and morphology of the lip. Although, Arachnangraecum sterrophyllum (Schltr.) Szlach., the lip of Arachnangraecum is devoid of any keel and Mytnik & Grochocka, comb. nov. the spur orifice is rather wide, enabling pollinators to Basionym: Angraecum sterrophyllum Schltr., Beih. insert the whole head inside the flower, and not only Bot. Centralbl. 34(2): 338. 1916. the proboscis. Very peculiar aspect of the flowers are Arachnangraecum teretifolium (Ridl.) Szlach., widely spread, caudate tepals, giving the flower a spider Mytnik & Grochocka, comb. nov. appearence. Basionym: Angraecum teretifolium Ridl., J. Linn. The genus includes 13 species. Soc., Bot. 21: 484. 1885.

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4. Boryangraceum (Schltr.) Szlach., Mytnik & Gro- Boryangraecum ramulicolum (H .Perrier) Szlach., chocka, stat. et gen. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum Bory sect. Boryangraecum Basionym: Angraecum ramulicolum H. Perrier, Schltr., Repert. Sp. Nov. Regni Veg., Beih. 33: 308. Notul. Syst. (Paris) 7: 123. 1938. 1925; L e c t o t y p e (Garay 1973: 502): Angraecum Boryangraecum sacciferum (Lindl.) Szlach., Myt- pumilio Schltr. [≡ Boryangraceum pumilio (Schltr.) nik & Grochocka, comb. nov. Szlach., Mytnik & Grochocka]. Basionym: Angraecum sacciferum Lindl., Com- Plants small. Stem hardly developed, usually much ab- panion Bot. Mag. 2: 205. 1836. breviated. Leaves linear to oblong-lanceolate, unequally ­ Boryangraecum sinuatiflorum (H. Perrier) Szlach., bilobed at the apex. Inflorescence elongate, several-flow- Mytnik & Grochocka, comb. nov. ered. Floral bracts prominent, amplexicaul. Pedicellate Basionym: Angraecum sinuatiflorum H. Perrier, ovary terete. Flowers small, nonresupinate, diaphanous, Notul. Syst. (Paris) 14: 163. 1951. yellowish, ochraceous, whittish, greenish. Sepals and pet- Boryangraecum tamarindicolum (Schltr.) Szlach., als dissimilar, acute. Lip rather fleshy, obscurely 3-lobed, Mytnik & Grochocka, comb. nov. transversely elliptic, cochleate, canaliculate, lateral lobes Basionym: Angraecum tamarindicolum Schltr., upcurved, apex acute to acuminate. Spur relatively short, Repert. Spec. Nov. Regni Veg. Beih. 33: 340. 1925. hardly exceeding pedicellate ovary, straight, swollen and Boryangraecum teres (Summerh.) Szlach., Mytnik blunt at the apex. Tegulae and viscidia double. & Grochocka, comb. nov. Boryangraceum appears to be similar to Lemuran- Basionym: Angraecum teres Summerh., Kew Bull. gis, but both genera are well separated by pollinarium 13: 265. 1958. structure. There are two pollinaria in the former and Boryangraecum vesiculiferum (Schltr.) Szlach., single in the latter genus. Additionally, the stem of Mytnik & Grochocka, comb. nov. Lemurangis is elongate with short leaves, while the Basionym: Angraecum vesiculiferum Schltr., stem of Boryangraecum is short, abbreviated with long Repert.­ Spec. Nov. Regni Veg. Beih. 33: 341. 1925. leaves. Boryangraceum shares similar type of pollinaria Boryangraecum xylopus (Rchb.f.) Szlach., Mytnik with Gomphocentrum, but our molecular analyses sug- & Grochocka, comb. nov. gest that it can be a result of homoplasy. It is coherent Basionym: Angraecum xylopus Rchb.f., Flora 68: with flower morphology of both genera. The flowers of 538. 1885. Gomphocentrum are resupinate with filiform spur with narrow entrance. 5. Dolabrifolia (Pfitzer) Szlach. & Romowicz Boryangraecum includes 13 species. Richardiana 7: 54. 2007. ≡ Mystacidium Lindl. sect. Boryangraecum aviceps (Schltr.) Szlach., Mytnik Dolabrifolia Pfitz.in Engl. & Prantl, Natürl. Pflanzen- & Grochocka, comb. nov. fam. 2(6): 216. 1889; G e n e r i t y p e : Angraecum dis- Basionym: Angraecum aviceps Schltr., Repert. tichum Lindl. [≡ Dolabrifolia disticha (Lindl.) Szlach. Spec. Nov. Regni Veg. Beih. 33: 335. 1925. & Romowicz] ≡ Angraecum Bory sect. Dolabrifolia Boryangraecum flavidum (Bosser) Szlach., Mytnik (Pfitz.) Garay, Kew Bull. 28(3): 499. 1973. & Grochocka, comb. nov. = Aeranthus Lindl. sect. Dolabrifolia Rchb.f., Walp., Basionym: Angraecum flavidum Bosser, Adanso- Ann. Bot. Syst. 6: 901. 1864, nom. nud. nia, n.s., 10: 100. 1970. = Epidorchis Thouars sect. Dolabraria Kuntze in Post Boryangraecum myrianthum (Schltr.) Szlach., & Kuntze, Lex. Gen. Phan.: 200. 1903, nom. illegit. Mytnik & Grochocka, comb. nov. Small, caespitose plant with elongate stem. Basionym: Angraecum myrianthum Schltr., Ann. Leaves distichous, equitant, basally imbricating, Inst. Bot.-Géol. Colon. Marseille, III, 1: 197. 1913. somewhat fleshy, thick. Inflorescence 1-flowered, Boryangraecum ochraceum (Schltr.) Szlach., Myt- almost sessile, peduncle very short. Flowers tiny, nik & Grochocka, comb. nov. nonresupinate, pure white. Floral bracts amplexicaul. Basionym: Mystacidium ochraceum Ridl., J. Linn. Pedicellate ovary terete. Sepals and petals dissimilar, Soc., Bot. 21: 488. 1885. petals smaller and narrower. Lip transversely elliptic, Boryangraecum pinifolium (Bosser) Szlach., Myt- more or less 3-lobed at the apex, acuminate apically, nik & Grochocka, comb. nov. cochleate and canaliculate, ecallose. Spur cylindrical, Basionym: Angraecum pinifolium Bosser, Adan- more or less as long as the pedicellate ovary, orifice sonia, n. s., 10: 99. 1970. rather narrow. Boryangraecum pumilio (Schltr.) Szlach., Mytnik The genus is easily distinguished from all other & Grochocka, comb. nov. Angraecinae­ by its peculiar habit, equitant, short leaves Basionym: Angraecum pumilio Schltr., Beih. Bot. and short-pedunculed inflorescence. It includes 4-5 spe- Centralbl. 34(2): 337. 1916. cies depending on the authors.

