Jacquelyn A. Kallunki

Phylogenetic relationships of Rutaceae: a cladistic analysis of the subfamilies using evidence from RBC and ATP sequence variation.

Sequence data for plastid rbcL and atpB from members of Anacardiaceae, Burseraceae, Cneoraceae, Meliaceae, Ptaeroxylaceae, Rutaceae, and Simaroubaceae were analyzed cladistically to evaluate the familial and subfamilial circumscriptions of Rutaceae. Taxa representing all subfamilies and tribes were sampled. The analysis shows that Rutaceae are paraphyletic, with Spathelia and Dictyoloma (Rutaceae), Harrisonia (Simaroubaceae), Cneorum (Cneoraceae), and Ptaeroxylon (Ptaeroxylaceae) forming a clade sister to all other Rutaceae. Circumscription of Rutaceae to include all of these taxa is recommended. This analysis indicates that Simaroubaceae and Meliaceae are the outgroups closest to Rutaceae. Correlation of the molecular phylogenies with biochemical data indicates that chemotaxonomic information is more reliable than fruit type as an indicator of familial and subfamilial circumscriptions. The subfamilial classification needs revision; none of the subfamilies of more than one genus is monophyletic.

Phylogeny of Rutaceae based on twononcoding regions from cpDNA

Primarily known only by the edible fruits of Citrus, Rutaceae comprise a large (c. 160 genera and 1900 species), morphologically diverse, cosmopolitan family. Of its extraordinary array of secondary chemical compounds, many have medicinal, antimicrobial, insecticidal, or herbicidal properties. To assist with the much-needed suprageneric reclassification and with studies of evolution of chemical compounds and biogeographic history of the family, here we included sequence data (from two noncoding regions of the chloroplast genome-rps16 intron and trnL-trnF region) from 65 species in 59 genera (more than one third of those in the family) that represented all subfamilies and tribes and more genera of Toddalioideae and of neotropical groups than previous studies. Results confirmed that Cneorum, Ptaeroxylon, Spathelia, and Dictyoloma form a clade sister to the remaining Rutaceae, none of the subfamilies with more than one genus (except Aurantioideae) is monophyletic, and characters of the ovary and fruit are not reliable for circumscription of subfamilies. Furthermore, clades are better correlated with geographic distributions of the genera than with ovary and fruit characters. Circumscriptions of subfamilies and tribes (and some subtribes of Rutoideae) must be reevaluated. Results are discussed in light of geographic distributions, caryology, chemotaxonomy, and other molecular studies.

Synopses of Angostura Roem. & Schult. and Conchocarpus J. C. Mikan (Rutaceae)

Angostura Roem. & Schult. as understood by Engler is defined more narrowly here. The species excluded from Angostura are recognized as species of ConchocarpusJ. C. Mikan. Three new species of Angostura (A. alipes Kallunki from Ecuador, A. quinquefolia Kallunki from Pari and Maranhao, Brazil, and A. simplex Kallunki from western Amazonas, Brazil, and San Martin, Peru) are described, and three new combinations in this genus are made. In Conchocarpus, 24 new combinations are made, and the following 21 new taxa are described: C. bellus Kallunki, C. cauliflorus Pirani, C. concinnus Kallunki, C. cuneifolius var. confertus Kallunki, C. cyrtanthus Kallunki, C. dasyanthus Kallunki, C. diadematus Pirani, C. fissicalyx Pirani, C. furcatus Kallunki, C. gaudichaudianus subsp. bahiensis Kallunki, C. grandis Kallunki, C. hirsutus Pirani, C. inopinatus Pirani, C. insignis Pirani, C. longipes Kallunki, C. mastigophorus Kallunki, C. modestus Kallunki, C. oppositifolius Kallunki, C. punctatus Kallunki, C. santosii Pirani & Kallunki, and C. sordidus Kallunki. With the exception of C. grandis from Amazonian Brazil, all are native to the coastal forests of eastern Brazil. As a result, seven species of Angostura and 45 of Conchocarpus are recognized. Keys to the taxa of both genera are provided. Lectotypes are designated for Cusparia grandiflora Engl., C. macrocarpa Engl., C. paniculata Engl., C. toxicaria Engl., Galipea odoratissima Lindl., and Lasiostemum silvestre Nees & Mart. and an epitype for Galipea elegans A. St.-Hil. The new combination, Rauia nodosa (Engl.) Kallunki, is made for Cusparia nodosa. The tribe Cusparieae DC. and subtribe Cuspariinae Engl., based on the illegitimate generic name Cusparia Humb., are renamed Galipeeae Kallunki and Galipeinae Kallunki.

