Tertiary Venation Research Articles

Hypericumliboense (Hypericaceae), a new species from Guizhou, China.

Hypericumliboense M.T.An & T.R.Wu, sp. nov. (Hypericaceae) is a newly described species found in the Maolan National Nature Reserve of Guizhou Province, where it grows in rocky habitats without soil on karst mountain tops. In this study, key morphological characters were compared between the new species and the other known Hypericum species of Hypericaceae. DNA sequences were extracted from the leaves of the new species, with nuclear gene sequences (ITS) generated to reconstruct phylogenetic trees and describe its phylogenetic position in relation to other species of Hypericum. Our results show that the proposed new species has the typical characteristics of the genus Hypericum in morphology being similar to Hypericummonogynum, but differing in its sessile and semi-clasped leaves, long elliptical to long circular leaf blades, thickly papery to thinly leathery, with entire and wavy leaf margins. The abaxial side of the leaves is covered with white powder, giving them a grey-white appearance. The main lateral veins of the leaves are 8-15-paired, and the midvein on both sides is convex. The main lateral veins and midvein branch are conspicuous, with tertiary venation forming a network on the leaf surface and appearing prominently sunken. The inflorescences are 1-3-flowered, with a large calyx and conspicuous veins. The molecular phylogenetic analysis (PP = 1.00) provided substantial evidence for the proposition of H.liboense as a new species within Hypericum. Morphological and molecular evidence is presented, corroborating the proposition of the new species, including a comprehensive account of the distinctive morphological attributes of H.liboense, along with its key distinguishing features from similar species.

Phylogeny and morphological evolution of the Neotropical genus Cuspidaria (Bignonieae, Bignoniaceae): Combining high‐throughput sequencing and targeted loci data

AbstractThe combined use of high‐throughput sequencing (HTS) with targeted loci data offers an excellent opportunity to reconstruct robust and comprehensive phylogenies at fine taxonomic scales using a hybrid approach. In this study, we infer the phylogeny of Cuspidaria (Bignonieae, Bignoniaceae), a diverse clade of lianas and shrubs centered in South America's wet and dry forests. We used HTS to obtain complete or nearly complete plastome sequences of seven individuals of Cuspidaria selected to represent the main clades in the genus. This sampling strategy aimed to reconstruct relationships at deeper nodes (i.e., backbone). We also used targeted loci data obtained through Sanger sequencing to obtain sequences of two chloroplast markers (ndhF, rpl32‐trnL) and one nuclear marker (PepC) for multiple individuals of 18 out of the 21 species of Cuspidaria recognized in the most recent treatment of the genus; only C. emmonsii, C. lachnaea, and C. simplicifolia were not sampled. This broad sampling strategy aimed to test the monophyly of individual species and reconstruct fine‐scale interspecific relationships within the genus. Both datasets were analyzed separately and combined using Bayesian inference and maximum likelihood approaches. The combined dataset includes 65 individuals representing 18 previously recognized species of Cuspidaria plus outgroup taxa. The analysis of the combined dataset recovered a monophyletic Cuspidaria, excluding C. bracteata, which was nested within the outgroup and is best treated elsewhere. Ancestral character state reconstructions provided novel insights into the evolution of morphology and identified multiple putative morphological synapomorphies for key lineages of Cuspidaria. Namely, anthers curved forward was reconstructed as a potential synapomorphy for the Cuspidaria clade, whereas interpetiolar gland fields in stems and inflorescence, subulate or inconspicuous prophylls of the axillary buds, biternate leaves, actinodromous venation, calyx cupulate or spathaceous, calyx apices dentate or irregularly lobed, fruits with winged valves, and fruit midrib limited by two longitudinal ridges were reconstructed as potential synapomorphies of different clades within the genus. Other traits such as habit, tertiary venation, inflorescence type, corolla color, number of ovule series, and pollen unit were highly labile. As circumscribed here, Cuspidaria is composed of eight main clades supported by molecular data and morphological synapomorphies: Sideropogon, Tetrastichella, Cinerea, Paracarpaea, Cremastus, Blepharitheca, Saldanhaea, and Cuspidaria s.str. clades. All species are monophyletic, except for C. pulchra and C. sceptrum, which form a species complex and are best treated as a single taxon. The necessary taxonomic changes are proposed, leading to the recognition of 19 species in Cuspidaria.