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Dolabrifolia aporoides (Summerh.) Szlach. & Ro- Schltr. by Schlechter (1918) and Garay (1973), which mowicz was typified by Garay (1973) byAngraecum ramosum Richardiana 7: 54. 2007. ≡ Angraecum aporoides Sum- Thouars, and intermixed along with such species as merh., Kew Bull. 17: 560. 1964. Angraecum conchiferum Lindl., A. teretifolium Ridl. Dolabrifolia bancoensis (Burg) Szlach & Romowicz and A. viguieri Schltr. All the taxa are characterised by Richardiana 7: 54. 2007. ≡ Angraecum bancoensis similar habit, but differ in the flower details, such as the Burg , Misc. Pap. Landbouwhogeschool, Wageningen lip form. The results of our molecular analyses clearly 19: 26. 1980. indicate that the section broadly defined is polyphyletic. Dolabrifolia disticha (Lindl.) Szlach. & Romowicz Eichlerangraecum birrimense (Rolfe) Szlach., Richardiana 7: 54. 2007. ≡ Lindl., Mytnik & Grochocka, comb. nov. Edwards’s Bot. Reg. 21: t. 1781. 1836. Basionym: Angraecum birrimense Rolfe, Bull. Dolabrifolia podochiloides (Schltr.) Szlach. & Ro- Misc. Inform. Kew 1914: 214. 1914. mowicz Eichlerangraecum eichlerianum (Kraenzl.) Richardiana 7: 54. 2007. ≡ Angraecum podochiloides Szlach., Mytnik & Grochocka, comb. nov. Schltr., Bot. Jahrb. Syst. 38: 162. 1906. Basionym: Angraecum eichlerianum Kraenzl., Dolabrifolia poppendickiana (Szlach. & Olszewski) Berliner Allg. Gartenzeitung 1: 434. 1882. Szlach. & Romowicz Eichlerangraecum eichlerianum (Kraenzl.) Szlach., Richardiana 7: 54. 2007. ≡ Angraecum poppendicki- Mytnik & Grochocka var. curvicalcaratum (Szlach. & anum Szlach. & Olszewski, Fl. Cameroun 36: 884. Olszewski) Szlach., Mytnik & Grochocka, comb. nov., 2001. Basionym: Angraecum eichlerianum Kraenzl. var curvicalcaratum Szlach. & Olszewski, Fl. Cameroun, 6. Eichlerangraecum Szlach., Mytnik & Grochocka, Orchid. 36: 898. 2001. gen. nov. Eichlerangraecum infundibulare (Lindl.) Szlach., Stem long, climbing, internodes elongate, leaves Mytnik & Grochocka, comb. nov. alternate. Inflorescence usually 1-4-flowered. Flowers Basionym: Angraecum infundibulare Lindl., J. resupinate, large, showy. Lip very large, transversely Proc. Linn. Soc., Bot. 6: 136. 1862. elliptic to obreniform, cochleate in the center, with Eichlerangraecum spectabile (Summerh.) Szlach., prominent keel along the midvein in the basal part. Spur Mytnik & Grochocka, comb. nov. very prominent, infundibular in the basal part. Tegula Basionym: Angraecum spectabile Summerh., Kew single, oblong-elliptic, viscidium single, lanceolate to Bull. 13: 274. 1958. ovate, oblong. Generitype: Angraecum eichlerianum Kraenzl. [≡ 7. Gomphocentrum (Benth.) Szlach., Mytnik & Gro- Eichlerangraecum eichlerianum (Kraenzl.) Szlach., chocka, stat. et gen. nov. Mytnik & Grochocka]. Basionym: Mystacidium Lindl. sect. Gompho- E t y m o l o g y : A combination of the specific epithet centrum Benth., J. Linn. Soc. Bot. 18: 337. 1881; of the generitype and the name Angraecum. Generitype: Angraecum caulescens Thouars [≡ Plants rather large. Stem long, climbing, internodes Gomphocentrum caulescens (Thouars) Szlach., Mytnik elongate, leaves alternate. Leaves oblong to elliptic, & Grochocka]. ≡ Angraecum sect. Gomphocentrum unequally bilobed at the apex. Inflorescence usually (Benth.) Garay, Kew Bull. 28(3): 501. 1973. 1-4-flowered. Flowers resupinate, large, showy. Floral Plants medium-sized. Stem elongate or more often bracts inconspicuous. Pedicellate ovary slender, terete. abbreviated, erect. Leaves lanceolate to oblong-lanceo- Sepals and petals subsimilar, narrowly lanceolate to late, attenuate and unequally bilobed at the apex, thin. linear-lanceolate, acute, spread. Lip very large, trans- Inflorescence elongate, peduncle thin, wiry, somewhat versely elliptic to obreniform, cochleate in the centre fractiflex, usually laxly several- to many-flowered. becoming conical towards the base, more or less apicu- Flowers small, inconspicuous, resupinate, greenish to late at the apex, more or less 3-lobed at the apex, with yellowish, thin in texture, rather diaphanous. Floral prominent keel along the midvein in the basal part. Spur bracts inconspicuous. Pedicellate ovary ribbed. Sepals very prominent, infundibular in the basal part, slender, and petals subsimilar, acute. Lip ovate, long-acuminate, cylindrical above, attenuate towards the apex. Tegula strongly cochleate, ecallose. Spur cylindrical, with nar- single, oblong-elliptic, viscidium single, lanceolate to row entrance, apex swollen or not. Viscidia 2, small, ovate, oblong. elliptic, tegulae 2, oblong-oblanceolate, thin. The genus includes four species known from conti- Molecular analyses indicate close relationships nental Africa, three of them occur in western Africa, the between this genus and Aeranthes Lindl. They share fourth, E. spectabile, has been noted in . They similar habit, leaf form and texture, but are essentially were classified within the section Arachnangraecum different in the flower morphology.

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The genus, as proposed here, includes 18 species. Basionym: Angraecum rhizanthium H. Perrier, Gomphocentrum acutipetalum (Schltr.) Szlach., Notul. Syst. (Paris) 14: 163. 1951. Mytnik & Grochocka, comb. nov. Gomphocentrum sacculatum (Schltr.) Szlach., Basionym: Angraecum acutipetalum Schltr., Beih. Mytnik & Grochocka, comb. nov. Bot. Centralbl. 34(2): 337. 1916. Basionym: Angraecum sacculatum Schltr., Repert. Gomphocentrum andringitranum (Schltr.) Szlach., Spec. Nov. Regni Veg. Beih. 33: 347. 1925. Mytnik & Grochocka, comb. nov. Gomphocentrum tenuipes (Summerh.) Szlach., Basionym: Angraecum andringitranum Schltr., Mytnik & Grochocka, comb. nov. Repert. Spec. Nov. Regni Veg. Beih. 33: 344. 1925. Basionym: Angraecum tenuipes Summerh., Kew Gomphocentrum calceolus (Thouars) Szlach., Myt- Bull. 6: 473. 1951 (publ. 1952). nik & Grochocka, comb. nov. Gomphocentrum undulatum (Schltr.) Szlach., Basionym: Angraecum calceolus Thouars, Hist. Mytnik & Grochocka, comb. nov. Orchid.: t. 77. 1822. Basionym: Mystacidium undulatum Cordem., Rev. Gomphocentrum caulescens (Thouars) Szlach., Gén. Bot. 11: 425. 1899. ≡ Angraecum undulatum (Cor- Mytnik & Grochocka, comb. nov. dem.) Schltr., Beih. Bot. Centralbl. 33(2): 348. 1915. Basionym: Angraecum caulescens Thouars, Hist. Gomphocentrum verecundum (Schltr.) Szlach., Orchid.: t. 75. 1822. Mytnik & Grochocka, comb. nov. Gomphocentrum cordemoyi (Schltr.) Szlach., Myt- Basionym: Angraecum verecundum Schltr., Repert. nik & Grochocka, comb. nov. Spec. Nov. Regni Veg. Beih. 33: 348. 1925. Basionym: Angraecum cordemoyi Schltr., Beih. Gomphocentrum vesiculatum (Schltr.) Szlach., Bot. Centralbl. 33(2): 432. 1915. Mytnik & Grochocka, comb. nov. Gomphocentrum cornucopiae (H. Perrier) Szlach., Basionym: Angraecum vesiculatum Schltr., Repert. Mytnik & Grochocka, comb. nov. Spec. Nov. Regni Veg. Beih. 33: 348. 1925. Basionym: Angraecum cornucopiae H. Perrier, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 265. 1955. 8. Hadrangis (Schltr.) Szlach., Mytnik & Grochocka, Gomphocentrum crassifolium (Schltr.) Szlach., stat. et gen. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum Bory sect. Hadrangis Basionym: Mystacidium crassifolium Cordem., Schltr., Beih. Bot. Centralbl. 36(2): 158. 