Phenology and Floral Biology of Gustavia superba (Lecythidaceae) in Central Panama

Leaf fall, leaf production, flower and fruit production, and floral biology of Gustavia superba (Kunth) Berg were studied for a year in the tropical moist forest of central Panama. Although leaves fall throughout the year, peak fall occurs in the first month of the wet season. Leaf production on any branch is possible throughout the year, but within the population studied two pronounced peaks were observed, one in the late wet-early dry season and the other after the first rains of the wet season. Flowering began about one month after the rains ceased and continued throughout the dry season into the first month of the following wet season. Fruits reached maturity at the end of the dry season and into the early wet season. The severity of the dry season of 1974-1975 may have altered the normal flowering pattern and reduced fruit production in this species. Observations revealed that the flowers are diurnal, without odor anid nectar, and are visited by bees for their pollen. A limited number of crosses suggest that this species is self-incompatible. SEVERAL PAPERS IN RECENT YEARS have emphasized the peak in flowering of trees in the seasonally dry forests of Central America in the dry season (Fournier and Salas 1966, Janzen 1967, Croat 1969, Frankie et al. 1974). Not only do more tree species bloom during this time of the year but those that do bloom tend to have larger, more spectacular flowers. In central Panama the large, bright-yellow flowers of Tabebuia guayacan (Seem.) Hemsl., T. ochracea (Cham.) Standl. subsp. neochrysantha (A. Gentry), A. Gen,try, Cespedesia macrophylla Seem., and Cochlospermum vitifolium (Willd.) Spreng., the golden-yellow flowers of Cassia moschata Benth., the duller yellow flowers olf Byrsonima crassifolia (L.) H.B.K., the pink flowers of Tazbebiaz rosea (Bertol.) DC., and the red ones of several species of Erythfina are among the more spectacular indicators of the dry season. The number of species with maturing fruit is also greater during the dry season (Janzen 1967, Croat 1969, Smythe 1970, Frankie et al. 1974). Smythe (1970) has demonstrated a pronounced peak of large animal-dispersed seeds (over 1.5 cm in greatest diameter) at the end of the dry season into the early wet season. In addition to flowering and fruiting, certain vegetative phenomena coincide with the dry season in the seasonal tropical forests of Central America. Frankie et al. (1974) and Daubenmire (1972) have shown a distinct dry-season peak in leaf fall for a number of species of trees studied in Guanacaste Province, Costa Rica. Production of new leaves at the beginning of the dry season has been demonstrated in several species by Rockwood (1975, table 8) and toward the end of the dry season and into the rainy season by Daubenmire (1972) and Frankie et al. (1974). Even though certain phenological patterns predominate in moist tropical forest with seasonal rainfall, there are, nonetheless, many deviations from these patterns. For example, most deciduous tree species lose their leaves in the dry season, but some species like Jacquinia pungens A. Gray (Janzen 1970) and Cordia alliodora (Ruiz & Pavon) Cham. (T. Croat, pers. colmm.) lose their leaves in the wet season. The large animal-dispersed seeds of Spondias mombin L. and S. radlkoferi J. Donn. Sm. ripen and fall in the midto late-wet season in contrast to the peak for this fruit type at the end of the dry season and early wet season (Smythe 1970, Croat 1974). Because of these and other exceptions to general phenological patterns, phenological studies of individual species of seasonal tropical forest are needed.