Papillaephyllum, a new genus of angiosperm foliage from the Cenomanian of the Czech Republic

Angiosperm foliage Papillaephyllum labutae Čepičková et J. Kvaček, gen. et sp. nov. is described from the Cenomanian, Late Cretaceous of the Peruc-Korycany Formation in the Bohemian Cretaceous Basin, Czech Republic. Papillaephyllum labutae has been erected for rarely-found fossils with basal-angiosperm signs, characterized by serrate, sometimes double-serrate margins with rounded chloranthoid teeth, each bearing a papilla. Venation is semicraspedodromous to festooned-semicraspedodromous with reticulate tertiary venation. Its micromorphology is distinctive in having the entire abaxial cuticle covered by papillae. Papillae of the subsidiary cells often form as visible wreaths around the stomata. Stomata are latero-cyclocytic and stephanocytic-bicyclic.Well-pronounced and numerous papillae of the external epidermal surface are an indicator of possible fresh-water stress, identifying the taxon as possibly meso- to xerophytic. Based on macro- and micromorphological comparison with fossil and recent material, affinity of Papillaephyllum labutae with the family Chloranthaceae is highly probable.

Nombrando lo conocido: una nueva especie de Desmopsis (Annonaceae) endémica de México

Antecedentes y Objetivos: Una especie de Desmopsis (Annonaceae) colectada frecuentemente en México ha permanecido indeterminada durante más de 30 años. Sus características florales son similares a las del género asiático Guamia (actualmente Meiogyne), y desde hace muchos años se le ha nombrado como “Guamia mexicana”, pero no ha sido descrita formalmente. Recientemente se demostró que “Guamia mexicana” está filogenéticamente emparentada con Desmopsis y Stenanona y no con Guamia. Sin embargo, y a pesar de ser colectada con frecuencia, no se ha aclarado su estatus taxonómico. En este artículo se estudiaron las relaciones filogenéticas y las características morfológicas y ecológicas que distinguen a “Guamia mexicana” de otras especies. Como resultado, esta especie nueva es aquí descrita. Métodos: El reconocimiento de la especie nueva se basó en la correlación de varios caracteres morfológicos, a través de comparaciones con especies morfológicamente similares y de la revisión de literatura. Las características de su polen se analizaron mediante microscopía electrónica de barrido. Sus relaciones filogenéticas se determinaron con base en filogenias publicadas previamente. Finalmente, a partir de sus localidades conocidas se calculó extensión de ocurrencia (EOO) y área de ocupación (AOO) en GeoCAT, y aplicando las categorías y criterios de la Lista Roja de la UICN se determinó su estado de conservación. Resultados clave: Desmopsis ibarrarum, una nueva especie de Annonaceae endémica de México, es descrita e ilustrada. Se distingue de otras especies de Desmopsis por la combinación de frutos largos y cilíndricos con muchas semillas, con una testa dura y leñosa y densamente cubierta por pelos seríceos muy pequeños (aterciopelados), por sus hojas glabras con venación terciaria percurrente, la ausencia de brácteas foliosas en los pedicelos, la diferenciación morfológica de los pétalos internos y externos, y por la fusión parcial de los sépalos. Conclusiones: Desmopsis ibarrarum esta únicamente relacionada a Desmopsis dubia y las dos especies se consideran aquí miembros de un linaje particular dentro del clado Desmopsis-Stenanona, el "grupo Desmopsis dubia".

Taxonomic novelties within a non‐monophyletic lineage: a new Mexican species of Desmopsis (Annonaceae)

A new species of Annonaceae endemic to Mexico is described and illustrated. Desmopsis dubia sp. nov. is placed within the Desmopsis–Stenanona clade and is clearly different from all other species of Neotropical Miliuseae by the combination of lower surface of leaves pubescent; tertiary venation strongly percurrent; leafy bracts absent at the base of the pedicels; sepals connate at the base and persistent in fruit; petals fleshy and stiff, elliptic to ovate with food bodies at the base of the inner ones; and cylindrical monocarps with ligneous exocarp. We discuss its phylogenetic relationships and placement within the genus Desmopsis and a key to the Mexican species of Desmopsis and Stenanona with yellow or greenish petals is provided.