1918; G e – Rev. Gén. Bot. 11: 422. 1899. ≡ Angraecum crassi- neritype: Angraecum striatum Thouars [≡ Hadran- folium (Cordem.) Schltr., Beih. Bot. Centralbl. 33(2): gis striata (Thouars) Szlach., Mytnik & Grochocka]. 433.1915. = Angraecum sect. Thouarsiangraecum Schltr., Repert. Gomphocentrum dauphinense (Schltr.) Szlach., Sp. Nov. Regni Veg, Beih. 33: 310. 1925; nom. illegit. Mytnik & Grochocka, comb. nov. Generitype: Angraecum striatum Thouars. Basionym: Mystacidium dauphinense Rolfe ex Stem short, stout, erect, leafy. Leaves ligulate to Scott-Elliot, J. Linn. Soc., Bot. 29: 55. 1891.≡ Angrae- oblong, stiff, unequally bilobed at the apex, coriaceous cum dauphinense (Rolfe) Schltr., Beih. Bot. Centralbl. or leathery. Inflorescence several-flowered. Flowers 33(2): 433. 1915. fleshy, resupinate, rather small to medium-sized. Floral Gomphocentrum guillauminii (H. Perrier) Szlach., bracts conspicuous, oblong-ovate. Ovary shortly pedi- Mytnik & Grochocka, comb. nov. cellate. Sepals and petals subsimilar, ovate-lanceolate, Basionym: Angraecum guillauminii H. Perrier, acute. Lip ovate-lanceolate, unlobed, long-acuminate, Bull. Mus. Natl. Hist. Nat., II, 22: 114. 1950. concave, ecallose. Spur short, conical to saccate, obtuse Gomphocentrum inapertum (Thouars) Szlach., at the apex, orifice wide. Mytnik & Grochocka, comb. nov. We had no access to any materials proper to molecu- Basionym: Angraecum inapertum Thouars, Hist. lar analyses representing this group. The species of the Orchid.: 50. 1822. genus appear to be similar to those of Lemurorchis. They Gomphocentrum laggiarae (Schltr.) Szlach., Mytnik share similar habit, long, multiflowered inflorescence & Grochocka, comb. nov. and general flower morphology, but differ clearly one Basionym: Angraecum laggiarae Schltr., Repert. from another by the lip and spur morphology. Lip of Spec. Nov. Regni Veg. 15: 337. 1918. Lemurochis is deeply 3-lobed, with both lateral lobes Gomphocentrum multiflorum (Thouars) Szlach., transversely elliptic, upcurved forming a kind of tube Mytnik & Grochocka, comb. nov. around gynostemium and spur entrance. It produces Basionym: Angraecum multiflorum Thouars, Hist. long, cylindrical spur. Lip of Hadrangis is concave and Orchid.: 74. 1822. spur is short, conical to saccate. These differencies are Gomphocentrum rhizanthium (H. Perrier) Szlach., probably result of adaptation to different pollination Mytnik & Grochocka, comb. nov. systems or different groups of pollinators.

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Hadrangis bracteosa (Balf. f. & S. Moore) Szlach., Hermansia pergracile (Schltr.) Szlach., Mytnik & Mytnik & Grochocka, comb. nov. Grochocka, comb. nov. Basionym: Angraecum bracteosum Balf. f. & S. Basionym: Angraecum pergracile Schltr., Repert. Moore, J. Bot. 14: 293. 1876. Spec. Nov. Regni Veg. Beih. 33: 337. 1925. Hadrangis cadetii (Bosser) Szlach., Mytnik & Gro- Hermansia rhynchoglossa (Schltr.) Szlach., Mytnik chocka, comb. nov. & Grochocka, comb. nov. Basionym: Angraecum cadetii Bosser, Bull. Mus. Basionym: Angraecum rhynchoglossum Schltr., Nation. Hist. Nat., B, Adansonia, Ser. 4, 9(3): 252. 1987. Repert. Spec. Nov. Regni Veg. Beih. 33: 339. 1925. Hadrangis striata (Thouars) Szlach., Mytnik & Hermansia setipes (Schltr.) Szlach., Mytnik & Grochocka, comb. nov. Grochocka, comb. nov. Basionym: Angraecum striatum Thouars, Hist. Basionym: Angraecum setipes Schltr., Repert. Orchid.: 72. 1822. Spec. Nov. Regni Veg. Beih. 33: 340. 1925.

9. Hermansia Szlach., Mytnik & Grochocka, nom. et 10. Humblotiangraecum (Schltr.) Szlach., Mytnik & stat. nov. Grochocka, stat. et gen. nov. Replaced synonym: Angraecum Bory sect. Basionym: Angraecum Bory sect. Humblotian- Acaulia Garay, Kew Bull. 28(3): 498. 1973; G e – graecum Schltr., Repert. Sp. Nov. Regni Veg., Beih. 33: neritype: Angraecum rhynchoglossum Schltr. [≡ 310. 1925; Generitype: Aeranthes leonis Rchb.f. Hermansia rhynchoglossa (Schltr.) Szlach., Mytnik [≡ Humblotiangraecum leonis (Rchb.f.) Szlach., Myt- & Grochocka]. nik & Grochocka]. Etymology: Dedicated to Johan Hermans, specia­ Plants almost stemless, with arrested internodes. list­ of angrecoid orchids. Leaves thick, fleshy, equitant, falcate, basally imbricat- Plants small. Stem abbreviated, very short. Leaves ing. Inflorescence few-flowered. Flowers large, showy, several, linear to oblong-ligulate, unequally bilobed at white, resupinate. Floral bracts prominent, amplexicaul. the apex. Inflorescence elongate, usually longer than Pedicellate ovary slender, elongate. Sepals and petals leaves, wiry, 1-2-flowered. Flowers small, resupinate, subsimilar, lanceolate or similar, acute. Lip large, el- yellowish-white to yellowish-green. Floral bracts small. liptic, concave, funnel-shaped in the lower half, keeled, Pedicellate ovary slender, terete. Sepals and petals acuminate towards the apex. Spur long, prominent, subsimilar, lanceolate, acute to acuminate. Lip ovate- filiform, attenuate towards the apex, orifice spacious, lanceolate to elliptic with acuminate apex, cochleate, broad. canaliculate, thin-textured. Spur filiform, straight, The genus includes a few species which are easily somewhat swollen at the apex, with narrow entrance. distinguishable from all other Angraecinae by their The section Acaulia has been described by Garay peculiar habit, i.e., abbreviated internodes of the stem, (1973) to accomodate the species of the section Bo- equitant leaves, which basally are usually imbricating, ryangraecum with 1-2-flowered inflorescence. Garay and broad spur orifice. The species of Humblotian- (1973) agreed with Schlechter (1918), who considered graecum and Rudolfangraecum are intermixed on 1-flowered inflorescence to be a mere state of reduction the phylogenetic tree presented here. However, we from the several-flowered condition. The species of both decided to keep them separately based on the habit sections, Acaulia and Boryangraceum, differ not only characters. in the number of flowers per inflorescence, but also in Humblotiangraecum aloifolium (Hermans & the length and form of the spur. Acaulia comprises spe- P.J.Cribb) Szlach., Mytnik & Grochocka, comb. nov. cies with filiform, elongate spur, and Boryangraecum Basionym: Angraecum aloifolium Hermans & P. J. – with relatively short spur, hardly exceeding pedicellate Cribb, Orchid Rev. 105: 108 (1997). ovary, which is swollen and blunt at the apex. In our Humblotiangraecum clareae (Hermans, la Croix & opinion, these differences indicate different pollination P. J. Cribb) Szlach., Mytnik & Grochocka, comb. nov. syndrome. Basionym: Angraecum clareae Hermans, la Croix Five species constitute this genus. & P. J. Cribb, Orchid Rev. 109: 43. 2001. Hermansia brachyrhopalon (Schltr.) Szlach., Myt- Humblotiangraecum equitans (Schltr.) Szlach., nik & Grochocka, comb. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum brachyrhopalon Schltr., Basionym: Angraecum equitans Schltr. Beih. Bot. Repert. Spec. Nov. Regni Veg. Beih. 33: 336. 1925. Centralbl. 34(2): 339. 1916. Hermansia chaetopoda (Schltr.) Szlach., Mytnik & Humblotiangraecum leonis (André) Szlach., Myt- Grochocka, comb. nov. nik & Grochocka, comb. nov. Basionym: Angraecum chaetopodum Schltr., Basionym: Aeranthes leonis Rchb. f., Gard. Chron., Repert.­ Spec. Nov. Regni Veg. Beih. 33: 336. 1925. n. s., 23: 726. 1885.