Pollen morphology of the subtribe Cuspariinae (Rutaceae)

The neotropical subtribe Cuspariinae (Rutaceae) comprises as many as 26 genera and over 125 species. Pollen grains from 111 collections representing 71 species and 24 genera were examined by LM, SEM, and TEM. The pollen morphology of this subtribe is very diverse. Grains are mostly 3–6-aperturate and colporate, rarely porate (Spiranthera) or pantocolporate (Almeidea). Exine sculpturing is most commonly reticulate, sometimes perforate, foveolate-perforate, foveolate, foveolate-reticulate, reticulate, striate-reticulate, echinate, clavate, or baculate. The exine structure is columellate and tectate-perforate, columellate and semitectate, or intectate and is stratified into ektexine and endexine. The exine ofLeptothyrsa is distinctive in that the ektexine of the mesocolpium is longitudinally deeply ridged. The pollen ofHortia, characterized by a psilate exine with rare perforations, a very thick foot-layer, and reduced columellae, is unlike that of any member of the Cuspariinae and offers no support for the transfer of this genus from the Toddalioideae. The pollen data correlate with macromorphological characters and are taxonomically useful.

Chilean Pitavia more closely related to Oceania and Old World Rutaceae than to Neotropical groups: evidence from two cpDNA non-coding regions, with a new subfamilial classification of the family

Abstract The position of the plant genus Pitavia within an infrafamilial phylogeny of Rutaceae (rue, or orange family) was investigated with the use of two non-coding regions from cpDNA, the trnL-trnF region and the rps16 intron. The only species of the genus, Pitavia punctata Molina, is restricted to the temperate forests of the Coastal Cordillera of Central-Southern Chile and threatened by loss of habitat. The genus traditionally has been treated as part of tribe Zanthoxyleae (subfamily Rutoideae) where it constitutes the monogeneric tribe Pitaviinae. This tribe and genus are characterized by fruits of 1 to 4 fleshy drupelets, unlike the dehiscent fruits typical of the subfamily. Fifty-five taxa of Rutaceae, representing 53 genera (nearly one-third of those in the family) and all subfamilies, tribes, and almost all subtribes of the family were included. Parsimony and Bayesian inference were used to infer the phylogeny; six taxa of Meliaceae, Sapindaceae, and Simaroubaceae, all members of Sapindales, were also used as out-groups. Results from both analyses were congruent and showed Pitavia as sister to Flindersia and Lunasia, both genera with species scattered through Australia, Philippines, Moluccas, New Guinea and the Malayan region, and phylogenetically far from other Neotropical Rutaceae, such as the Galipeinae (Galipeeae, Rutoideae) and Pteleinae (Toddalieae, former Toddalioideae). Additionally, a new circumscription of the subfamilies of Rutaceae is presented and discussed. Only two subfamilies (both monophyletic) are recognized: Cneoroideae (including Dictyolomatoideae, Spathelioideae, Cneoraceae, and Ptaeroxylaceae) and Rutoideae (including not only traditional Rutoideae but also Aurantioideae, Flindersioideae, and Toddalioideae). As a consequence, Aurantioideae (Citrus and allies) is reduced to tribal rank as Aurantieae.

REPRODUCTIVE BIOLOGY OF MIXED-SPECIES POPULATIONS OF GOODYERA (ORCHIDACEAE) IN NORTHERN MICHIGAN'

The reproductive biology ofGoodyera oblongifolia, G. repens var.ophioides andG. tesselata is discussed with emphasis on the reproductive isolating mechanisms operating in mixed-species populations. Cytology, phenology and pollination of the above three species were studied. Artificial hybridizations were made of these three species and ofG. pubescens. Results show that within mixed-species populations,G. oblongifolia,G. repens var.ophioides andG. tesselata are not completely isolated reproductively, and hybridization does occur. The deleterious effects of the loss of gametes to interspecific crosses are reduced by seasonal isolation, perennial growth and geitonogamous seed production which is encouraged by self-compatibility and clonal growth. Thus, hybridization in mixed-species populations is apparently kept to levels low enough to allow the species to maintain their identities.