Short Communication: Two new species and new record of Psydrax Gaertn. (Rubiaceae: Vanguerieae) in Borneo

Abstract. Mahyuni R, Chikmawati T, Ariyanti NS. 2019. Short Communication: Two new species and new record of Psydrax Gaertn. (Rubiaceae: Vanguerieae) in Borneo. Biodiversitas 20: 2011-2015. Psydrax sabahensis Mahyuni and P. wongii Mahyuni are two new species endemic to Borneo proposed in this present study with descriptions, illustrations and map of distribution provided. Psydrax sabahensis has blades ovate to elliptic, with angle 60º to the midrib, tertiary venation conspicuous on both side, petiole length 10-13 mm long, inflorescence sub-umbellate and pedicels 23-27 mm long. Psydrax wongii has blade elliptic, with angle 45º to midrib, apex acuminate to caudate, secondary veins 3-4 pairs, tertiary venation inconspicuous on upper side and slightly wrinkled on lower surface. The presence of P. maingayi (Hook.f.) Bridson in Borneo, previously thought to be a Malay Peninsula endemic species, is also confirmed as a new record.

A new fossil genus of angiosperm leaf from the Olmos Formation (upper Campanian), of northern Mexico

The Olmos Formation (upper Campanian) is located in the Sabinas Basin in northern Mexico. It represents a deltaic-fluvial system, and it is known by its abundance of fossil plants mainly represented by leaves and woods. In this contribution, we describe angiosperm fossil leaves, related to ?Violaceae. The new fossil genus has pinnate venation, with a major secondaries festooned semicraspedodromous, tertiary venation opposite percurrent, quaternary veins irregular reticulate, margin crenulated with tooth terminating in a glandular tissue, and fimbrial vein present. This mosaic of characteristics supports the creation of a new genus and species, Mascogophyllum elizondoa Centeno-González, Porras-Múzquiz et Estrada-Ruiz (?Violaceae). This record could be the oldest of the family; also, a new family is added to the very diverse fossil flora record that established during the Cretaceous in the northern of Mexico.

Enlarging the monotypic Monocarpieae (Annonaceae, Malmeoideae): recognition of a second genus from Vietnam informed by morphology and molecular phylogenetics

Chaowasku, T., A. Damthongdee, H. Jongsook, D.T. Ngo, H.T. Le, D.M. Tran & S. Suddee (2018). Enlarging the monotypic Monocarpieae (Annonaceae, Malmeoideae): recognition of a second genus from Vietnam informed by morphology and molecular phylogenetics. Candollea 73: 261–275. In English, English abstract. DOI: http://dx.doi.org/10.15553/c2018v732a11Recent botanical expeditions in central Vietnam yielded an unknown species of Annonaceae that could not be confidently identified to subfamily, tribe, and genus. Preliminary BLAST® searches based on plastid data have suggested that this taxon is genetically closely-related to the following tribes of subfamily Malmeoideae: Malmeeae, Fenerivieae, Maasieae, Phoenicantheae, Dendrokingstonieae, Monocarpieae, and Miliuseae. Using representatives of Piptostigmateae, another tribe of Malmeoideae, as outgroups and including representatives of all other tribes of Malmeoideae, molecular phylogenetic analyses of seven combined plastid markers (rbcL, matK, ndhF, ycf1 exons; trnL intron; trnL-trnF, psbA-trnH intergenic spacers) inferred the enigmatic Vietnamese taxon as belonging to the monotypic tribe Monocarpieae. Detailed morphological comparisons between this taxon and its sister group, Monocarpia Miq., warranted the recognition of a second genus of Monocarpieae to accommodate our unknown taxon: Leoheo Chaowasku with a single species, Leoheo domatiophorus Chaowasku, D.T. Ngo & H.T. Le. The morphology of the new genus agrees well with the diagnostic traits of Monocarpieae, e.g., a percurrent tertiary venation of the leaves, a highly reduced number of carpels per flower, enlarged and lobed stigmas, multiple ovules/seeds per ovary/monocarp, considerably large monocarps with a hardened pericarp when dry, and spiniform ruminations of the endosperm. However, the new genus does not exhibit two characteristic features of Monocarpia: terminal inflorescences and generally distinct intramarginal leaf veins. In addition, the new genus possesses three autapomorphic characters: hairy domatia on the lower leaf surface, longitudinal ridges on the monocarp surface, and subsessile monocarps with a stout stipe. The tribe Monocarpieae is consequently enlarged to include the genus Leoheo. The enlarged Monocarpieae, along with the recently established monotypic tribe Phoenicantheae and two other related tribes, Dendrokingstonieae and Miliuseae, are discussed.Received: November 4, 2017. Accepted: August 30, 2018. First published online: November 23, 2018.