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Synonym: (Rchb. f.) André, Rev. Basionym: Angraecum floribundum Bosser, Adan- Hort. 57: 294. 1885. sonia, n.s., 10: 102. 1970. Humblotiangraecum potamophilum (Schltr.) Lemurangis humile (Summerh.) Szlach., Mytnik & Szlach., Mytnik & Grochocka, comb. nov. Grochocka, comb. nov. Basionym: Angraecum potamophilum Schltr. Ann. Basionym: Angraecum humile Summerh., Kew Mus. Colon. Marseille, sér. 3, 1: 199, t. 23. 1913. Bull. 13: 269. 1958. Lemurangis longinoda (Frapp. ex Cordem.) Szlach., 11. Lemurangis (Garay) Szlach., Mytnik & Grochocka, Mytnik & Grochocka, comb. nov. stat. nov. Basionym: Angraecum longinode Frapp. ex Cor- Basionym: Angraecum Bory sect. Lemurangis dem., Fl. Réunion: 210. 1895. Garay, Kew Bull. 28(3): 501. 1973; G e n e r i t y p e : Lemurangis madagascariensis (Schltr.) Szlach., Macroplectrum madagascariense Finet. ≡ Angrae- Mytnik & Grochocka, comb. nov. cum madagascariense (Finet) Schltr. [≡ Lemuran- Basionym: Macroplectrum madagascariense Finet, gis madagascariensis (Finet) Szlach., Mytnik & Bull. Soc. Bot. France 54(9): 25. 1907. ≡ Angraecum Grochocka]. madagascariense (Finet) Schltr., Beih. Bot. Centralbl. Plants small. Stem elongate, leafy. Leaves oblong 36(2): 162. 1918. to ligulate, rather thick, fleshy, unequally bilobed at Lemurangis pseudopetiolata (Frapp. ex Cordem.) the apex, often falcately curved. Inflorescence usu- Szlach., Mytnik & Grochocka, comb. nov. ally 1-5-flowered, occasionally 1-flowered, mostly Basionym: Angraecum pseudopetiolatum Frapp. shorter than leaves, more or less fractiflex. Flowers ex Cordem., Fl. Réunion: 207. 1895. tiny or small, nonresupinate, greenish, yellowish or whittish, often diaphanous. Pedicellate ovary short. 12. Lepervenchea Cordem. Sepals and petals dissimilar, acute. Lip lanceolate- Rev. Gen. Bot. 11: 415.1899; Generitype: Angrae- ovate to broadly lanceolate, acute to acuminate, cum tenuifolium Frapp. ex Cordem. [≡ Lepervenchea concave at the base, canaliculate. Spur cylindrical tenuifolia (Frapp. ex Cordem.) Cordem.]. or saccate, blunt, usually shorter than pedicel and Stem prominent, elongate. Leaves numerous, ligu- ovary. Viscidium single, transversely elliptic, tegula late to linear, unequally bilobed at the apex, sometimes single, ligulate. falcately curved. Inflorescence as long as or longer than Molecular analyses clearly indicate that the genus leaves, racemose, rachis somewhat flexuos. Flowers small, is sister to Lemurorchis and Angraecum and together resupinate, thin in texture, subdiaphanous. Floral bracts form a monophyletic group. The species of the last two inconspicuous. Pedicellate ovary terete. Sepals and petals genera constituing this clade usually produce elongate, dissimilar, acute. Lip subcordate to oblong-lanceolate, many-flowered inflorescence. The exception are species more or less acuminate apically, occasionally obscurely of Lemurangis which flowers are born on 1-5-flowe­ 3-lobed, cochleate, ecallose. Spur relatively short, hardly red . All members of this group carry pollinia exceeding pedicellate ovary, usually much shorter, cylin- attached ­to single tegula and viscidium. drical to saccate, blunt, straight. Pollinaria double. Lemurangis includes nine species. Considering the habit, the species of Lepervenchea Lemurangis alleizettei (Schltr.) Szlach., Mytnik & may be confused with Gomphocentrum and Lemuran- Grochocka, comb. nov. gis. Due to the double pollinaria, the species of Leper- Basionym: Angraecum alleizettei Schltr., Repert. venchea are similar to Gomphocentrum, but both genera Spec. Nov. Regni Veg. 18: 325. 1922. differ in the spur form. In Gomphocentrum the spur is Lemurangis costata (Frapp. ex Cordem.) Szlach., narrowly cylindrical, adapted to penetration by insect’s Mytnik & Grochocka, comb. nov. long proboscis, whereas the spur in Lepervenchea is Basionym: Angraecum costatum Frapp. ex Cor- short and rather spacious. Basing on the spur and flower dem., Fl. Réunion: 211. 1895. morphology, we can speculate that both Lepervenchea Lemurangis decaryana (H. Perrier) Szlach., Mytnik and Lemurangis exploit the same group of insects as & Grochocka, comb. nov. pollinators. However, both genera are easily distinguish- Basionym: Angraecum decaryanum H. Perrier, able one from another by different number of pollinaria Notul. Syst. (Paris) 7: 129. 1938. produced in the flower, that is, single inLemurangis and Lemurangis falcifolia (Bosser) Szlach., Mytnik & double in Lepervenchea. Grochocka, comb. nov. The genus includes seven species. Basionym: Angraecum falcifolium Bosser, Adan- Lepervenchea appendiculoides (Schltr.) Szlach., sonia, n.s., 10: 104. 1970. Mytnik & Grochocka, comb. nov. Lemurangis floribunda (Bosser) Szlach., Mytnik & Basionym: Angraecum appendiculoides Schltr., Grochocka, comb. nov. Repert. Spec. Nov. Regni Veg. 18: 325. 1922.