A revision of Erythrochiton sensu lato (Cuspariinae, Rutaceae)

In the rutaceous subtribe Cuspariinae, species with relatively large, valvate, colored calyces have been assigned to Erythrochiton, but differences in arrangement of leaves, type of inflorescence, union of petals, of filaments, and of carpels, indument of corolla and testa, appendages of anthers, height of the intrastaminal disc, and exine of the pollen argue for the recognition of three genera. Erythrochiton s. str., characterized by often perennating inflorescences, connate, usually glabrous petals, free carpels, tomentulose seeds, and spinulose exine, consists of seven species of which four are new: E. fallax from the eastern flanks of the Andes from Colombia to Bolivia, E. odontoglossus from western Ecuador and adjacent Peru, E. trichanthus from eastern Peru, and E. gymnanthus from Costa Rica. The assignment to Toxosiphon of four species with woolly, coherent petals, connate carpels, glabrous seeds, and reticulate exine necessitates three new combinations: T. carinatus, T. macropodus, and T. trifoliatus. Recognition of a third unispecific genus with opposite simple leaves, sparsely pubescent, coherent, clawed petals, and spinulose exine requires a new genus name, Desmotes, and a new combination, D. incomparabilis.

Population studies in Goodyera (Orchidaceae) with emphasis on the hybrid origin of G. tesselata

Morphological, cytological, and paper Chromatographic studies of populations from northern Michigan and examination of herbarium specimens from throughout North America were used to clarify the relationships ofGoodyera oblongifolia, G. repens var.ophioides, andG. tesselata. A canonical analysis of morphological data from mixed populations of these three species depictsG. tesselata as intermediate betweenG. oblongifolia andG. repens var.ophioides. The latter two species are diploid (2n = 30) andG. tesselata is tetraploid (2n = 60). Triploids (2n = ca. 45) were found in two mixed-species populations in northern Michigan.Goodyera tesselata produces three phenolic compounds present inG. oblongifolia and five different compounds present inG. repens var.ophioides. The range ofG. tesselata is confined to glaciated territory (except for two stations) in northeastern North America where the postglacially produced ranges ofG. oblongifolia andG. repens var.ophioides overlap. However,G. tesselata is quite abundant in areas outside the region of sympatry of the other two species. Based on this evidence, it is postulated thatG. tesselata is an allotetraploid species which resulted from hybridization betweenG. oblongifolia andG. repens var.ophioides during early post-Pleistocene. The slightly earlier blooming season ofG. tesselata may have been selected for to provide a measure of reproductive isolation between the tetraploid and its parents and to adapt the new species to the rather short growing season of northeastern North America.

Chromosome numbers of Panamanian Lecythidaceae and their use in subfamilial classification

Nine species of Lecythidaceae subfamily Lecythidoideae in four genera whose chromosome numbers were previously unknown, have 17 as their basic chromosome number:Eschweilera pittieri, three other unidentified species ofEschweilera, Grias cauliflora, Gustavia dubia, G. superba, Lecythis minor, andL. tuyrana. All are diploid exceptGustavia superba, which is tetraploid.Couroupita guianensis, which was previously—and probably incorrectly—reported to have a gametic chromosome number of 18, also hasn = 17. The known chromosome numbers support recognizing at least three of Niedenzu’s subfamilies: Planchonioideae withx = 13, Napoleonaeoideae withx = 16, and Lecythidoideae withx = 17. His fourth subfamily, Foetidioideae, with one genus of five species, has not been counted. Cytological data have been and probably will be useful in indicating to what subfamily problematic genera belong and in showing interesting phytogeographic patterns within the family. On the other hand, cytological data provide no recognizable clues relating the Lecythidaceae to other families.