ANÁLISIS MORFOANATOMÍCO FOLIAR EN GRUPOS HORTÍCOLAS DE AGUACATE (Persea americana) DEPOSITADOS EN LA COLECCIÓN DEL INIA-CENIAP, VENEZUELA

<p>The avocado (<em>Persea americana</em> Mill.) is the most important species of Lauraceae in America due to its exploitation as food for pre-Columbian and modern cultures. It is a very important seasonal crop in Venezuela based on a perennial fruit tree management. From a selection of 76 accessions (45 cultivars) of avocados cultivated at the Germplasm Bank of INIA-CENIAP, a morphoanatomical analysis was performed to identify attributes of taxonomic resolution (diagnostic characters) which allow to characterize sets and / or <em>culta</em>. Morphological study was carried out from each accession herborized sample. Information was obtained by freehand transverse leaf sections (epidermis, mesophyll and midvein) as well as paradermic preparations, and observed data was recorded in DELTA System. New morphoanatomical characters and discriminating attributes between cultivars were identified and described, especially to discriminate the Mexican group, and a close relationship within West Indian and Guatemalan cultivars was observed due to the variability identified from the latter group. Indument- related attributes were highly informative to discriminate among cultivars, along with the outline, apical angle and projections at the base of the leaf blades, stem cross section and presence of anise odor, progress and joining of the secondary nerve branches, tertiary venation pattern, abaxial contour and thickness of the sclerenchymatous sheath and compaction of the phloem in the vascular bundle, adaxial contour of the median nerve, and thickness, outline and uniformity of the anticlinal walls of adaxial and abaxial epidermal cells. </p>

Novelties in neotropical Dilleniaceae

A new species of Doliocarpus (Dilleniaceae), Doliocarpus putumayensis Aymard from the Porvenir River (affluent of the lower Putumayo River), and the middle Caquetá River in the state of Amazonas was found during the examination of specimens for the Dilleniaceae treatment for the Flora de Colombia. This species is not clearly related to any other Doliocarpus species, however, it shows certain similarities with D. magnificus Sleumer in its leaves size, shape, and the fruit size and pubescence. Nonetheless, P. putumayensis differs in its tertiary venation, inflorescence size, sepal number and shape, anthers glabrous with a connective broadened, and fruit densely adpressed pubescent. An updated key to the species of Doliocarpus of Colombia is provided, and a previously described subspecies from Brazil is elevated to the rank of species (i.e., Doliocarpus brevipedicellatus Garcke subsp. hilarianus Kubitzki to D. hilarianus Aymard).

Characteristics of Cervical Mucus for Estrus Detection in Murrah Buffaloes (Bubalus bubalis)

The objective of the experiment was to study the changes in different characteristics of cervical mucus of Murrah buffaloes in relation to occurrence of estrus. The Murrah buffaloes maintained at Livestock Research Centre (LRC), ICAR-NDRI, Karnal were classified into three groups viz. heifers, primipara and pluripara. Cervical mucus was collected on the day of estrus prior to the artificial insemination by aspiration from the mid cervix using blue sterile sheath and Universal artificial insemination gun through recto-vaginal method. Immediately after collection, mucus sample was taken to the laboratory various physical and chemical parameters were assessed. From the present study it can be concluded that the changes in cervical mucus characteristics can be used for prediction of estrus time and subsequently time of artificial insemination. Murrah buffaloes exhibited higher proportion (70%) of copious discharge, 86% buffaloes had clean appearance and 68% of buffaloes showed moderate consistency of cervical mucus. The spinnbarkeit value of cervical mucus of 46% of estruses were in 8–16 cm and 58% showed the typical fern pattern predominantly with primary, secondary and tertiary venation. pH of cervical mucus were within 7.5–8.0 and conductivity in a range of 13.50–15.00 mS/cm.