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Lepervenchea caricifolia (H. Perrier) Szlach., Myt- The genus includes about 20 species. nik & Grochocka, comb. nov. Lesliegraecum andasibeense (H. Perrier) Szlach., Basionym: Angraecum caricifolium H. Perrier, Mytnik & Grochocka, comb. nov. Notul. Syst. (Paris) 7: 128. 1938. Basionym: Angraecum andasibeense H. Perrier, Lepervenchea montana (Piers) Szlach., Mytnik & Notul. Syst. (Paris) 7: 124. 1938. Grochocka, comb. nov. Lesliegraecum bemarivoense (Schltr.) Szlach., Basionym: Angraecum montanum Piers, Orchids Mytnik & Grochocka, comb. nov. E. Afr.: 244. 1968. Basionym: Angraecum bemarivoense Schltr., Rep- Lepervenchea musculifera (H. Perrier) Szlach., ert. Spec. Nov. Regni Veg. Beih. 33: 332. 1925. Mytnik & Grochocka, comb. nov. Lesliegraecum burchellii (Rchb.f.) Szlach., Mytnik Basionym: Angraecum musculiferum H. Perrier, & Grochocka, comb. nov. Notul. Syst. (Paris) 7: 129. 1938. Basionym: Angraecum burchellii Rchb.f., Flora Lepervenchea pauciramosa (Schltr.) Szlach., Myt- 50: 117. 1867. nik & Grochocka, comb. nov. Lesliegraecum (Schltr.) Szlach., Basionym: Angraecum pauciramosum Schltr., Mytnik & Grochocka, comb. nov. Repert. Spec. Nov. Regni Veg. Beih. 33: 350. 1925. Basionym: Angraecum chamaeanthus Schltr., Bot. Lepervenchea tenuifolia (Frapp. ex Cordem.) Jahrb. Syst. 53: 604. 1915. Szlach., Mytnik & Grochocka, comb. nov. Lesliegraecum decipiens (Summerh.) Szlach., Myt- Basionym: Angraecum tenuifolium Frapp. ex Cor- nik & Grochocka, comb. nov. dem., Fl. Réunion: 207. 1895. Basionym: Angraecum decipiens Summerh., Kew Lepervenchea tenuispica (Schltr.) Szlach., Mytnik Bull. 20: 189. 1966. & Grochocka, comb. nov. Lesliegraecum microcharis (Schltr.) Szlach., Myt- Basionym: Angraecum tenuispica Schltr., Repert. nik & Grochocka, comb. nov. Spec. Nov. Regni Veg. 15: 339. 1918. Basionym: Angraecum microcharis Schltr., Repert. Spec. Nov. Regni Veg. Beih. 33: 333. 1925. 13. Lesliegraecum Szlach., Mytnik & Grochocka, stat. Lesliegraecum minus (Summerh.) Szlach., Mytnik et nom. nov. & Grochocka, comb. nov. Replaced synonym: Mystacidium Lindl. sect. Basionym: Angraecum minus Summerh., Kew Nana Cordem., Rev. Gén. Bot. 11: 414. 1899; L e c t o - Bull. 13: 264. 1958. t y p e (Garay 1973: 502): Angraecum nanum Frapp. Lesliegraecum minutum (Frapp. ex Cordem.) [≡ Lesliegraecum nanum (Frapp.) Szlach., Mytnik & Szlach., Mytnik & Grochocka, comb. nov. Grochocka]. ≡ Angraecum Bory sect. Nana (Cordem.) Basionym: Angraecum minutum Frapp. ex Cor- Garay, Kew Bull. 28(3): 502. 1973. dem., Fl. Réunion: 209. 1895. E t y m o l o g y : Dedicated to Dr. Leslie Garay, an Lesliegraecum muscicolum (H.Perrier) Szlach., eminent American orchidologist. Mytnik & Grochocka, comb. nov. Plants small to tiny with abbreviated stem. Leaves Basionym: Angraecum muscicolum H.Perrier, few to several, linear-lanceolate, usually falcate, thick. Notul. Syst. (Paris) 7: 126. 1938. Inflorescence longer than leaves, laxly to subdensely Lesliegraecum nanum (Frapp. ex Cordem.) Szlach., few- to many-flowered, racemose. Flowers nonresupi- Mytnik & Grochocka, comb. nov. nate, tiny, inconspicuous, diaphanous, greenish, whit- Basionym: Angraecum nanum Frapp. ex Cordem., tish, yellowish to pinkish. Floral bracts prominent. Pedi- Fl. Réunion: 208. 1895. cellate ovary short. Sepals and petals dissimilar, acute Lesliegraecum oberonia (Finet) Szlach., Mytnik & to acuminate. Lip ovate-lanceolate to oblong-ovate, Grochocka, comb. nov. acute to acuminate, concave at the base, canaliculate. Basionym: Angraecum oberonia Finet, Bull. Soc. Spur saccate to conical, blunt, short. Viscidium single Bot. France 54(9): 10. 1907. oblong-elliptic, tegulae double, linear. Lesliegraecum onivense (H. Perrier) Szlach., Myt- We do not have access to any material of this group nik & Grochocka, comb. nov. appropriate to molecular analyses. Based on the plant Basionym: Angraecum onivense H. Perrier, Notul. habit and flower morphology, we can speculate thatLes - Syst. (Paris) 7: 122. 1938. liegraecum appears to be related to Lemurangis, how- Lesliegraecum parvulum (Ayres ex S.Moore) ever, a phylogenetic study is required. It is interesting­ to Szlach., Mytnik & Grochocka, comb. nov. note that both flower arrangement in the inflorescence Basionym: Angraecum parvulum Ayres ex S. and general flower architecture in Lesliegraecum and Moore in J. G. Baker, Fl. : 357. 1877. Schltr. are alike, what may suggest an adapta- Lesliegraecum perhumile (H. Perrier) Szlach., tion to similar pollination strategies. Mytnik & Grochocka, comb. nov.