Ormocarpopsis Anosyana Thulin & Razafim. (Fabaceae), a New Species from Southern Madagascar and Its Phylogenetic Position

Thulin, M. & S.G. Razafimandimbison (2016). Ormocarpopsis anosyana Thulin & Razafim. (Fabaceae), a new species from southern Madagascar and its phylogenetic position. Candollea 71 : 281–286. In English, English and French abstracts. DOI: http//dx.doi.org/10.15553/c2016v712a13Ormocarpopsis anosyana Thulin & Razafim. (Fabaceae), a new species from the Anosy Region of south-eastern Madagascar, is described and illustrated. According to phylogenetic analyses based on nuclear ribosomal ITS sequences, the new species is, with strong support, sister to Ormocarpopsis mandrarensis Dumaz-le-Grand, another species confined to southeastern Madagascar. Ormocarpopsis anosyana differs markedly from Ormocarpopsis mandrarensis by its generally smaller leaflets with the secondary and tertiary venation drying more or less blackish beneath, by its shorter hypanthium, by its smaller calyx with the lowest tooth about as long as the others, by its wing-petals that are much longer than the keel, by its glabrous ovary, and by its slightly articulated fruits. Ormocarpopsis anosyana is known only from two collections from a single patch of spiny dry forest east of Imonty. It is assigned a preliminary conservation status as “Endangered”.

Una especies nueva de Guarea (Meliaceae) del suroriente de Ecuador

A new species of Guarea (Meliaceae) from the rainforests of Ecuador, G. cauliracemosa, is described. It is distinguished by the following combination of characters: leaflets densely tomentulose, underside of leaves with conspicuously reticulate tertiary venation and inflorescence cauline-racemose. The species is known only from Cordillera del Cóndor, Zamora-Chinchipe province, south-eastern Ecuador.

Miocene Lonchocarpus (Papilionoideae) in San Esteban Tizatlán, Tlaxcala, Mexico

New leaflets from San Esteban Tizatlán, Tlaxcala, Mexico, are described and identified based on their leaf architecture. They have elliptic leaflets with pulvinate petiolule, entire margin, convex apex and base, well developed primary and craspedodromous secondary venation, with decurrent secondary veins with fused the mid-vein, a pair of obtuse basal veins, weak intersecundary veins, mixed percurrent tertiary venation dominated by the opposite type with sinuous course, alternate percurrent fourth-order venation, regular reticulate with few non ramified veinlets. All these characteristics are closely compared with members of sections Obtusifolii and Standleyi of Lonchocarpus (Leguminosae, Papillionoideae, Millettieae), but the mosaic of characters found in the fossil material granted recognition of a new species Lonchocarpus miocenicus sp. nov. This new record of Lonchocarpus from central Mexico further supports the presence of an evergreen tropical forest during the Miocene in the area, a plant community that developed from the Boreotropical Flora and contributed latter to the Neotropical Flora.

The type specimen of Debeya (Dewalquea) haldemiana rediscovered

The rediscovered specimen MGUWr 7536p, one of the syntypes of Dewalquea haldemiana, is described. It is selected as the lectotype of Debeya (Dewalquea) haldemiana (DEBEY ex SAPORTA et MARION 1873) HALAMSKI 2013 and of Debeya (Dewalquea) haldemiana var. haldemiana. It is the most complete and best preserved known specimen of the species. It allows supplementation of previous description of the following characters: brochidodromous secondary venation; acuminate leaflet apices; variation of the petiolule length from 0 (leaflets subsessile) to 30 mm; ramified tertiary venation. Other species considered up to now to belong to the same subgenus possess percurrent tertiary venation; a doubt is therefore expressed about the validity of the present circumscription of Debeya (Dewalquea). Additonally, validation of the previously published name Debeya (Dewalquea) haldemiana var. angustifolia (HOSIUS et VON DER MARCK 1880) HALAMSKI comb. nov. is presented.

The lianescent species ofGrewiaL. (Malvaceaes.l., formerly Tiliaceae) in Madagascar

ABSTRACT Three liana species of Grewia L. are described from Madagascar, bringing to five the members of the genus on the island to exhibit this habit. Grewia manomboensis G.E. Schatz, Randrian. & Lowry, sp. nov., G. rabehevitrae Randrian., Lowry & G.E. Schatz, sp. nov. and G. rufostellata Randrian., Lowry & G.E. Schatz, sp. nov., all three of which occur in low- to mid-elevation humid eastern forest, can be distinguished from one another by several features, including differences in the leaves (size, margin and prominence of the tertiary venation), pedicels (length, diameter and trichome density), sepal size, indument type and color, and petal shape. Illustrations are provided for each new taxon and photographs for two of them, along with a range map, preliminary assessments of the conservation status of each species, and an identification key in English and French.