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Basionym: Angraecum perhumile H. Perrier, Notul. Basionym: Angraecum bosseri Senghas, Die Orchi- Syst. (Paris) 7: 127. 1938. dee 24: 193. 1973. ≡ Angraecum sesquipedale Thouars Lesliegraecum perparvulum (H. Perrier) Szlach., var. angustifolium Bosser & Morat, Adansonia, n.s., Mytnik & Grochocka, comb. nov. 12: 77. 1972. Basionym:Angraecum perparvulum H. Perrier, Macroplectrum sesquipedale (Thouars) Szlach., Notul. Syst. (Paris) 7: 123. 1938. Mytnik & Grochocka, comb. nov. Lesliegraecum pusillum (Lindl.) Szlach., Mytnik Basionym: Angraecum sesquipedale Thouars, Hist. & Grochocka, comb. nov. Orchid.: 66. 1822. Basionym: Angraecum pusillum Lindl., Companion Macroplectrum sororium (Schltr.) Szlach., Mytnik Bot. Mag. 2: 205. 1836. & Grochocka, comb. nov. Lesliegraecum salazianum (Cordem.) Szlach., Basionym: Angraecum sororium Schltr., Repert. Mytnik & Grochocka, comb. nov. Spec. Nov. Regni Veg., Beih. 33: 360. 1925. Basionym: Mystacidium salazianum Cordem., Rev. Gén. Bot. 11: 423. 1899. ≡ Angraecum salazi- 15. Pectinariella Szlach., Mytnik & Grochocka, stat. anum (Cordem.) Schltr., Beih. Bot. Centralbl. 33(2): et. nom. nov. 427. 1915. Replaced synonym: Mystacidium Lindl. sect. Lesliegraecum spicatum (Cordem.) Szlach., Mytnik Benth. in Benth. & Hook.f., Gen. Pl. 3: & Grochocka, comb. nov. 585. 1883; Generitype: Angraecum pectinatum ­ Basionym: Mystacidium spicatum Cordem., Rev. Thouars­ [≡ Pectinariella pectinata (Thouars)]. ≡ An- Gén. Bot. 11: 423. 1899. ≡ Angraecum spicatum (Cor- graecum Bory sect. Pectinaria (Benth.) Schltr., Beih. dem.) Schltr., Beih. Bot. Centralbl. 33(2): 437. 1915. Bot. Centralbl. 36(2): 157. 1918, non Pectinaria Cor- Lesliegraecum tenellum (Ridl.) Szlach., Mytnik & dem., Rev. Gen. Bot. 11: 412. 1899, non Pectinaria Grochocka, comb. nov. Bernh., 1800 (Apiaceae); ­non Pectinaria Haw., 1819 Basionym: Mystacidium tenellum Ridl., J. Linn. (Apocynaceae); non Pectinaria (Benth.) Hack., 1887 Soc., Bot. 21: 489. 1885. ≡ Angraecum tenellum (Ridl.) (Poaceae). Schltr., Beih. Bot. Centralbl. 33(2): 438. 1915. Etymology: -ella – similar to Pectinaria; name of Lesliegraecum viridiflorum (Cordem.) Szlach., one of the sections of Angraecum. Raising the name Mytnik & Grochocka, comb. nov. Pectinaria to the generic rank is illegitimate due to ICN. Basionym: Angraecum viridiflorum Cordem., Rev. Stem long, usually pendent, branching, loosely leafy, Gén. Bot. 11: 9. 1899. internodes elongate. Leaves alternate, fleshy, linear to linear-lanceolate, shortly acuminate, never equitant, 14. Macroplectrum Pfitzer petiole twisted. Inflorescence always single-flowered. in Engl. & Prantl., Nat. Pflanzenfam. 2(6): 208. 1889; Flowers tiny, subresupinate, white, scented. Floral Generitype: Angraecum sesquipedale Thouars bracts small, amplexicaul. Pedicellate ovary short. Se- [≡ Macroplectrum sesquipedale (Thouars) Pfitzer]. pals and petals subsimilar, obtuse to subobtuse at the Stem erect, occasionally branching, stout. Leaves apex. Lip ligulate, oblong to transversely elliptic, cana- ligulate, unequally bilobed apically, leathery, stout, liculate, ecallose. Spur cylindrical to clavate, subequal usually gathered at the apex of the stem. Inflorescence in length to the ovary, sometimes shorter, occasionally few-flowered, racemose. Flowers large, showy, white, longer than pedicellate ovary, often swollen and blunt strongly fragrant, resupinate. Pedicellate ovary long, at the apex, orifice usually narrow. Pollinia pyriform, conspicuous. Sepals and petals subsimilar, attenuate tegula greatly reduced, viscidium elliptic, single. towards apex, acute, spread. Lip large, longer than The genus is easily distinguishable from all other wide, canaliculate, ecallose, apically long-acuminate. Angraecinae by its peculiar habit, that is fleshy leaves, Spur very conspicuous, filiform from narrowly conical basally twisted with almost sessile flowers forming base, acute. Tegula and viscidium separated, double, single-flowered inflorescence and ecallose lip. Pecti- hence, two pollinaria are produced. nariella is embedded in the clade comprising Angrae- The genus includes very spectacular species. It was coides and Eichlerangraceum. But the flower size and originally described by Pfitzer (1889) as a monotypic morphology indicate that they exploit different pollina- genus. Later his concept of the genus has been extended tors and evolved under strong pollinator’s pressure. by Finet (1907), who included in Macroplectrum several Pectinariella dasycarpa (Schltr.) Szlach., Mytnik & species from various sections. Grochocka, comb. nov. Macroplectrum includes only three species restricted Basionym: Angraecum dasycarpum Schltr., Repert. in their distribution to Madagascar. Spec. Nov. Regni Veg. 15: 337. 1918. Macroplectrum bosseri (Senghas) Szlach., Mytnik Pectinariella doratophylla (Summerh.) Szlach., & Grochocka, comb. nov. Mytnik & Grochocka, comb. nov.

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Basionym: Angraecum doratophyllum Summerh., Their similarity may result from an exploitation of the Bull. Misc. Inform. Kew 1937: 465. 1937. same group of pollinators. Both genera can be separated Pectinariella gabonense (Summerh.) Szlach., Myt- based on the number of flowers in the inflorescence and nik & Grochocka, comb. nov. the type of floral bracts. Basionym: Angraecum gabonense Summerh., Kew Perrierangraecum includes 34 species. Bull. 8: 587. 1953 (publ. 1954). Perrierangraecum ambrense (H. Perrier) Szlach., Pectinariella hermannii (Cordem.) Szlach., Mytnik Mytnik & Grochocka, comb. nov. & Grochocka, comb. nov. Basionym: Angraecum ambrense H. Perrier, Notul. Basionym: Mystacidium hermannii Cordem., Rev. Syst. (Paris) 7: 118. 1938. Gén. Bot. 11: 421. 1899. ≡ Angraecum hermannii (Cor- Perrierangraecum ankeranense (H. Perrier) dem.) Schltr., Beih. Bot. Centralbl. 33(2): 434. 1915. Szlach., Mytnik & Grochocka, comb. nov. Pectinariella humblotiana (Finet.) Szlach., Mytnik Basionym: Angraecum ankeranense H. Perrier, & Grochocka, comb. nov. Notul. Syst. (Paris) 7: 115. 1938. Basionym: Angraecum humblotianum (Finet) Perrierangraecum bicallosum (H. Perrier) Szlach., Schltr., Beih. Bot. Centralbl. 33(2): 434. 1915. Mytnik & Grochocka, comb. nov. Pectinariella pectinata (Thouars.) Szlach., Mytnik Basionym: Angraecum bicallosum H. Perrier, No- & Grochocka, comb. nov. tul. Syst. (Paris) 7: 111. 1938. Basionym: Angraecum pectinatum Thouars, Hist. Perrierangraecum breve (Schltr.) Szlach., Mytnik Orchid.: 51 .1822. & Grochocka, comb. nov. Pectinariella pungens (Schltr.) Szlach., Mytnik & Basionym: Angraecum breve Schltr., Repert. Spec. Grochocka, comb. nov. Nov. Regni Veg. Beih. 33: 324. 