<p class="HeadingRunIn"><strong>Characterization of <em>Hubera</em> (Annonaceae), a new genus segregated from <em>Polyalthia</em> and allied to <em>Miliusa</em></strong></p>

On the basis of molecular phylogenetics, pollen morphology and macromorphology, a new genus of the tribe Miliuseae, Hubera, segregrated from Polyalthia and allied to Miliusa, is established and described. It is characterized by the combination of reticulate tertiary venation of the leaves, axillary inflorescences, a single ovule per ovary and therefore single-seeded monocarps, seeds with a flat to slightly raised raphe, spiniform(-flattened peg) ruminations of the endosperm, and pollen with a finely and densely granular infratectum. Twenty-seven species are accordingly transferred to this new genus.

Aplicação de caracteres morfoanatômicos foliares na taxonomia de Campomanesia Ruiz & Pavón (Myrtaceae)

Para avaliar a utilidade de caracteres morfoanatômicos na identificação de materiais botânicos de Campomanesia Ruiz & Pavón, especialmente aqueles estéreis, o padrão de nervação foliar e a configuração do arco vascular no pecíolo de 15 táxons presentes no estado da Bahia foram analisados. O padrão de nervação secundário camptódromo-broquidódromo mostrou-se uniforme entre os táxons, assim como a nervação última marginal em arcos e a nervação terciária do tipo reticulada ortogonal. Diferenças foram observadas quando considerados a presença de nervuras intersecundárias e o tipo de aréolas na lâmina foliar, e a configuração do arco vascular e a natureza da bainha perivascular no pecíolo, podendo estes caracteres ser utilizados para fi ns taxonômicos.

Morfometria e nervação foliar em procedências de erva-mate (Ilex paraguariensis A. St. Hill.) (Aquifoliaceae)

The present research aimed to evaluate the foliar morphology (morphometry and leaf venation) of origins of maté collected in southern Brazil and Argentina. For the study of the morphometry, thirty leaves/plant were analyzed, and five measurements (leaf total length, maximum width, length from the base until maximum width, area and petiole length) and two indices were obtained (total leaf length on maximum width and length from basis until maximum width on total leaf length). The leaves were clarified by the clarification technique and examined in stereomicroscope. The highest and lowest values of leaf morphometry were, respectively, 118 and 67 mm for total leaf length, 51 and 34 mm for maximum width, 78 and 42 mm for length from the base until the maximum width, 16 and 10 mm for petiole length, 2.3 and 2 mm for total length on maximum width, and 0.65 and 0.6 for length from the base until the maximum width on total leaf length. In regards to leaf venation, differences between leaves of different origins were not observed, which presented pinnate primary venation; semicraspedodromous secondary venation, alternate percurrent and sinuous tertiary venation, regular polygonal reticulate quaternary venation and dichotomizing 5tha vein category

Menispermaceae from the Cerrejón Formation, middle to late Paleocene, Colombia

The origin and processes creating the high diversity of plant species in neotropical rain forests and their floristic composition and multistratitified forest structure are still uncertain. Here, we studied one of the most common leaf morphotypes of the Cerrejón flora (middle-late Paleocene, ca. 60-58 Ma), Guajira, Colombia, that contains one of the oldest records of neotropical rain forest floras. Fifty-seven leaf specimens were carefully examined with a focus on general morphology, venation patterns, and cuticular characteristics. The analysis allowed us to recognize four new species that were assigned to the fossil-leaf genus Menispermites on the basis of an ovate leaf shape with cordate to truncate bases, actinodromous primary venation, brochidodromous secondary venation, percurrent tertiary venation, regular polygonal reticulate fourth and fifth venation, well-developed polygonalareoles, entire margin, and the presence of a fimbrial vein. This set of characters suggests a possible affinity with the pantropical angiosperm family Menispermaceae. The predominantly climbing habit of this family suggests that the Cerrejón Paleocene tropical rain forest was already multistratified. These findings represent the earliest record for the family in northern South America.