1925. Basionym: Angraecum pungens Schltr., Bot. Jahrb. Perrierangraecum chimanimaniense (G. Will.) Syst. 38: 163. 1906. Szlach., Mytnik & Grochocka, comb. nov. Pectinariella subulata (Lindl.) Szlach., Mytnik & Basionym: Angraecum chimanimaniense G. Will., Grochocka, comb. nov. Kew Bull. 51(3): 557 (1996). 1996. Basionym: Angraecum subulatum Lindl., Compan- Perrierangraecum compactum (Schltr.) Szlach., ion Bot. Mag. 2: 206. 1836. Mytnik & Grochocka, comb. nov. Basionym: Angraecum compactum Schltr., Beih. 16. Perrierangraecum (Schltr.) Szlach., Mytnik & Bot. Centralbl. 34(2): 339. 1916. Grochocka, stat. et gen. nov. Perrierangraecum compressicaule (H. Perrier) Basionym: Angraecum Bory sect. Perrierangrae- Szlach., Mytnik & Grochocka, comb. nov. cum Schltr., Repert. Sp. Nov. Regni Veg., Beih. 33: 309. Basionym: Angraecum compressicaule H. Perrier, 1925; Generitype (Garay 1973: 499): Angraecum Notul. Syst. (Paris) 7: 118. 1938. triquetrum Thouars [≡ Perrierangraecum triquetrum Perrierangraecum cucullatum (Thouars) Szlach., (Thouars) Szlach., Mytnik & Grochocka]. Mytnik & Grochocka, comb. nov. Plants with usually erect, apically leafy stem. Leaves Basionym: Angraecum cucullatum Thouars, Hist. linear, leathery, occasionally thick, unequally bilobed Orchid.: 48. 1822. at the apex. Inflorescence usually 1-flowered, occa- Perrierangraecum curnowianum (Rchb. f.) Szlach., sionally few-flowered. Peduncle completely covered Mytnik & Grochocka, comb. nov. with 3-4 compressed sheaths. Flowers relatively large, Basionym: Aeranthes curnowiana Rchb. f., Gard. showy, resupinate. Pedicellate ovary often triquetros. Chron., n.s., 19: 306. 1883. ≡ Angraecum curnowianum Sepals and petals subsimilar, apically attenuate, long- (Rchb. f.) T. Durand & Schinz, Consp. Fl. Afric. 5: 41. acuminate. Lip distinctly larger than other segments, 1894. simple, ovate-lanceolate to broadly-lanceolate, attenuate Perrierangraecum curvicalcar (Schltr.) Szlach., to apex, long-acuminate, ecallose, canaliculate. Spur Mytnik & Grochocka, comb. nov. long, filiform, acute with very narrow base. Tegula Basionym: Angraecum curvicalcar Schltr., Repert. greatly reduced, viscidia double, separated, hence, two Spec. Nov. Regni Veg. Beih. 33: 325. 1925. pollinaria. Perrierangraecum didieri (Baill. ex Finet) Szlach., The genus, as proposed here, includes the species Mytnik & Grochocka, comb. nov. classified mostly within the section Perrierangrae- Basionym: Macroplectrum didieri Baill. ex Finet, cum so far. They may be confused with the species of Bull. Soc. Bot. France 54(9): 28. 1907. ≡ Angraecum Macroplectrum, because of similar habit, however the didieri (Baill. ex Finet) Schltr., Beih. Bot. Centralbl. representatives of the latter genus are generally smaller 33(2): 433. 1915. plants. The results of our molecular analyses clearly Perrierangraecum divaricatum (Frapp. ex Cor- indicate that both groups are only distantly related. dem..) Szlach., Mytnik & Grochocka, comb. nov.

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Basionym: Angraecum divaricatum Frapp. ex Basionym: Angraecum palmicolum Bosser, Bull. Cordem., Fl. Réunion: 177. 1895. Mus. Natl. Hist. Nat., B, Adansonia, Botanique Phy- Perrierangraecum drouhardii (H. Perrier) Szlach., tochimie 11: 376, f. 4. 1990. Mytnik & Grochocka, comb. nov. Perrierangraecum praestans (Schltr.) Szlach., Basionym: Angraecum drouhardii H. Perrier, No- Mytnik & Grochocka, comb. nov. tul. Syst. (Paris) 7: 117. 1938. Basionym: Angraecum praestans Schltr., Ann. Perrierangraecum dryadum (Schltr.) Szlach., Myt- Mus. Colon. Marseille, sér. 3, 1: 200, t. 21. 1913. nik & Grochocka, comb. nov. Perrierangraecum protensum (Schltr.) Szlach., Basionym: Angraecum dryadum Schltr., Repert. Mytnik & Grochocka, comb. nov. Spec. Nov. Regni Veg., Beih. 33: 326. 1925. Basionym: Angraecum protensum Schltr., Repert. Perrierangraecum elephantinum (Schltr.) Szlach., Spec. Nov. Regni Veg. Beih. 33: 358. 1925. Mytnik & Grochocka, comb. nov. Perrierangraecum pseudodidieri (H. Perrier) Basionym: Angraecum elephantinum Schltr., No- Szlach., Mytnik & Grochocka, comb. nov. tizbl. Bot. Gart. Berlin-Dahlem 7: 330. 1919. Basionym: Angraecum pseudodidieri H. Perrier, Perrierangraecum imerinense (Schltr.) Szlach., Notul. Syst. (Paris) 7: 113. 1937. Mytnik & Grochocka, comb. nov. Perrierangraecum rigidifolium (H. Perrier) Szlach., Basionym: Angraecum imerinense Schltr., Repert. Mytnik & Grochocka, comb. nov. Spec. Nov. Regni Veg., Beih. 33: 328. 1925. Basionym: Angraecum rigidifolium H. Perrier, Perrierangraecum kranzlinianum (H. Perrier) Notul. Syst. (Paris) 7: 116. 1938. Szlach., Mytnik & Grochocka, comb. nov. Perrierangraecum rutenbergianum (Kraenzl.) Basionym: Angraecum kranzlinianum H. Perrier, Szlach., Mytnik & Grochocka, comb. nov. Fl. Madag., Orchid. 4(2): 276. 1941. Basionym: Angraecum rutenbergianum Kraenzl., Perrierangraecum lecomtei (H. Perrier) Szlach., Abh. Naturwiss. Vereine Bremen 7: 257. 1882. Mytnik & Grochocka, comb. nov. Perrierangraecum sambiranoense (Schltr.) Szlach., Basionym: Angraecum lecomtei H. Perrier, Notul. Mytnik & Grochocka, comb. nov. Syst. (Paris) 7: 119. 1938. Basionym: Angraecum sambiranoense Schltr., Perrierangraecum letouzeyi (Bosser) Szlach., Myt- Repert. Spec. Nov. Regni Veg., Beih. 33: 329. 1925. nik & Grochocka, comb. nov. Perrierangraecum stella-africae (P. J. Cribb) Basionym: Angraecum letouzeyi Bosser, Bull. Mus. Szlach., Mytnik & Grochocka, comb. nov. Natl. Hist. Nat., B, Adansonia, Botanique Phytochimie Basionym: Angraecum stella-africae P.J.Cribb, 11(4): 374, f. 3. 1990. Malawi Orch. 1: 134. 1983. Perrierangraecum liliodorum (Frapp. ex Cordem.) Perrierangraecum triquetrum (Thouars) Szlach., Szlach., Mytnik & Grochocka, comb. nov. Mytnik & Grochocka, comb. nov. Basionym: Angraecum liliodorum Frapp. ex Cor- Basionym: Angraecum triquetrum Thouars, Hist. dem., Fl. Réunion: 198. 1895. Orchid.: 49. 1822. Perrierangraecum litorale (Schltr.) Szlach., Mytnik Perrierangraecum urschianum (Toill.-Gen. & & Grochocka, comb. nov. Bosser) Szlach., Mytnik & Grochocka, comb. nov. Basionym: Angraecum litorale Schltr., Repert. Basionym: Angraecum urschianum Toill.-Gen. & Spec. Nov. Regni Veg. Beih. 33: 327. 1925. Bosser, Adansonia, n.s., 1: 101. 1961. Perrierangraecum longicaule (H. Perrier) Szlach., Mytnik & Grochocka, comb. nov. 17. Pseudojumellea (Schltr.) Szlach., Mytnik & Gro- Basionym: Angraecum longicaule H. Perrier, chocka, stat. et gen. nov. Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 266. Basionym: Angraecum Bory sect. Pseudojumellea 1955. Schltr., Beih. Bot. Centralbl. 36(2): 157. 1918; Perrierangraecum obesum (H.Perrier) Szlach., Lectotype (Garay 1973: 500): Angraecum mauri- Mytnik & Grochocka, comb. nov. tianum (Poir.) Frapp. [≡ Pseudojumellea mauritiana Basionym: Angraecum obesum H.Perrier, Notul. (Poir.) Szlach., Mytnik & Grochocka]. Syst. (Paris) 7: 114. 1938. = Angraecum Bory sect. Filangis Garay, Kew Bull. Perrierangraecum oblongifolium (Toilliez-Ge- 28(3): 500. 1973; Type species: Angraecum filicornu noud & Bosser) Szlach., Mytnik & Grochocka, comb. Thouars nov. Plants with elongate stem with prominent internodes. Basionym:Angraecum oblongifolium Toilliez- Leaves linear to ligulate, occasionally oblong-elliptic, Genoud & Bosser, Nat. Malgache 12: 13. 1960. conduplicate, unequally bilobed at apex, sheaths tightly Perrierangraecum palmicolum (Bosser) Szlach., adnate to the stem. Inflorescence always single-flowe­ Mytnik & Grochocka, comb. nov. red. Peduncle prominent, slender, covered at base with

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1-2 adpressed sheaths. Flowers rather large to medium- Basionym: Angraecum implicatum Thouars, Hist. sized, resupinate, showy. Sepals and petals subsimilar, Orchid.: 58. 1822. widely spread, giving the flowers a star appearence. Lip Pseudojumellea mauritiana (Poir.) Szlach., Mytnik oblong-ovate to ovate-lanceolate, widest at the base, & Grochocka, comb. nov. attenuate towards the apex, acute, with more or less Basionym: Orchis mauritiana Poir. in J. B. A .M. conspicuous keel in the basal part. Spur very long, slen- de Lamarck, Encycl. 4: 601. 1798. ≡ Angraecum mau- der, filiform, acute, narrow at the base. Viscidia double, ritianum (Poir.) Frapp., Cat. Orchid. Reunion: 13. 1889. each oblong-elliptic, thin, tegulae double filiform. Pseudojumellea meirax (Rchb. f.) Szlach., Mytnik We did not have an access to any DNA materials & Grochocka, comb. nov. of the Pseudojumellea species. Therefore, our conclu- Basionym: Aeranthes meirax Rchb. f., Flora 68: sions are based on the detailed morphological studies 540. 1885. ≡ Angraecum meirax (Rchb. f.) H. Perrier, only. These species of the two Garay’s (1973) sections, Notul. Syst. (Paris) 7: 120. 1938. that is Filangis and Pseudojumellea, are not separable Pseudojumellea melanosticta (Schltr.) Szlach., one from another. According to Garay (1973), the key Mytnik & Grochocka, comb. nov. character distinguishing the two taxa is the length of Basionym: Angraecum melanostictum Schltr., peduncle, which should be short in most Filangis spe- Repert. Spec. Nov. Regni Veg. 15: 338. 1918. cies and usually long and prominent in Pseudojumellea. Pseudojumellea moratii (Bosser) Szlach., Mytnik In our opinion the taxonomic value and constancy of & Grochocka, comb. nov. this feature is questionable. Basionym: Angraecum moratii Bosser, Adansonia, The species of the genus, as a generic name suggests, n.s., 10: 106. 1970. are like those of Jumellea, but the lip base is broad and Pseudojumellea trichoplectron (Rchb.f.) Szlach., overlaps the gynostemium, in contrast to Jumellea, in Mytnik & Grochocka, comb. nov. which the lip is narrow at the base and does not overlap Basionym: Aeranthes trichoplectron Rchb.f., Gard. the gynostemium. The Pseudojumellea species might be Chron. 1888: 264. 1888. ≡ Angraecum trichoplectron confused with those of Rudolfangraecum, but the former (Rchb.f.) Schltr., Beih. Bot. Centralbl. 33(2): 348. 1915. is characterized by a narrow spur entrance and presence of two separate pollinaria produced in each gynoste- 18. Rudolfangraecum Szlach., Mytnik & Grochocka, mium. Rudolfangraecum possesses broad spur entrance gen. nov. and single pollinarium, that is, pollinia are attached to Stem elongate or relatively short. Leaves condu- V-shaped tegula connected to a single viscidium.­ This plicate, ligulate to linear, unequally bilobed at the differences could emerge as an adaptation to different apex, leathery to fleshy. Inflorescence 1-2-flowered. pollination strategies. Flowers resupinate, showy, scenty. Lip large, unlobed, Pseudojumellea is represented by eleven species. long-acuminate apically, ecallose. Spur elongate, in- Pseudojumellea amplexicaule (Toill.-Gen. & fundibuliform basally, filiform above. Viscidia double, Bosser) Szlach., Mytnik & Grochocka, comb. nov. but connate marginally to one another, forming a single, Basionym: Angraecum amplexicaule Toill.-Gen. very large, more or less reniform structure, tegula much & Bosser, Nat. Malgache 12: 11. 1961. smaller, single. Pseudojumellea cornigera (Cordem.) Szlach., Myt- Generitype: Angraecum magdalenae Schltr. & H. nik & Grochocka, comb. nov. Perrier [≡ Rudolfangraecum magdalenae (Schltr. & H. Basionym: Angraecum cornigerum Cordem., Rev. Perrier) Szlach., Mytnik & Grochocka]. Gén. Bot. 11: 418. 1899. E t y m o l o g y : Dedicated to Dr. Rudolf Schlechter, a Pseudojumellea coutrixii (Bosser) Szlach., Mytnik founder of modern orchidology. & Grochocka, comb. nov. Undoubtedly, the species of this genus are closely Basionym: Angraecum coutrixii Bosser, Adansonia, related to Humblotiangraecum, with which they share n. s., 10: 107. 1970. general flower structure, but are distinguished from the Pseudojumellea filicornu (Thouars) Szlach., Mytnik latter by the habit. They produce more or less elongate & Grochocka, comb. nov. stem with conduplicate, more or less leathery leaves. Basionym: Angraecum filicornu Thouars, Hist. The species of this genus can be misidentified with the Orchid.: 52. 1822. Perrierangraecum species, because they are similar in Pseudojumellea florulenta (Rchb.f.) Szlach., Myt- habit. The clear border line between two genera is the nik & Grochocka, comb. nov. mouth of the spur, in Perrierangraceum it is narrow Basionym: Angraecum florulentumRchb. f., Gard. from the base, in Rudolfangraecum it is infundibuliform. Chron. I. 787. 1885. The genus includes three species. Pseudojumellea implicata (Thouars) Szlach., Myt- Rudolfangraecum dollii (Senghas) Szlach., Mytnik nik & Grochocka, comb. nov. & Grochocka, comb. nov.

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Basionym: Angraecum dollii Senghas, J. Orchi- Rudolfangraecum viguieri (Schltr.) Szlach., Mytnik deenfreund 4: 16. 1997. & Grochocka, comb. nov. Rudolfangraecum magdalenae (Schltr. & H. Per- Basionym: Schltr. Repert. rier) Szlach., Mytnik & Grochocka, comb. nov. Spec. Nov. Regni Veg. 18: 326. 1922. Basionym: Angraecum magdalenae Schltr. & H.Perrier, Repert. Spec. Nov. Regni Veg. Beih. 33: 354. 1925.

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